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<document id="9BE71E8BFBA2CF53A2C0968EA96F1BA8" ID-CLB-Dataset="305075" ID-DOI="10.1002/tax.12818" ID-GBIF-Dataset="771a24d1-838a-474b-a385-958f0b997524" ID-ISSN="1996-8175" ID-Zenodo-Dep="14047522" IM.bibliography_approvedBy="julia" IM.illustrations_approvedBy="julia" IM.materialsCitations_approvedBy="julia" IM.metadata_approvedBy="julia" IM.tables_approvedBy="julia" IM.taxonomicNames_approvedBy="julia" IM.treatmentCitations_approvedBy="julia" IM.treatments_approvedBy="julia" checkinTime="1730918127147" checkinUser="felipe" docAuthor="Mátis, Attila, Malkócs, Tamás, Kuhn, Thomas, Laczkó, Levente, Moysiyenko, Ivan, Szabó, Anna, Bădărău, Alexandru S. &amp; Sramkó, Gábor" docDate="2023" docId="450487C0FFC5FF9F212B9665FAC6FD90" docLanguage="en" docName="TAXON.72.1.78-97.pdf" docOrigin="TAXON 72 (1)" docSource="http://dx.doi.org/10.1002/tax.12818" docStyle="DocumentStyle:239A75409313D6488D9D82529296B06F.1:TAXON.2020-.journal_article.open" docStyleId="239A75409313D6488D9D82529296B06F" docStyleName="TAXON.2020-.journal_article.open" docStyleVersion="1" docTitle="Salvia austriaca" docType="treatment" docVersion="3" lastPageNumber="84" masterDocId="B93DFFB8FFC1FF992269971EFFD6FFC2" masterDocTitle="Hiding in plain sight: Integrative analyses uncover a cryptic Salvia species in Europe" masterLastPageNumber="97" masterPageNumber="78" pageNumber="82" updateTime="1731091457251" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-NC-ND-4.0">
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<mods:title id="2FB7F0F9DA803CF650B80F7842AA8A5D">Hiding in plain sight: Integrative analyses uncover a cryptic Salvia species in Europe</mods:title>
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<mods:namePart id="03EC85B8D2D2715CD8AADBDE53E8F49B">Mátis, Attila</mods:namePart>
<mods:affiliation id="25D9CDB28028D6F8F99330B30D782968">* Tamás Malkócs, &amp; Levente Laczkó, &amp; Alexandru S. Bădărău &amp; Hungarian Department of Biology and Ecology, Faculty of Biology and Geology, Babeş-Bolyai University, &amp; 4032 Debrecen, Hungary</mods:affiliation>
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<mods:namePart id="973112F9A4E8A8274547F83DC2E82BEA">Malkócs, Tamás</mods:namePart>
<mods:affiliation id="FFE637141608E00F5BCA7B0E3E0A3B8D">* Thomas Kuhn, &amp; Ivan Moysiyenko, &amp; Department of Botany, Institute of Biology and Ecology, University of Debrecen, Egyetem tér &amp; * Thomas Kuhn, &amp; Institute of Aquatic Ecology, Centre for Ecological Research, 4026 Debrecen, Hungary</mods:affiliation>
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<mods:affiliation id="63628FFA6B954AC6A3E81E7AA913A76B">* Tamás Malkócs, &amp; Levente Laczkó, &amp; Alexandru S. Bădărău &amp; Hungarian Department of Biology and Ecology, Faculty of Biology and Geology, Babeş-Bolyai University, &amp; 4032 Debrecen, Hungary</mods:affiliation>
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<mods:namePart id="4E41E28C34504B85434DF74F479D566A">Laczkó, Levente</mods:namePart>
<mods:affiliation id="919CF96BCF45E833EF3C4834AAFCB452">* Thomas Kuhn, &amp; Ivan Moysiyenko, &amp; Department of Botany, Institute of Biology and Ecology, University of Debrecen, Egyetem tér &amp; Ivan Moysiyenko, &amp; MTA-DE “ Lendület ” Evolutionary Phylogenomics Research Group, Debrecen, Hungary &amp; Ivan Moysiyenko, &amp; - &amp; ELKH-DE Conservation Biology Research Group, Debrecen, Hungary</mods:affiliation>
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<mods:namePart id="C79E440186A632AEA8913D4A63D70A92">Szabó, Anna</mods:namePart>
<mods:affiliation id="D50E03FE0A52F67D9EC232CF4D3AED34">* Tamás Malkócs, &amp; Levente Laczkó, &amp; Alexandru S. Bădărău &amp; Hungarian Department of Biology and Ecology, Faculty of Biology and Geology, Babeş-Bolyai University, &amp; 4032 Debrecen, Hungary</mods:affiliation>
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<mods:namePart id="93ACAB16F63CC630D44F026FC32E3FA8">Bădărău, Alexandru S.</mods:namePart>
<mods:affiliation id="D694EFC03B861EFBA524C095EDCE261A">&amp; Gábor Sramkó &amp; Clinicilor Street, &amp; Faculty of Environmental Sciences and Engineering, Babeş-Bolyai University, 30 Fântânele Street, 400294 Cluj-Napoca, Romania</mods:affiliation>
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<mods:affiliation id="FA07F9E895DDD63256B2A37CD09972C2">* Thomas Kuhn, &amp; Ivan Moysiyenko, &amp; Department of Botany, Institute of Biology and Ecology, University of Debrecen, Egyetem tér &amp; Ivan Moysiyenko, &amp; MTA-DE “ Lendület ” Evolutionary Phylogenomics Research Group, Debrecen, Hungary &amp; Ivan Moysiyenko, &amp; - &amp; ELKH-DE Conservation Biology Research Group, Debrecen, Hungary</mods:affiliation>
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<mods:title id="96FD6BCAE4EA377B0356CADD4901D020">TAXON</mods:title>
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<mods:date id="0844CFE027887E1B5ADF556198CB7971">2023</mods:date>
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<mods:number id="0265A1D16F244930BED7126F4D4F4BF1">2023-02-28</mods:number>
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<mods:identifier id="0D23CF3C8343662D6B0F47C78988B40A" type="DOI">10.1002/tax.12818</mods:identifier>
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<treatment id="450487C0FFC5FF9F212B9665FAC6FD90" ID-DOI="http://doi.org/10.5281/zenodo.14057458" ID-Zenodo-Dep="14057458" LSID="urn:lsid:plazi:treatment:450487C0FFC5FF9F212B9665FAC6FD90" httpUri="http://treatment.plazi.org/id/450487C0FFC5FF9F212B9665FAC6FD90" lastPageId="6" lastPageNumber="84" pageId="4" pageNumber="82">
<subSubSection id="85B7655DFFC5FF9D212B9665FA4BFE50" box="[834,1437,379,403]" pageId="4" pageNumber="82" type="nomenclature">
<paragraph id="CD1236D6FFC5FF9D212B9665FA4BFE50" blockId="4.[789,1438,379,1970]" box="[834,1437,379,403]" pageId="4" pageNumber="82">
<emphasis id="FFD9EAC4FFC5FF9D212B9665FA4BFE50" bold="true" box="[834,1437,379,403]" pageId="4" pageNumber="82">
Distribution of the taxa of the
<taxonomicName id="0AAD4D55FFC5FF9D26E19662FAF9FE50" ID-CoL="79FFL" box="[1160,1327,380,402]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="4" pageNumber="82" phylum="Tracheophyta" rank="species" species="austriaca">
<emphasis id="FFD9EAC4FFC5FF9D26E19662FAF9FE50" bold="true" box="[1160,1327,380,402]" italics="true" pageId="4" pageNumber="82">Salvia austriaca</emphasis>
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group.
