567 lines
74 KiB
XML
567 lines
74 KiB
XML
<document id="30B0DC40F58E9DE526AA05F911AE6254" ID-DOI="10.1002/tax.13123" ID-ISSN="1996-8175" ID-Zenodo-Dep="14047584" IM.illustrations_approvedBy="julia" IM.tables_requiresApprovalFor="existingObjects,operationResults" IM.taxonomicNames_approvedBy="julia" IM.treatments_approvedBy="julia" checkinTime="1730918342851" checkinUser="felipe" docAuthor="García, Miguel A., Mucina, Ladislav & Nickrent, Daniel L." docDate="2024" docId="F275A6752C574028FF70A72EFB1068C2" docLanguage="en" docName="TAXON.73.1.190-236.pdf" docOrigin="TAXON 73 (1)" docSource="http://dx.doi.org/10.1002/tax.13123" docStyle="DocumentStyle:239A75409313D6488D9D82529296B06F.1:TAXON.2020-.journal_article.open" docStyleId="239A75409313D6488D9D82529296B06F" docStyleName="TAXON.2020-.journal_article.open" docStyleVersion="1" docTitle="Thesium García & Mucina & Nickrent 2024" docType="treatment" docVersion="1" lastPageNumber="228" masterDocId="0E4CDE0D2C73400EFFE4A175FFF36E50" masterDocTitle="A tough nutlet to crack: Resolving the phylogeny of Thesium (Thesiaceae), the largest genus in Santalales" masterLastPageNumber="236" masterPageNumber="190" pageNumber="226" updateTime="1731336947450" updateUser="julia" zenodo-license-document="CC-BY-NC-ND-4.0">
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<mods:title id="A84AEE696AB10F1DC133DFAA1E928298">A tough nutlet to crack: Resolving the phylogeny of Thesium (Thesiaceae), the largest genus in Santalales</mods:title>
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<mods:name id="367FFE33E31E8FC1C5B4B2DE251E2DF0" type="personal">
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<mods:namePart id="EE6A638790F99E60B2393D5F8188CCBA">García, Miguel A.</mods:namePart>
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<mods:affiliation id="44427EF08E37EC73D787F8E799D65B39">Ladislav Mucina & Real Jardín Botánico-CSIC, Plaza de Murillo</mods:affiliation>
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<mods:namePart id="29338CBFBA9C7B4467F1BE24E46F6113">Mucina, Ladislav</mods:namePart>
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<mods:affiliation id="A0E0378F8671A0681EE2E7BEAB240B42">& Daniel L. Nickrent & 28014 Madrid, Spain & Harry Butler Institute, Murdoch University, 90 South Street, Murdoch 6150, Perth, Australia & & Daniel L. Nickrent & Department of Geography and Environmental Studies, Stellenbosch University, Stellenbosch, South Africa</mods:affiliation>
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<mods:namePart id="89A716987429AE30775EC37090ED5BFA">Nickrent, Daniel L.</mods:namePart>
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<mods:affiliation id="5AB2592FB5AD4370E3DDBE5614BF45E2">Plant Biology Section, School of Integrative Plant Science, 332 Emerson Hall, College of Agriculture and Life Science,</mods:affiliation>
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<mods:title id="90885AF21A13295EAEE4EDBB9EE265D1">TAXON</mods:title>
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<mods:part id="8806ED976A4447D6A3C7FB2FB1A66BF8">
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<mods:date id="37B8DDAFF5385887021543F193EE6018">2024</mods:date>
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<mods:number id="CA84237A0ABB52CC0061B2F580AEE3DF">2024-02-29</mods:number>
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<mods:number id="FB223B3D548C3691834CCDE1428F6B3F">73</mods:number>
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<mods:url id="0F53E6659C9428B9FD9BFA679712655C">http://dx.doi.org/10.1002/tax.13123</mods:url>
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<treatment id="F275A6752C574028FF70A72EFB1068C2" LSID="urn:lsid:plazi:treatment:F275A6752C574028FF70A72EFB1068C2" httpUri="http://treatment.plazi.org/id/F275A6752C574028FF70A72EFB1068C2" lastPageId="38" lastPageNumber="228" pageId="36" pageNumber="226">
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<subSubSection id="32C644E82C57402AFF70A72EFE5C6822" box="[148,431,1627,1651]" pageId="36" pageNumber="226" type="nomenclature">
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<paragraph id="7A6317632C57402AFF70A72EFE5C6822" blockId="36.[148,431,1627,1651]" box="[148,431,1627,1651]" pageId="36" pageNumber="226">
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<heading id="212BA00F2C57402AFF70A72EFE5C6822" bold="true" box="[148,431,1627,1651]" fontSize="10" level="2" pageId="36" pageNumber="226" reason="2">
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<emphasis id="48A8CB712C57402AFF70A72EFE5C6822" bold="true" box="[148,431,1627,1651]" pageId="36" pageNumber="226">
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Biogeography of
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<taxonomicName id="BDDC6CE02C57402AFEB1A729FE5C6822" ID-CoL="7VVV" authorityName="García & Mucina & Nickrent" authorityYear="2024" box="[341,431,1628,1650]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="36" pageNumber="226" phylum="Tracheophyta" rank="genus">
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<emphasis id="48A8CB712C57402AFEB1A729FE5C6822" bold="true" box="[341,431,1628,1650]" italics="true" pageId="36" pageNumber="226">Thesium</emphasis>
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</taxonomicName>
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</emphasis>
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</heading>
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</paragraph>
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</subSubSection>
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<subSubSection id="32C644E82C574028FF70A7EEFB1068C2" lastPageId="38" lastPageNumber="228" pageId="36" pageNumber="226" type="discussion">
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<paragraph id="7A6317632C57402AFF70A7EEFD986922" blockId="36.[103,751,1691,1970]" pageId="36" pageNumber="226">
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The extensive sampling in this study throughout the entire distributional area of
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<taxonomicName id="BDDC6CE02C57402AFEA8A7CEFE556882" authorityName="García & Mucina & Nickrent" authorityYear="2024" box="[332,422,1723,1746]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="36" pageNumber="226" phylum="Tracheophyta" rank="genus">
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<emphasis id="48A8CB712C57402AFEA8A7CEFE556882" box="[332,422,1723,1746]" italics="true" pageId="36" pageNumber="226">Thesium</emphasis>
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</taxonomicName>
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(
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<figureCitation id="E2E70BE62C57402AFE51A7CEFE076882" box="[437,500,1723,1746]" captionStart="Fig" captionStartId="1.[151,183,1921,1942]" captionTargetBox="[151,1483,1231,1887]" captionTargetId="figure-652@1.[151,1483,1231,1887]" captionTargetPageId="1" captionText="Fig. 1. Map showing the distribution of Thesium worldwide. Data from GBIF were vetted and used for generating the first iteration. Additional information was obtained from specimen records given in various Floras." figureDoi="http://doi.org/10.5281/zenodo.14047586" httpUri="https://zenodo.org/record/14047586/files/figure.png" pageId="36" pageNumber="226">Fig. 1</figureCitation>
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) allows the reconstruction of the historical biogeography of the genus. Further studies including estimation of absolute divergence times will help to understand the dispersal and speciation events within the genus and allow comparison with the large-scale floristic patterns observed in other groups of African origin.
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</paragraph>
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<paragraph id="7A6317632C57402AFF70A60EFAB16C02" blockId="36.[103,751,1691,1970]" lastBlockId="36.[789,1437,187,1970]" pageId="36" pageNumber="226">
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The monotypic genus
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<taxonomicName id="BDDC6CE02C57402AFE6CA60EFDD769C2" box="[392,548,1915,1938]" class="Magnoliopsida" family="Thesiaceae" genus="Lacomucinaea" kingdom="Plantae" order="Santalales" pageId="36" pageNumber="226" phylum="Tracheophyta" rank="genus">
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<emphasis id="48A8CB712C57402AFE6CA60EFDD769C2" box="[392,548,1915,1938]" italics="true" pageId="36" pageNumber="226">Lacomucinaea</emphasis>
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</taxonomicName>
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has a SW African distribution (
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<bibRefCitation id="1E4D6A922C57402AFF14A6EEFE0E69E2" author="Nickrent, D. L. & Garcia, M. A." box="[240,509,1947,1970]" pageId="36" pageNumber="226" pagination="173 - 184" refId="ref44227" refString="Nickrent, D. L. & Garcia, M. A. 2015. Lacomucinaea, a new monotypic genus in Thesiaceae (Santalales). Phytotaxa 224: 173 - 184. https: // doi. org / 10.11646 / phytotaxa. 224.2.4" type="journal article" year="2015">Nickrent & García, 2015</bibRefCitation>
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||
) and
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<taxonomicName id="BDDC6CE02C57402AFDD9A6EEFD2769E2" box="[573,724,1947,1970]" class="Magnoliopsida" family="Thesiaceae" genus="Osyridicarpos" kingdom="Plantae" order="Santalales" pageId="36" pageNumber="226" phylum="Tracheophyta" rank="genus">
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<emphasis id="48A8CB712C57402AFDD9A6EEFD2769E2" box="[573,724,1947,1970]" italics="true" pageId="36" pageNumber="226">Osyridicarpos</emphasis>
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</taxonomicName>
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is more common in SE Africa with populations extending north to
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<collectingCountry id="02CB57F32C57402AFCD9A1AEFC656EA2" box="[829,918,219,242]" name="Ethiopia" pageId="36" pageNumber="226">Ethiopia</collectingCountry>
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along the Eastern Arc African Mountains (EAAM). Both genera are the extant lineages most closely related to
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<taxonomicName id="BDDC6CE02C57402AFC6AA06EFC1B6F62" authorityName="García & Mucina & Nickrent" authorityYear="2024" box="[910,1000,283,306]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="36" pageNumber="226" phylum="Tracheophyta" rank="genus">
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<emphasis id="48A8CB712C57402AFC6AA06EFC1B6F62" box="[910,1000,283,306]" italics="true" pageId="36" pageNumber="226">Thesium</emphasis>
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</taxonomicName>
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.
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<bibRefCitation id="1E4D6A922C57402AFC18A06EFB176F62" author="Moore, T. E. & Verboom, G. A. & Forest, F." box="[1020,1252,283,306]" pageId="36" pageNumber="226" pagination="435 - 452" refId="ref43881" refString="Moore, T. E., Verboom, G. A. & Forest, F. 2010. Phylogenetics and biogeography of the parasitic genus Thesium L. (Santalaceae), with an emphasis on the Cape of South Africa. Bot. J. Linn. Soc. 162: 435 - 452. https: // doi. org / 10.1111 / j. 1095 - 8339.2010. 01032. x" type="journal article" year="2010">Moore & al. (2010)</bibRefCitation>
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, using ancestral range reconstruction, suggested a southern African origin of
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||
<taxonomicName id="BDDC6CE02C57402AFCD8A02EFC656F22" authorityName="García & Mucina & Nickrent" authorityYear="2024" box="[828,918,347,370]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="36" pageNumber="226" phylum="Tracheophyta" rank="genus">
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||
<emphasis id="48A8CB712C57402AFCD8A02EFC656F22" box="[828,918,347,370]" italics="true" pageId="36" pageNumber="226">Thesium</emphasis>
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</taxonomicName>
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in the Late Eocene, with a crown age of 39.1 ± 11.9 million years.
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<taxonomicName id="BDDC6CE02C57402AFBD6A00EFB3D6FC2" box="[1074,1230,379,402]" class="Magnoliopsida" family="Thesiaceae" genus="Lacomucinaea" kingdom="Plantae" order="Santalales" pageId="36" pageNumber="226" phylum="Tracheophyta" rank="genus">
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<emphasis id="48A8CB712C57402AFBD6A00EFB3D6FC2" box="[1074,1230,379,402]" italics="true" pageId="36" pageNumber="226">Lacomucinaea</emphasis>
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</taxonomicName>
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and
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<taxonomicName id="BDDC6CE02C57402AFAE1A00EFA6F6FC2" box="[1285,1436,379,402]" class="Magnoliopsida" family="Thesiaceae" genus="Osyridicarpos" kingdom="Plantae" order="Santalales" pageId="36" pageNumber="226" phylum="Tracheophyta" rank="genus">
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<emphasis id="48A8CB712C57402AFAE1A00EFA6F6FC2" box="[1285,1436,379,402]" italics="true" pageId="36" pageNumber="226">Osyridicarpos</emphasis>
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</taxonomicName>
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were not included in their analyses, but their current distribution supports their common origin in southern Africa. The topology of both ITS and chloroplast trees resolves, with the highest support in all the analyses, two main lineages in the genus, the CMPB lineage (clade 2), mostly including the Eurasian species, and the African lineage (clade 26).
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</paragraph>
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<paragraph id="7A6317632C57402AFCA6A32FFBF66D82" blockId="36.[789,1437,187,1970]" pageId="36" pageNumber="226">
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<emphasis id="48A8CB712C57402AFCA6A32FFB026C22" bold="true" box="[834,1265,602,626]" pageId="36" pageNumber="226">
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Biogeography of the CMPB lineage.
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<emphasis id="48A8CB712C57402AFB32A328FB026C22" bold="true" box="[1238,1265,605,626]" italics="true" pageId="36" pageNumber="226">—</emphasis>
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</emphasis>
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The CMPB lineage has a remarkable disjunct distribution pattern because it includes the
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<collectingRegion id="B818D9812C57402AFC78A3EEFC216CE2" box="[924,978,667,690]" country="South Africa" name="Western Cape" pageId="36" pageNumber="226">Cape</collectingRegion>
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endemic species of
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<taxonomicName id="BDDC6CE02C57402AFB4FA3EEFC946C82" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="36" pageNumber="226" phylum="Tracheophyta" rank="subGenus" subGenus="Hagnothesium">
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<emphasis id="48A8CB712C57402AFB4FA3EEFAF76CE2" box="[1195,1284,667,690]" italics="true" pageId="36" pageNumber="226">Thesium</emphasis>
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subg.
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<emphasis id="48A8CB712C57402AFAA8A3EEFC946C82" italics="true" pageId="36" pageNumber="226">Hagnothesium</emphasis>
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</taxonomicName>
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(former genus
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<taxonomicName id="BDDC6CE02C57402AFBF6A3CEFB736C82" baseAuthorityName="Sonder" baseAuthorityYear="1857" box="[1042,1152,699,722]" class="Magnoliopsida" family="Thesiaceae" genus="Thesidium" kingdom="Plantae" order="Santalales" pageId="36" pageNumber="226" phylum="Tracheophyta" rank="genus">
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<emphasis id="48A8CB712C57402AFBF6A3CEFB736C82" box="[1042,1152,699,722]" italics="true" pageId="36" pageNumber="226">Thesidium</emphasis>
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</taxonomicName>
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) nested within Mediterranean, Canarian, and Eurasian species (
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<emphasis id="48A8CB712C57402AFB4AA3AEFB4E6CA2" box="[1198,1213,731,754]" italics="true" pageId="36" pageNumber="226">T</emphasis>
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||
. subg.