<emphasis id="FFD9EAC4FFC5FF9D27EB9663FA4BFE50" bold="true" box="[1410,1437,381,402]" italics="true" pageId="4" pageNumber="82">—</emphasis>
</emphasis>
</paragraph>
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<subSubSection id="85B7655DFFC5FF9D217C9685FB5CFCF0" pageId="4" pageNumber="82" type="distribution">
<paragraph id="CD1236D6FFC5FF9D217C9685FB5CFCF0" blockId="4.[789,1438,379,1970]" pageId="4" pageNumber="82">
Our collection of occurrence data for taxa of the
<taxonomicName id="0AAD4D55FFC5FF9D27669685FCEDFE10" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="4" pageNumber="82" phylum="Tracheophyta" rank="species" species="austriaca">
<emphasis id="FFD9EAC4FFC5FF9D27669685FCEDFE10" italics="true" pageId="4" pageNumber="82">Salvia austriaca</emphasis>
</taxonomicName>
group with two different pollen deposition modes (i.e., dorsal and lateral deposition) indicates strong eco-geographical differentiation with an almost complete vicariance of the two forms (
<figureCitation id="55962A53FFC5FF9D21FA9505FC02FDF0" box="[915,980,539,562]" captionStart="Fig" captionStartId="5.[151,183,1160,1181]" captionTargetBox="[152,1483,187,1126]" captionTargetId="figure-0@5.[152,1483,187,1126]" captionTargetPageId="5" captionText="Fig. 3. Distribution of Salvia FDPD and S. austriaca Jacq. Delimitation of the steppe zone follows Lavrenko &amp; al. (1991). Delimitation of the Crimean Submediterranean forest ecoregion follows Olson &amp; al. (2001). For geographic coordinates of each observation, see suppl. Table S1." figureDoi="http://doi.org/10.5281/zenodo.14047528" httpUri="https://zenodo.org/record/14047528/files/figure.png" pageId="4" pageNumber="82">Fig. 3</figureCitation>
). The taxon with lateral pollen deposition (i.e.,
<taxonomicName id="0AAD4D55FFC5FF9D21399525FC05FD90" box="[848,979,571,594]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="4" pageNumber="82" phylum="Tracheophyta" rank="species" species="austriaca">
<emphasis id="FFD9EAC4FFC5FF9D21399525FC05FD90" box="[848,979,571,594]" italics="true" pageId="4" pageNumber="82">S. austriaca</emphasis>
</taxonomicName>
<taxonomicNameLabel id="E4EA57BFFFC5FF9D21B79525FBD9FD90" box="[990,1039,571,594]" pageId="4" pageNumber="82" sensu="stricto">s.str.</taxonomicNameLabel>
) occupies the forest-steppe zone of the Pannonian basin, Transylvanian basin, the Pontic steppe, the SE part of Crimea, and the Balkans, whereas
<taxonomicName id="0AAD4D55FFC5FF9D27789565FA4AFD50" authority="FDPD" authorityName="FDPD" box="[1297,1436,635,658]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="4" pageNumber="82" phylum="Tracheophyta" rank="genus">
<emphasis id="FFD9EAC4FFC5FF9D27789565FA85FD50" box="[1297,1363,635,658]" italics="true" pageId="4" pageNumber="82">Salvia</emphasis>
FDPD
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is confined to the short-grass steppe zone (or true steppe zone) as defined by
<bibRefCitation id="A93C4B27FFC5FF9D21CE95A5FB45FD10" author="Lavrenko, E. M. &amp; Karamysheva, Z. V. &amp; Nikulina, R. I." box="[935,1171,699,722]" pageId="4" pageNumber="82" refId="ref15249" refString="Lavrenko, E. M., Karamysheva, Z. V. &amp; Nikulina, R. I. 1991. Stepi Evrazii [Steppes of Eurasia]. Leningrad: Nauka." type="book" year="1991">Lavrenko &amp; al. (1991)</bibRefCitation>
of the Pontic steppe. The westernmost observations of
<taxonomicName id="0AAD4D55FFC5FF9D262595C5FB0FFD30" authority="FDPD" authorityName="FDPD" box="[1100,1241,731,754]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="4" pageNumber="82" phylum="Tracheophyta" rank="genus">
<emphasis id="FFD9EAC4FFC5FF9D262595C5FB58FD30" box="[1100,1166,731,754]" italics="true" pageId="4" pageNumber="82">Salvia</emphasis>
FDPD
</taxonomicName>
are from northern
<collectingCountry id="B5BA7646FFC5FF9D217C95E5FCA6FCD0" box="[789,880,763,786]" name="Bulgaria" pageId="4" pageNumber="82">Bulgaria</collectingCountry>
(settlement: Bozhurluka), whereas the easternmost is from the city of
<collectingRegion id="0F69F834FFC5FF9D21B39405FBF2FCF0" box="[986,1060,795,818]" country="Russia" name="Rostov" pageId="4" pageNumber="82">Rostov</collectingRegion>
(
<collectingCountry id="B5BA7646FFC5FF9D265A9405FBA9FCF0" box="[1075,1151,795,818]" name="Russia" pageId="4" pageNumber="82">Russia</collectingCountry>
).
</paragraph>
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<emphasis id="FFD9EAC4FFC5FF9D212B9425FBC9FCB0" bold="true" pageId="4" pageNumber="82">
Morphological differentiation between
<taxonomicName id="0AAD4D55FFC5FF9D27619422FCEDFCB0" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="4" pageNumber="82" phylum="Tracheophyta" rank="species" species="austriaca">
<emphasis id="FFD9EAC4FFC5FF9D27619422FCEDFCB0" bold="true" italics="true" pageId="4" pageNumber="82">Salvia austriaca</emphasis>
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and
<taxonomicName id="0AAD4D55FFC5FF9D211B9442FC28FCB1" authority="FDPD." authorityName="FDPD." box="[882,1022,859,883]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="4" pageNumber="82" phylum="Tracheophyta" rank="genus">
<emphasis id="FFD9EAC4FFC5FF9D211B9442FC65FCB0" bold="true" box="[882,947,860,882]" italics="true" pageId="4" pageNumber="82">Salvia</emphasis>
FDPD.
</taxonomicName>
<emphasis id="FFD9EAC4FFC5FF9D266D9443FBC9FCB0" bold="true" box="[1028,1055,861,882]" italics="true" pageId="4" pageNumber="82">—</emphasis>
</emphasis>
While processing the collected samples, we noted that several individuals exhibited reduced, shortened staminal levers, indicating that
<taxonomicName id="0AAD4D55FFC5FF9D26A59485FA81FC70" authority="FDPD" authorityName="FDPD" box="[1228,1367,923,946]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="4" pageNumber="82" phylum="Tracheophyta" rank="genus">
<emphasis id="FFD9EAC4FFC5FF9D26A59485FADBFC70" box="[1228,1293,923,946]" italics="true" pageId="4" pageNumber="82">Salvia</emphasis>
FDPD
</taxonomicName>
might be exhibiting gynodioecy, a dimorphic sexual system where populations are composed of both hermaphroditic and female individuals. Gynodioecy is known in
<taxonomicName id="0AAD4D55FFC5FF9D26F094E5FAC3FBD0" box="[1177,1301,1019,1042]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="4" pageNumber="82" phylum="Tracheophyta" rank="species" species="austriaca">
<emphasis id="FFD9EAC4FFC5FF9D26F094E5FAC3FBD0" box="[1177,1301,1019,1042]" italics="true" pageId="4" pageNumber="82">S. austriaca</emphasis>
</taxonomicName>
,
<taxonomicName id="0AAD4D55FFC5FF9D274B94E5FC90FBF0" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="4" pageNumber="82" phylum="Tracheophyta" rank="species" species="candidissima">
<emphasis id="FFD9EAC4FFC5FF9D274B94E5FC90FBF0" italics="true" pageId="4" pageNumber="82">S. candidissima</emphasis>
</taxonomicName>
,
<taxonomicName id="0AAD4D55FFC5FF9D21389305FC09FBF0" box="[849,991,1051,1074]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="4" pageNumber="82" phylum="Tracheophyta" rank="species" species="cyanescens">
<emphasis id="FFD9EAC4FFC5FF9D21389305FC09FBF0" box="[849,991,1051,1074]" italics="true" pageId="4" pageNumber="82">S. cyanescens</emphasis>
</taxonomicName>
and
<taxonomicName id="0AAD4D55FFC5FF9D26679305FB52FBF0" box="[1038,1156,1051,1074]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="4" pageNumber="82" phylum="Tracheophyta" rank="species" species="pratensis">
<emphasis id="FFD9EAC4FFC5FF9D26679305FB52FBF0" box="[1038,1156,1051,1074]" italics="true" pageId="4" pageNumber="82">S. pratensis</emphasis>
</taxonomicName>
as well (