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<taxonomicName id="BDDC6CE02C57402AFAEFA3AEFA976CA2" authorityName="García & Mucina & Nickrent" authorityYear="2024" box="[1291,1380,731,754]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="36" pageNumber="226" phylum="Tracheophyta" rank="genus">
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<emphasis id="48A8CB712C57402AFAEFA3AEFA976CA2" box="[1291,1380,731,754]" italics="true" pageId="36" pageNumber="226">Thesium</emphasis>
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</taxonomicName>
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), not within the main African lineage. Both chloroplast and ITS phylogenies highly support this relationship.
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<bibRefCitation id="1E4D6A922C57402AFAEAA26EFCAF6D02" author="Moore, T. E. & Verboom, G. A. & Forest, F." pageId="36" pageNumber="226" pagination="435 - 452" refId="ref43881" refString="Moore, T. E., Verboom, G. A. & Forest, F. 2010. Phylogenetics and biogeography of the parasitic genus Thesium L. (Santalaceae), with an emphasis on the Cape of South Africa. Bot. J. Linn. Soc. 162: 435 - 452. https: // doi. org / 10.1111 / j. 1095 - 8339.2010. 01032. x" type="journal article" year="2010">Moore & al. (2010)</bibRefCitation>
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dated the divergence of both subgenera to the Oligocene (33.7–23.8 million years ago), preceding other plant groups with origin in southern Africa that dispersed north via East Africa with the Late-Miocene uplift of the EAAM (
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<bibRefCitation id="1E4D6A922C57402AFCF8A2CEFC066D82" author="Pokorny, L. & Riina, R. & Mairal, M. & Meseguer, A. S. & Culshaw, V. & Cendoya, J. & Serrano, M. & Carbajal, R. & Ortiz, S. & Heuertz, M. & Sanmartin, I." box="[796,1013,955,978]" pageId="36" pageNumber="226" refId="ref44646" refString="Pokorny, L., Riina, R., Mairal, M., Meseguer, A. S., Culshaw, V., Cendoya, J., Serrano, M., Carbajal, R., Ortiz, S., Heuertz, M. & Sanmartin, I. 2015. Living on the edge: Timing of Rand Flora disjunctions congruent with ongoing aridification in Africa. Frontiers Genet. 6: 154. https: // doi. org / 10.3389 / fgene. 2015.00154" type="journal volume" year="2015">Pokorny & al., 2015</bibRefCitation>
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||
).
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</paragraph>
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<paragraph id="7A6317632C57402AFCA6A2AEFC726B62" blockId="36.[789,1437,187,1970]" pageId="36" pageNumber="226">
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The early-diverging lineages in the
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<typeStatus id="A567A9C12C57402AFB50A2AEFB116DA2" box="[1204,1250,987,1010]" pageId="36" pageNumber="226">type</typeStatus>
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subgenus include the NW African
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<taxonomicName id="BDDC6CE02C57402AFC28A28EFB316A42" box="[972,1218,1019,1042]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="36" pageNumber="226" phylum="Tracheophyta" rank="species" species="mauritanicum">
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<emphasis id="48A8CB712C57402AFC28A28EFB316A42" box="[972,1218,1019,1042]" italics="true" pageId="36" pageNumber="226">Thesium mauritanicum</emphasis>
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</taxonomicName>
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and
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<emphasis id="48A8CB712C57402AFB1FA28EFAF96A42" box="[1275,1290,1019,1042]" italics="true" pageId="36" pageNumber="226">T</emphasis>
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. sect.
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<taxonomicName id="BDDC6CE02C57402AFAB5A28EFCB86A62" authorityName="J. C. Manning & F. Forest" baseAuthorityName="Stearn" baseAuthorityYear="1972" class="Magnoliopsida" family="Thesiaceae" genus="Kunkeliella" kingdom="Plantae" order="Santalales" pageId="36" pageNumber="226" phylum="Tracheophyta" rank="genus">
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||
<emphasis id="48A8CB712C57402AFAB5A28EFCB86A62" italics="true" pageId="36" pageNumber="226">Kunkeliella</emphasis>
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</taxonomicName>
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, endemic to the
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<collectingRegion id="B818D9812C57402AFC11A56EFB666A62" box="[1013,1173,1051,1074]" country="Spain" name="Canarias" pageId="36" pageNumber="226">Canary Islands</collectingRegion>
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. Even though
|
||
<taxonomicName id="BDDC6CE02C57402AFACCA56EFCA56A02" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="36" pageNumber="226" phylum="Tracheophyta" rank="species" species="mauritanicum">
|
||
<emphasis id="48A8CB712C57402AFACCA56EFCA56A02" italics="true" pageId="36" pageNumber="226">T. mauritanicum</emphasis>
|
||
</taxonomicName>
|
||
might be the closest extant relative of
|
||
<emphasis id="48A8CB712C57402AFAA2A54EFAA66A02" box="[1350,1365,1083,1106]" italics="true" pageId="36" pageNumber="226">T</emphasis>
|
||
. sect.
|
||
<taxonomicName id="BDDC6CE02C57402AFCF1A52EFC7D6A22" authorityName="J. C. Manning & F. Forest" baseAuthorityName="Stearn" baseAuthorityYear="1972" box="[789,910,1115,1138]" class="Magnoliopsida" family="Thesiaceae" genus="Kunkeliella" kingdom="Plantae" order="Santalales" pageId="36" pageNumber="226" phylum="Tracheophyta" rank="genus">
|
||
<emphasis id="48A8CB712C57402AFCF1A52EFC7D6A22" box="[789,910,1115,1138]" italics="true" pageId="36" pageNumber="226">Kunkeliella</emphasis>
|
||
</taxonomicName>
|
||
, their sister relationship is not resolved in our phylogenies. Our data indicate that
|
||
<emphasis id="48A8CB712C57402AFB8DA50EFB8B6AC2" box="[1129,1144,1147,1170]" italics="true" pageId="36" pageNumber="226">T</emphasis>
|
||
. sect.
|
||
<taxonomicName id="BDDC6CE02C57402AFB5AA50EFAC46AC2" authorityName="J. C. Manning & F. Forest" baseAuthorityName="Stearn" baseAuthorityYear="1972" box="[1214,1335,1147,1170]" class="Magnoliopsida" family="Thesiaceae" genus="Kunkeliella" kingdom="Plantae" order="Santalales" pageId="36" pageNumber="226" phylum="Tracheophyta" rank="genus">
|
||
<emphasis id="48A8CB712C57402AFB5AA50EFAC46AC2" box="[1214,1335,1147,1170]" italics="true" pageId="36" pageNumber="226">Kunkeliella</emphasis>
|
||
</taxonomicName>
|
||
is monophyletic and its presence in the
|
||
<collectingRegion id="B818D9812C57402AFB97A5EEFAE76AE2" box="[1139,1300,1179,1202]" country="Spain" name="Canarias" pageId="36" pageNumber="226">Canary Islands</collectingRegion>
|
||
is the result of a single dispersal event followed by local
|
||
<emphasis id="48A8CB712C57402AFB3BA5CEFAEC6A82" box="[1247,1311,1211,1234]" italics="true" pageId="36" pageNumber="226">in situ</emphasis>
|
||
diversification in the islands with one species in Gran Canaria, three in Tenerife and one in La Palma (
|
||
<bibRefCitation id="1E4D6A922C57402AFB76A58EFC826B62" author="Rodriguez-Rodriguez, P. & Fernandez de Castro, A. G. & Perez de Paz, P. L. & Curbelo, L. & Palomares, A. & Mesa, R. & Acevedo, A. & Sosa, P. A." pageId="36" pageNumber="226" pagination="419 - 436" refId="ref45152" refString="Rodriguez-Rodriguez, P., Fernandez de Castro, A. G., Perez de Paz, P. L., Curbelo, L., Palomares, A., Mesa, R., Acevedo, A., & Sosa, P. A. 2022. Evolution and conservation genetics of an insular hemiparasitic plant lineage at the limit of survival: The case of Thesium sect. Kunkeliella in the Canary Islands. Amer. J. Bot. 109: 419 - 436. https: // doi. org / 10.1002 / ajb 2.1830" type="journal article" year="2022">Rodríguez-Rodríguez & al., 2022</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="7A6317632C57402AFCA6A44EFB8069E2" blockId="36.[789,1437,187,1970]" pageId="36" pageNumber="226">
|
||
A second lineage, the Paleoboreal clade (clade 7), is well-supported with both ITS and chloroplast markers and includes the remainder species of
|
||
<taxonomicName id="BDDC6CE02C57402AFBA4A40EFAB86BC2" box="[1088,1355,1403,1426]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="36" pageNumber="226" phylum="Tracheophyta" rank="subGenus" subGenus="Thesium">
|
||
<emphasis id="48A8CB712C57402AFBA4A40EFB696BC2" box="[1088,1178,1403,1426]" italics="true" pageId="36" pageNumber="226">Thesium</emphasis>
|
||
subg.
|
||
<emphasis id="48A8CB712C57402AFB15A40EFAB86BC2" box="[1265,1355,1403,1426]" italics="true" pageId="36" pageNumber="226">Thesium</emphasis>
|
||
</taxonomicName>
|
||
. Well-supported backbone relationships in this lineage are not always seen, but among the well-supported clades the Core Eurasian clade (12), that includes most of the diversity in terms of number of species, emerges as an independent lineage with a mostly W Asian distribution. The ITS phylogeny shows that
|
||
<taxonomicName id="BDDC6CE02C57402AFCF1A74EFC896802" box="[789,890,1595,1618]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="36" pageNumber="226" phylum="Tracheophyta" rank="species" species="humile">
|
||
<emphasis id="48A8CB712C57402AFCF1A74EFC896802" box="[789,890,1595,1618]" italics="true" pageId="36" pageNumber="226">T. humile</emphasis>
|
||
</taxonomicName>
|
||
, a circum-Mediterranean and Canarian species, is moderately supported as sister to the Macranthia clade (9), which includes species distributed in the E Mediterranean region and W Asia. A second lineage, the Procumbens clade (11), is present in W Asia and SE Europe and finally, the C Asian
|
||
<taxonomicName id="BDDC6CE02C57402AFCB9A7AEFBE868A2" box="[861,1051,1755,1778]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="36" pageNumber="226" phylum="Tracheophyta" rank="species" species="minkwitzianum">
|
||
<emphasis id="48A8CB712C57402AFCB9A7AEFBE868A2" box="[861,1051,1755,1778]" italics="true" pageId="36" pageNumber="226">T. minkwitzianum</emphasis>
|
||
</taxonomicName>
|
||
whose closest relatives are not resolved. This scenario suggests that after the migration from the Cape region to the Mediterranean, one or several dispersals to W Asia and subsequent speciation generated the extant diversity of species of
|
||
<taxonomicName id="BDDC6CE02C57402AFBA3A62EFB526922" authorityName="García & Mucina & Nickrent" authorityYear="2024" box="[1095,1185,1883,1906]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="36" pageNumber="226" phylum="Tracheophyta" rank="genus">
|
||
<emphasis id="48A8CB712C57402AFBA3A62EFB526922" box="[1095,1185,1883,1906]" italics="true" pageId="36" pageNumber="226">Thesium</emphasis>
|
||
</taxonomicName>
|
||
with mostly E Mediterranean and W Asian distributions, especially in
|
||
<collectingCountry id="02CB57F32C57402AFAF0A60EFA9C69C2" box="[1300,1391,1915,1938]" name="Turkey" pageId="36" pageNumber="226">Anatolia</collectingCountry>
|
||
and other regions of the Middle East.
|
||
</paragraph>
|
||
<paragraph id="7A6317632C56402BFF20A1CEFDC86C22" blockId="37.[151,799,187,1970]" pageId="37" pageNumber="227">
|
||
The Core Eurasian clade (12) represents the dispersal and subsequent speciation in Europe that produced many of the endemic species of the continent, especially in southern mountain areas. In the ITS phylogeny, the Linophylla clade (17) together with
|
||
<taxonomicName id="BDDC6CE02C56402BFE80A04EFDE36F02" box="[356,528,315,338]" class="Magnoliopsida" family="Thesiaceae" genus="Thesidium" kingdom="Plantae" order="Santalales" pageId="37" pageNumber="227" phylum="Tracheophyta" rank="species" species="catalaunicum">
|
||
<emphasis id="48A8CB712C56402BFE80A04EFDE36F02" box="[356,528,315,338]" italics="true" pageId="37" pageNumber="227">T. catalaunicum</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="BDDC6CE02C56402BFDADA04EFD386F02" box="[585,715,315,338]" class="Magnoliopsida" family="Thesiaceae" genus="Thesidium" kingdom="Plantae" order="Santalales" pageId="37" pageNumber="227" phylum="Tracheophyta" rank="species" species="rostratum">
|
||
<emphasis id="48A8CB712C56402BFDADA04EFD386F02" box="[585,715,315,338]" italics="true" pageId="37" pageNumber="227">T. rostratum</emphasis>
|
||
</taxonomicName>
|
||
are the closest European relatives of a group of species of Asian distribution. However, chloroplast markers resolve the Linophylla clade as being more closely related to other European species. Independent of the markers used, the phylogenies within the Core Eurasian clade are congruent with the concept of one to several dispersals from Europe into Asia and subsequent speciation in Siberia, C Asia,
|
||
<collectingCountry id="02CB57F32C56402BFDEFA36EFDBE6C62" box="[523,589,539,562]" name="China" pageId="37" pageNumber="227">China</collectingCountry>
|
||
, and the Himalayas. This represents the second Asian radiation and subsequently migration into E
|
||
<collectingCountry id="02CB57F32C56402BFEA8A32EFE5F6C22" box="[332,428,603,626]" name="Australia" pageId="37" pageNumber="227">Australia</collectingCountry>
|
||
(
|
||
<taxonomicName id="BDDC6CE02C56402BFE58A32EFDD86C22" box="[444,555,603,626]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="37" pageNumber="227" phylum="Tracheophyta" rank="species" species="australe">
|
||
<emphasis id="48A8CB712C56402BFE58A32EFDD86C22" box="[444,555,603,626]" italics="true" pageId="37" pageNumber="227">T. australe</emphasis>
|
||
</taxonomicName>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="7A6317632C56402BFF20A30EFD416AE2" blockId="37.[151,799,187,1970]" pageId="37" pageNumber="227">
|
||
<emphasis id="48A8CB712C56402BFF20A30EFD856CC2" bold="true" box="[196,630,635,659]" pageId="37" pageNumber="227">
|
||
Biogeography of the African lineage.