<bibRefCitation id="A93C4B27FFC5FF9D26B29305FA9BFBF0" author="Kaul, M. L. H." box="[1243,1357,1051,1074]" pageId="4" pageNumber="82" refId="ref14768" refString="Kaul, M. L. H. 1988. Male sterility in higher plants. Monographs on Theoretical and Applied Genetics 10. Berlin &amp; Heidelberg: Springer. https: // doi. org / 10.1007 / 978 - 3 - 642 - 83139 - 3" type="journal volume" year="1988">Kaul, 1988</bibRefCitation>
;
<bibRefCitation id="A93C4B27FFC5FF9D27339305FBF0FB90" author="Zhang, B. &amp; Classen-Bockhoff, R." pageId="4" pageNumber="82" refId="ref19127" refString="Zhang, B. &amp; Classen-Bockhoff, R. 2019. Sex-differential reproduction success and selection on floral traits in gynodioecious Salvia pratensis. B. M. C. Pl. Biol. 19: 375. https: // doi. org / 10.1186 / s 12870 - 019 - 1972 - y" type="journal volume" year="2019">Zhang &amp; Classen-Bockhoff, 2019</bibRefCitation>
). Measurements of quantitative floral traits were performed only on hermaphroditic flowers. Based on the results of the linear mixed-effects models, five of the six investigated floral traits differed significantly between the two taxa: calyx length (
<emphasis id="FFD9EAC4FFC5FF9D262B93A5FB98FB10" box="[1090,1102,1211,1234]" italics="true" pageId="4" pageNumber="82">χ</emphasis>
<superScript id="3AD89B9EFFC5FF9D262693ABFB8FFB07" attach="left" box="[1103,1113,1205,1221]" fontSize="7" pageId="4" pageNumber="82">2</superScript>
= 34.70,
<emphasis id="FFD9EAC4FFC5FF9D26A093A5FB00FB10" box="[1225,1238,1211,1234]" italics="true" pageId="4" pageNumber="82">p</emphasis>
&lt;0.0001), corolla height (
<emphasis id="FFD9EAC4FFC5FF9D210193C5FCA2FB30" box="[872,884,1243,1266]" italics="true" pageId="4" pageNumber="82">χ</emphasis>
<superScript id="3AD89B9EFFC5FF9D211C93CBFCA9FB27" attach="left" box="[885,895,1237,1253]" fontSize="7" pageId="4" pageNumber="82">2</superScript>
= 6.18,
<emphasis id="FFD9EAC4FFC5FF9D218993C5FC3BFB30" box="[992,1005,1243,1266]" italics="true" pageId="4" pageNumber="82">p</emphasis>
= 0.0129), corolla length (
<emphasis id="FFD9EAC4FFC5FF9D277D93C5FAF6FB30" box="[1300,1312,1243,1266]" italics="true" pageId="4" pageNumber="82">χ</emphasis>
<superScript id="3AD89B9EFFC5FF9D274893CBFAFDFB27" attach="left" box="[1313,1323,1237,1253]" fontSize="7" pageId="4" pageNumber="82">2</superScript>
= 107.64,
<emphasis id="FFD9EAC4FFC5FF9D217C93E5FCF4FAD0" box="[789,802,1275,1298]" italics="true" pageId="4" pageNumber="82">p</emphasis>
&lt;0.0001), absolute length of the abaxial lever arm (
<emphasis id="FFD9EAC4FFC5FF9D270293E5FAA1FAD0" box="[1387,1399,1275,1298]" italics="true" pageId="4" pageNumber="82">χ</emphasis>
<superScript id="3AD89B9EFFC5FF9D271193EBFA54FAC7" attach="left" box="[1400,1410,1269,1285]" fontSize="7" pageId="4" pageNumber="82">2</superScript>
= 269.67,
<emphasis id="FFD9EAC4FFC5FF9D21049205FCACFAF0" box="[877,890,1307,1330]" italics="true" pageId="4" pageNumber="82">p</emphasis>
&lt;0.0001), and absolute length of the adaxial lever arm (
<emphasis id="FFD9EAC4FFC5FF9D21279225FC8CFA90" box="[846,858,1339,1362]" italics="true" pageId="4" pageNumber="82">χ</emphasis>
<superScript id="3AD89B9EFFC5FF9D2132922BFCB3FA87" attach="left" box="[859,869,1333,1349]" fontSize="7" pageId="4" pageNumber="82">2</superScript>
= 10.25,
<emphasis id="FFD9EAC4FFC5FF9D21A79225FC0DFA90" box="[974,987,1339,1362]" italics="true" pageId="4" pageNumber="82">p</emphasis>
= 0.0014) had significantly greater values for
<taxonomicName id="0AAD4D55FFC5FF9D21529245FC11FAB0" authority="FDPD" authorityName="FDPD" box="[827,967,1371,1394]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="4" pageNumber="82" phylum="Tracheophyta" rank="genus">
<emphasis id="FFD9EAC4FFC5FF9D21529245FCAAFAB0" box="[827,892,1371,1394]" italics="true" pageId="4" pageNumber="82">Salvia</emphasis>
FDPD
</taxonomicName>
compared to
<taxonomicName id="0AAD4D55FFC5FF9D26339245FB0EFAB0" box="[1114,1240,1371,1394]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="4" pageNumber="82" phylum="Tracheophyta" rank="species" species="austriaca">
<emphasis id="FFD9EAC4FFC5FF9D26339245FB0EFAB0" box="[1114,1240,1371,1394]" italics="true" pageId="4" pageNumber="82">S. austriaca</emphasis>
</taxonomicName>
<taxonomicNameLabel id="E4EA57BFFFC5FF9D26899245FAC6FAB0" box="[1248,1296,1371,1394]" pageId="4" pageNumber="82" sensu="stricto">s.str.</taxonomicNameLabel>
(
<tableCitation id="802F036DFFC5FF9D27779245FAB9FAB0" box="[1310,1391,1371,1394]" captionStart="Table 1" captionStartId="5.[151,204,1348,1369]" captionTargetPageId="5" captionText="Table 1. Biometric data (floral and vegetative) of morphological characters for Salvia austriaca Jacq. (N = 29) and Salvia FDPD (N = 35)." httpUri="http://table.plazi.org/id/99D2665EFFC4FF9C22FE925AFA42FA9A" pageId="4" pageNumber="82" tableUuid="99D2665EFFC4FF9C22FE925AFA42FA9A">Table 1</tableCitation>
). In contrast, we only observed a marginally significant difference between the number of stem leaf pairs (
<emphasis id="FFD9EAC4FFC5FF9D26839285FB20FA70" box="[1258,1270,1435,1458]" italics="true" pageId="4" pageNumber="82">χ</emphasis>
<superScript id="3AD89B9EFFC5FF9D269E928BFAD7FA67" attach="left" box="[1271,1281,1429,1445]" fontSize="7" pageId="4" pageNumber="82">2</superScript>
= 6.11,
<emphasis id="FFD9EAC4FFC5FF9D27189285FAA8FA70" box="[1393,1406,1435,1458]" italics="true" pageId="4" pageNumber="82">p</emphasis>
= 0.0134), where the higher values also belonged to
<taxonomicName id="0AAD4D55FFC5FF9D273592A5FCB6FA30" authority="FDPD." authorityName="FDPD." class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="4" pageNumber="82" phylum="Tracheophyta" rank="genus">
<emphasis id="FFD9EAC4FFC5FF9D273592A5FA4BFA10" box="[1372,1437,1467,1490]" italics="true" pageId="4" pageNumber="82">Salvia</emphasis>
FDPD.
</taxonomicName>
There was no difference in the other vegetative traits between the two taxa (data not shown).
</paragraph>
<paragraph id="CD1236D6FFC5FF9D212B9105FBB4F950" blockId="4.[789,1438,379,1970]" pageId="4" pageNumber="82">
In the PCA, 74.6% of total variation in flower-traits was explained by the first two PC-axes (PC1 – 54.0%, PC2 – 20.6%). However, the two focal taxa were differentiated only along the first PC-axis (
<figureCitation id="55962A53FFC5FF9D267A9165FB84F950" box="[1043,1106,1659,1682]" captionStart="Fig" captionStartId="6.[103,135,1893,1914]" captionTargetBox="[150,1390,1076,1854]" captionTargetId="figure-486@6.[146,1393,1076,1858]" captionTargetPageId="6" captionText="Fig. 4. PCA biplot of Salvia L. samples based on six quantitative floral characters, where the arrows represent loadings. Blue circles, S. austriaca Jacq. (N = 16); red triangles, Salvia FDPD (N = 26); ch, corolla height; cl, corolla length; cal, calyx length; a, absolute length of the abaxial lever arm; b, distance between joint and pollen-sacs; c, absolute length of the adaxial lever arm." figureDoi="http://doi.org/10.5281/zenodo.14047530" httpUri="https://zenodo.org/record/14047530/files/figure.png" pageId="4" pageNumber="82">Fig. 4</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="85B7655DFFC5FF9F212B9185FAC6FD90" lastPageId="6" lastPageNumber="84" pageId="4" pageNumber="82" type="discussion">
<paragraph id="CD1236D6FFC5FF9F212B9185FEC8FED0" blockId="4.[789,1438,379,1970]" lastBlockId="6.[103,751,187,1010]" lastPageId="6" lastPageNumber="84" pageId="4" pageNumber="82">
<emphasis id="FFD9EAC4FFC5FF9D212B9185FC51F910" bold="true" pageId="4" pageNumber="82">
Phylogenomics, species delimitation and divergence dating.
<emphasis id="FFD9EAC4FFC5FF9D210591A3FC51F910" bold="true" box="[876,903,1725,1746]" italics="true" pageId="4" pageNumber="82">—</emphasis>
</emphasis>
Our maximum likelihood (ML) phylogenetic tree reconstruction based on unlinked genome-wide SNPs produced a well-resolved phylogram (
<figureCitation id="55962A53FFC5FF9D26ED91E5FB39F8D0" box="[1156,1263,1787,1810]" captionStart="Fig" captionStartId="7.[151,183,1781,1802]" captionTargetBox="[250,1385,716,1745]" captionTargetId="figure-251@7.[250,1386,712,1746]" captionTargetPageId="7" captionText="Fig.5. Phylogenetic relationships between the studied taxa. A, Phylogenetic tree shown as a phylogram resulting from a ML search in RAxML using the ascertainment bias correction option. Circles at nodes represent maximum bootstrap support (BS = 100), thick branches represent BS ≥70, and thin branches represent BS &lt;70, where branch thickness increases in proportion to bootstrap support. The two main internal branches were shortened to condense the figure. B, Topology of the untruncated ML phylogram. C, Posterior membership probabilities based on the first two discriminant functions of four PCs using group assignment from k-means clustering at K = 5 in a DAPC analysis. The order of individuals follows the order of the tips of the phylogenetic tree. The beginning of an individual’s ID refers to the taxon: rev, Salvia FDPD; aus, S. austriaca Jacq. s.str.; prat, S. pratensis L.; nemo, S. nemorosa L.; nut, S. nutans L." figureDoi="http://doi.org/10.5281/zenodo.14047532" httpUri="https://zenodo.org/record/14047532/files/figure.png" pageId="4" pageNumber="82">Fig. 5A,B</figureCitation>
), in which samples of the same a priori classified taxa are monophyletic and strongly supported (BS = 100), but some of the tip nodes received moderate support (i.e., 70 ≤ BS ≤ 99). The topology strongly supports the separation of the
<taxonomicName id="0AAD4D55FFC5FF9D26C59065FA8EF850" box="[1196,1368,1915,1938]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="4" pageNumber="82" phylum="Tracheophyta" rank="species" species="austriaca">
<emphasis id="FFD9EAC4FFC5FF9D26C59065FA8EF850" box="[1196,1368,1915,1938]" italics="true" pageId="4" pageNumber="82">Salvia austriaca</emphasis>
</taxonomicName>
group from the rest of the samples (our a priori outgroup) along the longest branch on the tree (
<figureCitation id="55962A53FFC7FF9F23E797A5FE37FF10" box="[398,481,187,210]" captionStart="Fig" captionStartId="7.[151,183,1781,1802]" captionTargetBox="[250,1385,716,1745]" captionTargetId="figure-251@7.[250,1386,712,1746]" captionTargetPageId="7" captionText="Fig.5. Phylogenetic relationships between the studied taxa. A, Phylogenetic tree shown as a phylogram resulting from a ML search in RAxML using the ascertainment bias correction option. Circles at nodes represent maximum bootstrap support (BS = 100), thick branches represent BS ≥70, and thin branches represent BS &lt;70, where branch thickness increases in proportion to bootstrap support. The two main internal branches were shortened to condense the figure. B, Topology of the untruncated ML phylogram. C, Posterior membership probabilities based on the first two discriminant functions of four PCs using group assignment from k-means clustering at K = 5 in a DAPC analysis. The order of individuals follows the order of the tips of the phylogenetic tree. The beginning of an individual’s ID refers to the taxon: rev, Salvia FDPD; aus, S. austriaca Jacq. s.str.; prat, S. pratensis L.; nemo, S. nemorosa L.; nut, S. nutans L." figureDoi="http://doi.org/10.5281/zenodo.14047532" httpUri="https://zenodo.org/record/14047532/files/figure.png" pageId="6" pageNumber="84">Fig. 5A</figureCitation>
). Within the latter clade,
<taxonomicName id="0AAD4D55FFC7FF9F220E97C5FF1CFF30" box="[103,202,219,242]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="84" phylum="Tracheophyta" rank="species" species="nutans">
<emphasis id="FFD9EAC4FFC7FF9F220E97C5FF1CFF30" box="[103,202,219,242]" italics="true" pageId="6" pageNumber="84">S. nutans</emphasis>
</taxonomicName>
is sister to the crown-clade formed by
<taxonomicName id="0AAD4D55FFC7FF9F201B97C5FD38FF30" box="[626,750,219,242]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="84" phylum="Tracheophyta" rank="species" species="pratensis">
<emphasis id="FFD9EAC4FFC7FF9F201B97C5FD38FF30" box="[626,750,219,242]" italics="true" pageId="6" pageNumber="84">S. pratensis</emphasis>
</taxonomicName>
and
<taxonomicName id="0AAD4D55FFC7FF9F22FD97E5FEC1FED0" box="[148,279,251,274]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="84" phylum="Tracheophyta" rank="species" species="nemorosa">
<emphasis id="FFD9EAC4FFC7FF9F22FD97E5FEC1FED0" box="[148,279,251,274]" italics="true" pageId="6" pageNumber="84">S. nemorosa</emphasis>
</taxonomicName>
.
</paragraph>
<caption id="99D2665EFFC4FF9C22FE9396FA64FB7B" ID-DOI="http://doi.org/10.5281/zenodo.14047528" ID-Zenodo-Dep="14047528" httpUri="https://zenodo.org/record/14047528/files/figure.png" pageId="5" pageNumber="83" startId="5.[151,183,1160,1181]" targetBox="[152,1483,187,1126]" targetPageId="5" targetType="figure">
<paragraph id="CD1236D6FFC4FF9C22FE9396FA64FB7B" blockId="5.[151,1484,1160,1209]" pageId="5" pageNumber="83">
<emphasis id="FFD9EAC4FFC4FF9C22FE9396FF05FB5F" bold="true" box="[151,211,1160,1181]" pageId="5" pageNumber="83">Fig. 3.</emphasis>
Distribution of
<taxonomicName id="0AAD4D55FFC4FF9C23029397FE30FB5F" authority="FDPD" authorityName="FDPD" box="[363,486,1161,1181]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="5" pageNumber="83" phylum="Tracheophyta" rank="genus">
<emphasis id="FFD9EAC4FFC4FF9C23029397FE72FB5F" box="[363,420,1161,1181]" italics="true" pageId="5" pageNumber="83">Salvia</emphasis>
FDPD
</taxonomicName>
and
<taxonomicName id="0AAD4D55FFC4FF9C207F9397FCE1FB5F" authority="Jacq. Delimitation" authorityName="Jacq. Delimitation" box="[534,823,1161,1181]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="5" pageNumber="83" phylum="Tracheophyta" rank="species" species="austriaca">
<emphasis id="FFD9EAC4FFC4FF9C207F9397FD50FB5F" box="[534,646,1161,1181]" italics="true" pageId="5" pageNumber="83">S. austriaca</emphasis>
Jacq. Delimitation
</taxonomicName>
of the steppe zone follows
<bibRefCitation id="A93C4B27FFC4FF9C26529397FAC6FB5F" author="Lavrenko, E. M. &amp; Karamysheva, Z. V. &amp; Nikulina, R. I." box="[1083,1296,1161,1181]" pageId="5" pageNumber="83" refId="ref15249" refString="Lavrenko, E. M., Karamysheva, Z. V. &amp; Nikulina, R. I. 1991. Stepi Evrazii [Steppes of Eurasia]. Leningrad: Nauka." type="book" year="1991">Lavrenko &amp; al. (1991)</bibRefCitation>
. Delimitation of the Crimean Submediterranean forest ecoregion follows
<bibRefCitation id="A93C4B27FFC4FF9C201793BBFCF8FB7B" author="Olson, D. M. &amp; Dinerstein, E. &amp; Wikramanayake, E. D. &amp; Burgess, N. D. &amp; Powell, G. V. N. &amp; Underwood, E. C. &amp; J. A &amp; Itoua, I. &amp; Strand, H. E. &amp; Morrison, J. C. &amp; Loucks, C. J. &amp; Allnutt, T. F. &amp; Ricketts, T. H. &amp; Kura, Y. &amp; Lamoreux, J. F. &amp; Wettengel, W. W. &amp; Hedao, P. &amp; Kassem, K. R." box="[638,814,1189,1209]" pageId="5" pageNumber="83" pagination="933 - 938" refId="ref16518" refString="Olson, D. M., Dinerstein, E., Wikramanayake, E. D., Burgess, N. D., Powell, G. V. N., Underwood, E. C., D' amico, J. A., Itoua, I., Strand, H. E., Morrison, J. C., Loucks, C. J., Allnutt, T. F., Ricketts, T. H., Kura, Y., Lamoreux, J. F., Wettengel, W. W., Hedao, P. &amp; Kassem, K. R. 2001. Terrestrial ecoregions of the world: A new map of life on Earth: A new global map of terrestrial ecoregions provides an innovative tool for conserving biodiversity. Bioscience 51: 933 - 938. https: // doi. org / 10.1641 / 0006 - 3568 (2001) 051 [0933: TEOTWA] 2.0. CO; 2" type="journal article" year="2001">Olson &amp; al. (2001)</bibRefCitation>
. For geographic coordinates of each observation, see suppl. Table S1.