|
||
<emphasis id="48A8CB712C56402BFDBFA308FD856CC2" bold="true" box="[603,630,637,658]" italics="true" pageId="37" pageNumber="227">—</emphasis>
|
||
</emphasis>
|
||
The topology of ITS and chloroplast trees resolves three early-diverging lineages (clades 28–30) including species with current distributions in the Succulent Karoo and Nama-Karoo semi-deserts (clade 29) and Fynbos (clades 28 and 30). One species,
|
||
<taxonomicName id="BDDC6CE02C56402BFD62A38EFF326D62" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="37" pageNumber="227" phylum="Tracheophyta" rank="species" species="triflorum">
|
||
<emphasis id="48A8CB712C56402BFD62A38EFF326D62" italics="true" pageId="37" pageNumber="227">Thesium triflorum</emphasis>
|
||
</taxonomicName>
|
||
, is also present in areas of the East African mountains, probably as the result of a later dispersal to the north from
|
||
<collectingCountry id="02CB57F32C56402BFF73A22EFEEC6D22" box="[151,287,859,882]" name="South Africa" pageId="37" pageNumber="227">South Africa</collectingCountry>
|
||
(
|
||
<bibRefCitation id="1E4D6A922C56402BFECAA22EFE2B6D22" author="Brummitt, R. K." box="[302,472,859,882]" pageId="37" pageNumber="227" refId="ref41290" refString="Brummitt, R. K. 1976. Notes arising from the Wye College Expedition to Malawi. 13. Thesium triflorum (Santalaceae) in the Flora Zambesiaca area. Kew Bull. 31: 176. https: // doi. org / 10.2307 / 4109002" type="journal volume" year="1976">Brummitt, 1976</bibRefCitation>
|
||
). Mountains of the
|
||
<collectingRegion id="B818D9812C56402BFD54A22EFD156D22" box="[688,742,859,882]" country="South Africa" name="Western Cape" pageId="37" pageNumber="227">Cape</collectingRegion>
|
||
Fold Belt were important for the further diversification of
|
||
<taxonomicName id="BDDC6CE02C56402BFD22A20EFCED6DC2" authorityName="García & Mucina & Nickrent" authorityYear="2024" box="[710,798,891,914]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="37" pageNumber="227" phylum="Tracheophyta" rank="genus">
|
||
<emphasis id="48A8CB712C56402BFD22A20EFCED6DC2" box="[710,798,891,914]" italics="true" pageId="37" pageNumber="227">Thesium</emphasis>
|
||
</taxonomicName>
|
||
in the
|
||
<collectingRegion id="B818D9812C56402BFF0EA2EEFED36DE2" box="[234,288,923,946]" country="South Africa" name="Western Cape" pageId="37" pageNumber="227">Cape</collectingRegion>
|
||
as revealed by the phylogenetic position of
|
||
<taxonomicName id="BDDC6CE02C56402BFF73A2CEFEA96D82" box="[151,346,955,978]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="37" pageNumber="227" phylum="Tracheophyta" rank="species" species="nautimontanum">
|
||
<emphasis id="48A8CB712C56402BFF73A2CEFEA96D82" box="[151,346,955,978]" italics="true" pageId="37" pageNumber="227">T. nautimontanum</emphasis>
|
||
</taxonomicName>
|
||
, an endemic from the Matroosberg Mt. (
|
||
<bibRefCitation id="1E4D6A922C56402BFF7AA2AEFE996DA2" author="Garcia, M. A. & Nickrent, D. L. & Mucina, L." box="[158,362,987,1010]" pageId="37" pageNumber="227" pagination="41 - 51" refId="ref42161" refString="Garcia, M. A., Nickrent, D. L. & Mucina, L. 2018. Thesium nautimontanum, a new species of Thesiaceae (Santalales) from South Africa. PhytoKeys 109: 41 - 51. https: // doi. org / 10.3897 / phytokeys. 109.28607" type="journal article" year="2018">García & al., 2018</bibRefCitation>
|
||
), as sister to the entire African clade in the ITS phylogeny or in the chloroplast phylogeny as sister to the African species (with the exception of clades 28–30 that were resolved as earlier diverging lineages). The rapid diversification in the fynbos between the Great Escarpment and the coast generated the current diversity of many endemic species, all of which are included
|
||
<emphasis id="48A8CB712C56402BFE44A5EEFE5D6AE2" box="[416,430,1179,1202]" italics="true" pageId="37" pageNumber="227">T</emphasis>
|
||
. subg.
|
||
<taxonomicName id="BDDC6CE02C56402BFE1EA5EEFDCF6AE2" box="[506,572,1179,1202]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="37" pageNumber="227" phylum="Tracheophyta" rank="species" species="frisea">
|
||
<emphasis id="48A8CB712C56402BFE1EA5EEFDCF6AE2" box="[506,572,1179,1202]" italics="true" pageId="37" pageNumber="227">Frisea</emphasis>
|
||
</taxonomicName>
|
||
(clade 32).
|
||
</paragraph>
|
||
<paragraph id="7A6317632C56402BFF20A5CEFED468C2" blockId="37.[151,799,187,1970]" pageId="37" pageNumber="227">
|
||
Both nuclear and chloroplast phylogenies resolved
|
||
<taxonomicName id="BDDC6CE02C56402BFD0AA5CEFE646AA2" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="37" pageNumber="227" phylum="Tracheophyta" rank="subGenus" subGenus="Psilothesium">
|
||
<emphasis id="48A8CB712C56402BFD0AA5CEFF3A6AA2" italics="true" pageId="37" pageNumber="227">Thesium</emphasis>
|
||
subg.
|
||
<emphasis id="48A8CB712C56402BFEEBA5AEFE646AA2" box="[271,407,1243,1266]" italics="true" pageId="37" pageNumber="227">Psilothesium</emphasis>
|
||
</taxonomicName>
|
||
(clade 48) as sister to the Core
|
||
<collectingRegion id="B818D9812C56402BFD0CA5AEFCED6AA2" box="[744,798,1243,1266]" country="South Africa" name="Western Cape" pageId="37" pageNumber="227">Cape</collectingRegion>
|
||
clade, representing the broad diversification of the genus in the grassy biomes of E and N
|
||
<collectingCountry id="02CB57F32C56402BFE3FA46EFD976B62" box="[475,612,1307,1330]" name="South Africa" pageId="37" pageNumber="227">South Africa</collectingCountry>
|
||
and the radiation into tropical Africa. The ITS topology resolves the Gnidiaceum clade (49) as sister to the rest of the subgenus. This clade includes species from the SW ranges of the
|
||
<collectingRegion id="B818D9812C56402BFD9DA40EFD5C6BC2" box="[633,687,1403,1426]" country="South Africa" name="Western Cape" pageId="37" pageNumber="227">Cape</collectingRegion>
|
||
Fold Belt (
|
||
<taxonomicName id="BDDC6CE02C56402BFF44A4EEFEC66BE2" box="[160,309,1435,1458]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="37" pageNumber="227" phylum="Tracheophyta" rank="species" species="oresigenum">
|
||
<emphasis id="48A8CB712C56402BFF44A4EEFEC66BE2" box="[160,309,1435,1458]" italics="true" pageId="37" pageNumber="227">T. oresigenum</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="BDDC6CE02C56402BFE8DA4EEFE306BE2" authority="García & Mucina & Nickrent, 2024" authorityName="García & Mucina & Nickrent" authorityYear="2024" box="[361,451,1435,1458]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="37" pageNumber="227" phylum="Tracheophyta" rank="genus" status="sp. nov.">