</paragraph>
</caption>
<caption id="99D2665EFFC4FF9C22FE925AFA42FA9A" ID-Table-UUID="99D2665EFFC4FF9C22FE925AFA42FA9A" box="[151,1428,1348,1369]" httpUri="http://table.plazi.org/id/99D2665EFFC4FF9C22FE925AFA42FA9A" pageId="5" pageNumber="83" startId="5.[151,204,1348,1369]" targetBox="[162,1469,1392,1900]" targetIsTable="true" targetPageId="5" targetType="table">
<paragraph id="CD1236D6FFC4FF9C22FE925AFA42FA9A" blockId="5.[151,1428,1348,1369]" box="[151,1428,1348,1369]" pageId="5" pageNumber="83">
<emphasis id="FFD9EAC4FFC4FF9C22FE925AFF32FA9B" bold="true" box="[151,228,1348,1369]" pageId="5" pageNumber="83">Table 1.</emphasis>
Biometric data (floral and vegetative) of morphological characters for
<taxonomicName id="0AAD4D55FFC4FF9C2119925AFBEAFA9A" authority="Jacq." authorityName="Jacq." box="[880,1084,1348,1368]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="5" pageNumber="83" phylum="Tracheophyta" rank="species" species="austriaca">
<emphasis id="FFD9EAC4FFC4FF9C2119925AFBD0FA9A" box="[880,1030,1348,1368]" italics="true" pageId="5" pageNumber="83">Salvia austriaca</emphasis>
Jacq.
</taxonomicName>
(N = 29) and
<taxonomicName id="0AAD4D55FFC4FF9C26D7925AFAEEFA9A" authority="FDPD" authorityName="FDPD" box="[1214,1336,1348,1368]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="5" pageNumber="83" phylum="Tracheophyta" rank="genus">
<emphasis id="FFD9EAC4FFC4FF9C26D7925AFB21FA9A" box="[1214,1271,1348,1368]" italics="true" pageId="5" pageNumber="83">Salvia</emphasis>
FDPD
</taxonomicName>
(N = 35).
</paragraph>
</caption>
<paragraph id="CD1236D6FFC4FF9C22CB926EFA42F8AE" pageId="5" pageNumber="83">
<table id="BFADC476FFC4006622CB926EFA6BF8AE" box="[162,1469,1392,1900]" gridcols="3" gridrows="11" pageId="5" pageNumber="83">
<tr id="739D3494FFC4006622CB926EFA6BFA6F" box="[162,1469,1392,1453]" gridrow="0" pageId="5" pageNumber="83">
<th id="304C5DE8FFC4006622CB926EFD99FA6F" box="[162,591,1392,1453]" gridcol="0" gridrow="0" pageId="5" pageNumber="83">Variable Floral traits</th>
<th id="304C5DE8FFC400662169926EFBDAFA6F" box="[768,1036,1392,1453]" gridcol="1" gridrow="0" pageId="5" pageNumber="83">
<taxonomicName id="0AAD4D55FFC4FF9C216A926EFC95FA6F" authority="Values" authorityName="Values" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="5" pageNumber="83" phylum="Tracheophyta" rank="species" species="austriaca">
<emphasis id="FFD9EAC4FFC4FF9C216A926EFC4FFA46" box="[771,921,1392,1412]" italics="true" pageId="5" pageNumber="83">Salvia austriaca</emphasis>
Values
</taxonomicName>
in mm
</th>
<th id="304C5DE8FFC4006626D6926EFA6BFA6F" box="[1215,1469,1392,1453]" gridcol="2" gridrow="0" pageId="5" pageNumber="83">
<taxonomicName id="0AAD4D55FFC4FF9C26D6926EFB29FA6F" authority="FDPD Values" authorityName="FDPD Values" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="5" pageNumber="83" phylum="Tracheophyta" rank="genus">
<emphasis id="FFD9EAC4FFC4FF9C26D6926EFB2EFA46" box="[1215,1272,1392,1412]" italics="true" pageId="5" pageNumber="83">Salvia</emphasis>
FDPD Values
</taxonomicName>
in mm
</th>
</tr>
<tr id="739D3494FFC4006622CB92DAFA6BFA1B" box="[162,1469,1476,1497]" gridrow="1" pageId="5" pageNumber="83">
<th id="304C5DE8FFC4006622CB92DAFD99FA1B" box="[162,591,1476,1497]" gridcol="0" gridrow="1" pageId="5" pageNumber="83">ch* (corolla height)</th>
<td id="304C5DE8FFC40066216992DAFBDAFA1B" box="[768,1036,1476,1497]" gridcol="1" gridrow="1" pageId="5" pageNumber="83">(6.24) 10.69 ± 0.55 (17)</td>
<td id="304C5DE8FFC4006626D692DAFA6BFA1B" box="[1215,1469,1476,1497]" gridcol="2" gridrow="1" pageId="5" pageNumber="83">(8.36) 12.38 ± 0.54 (21.33)</td>
</tr>
<tr id="739D3494FFC4006622CB92F2FA6BF9C3" box="[162,1469,1516,1537]" gridrow="2" pageId="5" pageNumber="83">
<th id="304C5DE8FFC4006622CB92F2FD99F9C3" box="[162,591,1516,1537]" gridcol="0" gridrow="2" pageId="5" pageNumber="83">cl* (corolla length)</th>
<td id="304C5DE8FFC40066216992F2FBDAF9C3" box="[768,1036,1516,1537]" gridcol="1" gridrow="2" pageId="5" pageNumber="83">(14.27) 17.7 ± 0.26 (20.4)</td>
<td id="304C5DE8FFC4006626D692F2FA6BF9C3" box="[1215,1469,1516,1537]" gridcol="2" gridrow="2" pageId="5" pageNumber="83">(19) 22.75 ± 0.4 (26.34)</td>
</tr>
<tr id="739D3494FFC4006622CB910AFA6BF9EB" box="[162,1469,1556,1577]" gridrow="3" pageId="5" pageNumber="83">
<th id="304C5DE8FFC4006622CB910AFD99F9EB" box="[162,591,1556,1577]" gridcol="0" gridrow="3" pageId="5" pageNumber="83">a* (absolute length of the abaxial lever arm)</th>
<td id="304C5DE8FFC400662169910AFBDAF9EB" box="[768,1036,1556,1577]" gridcol="1" gridrow="3" pageId="5" pageNumber="83">(12.79) 14.44 ± 0.22 (17.23)</td>
<td id="304C5DE8FFC4006626D6910AFA6BF9EB" box="[1215,1469,1556,1577]" gridcol="2" gridrow="3" pageId="5" pageNumber="83">(16.04) 20.71 ± 0.31 (24)</td>
</tr>
<tr id="739D3494FFC4006622CB9123FA6BF993" box="[162,1469,1597,1617]" gridrow="4" pageId="5" pageNumber="83">
<th id="304C5DE8FFC4006622CB9123FD99F993" box="[162,591,1597,1617]" gridcol="0" gridrow="4" pageId="5" pageNumber="83">b (distance between joint and pollen-sacs)</th>
<td id="304C5DE8FFC4006621699123FBDAF993" box="[768,1036,1597,1617]" gridcol="1" gridrow="4" pageId="5" pageNumber="83">(11.25) 14.06 ± 0.35 (18.57)</td>
<td id="304C5DE8FFC4006626D69123FA6BF993" box="[1215,1469,1597,1617]" gridcol="2" gridrow="4" pageId="5" pageNumber="83">(9.69) 13.69 ± 0.39 (18.27)</td>
</tr>
<tr id="739D3494FFC4006622CB917AFA6BF9BB" box="[162,1469,1636,1657]" gridrow="5" pageId="5" pageNumber="83">
<th id="304C5DE8FFC4006622CB917AFD99F9BB" box="[162,591,1636,1657]" gridcol="0" gridrow="5" pageId="5" pageNumber="83">c* (absolute length of the adaxial lever arm)</th>
<td id="304C5DE8FFC400662169917AFBDAF9BB" box="[768,1036,1636,1657]" gridcol="1" gridrow="5" pageId="5" pageNumber="83">(2.15) 2.72 ± 0.07 (3.54)</td>
<td id="304C5DE8FFC4006626D6917AFA6BF9BB" box="[1215,1469,1636,1657]" gridcol="2" gridrow="5" pageId="5" pageNumber="83">(2.23) 3.06 ± 0.09 (4.04)</td>
</tr>
<tr id="739D3494FFC4006622CB9192FA6BF90B" box="[162,1469,1676,1737]" gridrow="6" pageId="5" pageNumber="83">
<th id="304C5DE8FFC4006622CB9192FD99F90B" box="[162,591,1676,1737]" gridcol="0" gridrow="6" pageId="5" pageNumber="83">cal* (calyx length) Vegetative traits</th>
<td id="304C5DE8FFC4006621699192FBDAF90B" box="[768,1036,1676,1737]" gridcol="1" gridrow="6" pageId="5" pageNumber="83">(7.4) 8.22 ± 0.15 (9.86) Values in cm, except stlp</td>
<td id="304C5DE8FFC4006626D69192FA6BF90B" box="[1215,1469,1676,1737]" gridcol="2" gridrow="6" pageId="5" pageNumber="83">(8.05) 9.54 ± 0.15 (10.82) Values in cm, except stlp</td>
</tr>
<tr id="739D3494FFC4006622CB91FEFA6BF936" box="[162,1469,1760,1780]" gridrow="7" pageId="5" pageNumber="83">
<th id="304C5DE8FFC4006622CB91FEFD99F936" box="[162,591,1760,1780]" gridcol="0" gridrow="7" pageId="5" pageNumber="83">sth (stem height)</th>
<td id="304C5DE8FFC40066216991FEFBDAF936" box="[768,1036,1760,1780]" gridcol="1" gridrow="7" pageId="5" pageNumber="83">(24) 56.23 ± 2.27 (79)</td>
<td id="304C5DE8FFC4006626D691FEFA6BF936" box="[1215,1469,1760,1780]" gridcol="2" gridrow="7" pageId="5" pageNumber="83">(8) 61.2 ± 3.65 (104)</td>
</tr>
<tr id="739D3494FFC4006622CB9016FA6BF8DF" box="[162,1469,1800,1821]" gridrow="8" pageId="5" pageNumber="83">
<th id="304C5DE8FFC4006622CB9016FD99F8DF" box="[162,591,1800,1821]" gridcol="0" gridrow="8" pageId="5" pageNumber="83">stlp* (number of stem leaf pairs)</th>
<td id="304C5DE8FFC4006621699016FBDAF8DF" box="[768,1036,1800,1821]" gridcol="1" gridrow="8" pageId="5" pageNumber="83">(1) 1.53 ± 0.14 (3)</td>
<td id="304C5DE8FFC4006626D69016FA6BF8DF" box="[1215,1469,1800,1821]" gridcol="2" gridrow="8" pageId="5" pageNumber="83">(1) 1.69 ± 0.09 (3)</td>
</tr>
<tr id="739D3494FFC4006622CB902FFA6BF887" box="[162,1469,1841,1861]" gridrow="9" pageId="5" pageNumber="83">
<th id="304C5DE8FFC4006622CB902FFD99F887" box="[162,591,1841,1861]" gridcol="0" gridrow="9" pageId="5" pageNumber="83">lpet (length of petiole of rosette leaf)</th>
<td id="304C5DE8FFC400662169902FFBDAF887" box="[768,1036,1841,1861]" gridcol="1" gridrow="9" pageId="5" pageNumber="83">(1.8) 3.67 ± 0.32 (7.5)</td>
<td id="304C5DE8FFC4006626D6902FFA6BF887" box="[1215,1469,1841,1861]" gridcol="2" gridrow="9" pageId="5" pageNumber="83">(1) 4.46 ± 0.44 (12)</td>
</tr>
<tr id="739D3494FFC4006622CB9046FA6BF8AE" box="[162,1469,1880,1900]" gridrow="10" pageId="5" pageNumber="83">
<th id="304C5DE8FFC4006622CB9046FD99F8AE" box="[162,591,1880,1900]" gridcol="0" gridrow="10" pageId="5" pageNumber="83">lobl (length of leaf blade of rosette leaf)</th>
<td id="304C5DE8FFC4006621699046FBDAF8AE" box="[768,1036,1880,1900]" gridcol="1" gridrow="10" pageId="5" pageNumber="83">(5.2) 11.61 ± 0.53 (14.5)</td>
<td id="304C5DE8FFC4006626D69046FA6BF8AE" box="[1215,1469,1880,1900]" gridcol="2" gridrow="10" pageId="5" pageNumber="83">(8.5) 13.83 ± 0.77 (25)</td>
</tr>
</table>
</paragraph>
<paragraph id="CD1236D6FFC4FF9C22FE909CFCC1F873" blockId="5.[151,791,1922,1970]" pageId="5" pageNumber="83">
<tableNote id="304B3758FFC4FF9C22FE909CFCC1F873" pageId="5" pageNumber="83" targetBox="[162,1469,1392,1900]" targetPageId="5">
Values represent (minimum) mean ± standard error (maximum). * indicates significant differences between measurements (
<emphasis id="FFD9EAC4FFC4FF9C20D29083FD11F873" box="[699,711,1949,1969]" italics="true" pageId="5" pageNumber="83">p</emphasis>
&lt;0.05).
</tableNote>
</paragraph>
<paragraph id="CD1236D6FFC7FF9F22FD9605FDC1FD10" blockId="6.[103,751,187,1010]" pageId="6" pageNumber="84">
The species tree reconstruction based on coalescence analysis (
<figureCitation id="55962A53FFC7FF9F22A79625FEC7FE90" box="[206,273,315,338]" captionStart="Fig" captionStartId="8.[103,135,1518,1539]" captionTargetBox="[116,1436,519,1492]" captionTargetPageId="8" captionText="Fig. 6. The dated species tree of the studied Salvia L. taxa based on the best model of species delimitation as inferred by SNAPP. Blue bars represent 95% confidence intervals. The geological time chart below the chronogram is presented according to the International Union of Stratigraphy." figureDoi="http://doi.org/10.5281/zenodo.14057462" httpUri="https://zenodo.org/record/14057462/files/figure.png" pageId="6" pageNumber="84">Fig. 6</figureCitation>
) yielded a fully congruent topology to the ML-tree, in which taxa of the
<taxonomicName id="0AAD4D55FFC7FF9F23F49645FD9EFEB0" box="[413,584,347,370]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="84" phylum="Tracheophyta" rank="species" species="austriaca">
<emphasis id="FFD9EAC4FFC7FF9F23F49645FD9EFEB0" box="[413,584,347,370]" italics="true" pageId="6" pageNumber="84">Salvia austriaca</emphasis>
</taxonomicName>
group are sister to the lineage formed by
<taxonomicName id="0AAD4D55FFC7FF9F231B9665FE02FE50" box="[370,468,379,402]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="84" phylum="Tracheophyta" rank="species" species="nutans">
<emphasis id="FFD9EAC4FFC7FF9F231B9665FE02FE50" box="[370,468,379,402]" italics="true" pageId="6" pageNumber="84">S. nutans</emphasis>
</taxonomicName>
,
<taxonomicName id="0AAD4D55FFC7FF9F238B9665FD8BFE50" box="[482,605,379,402]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="84" phylum="Tracheophyta" rank="species" species="pratensis">
<emphasis id="FFD9EAC4FFC7FF9F238B9665FD8BFE50" box="[482,605,379,402]" italics="true" pageId="6" pageNumber="84">S. pratensis</emphasis>
</taxonomicName>
and
<taxonomicName id="0AAD4D55FFC7FF9F20F89665FF43FE70" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="84" phylum="Tracheophyta" rank="species" species="nemorosa">
<emphasis id="FFD9EAC4FFC7FF9F20F89665FF43FE70" italics="true" pageId="6" pageNumber="84">S. nemorosa</emphasis>
</taxonomicName>
. The next split within this lineage is much younger and is represented by
<taxonomicName id="0AAD4D55FFC7FF9F234196A5FE5AFE10" box="[296,396,443,466]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="84" phylum="Tracheophyta" rank="species" species="nutans">
<emphasis id="FFD9EAC4FFC7FF9F234196A5FE5AFE10" box="[296,396,443,466]" italics="true" pageId="6" pageNumber="84">S. nutans</emphasis>
</taxonomicName>
, which separated from the other two species during the Middle Pleistocene. The following isolation event is marked by the separation of the two taxa of the
<taxonomicName id="0AAD4D55FFC7FF9F220E9505FEC0FDF0" box="[103,278,539,562]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="84" phylum="Tracheophyta" rank="species" species="austriaca">
<emphasis id="FFD9EAC4FFC7FF9F220E9505FEC0FDF0" box="[103,278,539,562]" italics="true" pageId="6" pageNumber="84">Salvia austriaca</emphasis>
</taxonomicName>
group, which was followed by the split between
<taxonomicName id="0AAD4D55FFC7FF9F22C29525FEFBFD90" box="[171,301,571,594]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="84" phylum="Tracheophyta" rank="species" species="nemorosa">
<emphasis id="FFD9EAC4FFC7FF9F22C29525FEFBFD90" box="[171,301,571,594]" italics="true" pageId="6" pageNumber="84">S. nemorosa</emphasis>
</taxonomicName>
and
<taxonomicName id="0AAD4D55FFC7FF9F23369525FE0FFD90" box="[351,473,571,594]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="84" phylum="Tracheophyta" rank="species" species="pratensis">