|
||
<emphasis id="48A8CB712C56402BFE8DA4EEFE306BE2" box="[361,451,1435,1458]" italics="true" pageId="37" pageNumber="227">Thesium</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="539B760A2C56402BFE2EA4EEFDE86BE2" box="[458,539,1435,1458]" pageId="37" pageNumber="227" rank="species">sp. nov.</taxonomicNameLabel>
|
||
5574) as sister to others from the SE Great Escarpment and E
|
||
<collectingRegion id="B818D9812C56402BFDF8A4CEFDA16B82" box="[540,594,1467,1490]" country="South Africa" name="Western Cape" pageId="37" pageNumber="227">Cape</collectingRegion>
|
||
Fold Belt, supporting the biogeographical link between both areas (
|
||
<bibRefCitation id="1E4D6A922C56402BFD06A4AEFEE36842" author="Clark, V. R. & Barker, N. P. & Mucina, L." pageId="37" pageNumber="227" pagination="2543 - 2561" refId="ref41523" refString="Clark, V. R., Barker, N. P. & Mucina, L. 2011. The Great Escarpment of southern Africa: A new frontier for biodiversity exploration. Biodivers. & Conservation 20: 2543 - 2561. https: // doi. org / 10. 1007 / s 10531 - 011 - 0103 - 3" type="journal article" year="2011">Clark & al., 2011</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="1E4D6A922C56402BFEFBA48EFEA76842" author="Clark, V. R. & Barker, N. P. & Mucina, L." box="[287,340,1531,1554]" pageId="37" pageNumber="227" pagination="313 - 328" refId="ref41586" refString="Clark, V. R., Barker, N. P. & Mucina, L. 2012. Taking the scenic route - The southern Great Escarpment (South Africa) as part of the Cape to Cairo floristic highway. Pl. Ecol. Divers. 4: 313 - 328. https: // doi. org / 10.1080 / 17550874.2011.619584" type="journal article" year="2012">2012</bibRefCitation>
|
||
). Although the topology of the chloroplast phylogeny does not place
|
||
<taxonomicName id="BDDC6CE02C56402BFE05A76EFD856862" box="[481,630,1563,1586]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="37" pageNumber="227" phylum="Tracheophyta" rank="species" species="oresigenum">
|
||
<emphasis id="48A8CB712C56402BFE05A76EFD856862" box="[481,630,1563,1586]" italics="true" pageId="37" pageNumber="227">T. oresigenum</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="BDDC6CE02C56402BFD21A76EFCEC6862" authority="García & Mucina & Nickrent, 2024" authorityName="García & Mucina & Nickrent" authorityYear="2024" box="[709,799,1563,1586]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="37" pageNumber="227" phylum="Tracheophyta" rank="genus" status="sp. nov.">
|
||
<emphasis id="48A8CB712C56402BFD21A76EFCEC6862" box="[709,799,1563,1586]" italics="true" pageId="37" pageNumber="227">Thesium</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="539B760A2C56402BFF73A74EFF1B6802" box="[151,232,1595,1618]" pageId="37" pageNumber="227" rank="species">sp. nov.</taxonomicNameLabel>
|
||
<quantity id="BD24BA862C56402BFF0BA74EFEB26802" box="[239,321,1595,1618]" metricMagnitude="2" metricUnit="m" metricValue="1.4157959999999998" pageId="37" pageNumber="227" unit="in" value="5574.0">5574 in</quantity>
|
||
the Gnidiaceum clade, both species were resolved as successive sisters to the grassland and tropical African radiation.
|
||
</paragraph>
|
||
<paragraph id="7A6317632C56402BFF20A7EEFA3F6F02" blockId="37.[151,799,187,1970]" lastBlockId="37.[837,1485,187,1298]" pageId="37" pageNumber="227">
|
||
Subsequent radiation in the tropics is, in part, represented by clades 52 and 60 that include species lacking an apical beard and mostly distributed in savannas in regions spanning East and West Africa. Our data suggest at least two long-distance dispersals in clade 52, one to South America and at least one to
|
||
<collectingCountry id="02CB57F32C56402BFEF3A64EFE6A6902" box="[279,409,1851,1874]" name="Madagascar" pageId="37" pageNumber="227">Madagascar</collectingCountry>
|
||
. The monophyly of the South American species suggests a single long-distance dispersal from tropical Africa. Although we only have sequence data of
|
||
<taxonomicName id="BDDC6CE02C56402BFD0AA60EFE7A69E2" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="37" pageNumber="227" phylum="Tracheophyta" rank="species" species="madagascariense">
|
||
<emphasis id="48A8CB712C56402BFD0AA60EFE7A69E2" italics="true" pageId="37" pageNumber="227">Thesium madagascariense</emphasis>
|
||
</taxonomicName>
|
||
, the morphological features typical of the Brachyblast clade (54) suggest that another Madagascan endemic species,
|
||
<taxonomicName id="BDDC6CE02C56402BFBE7A1AEFB836EA2" box="[1027,1136,219,242]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="37" pageNumber="227" phylum="Tracheophyta" rank="species" species="perrieri">
|
||
<emphasis id="48A8CB712C56402BFBE7A1AEFB836EA2" box="[1027,1136,219,242]" italics="true" pageId="37" pageNumber="227">T. perrieri</emphasis>
|
||
</taxonomicName>
|
||
, belongs in this group. Because the latter could not be sampled it is unknown whether the presence of these two species in
|
||
<collectingCountry id="02CB57F32C56402BFB88A06EFB186F62" box="[1132,1259,283,306]" name="Madagascar" pageId="37" pageNumber="227">Madagascar</collectingCountry>
|
||
is the result of one or two independent dispersal events from the African continent.
|
||
</paragraph>
|
||
<paragraph id="7A6317632C56402BFC96A02EFAB16B42" blockId="37.[837,1485,187,1298]" pageId="37" pageNumber="227">
|
||
Although not well-supported, the topologies of ITS and chloroplast trees suggest an independent diversification of the leafy species of
|
||
<taxonomicName id="BDDC6CE02C56402BFBFDA0EEFAB06FE2" box="[1049,1347,411,434]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="37" pageNumber="227" phylum="Tracheophyta" rank="subGenus" subGenus="Psilothesium">
|
||
<emphasis id="48A8CB712C56402BFBFDA0EEFB806FE2" box="[1049,1139,411,434]" italics="true" pageId="37" pageNumber="227">Thesium</emphasis>
|
||
subg.
|
||
<emphasis id="48A8CB712C56402BFB5FA0EEFAB06FE2" box="[1211,1347,411,434]" italics="true" pageId="37" pageNumber="227">Psilothesium</emphasis>
|
||
</taxonomicName>
|
||
in the grassdominated biomes of southern Africa with further range expansion to the tropical Africa. The chloroplast phylogenies highly support these species as monophyletic and with lower support its sister relationship with
|
||
<emphasis id="48A8CB712C56402BFB21A36EFB276C62" box="[1221,1236,539,562]" italics="true" pageId="37" pageNumber="227">T</emphasis>
|
||
. sect.