<emphasis id="FFD9EAC4FFC7FF9F23369525FE0FFD90" box="[351,473,571,594]" italics="true" pageId="6" pageNumber="84">S. pratensis</emphasis>
</taxonomicName>
. All nodes were fully supported in the species tree (posterior probability [PP] = 1.0). The divergence dating analysis indicated that
<taxonomicName id="0AAD4D55FFC7FF9F20039565FD38FD50" box="[618,750,635,658]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="84" phylum="Tracheophyta" rank="species" species="austriaca">
<emphasis id="FFD9EAC4FFC7FF9F20039565FD38FD50" box="[618,750,635,658]" italics="true" pageId="6" pageNumber="84">S. austriaca</emphasis>
</taxonomicName>
<taxonomicNameLabel id="E4EA57BFFFC7FF9F220E9585FF4EFD70" box="[103,152,667,690]" pageId="6" pageNumber="84" sensu="stricto">s.str.</taxonomicNameLabel>
and
<taxonomicName id="0AAD4D55FFC7FF9F22A59585FE8FFD70" authority="FDPD" authorityName="FDPD" box="[204,345,667,690]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="84" phylum="Tracheophyta" rank="genus">
<emphasis id="FFD9EAC4FFC7FF9F22A59585FED8FD70" box="[204,270,667,690]" italics="true" pageId="6" pageNumber="84">Salvia</emphasis>
FDPD
</taxonomicName>
diverged 0.197 mya (95% confidence interval [CI]:
<geoCoordinate id="A8995011FFC7FF9F229195A5FEE2FD10" box="[248,308,699,722]" degrees="0.302" direction="north" orientation="latitude" pageId="6" pageNumber="84" precision="55" value="0.302">0.302</geoCoordinate>
<geoCoordinate id="A8995011FFC7FF9F235D95A4FEABFD10" box="[308,381,698,722]" degrees="0.087" direction="west" orientation="longitude" pageId="6" pageNumber="84" precision="55" value="-0.087">–0.087</geoCoordinate>
mya) (
<figureCitation id="55962A53FFC7FF9F23A095A5FDD1FD10" box="[457,519,699,722]" captionStart="Fig" captionStartId="8.[103,135,1518,1539]" captionTargetBox="[116,1436,519,1492]" captionTargetPageId="8" captionText="Fig. 6. The dated species tree of the studied Salvia L. taxa based on the best model of species delimitation as inferred by SNAPP. Blue bars represent 95% confidence intervals. The geological time chart below the chronogram is presented according to the International Union of Stratigraphy." figureDoi="http://doi.org/10.5281/zenodo.14057462" httpUri="https://zenodo.org/record/14057462/files/figure.png" pageId="6" pageNumber="84">Fig. 6</figureCitation>
).
</paragraph>
<paragraph id="CD1236D6FFC7FF9F22FD95C5FAFCFED0" blockId="6.[103,751,187,1010]" lastBlockId="6.[789,1437,187,594]" pageId="6" pageNumber="84">
The DAPC analysis attained the lowest BIC scores to models with 5 and 7 clusters (suppl.
<figureCitation id="55962A53FFC7FF9F238195E5FDE2FCD0" box="[488,564,763,786]" captionStart="Fig" captionStartId="2.[103,135,1893,1914]" captionTargetBox="[104,1435,980,1859]" captionTargetId="figure-404@2.[104,1435,980,1859]" captionTargetPageId="2" captionText="Fig. 1. A, Spatial distribution of sampled Salvia L. populations; B, Study area highlighted in red on a map of Europe; C, Transylvanian sites enlarged; D, Dobrogean sites enlarged. — White, Salvia austriaca Jacq. s.str.; black, Salvia FDPD; circle, genetic samples; diamond, morphometric samples; square: both genetic and morphometric samples; white lines represent country borders." figureDoi="http://doi.org/10.5281/zenodo.14047524" httpUri="https://zenodo.org/record/14047524/files/figure.png" pageId="6" pageNumber="84">Fig. S1</figureCitation>
). At
<emphasis id="FFD9EAC4FFC7FF9F200395E5FDAAFCD0" box="[618,636,763,786]" italics="true" pageId="6" pageNumber="84">K</emphasis>
= 5, all individuals were assigned into a cluster—according to existing classification and splitting the taxa of the
<taxonomicName id="0AAD4D55FFC7FF9F20579425FD38FC90" box="[574,750,827,850]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="84" phylum="Tracheophyta" rank="species" species="austriaca">
<emphasis id="FFD9EAC4FFC7FF9F20579425FD38FC90" box="[574,750,827,850]" italics="true" pageId="6" pageNumber="84">Salvia austriaca</emphasis>
</taxonomicName>
group—with full posterior probability (
<figureCitation id="55962A53FFC7FF9F206B9445FD87FCB0" box="[514,593,859,882]" captionStart="Fig" captionStartId="7.[151,183,1781,1802]" captionTargetBox="[250,1385,716,1745]" captionTargetId="figure-251@7.[250,1386,712,1746]" captionTargetPageId="7" captionText="Fig.5. Phylogenetic relationships between the studied taxa. A, Phylogenetic tree shown as a phylogram resulting from a ML search in RAxML using the ascertainment bias correction option. Circles at nodes represent maximum bootstrap support (BS = 100), thick branches represent BS ≥70, and thin branches represent BS &lt;70, where branch thickness increases in proportion to bootstrap support. The two main internal branches were shortened to condense the figure. B, Topology of the untruncated ML phylogram. C, Posterior membership probabilities based on the first two discriminant functions of four PCs using group assignment from k-means clustering at K = 5 in a DAPC analysis. The order of individuals follows the order of the tips of the phylogenetic tree. The beginning of an individual’s ID refers to the taxon: rev, Salvia FDPD; aus, S. austriaca Jacq. s.str.; prat, S. pratensis L.; nemo, S. nemorosa L.; nut, S. nutans L." figureDoi="http://doi.org/10.5281/zenodo.14047532" httpUri="https://zenodo.org/record/14047532/files/figure.png" pageId="6" pageNumber="84">Fig. 5C</figureCitation>
). Interestingly, models with higher number of clusters assigned individuals from
<taxonomicName id="0AAD4D55FFC7FF9F22CE9485FEE6FC70" box="[167,304,923,946]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="84" phylum="Tracheophyta" rank="species" species="nemorosa">
<emphasis id="FFD9EAC4FFC7FF9F22CE9485FEE6FC70" box="[167,304,923,946]" italics="true" pageId="6" pageNumber="84">S. nemorosa</emphasis>
</taxonomicName>
and
<taxonomicName id="0AAD4D55FFC7FF9F23199485FE27FC70" box="[368,497,923,946]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="84" phylum="Tracheophyta" rank="species" species="pratensis">
<emphasis id="FFD9EAC4FFC7FF9F23199485FE27FC70" box="[368,497,923,946]" italics="true" pageId="6" pageNumber="84">S. pratensis</emphasis>
</taxonomicName>
into separate clusters, whereas the clusters formed by samples of the taxa of the
<taxonomicName id="0AAD4D55FFC7FF9F220E94C5FECFFC30" box="[103,281,987,1010]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="84" phylum="Tracheophyta" rank="species" species="austriaca">
<emphasis id="FFD9EAC4FFC7FF9F220E94C5FECFFC30" box="[103,281,987,1010]" italics="true" pageId="6" pageNumber="84">Salvia austriaca</emphasis>
</taxonomicName>
group were not divided (suppl.
<figureCitation id="55962A53FFC7FF9F20E494C5FD34FC30" box="[653,738,987,1010]" captionStart="Fig" captionStartId="3.[1156,1188,1809,1830]" captionTargetBox="[151,1133,1490,1965]" captionTargetId="figure-732@3.[151,1134,1490,1965]" captionTargetPageId="3" captionText="Fig. 2. Quantitative floral traits measured: absolute length of the abaxial lever arm (a), distance between joint and pollen-sacs (b), absolute length of the adaxial lever arm (c). Photo: Gabriel Gigea." figureDoi="http://doi.org/10.5281/zenodo.14047526" httpUri="https://zenodo.org/record/14047526/files/figure.png" pageId="6" pageNumber="84">Fig. S2</figureCitation>
). Similarly, at
<emphasis id="FFD9EAC4FFC7FF9F21F397A5FC7AFF10" box="[922,940,187,210]" italics="true" pageId="6" pageNumber="84">K</emphasis>
= 4,
<taxonomicName id="0AAD4D55FFC7FF9F218897A5FBB5FF10" box="[993,1123,187,210]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="84" phylum="Tracheophyta" rank="species" species="nemorosa">
<emphasis id="FFD9EAC4FFC7FF9F218897A5FBB5FF10" box="[993,1123,187,210]" italics="true" pageId="6" pageNumber="84">S. nemorosa</emphasis>
</taxonomicName>
and
<taxonomicName id="0AAD4D55FFC7FF9F26FC97A5FAD9FF10" box="[1173,1295,187,210]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="84" phylum="Tracheophyta" rank="species" species="pratensis">
<emphasis id="FFD9EAC4FFC7FF9F26FC97A5FAD9FF10" box="[1173,1295,187,210]" italics="true" pageId="6" pageNumber="84">S. pratensis</emphasis>
</taxonomicName>
were lumped together, whereas
<taxonomicName id="0AAD4D55FFC7FF9F21B297C5FB8AFF30" box="[987,1116,219,242]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="84" phylum="Tracheophyta" rank="species" species="austriaca">
<emphasis id="FFD9EAC4FFC7FF9F21B297C5FB8AFF30" box="[987,1116,219,242]" italics="true" pageId="6" pageNumber="84">S. austriaca</emphasis>
</taxonomicName>
<taxonomicNameLabel id="E4EA57BFFFC7FF9F260F97C5FB41FF30" box="[1126,1175,219,242]" pageId="6" pageNumber="84" sensu="stricto">s.str.</taxonomicNameLabel>
and
<taxonomicName id="0AAD4D55FFC7FF9F26B897C5FAB6FF30" authority="FDPD" authorityName="FDPD" box="[1233,1376,219,242]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="84" phylum="Tracheophyta" rank="genus">
<emphasis id="FFD9EAC4FFC7FF9F26B897C5FAC5FF30" box="[1233,1299,219,242]" italics="true" pageId="6" pageNumber="84">Salvia</emphasis>
FDPD
</taxonomicName>
were still conceived as separate groups (suppl.
<figureCitation id="55962A53FFC7FF9F26A797E5FACBFED0" box="[1230,1309,251,274]" captionStart="Fig" captionStartId="3.[1156,1188,1809,1830]" captionTargetBox="[151,1133,1490,1965]" captionTargetId="figure-732@3.[151,1134,1490,1965]" captionTargetPageId="3" captionText="Fig. 2. Quantitative floral traits measured: absolute length of the abaxial lever arm (a), distance between joint and pollen-sacs (b), absolute length of the adaxial lever arm (c). Photo: Gabriel Gigea." figureDoi="http://doi.org/10.5281/zenodo.14047526" httpUri="https://zenodo.org/record/14047526/files/figure.png" pageId="6" pageNumber="84">Fig. S2</figureCitation>
).
</paragraph>
<paragraph id="CD1236D6FFC7FF9F212B9605FAC6FD90" blockId="6.[789,1437,187,594]" pageId="6" pageNumber="84">
The BFD favored the alternative model, which considered
<taxonomicName id="0AAD4D55FFC7FF9F217C9625FC6AFE90" box="[789,956,315,338]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="84" phylum="Tracheophyta" rank="species" sensu="stricto" species="austriaca">
<emphasis id="FFD9EAC4FFC7FF9F217C9625FC6AFE90" box="[789,956,315,338]" italics="true" pageId="6" pageNumber="84">Salvia austriaca</emphasis>
</taxonomicName>
<taxonomicNameLabel id="E4EA57BFFFC7FF9F21A89625FC26FE90" box="[961,1008,315,338]" pageId="6" pageNumber="84" sensu="stricto">s.str.</taxonomicNameLabel>
and
<taxonomicName id="0AAD4D55FFC7FF9F26769625FB71FE90" authority="FDPD" authorityName="FDPD" box="[1055,1191,315,338]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="84" phylum="Tracheophyta" rank="genus">
<emphasis id="FFD9EAC4FFC7FF9F26769625FBB6FE90" box="[1055,1120,315,338]" italics="true" pageId="6" pageNumber="84">Salvia</emphasis>
FDPD
</taxonomicName>
as separate entities. The null model, grouping all individuals into one species within the
<taxonomicName id="0AAD4D55FFC7FF9F217C9665FC6BFE50" box="[789,957,379,402]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="84" phylum="Tracheophyta" rank="species" species="austriaca">
<emphasis id="FFD9EAC4FFC7FF9F217C9665FC6BFE50" box="[789,957,379,402]" italics="true" pageId="6" pageNumber="84">Salvia austriaca</emphasis>
</taxonomicName>
group (i.e.,
<taxonomicName id="0AAD4D55FFC7FF9F26539665FB63FE50" box="[1082,1205,379,402]" class="Magnoliopsida" family="Lamiaceae" genus="Salvia" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="84" phylum="Tracheophyta" rank="species" species="austriaca">
<emphasis id="FFD9EAC4FFC7FF9F26539665FB63FE50" box="[1082,1205,379,402]" italics="true" pageId="6" pageNumber="84">S. austriaca</emphasis>
</taxonomicName>
<taxonomicNameLabel id="E4EA57BFFFC7FF9F26D29665FB0EFE50" box="[1211,1240,379,402]" pageId="6" pageNumber="84" sensu="lato">s.l.</taxonomicNameLabel>
), performed worse in both repetitions with a mean BF value of
<quantity id="0A559B33FFC7FF9F26889685FA93FE70" box="[1249,1349,411,434]" metricMagnitude="0" metricUnit="m" metricValue="8.856218" pageId="6" pageNumber="84" unit="in" value="348.67">348.67 in</quantity>
favor of the alternative model (
<tableCitation id="802F036DFFC7FF9F219596A5FB9AFE10" box="[1020,1100,443,466]" captionStart="Table 2" captionStartId="8.[103,156,1673,1694]" captionTargetId="graphics-365@8.[103,1436,1813,1853]" captionTargetPageId="8" captionText="Table 2. Marginal likelihood estimates (lnMLE) and Bayes factors (2lnBF) for alternative delimitation hypotheses: grouping Salvia austriaca Jacq. and Salvia FDPD individuals into one species, S. austriaca s.l., or splitting them into two species." httpUri="http://table.plazi.org/id/99D2665EFFC9FF91220E9197FC3BF97A" pageId="6" pageNumber="84" tableUuid="99D2665EFFC9FF91220E9197FC3BF97A">Table 2</tableCitation>
) across the two repetitions. This is considered decisive according to
<bibRefCitation id="A93C4B27FFC7FF9F26F296C5FA4EFE30" author="Kass, R. E. &amp; Raftery, A. E." box="[1179,1432,475,498]" pageId="6" pageNumber="84" pagination="773 - 795" refId="ref14685" refString="Kass, R. E. &amp; Raftery, A. E. 1995. Bayes factors. J. Amer. Statist. Assoc. 90: 773 - 795. https: // doi. org / 10.1080 / 01621459.1995.104 76572" type="journal article" year="1995">Kass &amp; Raftery (1995)</bibRefCitation>
. Mean ESS was 336.34 (standard deviation = 200.74) across both models and all steps and repetitions, implying that the chain length was adequate to reach convergence.
</paragraph>
</subSubSection>
</treatment>
</document>
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