|
||
<taxonomicName id="BDDC6CE02C56402BFAFBA36EFA3F6C62" box="[1311,1484,539,562]" pageId="37" pageNumber="227">
|
||
<emphasis id="48A8CB712C56402BFAFBA36EFA856C62" box="[1311,1398,539,562]" italics="true" pageId="37" pageNumber="227">Barbata</emphasis>
|
||
species
|
||
</taxonomicName>
|
||
of the Gnidiaceum clade and other species mostly distributed around the Drakensberg mountains. This area has been considered a “stepping-stone” between the CFR and the so-called Afrotemperate Region (
|
||
<bibRefCitation id="1E4D6A922C56402BFBA5A3EEFB356CE2" author="White, F." box="[1089,1222,667,690]" pageId="37" pageNumber="227" pagination="463 - 513" refId="ref45958" refString="White, F. 1978. The Afromontane Region. Pp. 463 - 513 in: Werger, M. J. A. & Van Bruggen, A. C. (eds.), Biogeography and ecology of southern Africa. The Hague: Dr W. Junk." type="book chapter" year="1978">White, 1978</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="1E4D6A922C56402BFB31A3EEFA696CE2" author="Galley, C. & Bytebier, B. & Bellstedt, D. U. & Linder, H. P." box="[1237,1434,667,690]" pageId="37" pageNumber="227" pagination="535 - 543" refId="ref42088" refString="Galley, C., Bytebier, B., Bellstedt, D. U. & Linder, H. P. 2007. The Cape element in the Afrotemperate flora: From Cape to Cairo? Proc. Roy. Soc. London, Ser. B., Biol. Sci. 274: 535 - 543. https: // doi. org / 10.1098 / rspb. 2006.0046" type="journal article" year="2007">Galley & al., 2007</bibRefCitation>
|
||
) encompassing high elevations of mountain ranges of southern and East Africa (and to a minor extent Central and West Africa). Migration northwards is supported by the presence in this clade of species endemic to East African mountains and highlands such as
|
||
<taxonomicName id="BDDC6CE02C56402BFBFDA24EFAF76D02" box="[1049,1284,827,850]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="37" pageNumber="227" phylum="Tracheophyta" rank="species" species="kilimandscharicum">
|
||
<emphasis id="48A8CB712C56402BFBFDA24EFAF76D02" box="[1049,1284,827,850]" italics="true" pageId="37" pageNumber="227">T. kilimandscharicum</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="BDDC6CE02C56402BFAF3A24EFA356D02" box="[1303,1478,827,850]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="37" pageNumber="227" phylum="Tracheophyta" rank="species" species="dolichomeras">
|
||
<emphasis id="48A8CB712C56402BFAF3A24EFA356D02" box="[1303,1478,827,850]" italics="true" pageId="37" pageNumber="227">T. dolichomeras</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="BDDC6CE02C56402BFCA1A22EFC2A6D22" box="[837,985,859,882]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="37" pageNumber="227" phylum="Tracheophyta" rank="species" species="whyteanum">
|
||
<emphasis id="48A8CB712C56402BFCA1A22EFC2A6D22" box="[837,985,859,882]" italics="true" pageId="37" pageNumber="227">T. whyteanum</emphasis>
|
||
</taxonomicName>
|
||
, etc. and its expansion to
|
||
<collectingCountry id="02CB57F32C56402BFB18A22EFAA66D22" box="[1276,1365,859,882]" name="Ethiopia" pageId="37" pageNumber="227">Ethiopia</collectingCountry>
|
||
and SE of the Arabia Peninsula (species such as
|
||
<taxonomicName id="BDDC6CE02C56402BFAC5A20EFA576DC2" box="[1313,1444,891,914]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="37" pageNumber="227" phylum="Tracheophyta" rank="species" species="radicans">
|
||
<emphasis id="48A8CB712C56402BFAC5A20EFA576DC2" box="[1313,1444,891,914]" italics="true" pageId="37" pageNumber="227">T. radicans</emphasis>
|
||
</taxonomicName>
|
||
or
|
||
<taxonomicName id="BDDC6CE02C56402BFCA1A2EEFC3C6DE2" box="[837,975,923,946]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="37" pageNumber="227" phylum="Tracheophyta" rank="species" species="hararensis">
|
||
<emphasis id="48A8CB712C56402BFCA1A2EEFC3C6DE2" box="[837,975,923,946]" italics="true" pageId="37" pageNumber="227">T. hararensis</emphasis>
|
||
</taxonomicName>
|
||
). Expansion and diversification took place also in lowland areas of the subtropical and tropical grasslands of SE Africa and the tropical Central and West Africa as well as more recent
|
||
<emphasis id="48A8CB712C56402BFC0BA28EFBC76A42" box="[1007,1076,1019,1042]" italics="true" pageId="37" pageNumber="227">in situ</emphasis>
|
||
speciations suggested by the overlap of morphological characters among species in this taxonomically difficult group. Further events of long-distance dispersal are seen in clade 64, one to
|
||
<collectingCountry id="02CB57F32C56402BFB8FA52EFB196A22" box="[1131,1258,1115,1138]" name="Madagascar" pageId="37" pageNumber="227">Madagascar</collectingCountry>
|
||
(
|
||
<taxonomicName id="BDDC6CE02C56402BFB1EA52EFA646A22" box="[1274,1431,1115,1138]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="37" pageNumber="227" phylum="Tracheophyta" rank="species" species="decaryanum">
|
||
<emphasis id="48A8CB712C56402BFB1EA52EFA646A22" box="[1274,1431,1115,1138]" italics="true" pageId="37" pageNumber="227">T. decaryanum</emphasis>
|
||
</taxonomicName>
|
||
) and another one to E and SE Asia (
|
||
<taxonomicName id="BDDC6CE02C56402BFB69A50EFAEA6AC2" box="[1165,1305,1147,1170]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="37" pageNumber="227" phylum="Tracheophyta" rank="species" species="psilotoides">
|
||
<emphasis id="48A8CB712C56402BFB69A50EFAEA6AC2" box="[1165,1305,1147,1170]" italics="true" pageId="37" pageNumber="227">T. psilotoides</emphasis>
|
||
</taxonomicName>
|
||
).
|
||
<taxonomicName id="BDDC6CE02C56402BFACBA50EFC7C6AE2" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="37" pageNumber="227" phylum="Tracheophyta" rank="species" species="wightianum">
|
||
<emphasis id="48A8CB712C56402BFACBA50EFC7C6AE2" italics="true" pageId="37" pageNumber="227">Thesium wightianum</emphasis>
|
||
</taxonomicName>
|
||
(unsampled) from SE
|
||
<collectingCountry id="02CB57F32C56402BFB7DA5EEFB216AE2" box="[1177,1234,1179,1202]" name="India" pageId="37" pageNumber="227">India</collectingCountry>
|
||
, shares morphological characters with
|
||
<taxonomicName id="BDDC6CE02C56402BFC08A5CEFB916A82" box="[1004,1122,1211,1234]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="37" pageNumber="227" phylum="Tracheophyta" rank="species" species="radicans">
|
||
<emphasis id="48A8CB712C56402BFC08A5CEFB916A82" box="[1004,1122,1211,1234]" italics="true" pageId="37" pageNumber="227">T. radicans</emphasis>
|
||
</taxonomicName>
|
||
and molecular data from this species might reveal another case of dispersal to Asia from NE Africa in this clade (see discussion of clade 64).
|
||
</paragraph>
|
||
<paragraph id="7A6317632C56402BFC96A44EFA696B03" blockId="37.[882,1434,1339,1363]" box="[882,1434,1339,1363]" pageId="37" pageNumber="227">
|
||
<heading id="212BA00F2C56402BFC96A44EFA696B03" bold="true" box="[882,1434,1339,1363]" centered="true" fontSize="10" level="2" pageId="37" pageNumber="227" reason="2">
|
||
<emphasis id="48A8CB712C56402BFC96A44EFA696B03" bold="true" box="[882,1434,1339,1363]" pageId="37" pageNumber="227">Conflict between the ITS and chloroplast datasets</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph id="7A6317632C56402BFC96A40EFA8A69E2" blockId="37.[837,1485,1403,1970]" pageId="37" pageNumber="227">
|
||
Although gene trees derived from the two subcellular compartments were generally congruent, ITS and chloroplast trees showed remarkable incongruences at the specific or even infraspecific levels. These incongruences are not observed equally along the main lineages. The Eurasian, grassland and tropical African lineages show, in general, high congruence between datasets with particular exceptions previously discussed. However, and in agreement with the results obtained by
|
||
<bibRefCitation id="1E4D6A922C56402BFA66A72EFC3768C2" author="Zhigila, D. A. & Verboom, G. A. & Muasya, A. M." pageId="37" pageNumber="227" pagination="100 - 123" refId="ref46295" refString="Zhigila, D. A., Verboom, G. A. & Muasya, A. M. 2020. An infrageneric classification of Thesium (Santalaceae) based on molecular phylogenetic data. Taxon 69: 100 - 123. https: // doi. org / 10.1002 / tax. 12202" type="journal article" year="2020">Zhigila & al. (2020)</bibRefCitation>
|
||
, the
|
||
<collectingRegion id="B818D9812C56402BFC17A70EFBDA68C2" box="[1011,1065,1659,1682]" country="South Africa" name="Western Cape" pageId="37" pageNumber="227">Cape</collectingRegion>
|
||
radiation (
|
||
<taxonomicName id="BDDC6CE02C56402BFB73A70EFA8068C2" box="[1175,1395,1659,1682]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="37" pageNumber="227" phylum="Tracheophyta" rank="subGenus" subGenus="Frisea">
|
||
<emphasis id="48A8CB712C56402BFB73A70EFB1C68C2" box="[1175,1263,1659,1682]" italics="true" pageId="37" pageNumber="227">Thesium</emphasis>
|
||
subg.
|
||
<emphasis id="48A8CB712C56402BFAD5A70EFA8068C2" box="[1329,1395,1659,1682]" italics="true" pageId="37" pageNumber="227">Frisea</emphasis>
|
||
</taxonomicName>
|
||
) is characterized by the high incongruence of chloroplast and nuclear trees. Within the subgenus this incongruence is not detected in the monophyletic
|
||
<emphasis id="48A8CB712C56402BFBC5A7AEFBC368A2" box="[1057,1072,1755,1778]" italics="true" pageId="37" pageNumber="227">
|
||
<collectionCode id="1CCD8FA62C56402BFBC5A7AEFBC368A2" box="[1057,1072,1755,1778]" name="Tavera, Department of Geology and Geophysics" pageId="37" pageNumber="227">T</collectionCode>
|
||
</emphasis>
|
||
. sect.
|
||
<taxonomicName id="BDDC6CE02C56402BFB92A7AEFB4B68A2" box="[1142,1208,1755,1778]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="37" pageNumber="227" phylum="Tracheophyta" rank="species" species="frisea">
|
||
<emphasis id="48A8CB712C56402BFB92A7AEFB4B68A2" box="[1142,1208,1755,1778]" italics="true" pageId="37" pageNumber="227">Frisea</emphasis>
|
||
</taxonomicName>
|
||
(clade 39) because of the lack of internal resolution in this clade but it is interesting that none of the species of this section are resolved among other clades of the
|
||
<collectingRegion id="B818D9812C56402BFC3FA64EFBE26902" box="[987,1041,1851,1874]" country="South Africa" name="Western Cape" pageId="37" pageNumber="227">Cape</collectingRegion>
|
||
radiation on the chloroplast trees except for accession 5460. Biological factors known to cause phylogenetic incongruence in plants include paralogy, hybridization, incomplete lineage sorting, and horizontal gene transfer.
|
||
</paragraph>
|
||
<paragraph id="7A6317632C554028FF70A1CEFE8C6FE2" blockId="38.[103,751,187,1970]" pageId="38" pageNumber="228">
|
||
<emphasis id="48A8CB712C554028FF70A1CEFEEC6E82" bold="true" box="[148,287,187,211]" pageId="38" pageNumber="228">
|
||
Paralogy.
|
||
<emphasis id="48A8CB712C554028FEE0A1C8FEEC6E82" bold="true" box="[260,287,189,210]" italics="true" pageId="38" pageNumber="228">—</emphasis>
|
||
</emphasis>
|
||
Paralogy cannot explain the incongruences detected because there is very low ITS intraspecific variation and in those cases with polymorphic ribotypes the paralogous sequences obtained after cloning were closely related to each other. Moreover, clades resolved for
|
||
<taxonomicName id="BDDC6CE02C554028FE0AA04EFD216F02" box="[494,722,315,338]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="38" pageNumber="228" phylum="Tracheophyta" rank="subGenus" subGenus="Frisea">
|
||
<emphasis id="48A8CB712C554028FE0AA04EFDBB6F02" box="[494,584,315,338]" italics="true" pageId="38" pageNumber="228">Thesium</emphasis>
|
||
subg.
|
||
<emphasis id="48A8CB712C554028FD6BA04EFD216F02" box="[655,722,315,338]" italics="true" pageId="38" pageNumber="228">Frisea</emphasis>
|
||
</taxonomicName>
|
||
in the ITS trees largely correspond with morphological features and species are mostly resolved as monophyletic, unlike in the chloroplast phylogeny.
|
||
</paragraph>
|
||
<paragraph id="7A6317632C554028FF70A0CEFD1C6B02" blockId="38.[103,751,187,1970]" pageId="38" pageNumber="228">
|
||
<emphasis id="48A8CB712C554028FF70A0CEFEA16F82" bold="true" box="[148,338,443,467]" pageId="38" pageNumber="228">
|
||
Hybridization.
|
||
<emphasis id="48A8CB712C554028FED3A0C8FEA16F82" bold="true" box="[311,338,445,466]" italics="true" pageId="38" pageNumber="228">—</emphasis>
|
||
</emphasis>
|
||
Hybridization has been identified as one of the most common sources of incongruence between nuclear and plastid phylogenies. Although hybridization probably occurs in
|
||
<taxonomicName id="BDDC6CE02C554028FF64A36EFF256C62" authorityName="García & Mucina & Nickrent" authorityYear="2024" box="[128,214,539,562]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="38" pageNumber="228" phylum="Tracheophyta" rank="genus">
|
||
<emphasis id="48A8CB712C554028FF64A36EFF256C62" box="[128,214,539,562]" italics="true" pageId="38" pageNumber="228">Thesium</emphasis>
|
||
</taxonomicName>
|
||
, it has yet to be demonstrated and by itself cannot explain the widespread incongruences found in the
|
||
<collectingRegion id="B818D9812C554028FD90A34EFD5A6C02" box="[628,681,571,594]" country="South Africa" name="Western Cape" pageId="38" pageNumber="228">Cape</collectingRegion>
|
||
clade.
|
||
<bibRefCitation id="1E4D6A922C554028FF83A32EFEE66C22" author="Hendrych, R." box="[103,277,603,626]" pageId="38" pageNumber="228" pagination="293 - 358" refId="ref42556" refString="Hendrych, R. 1972. The natural history and systematic of the genus Thesium L. Acta Univ. Carol., Biol. 1970: 293 - 358." type="journal article" year="1972">Hendrych (1972)</bibRefCitation>
|
||
discussed three cases described in the literature as possible interspecific hybrids and his conclusion was that hybrid origin could not be proven. We sequenced cpDNA from two or more individuals in 25 species of
|
||
<emphasis id="48A8CB712C554028FE02A3CEFE076C82" box="[486,500,699,722]" italics="true" pageId="38" pageNumber="228">
|
||
<collectionCode id="1CCD8FA62C554028FE02A3CEFE076C82" box="[486,500,699,722]" name="Tavera, Department of Geology and Geophysics" pageId="38" pageNumber="228">T</collectionCode>
|
||
</emphasis>
|
||
. subg.
|
||
<taxonomicName id="BDDC6CE02C554028FDA5A3CEFD716C82" box="[577,642,699,722]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="38" pageNumber="228" phylum="Tracheophyta" rank="species" species="frisea">
|
||
<emphasis id="48A8CB712C554028FDA5A3CEFD716C82" box="[577,642,699,722]" italics="true" pageId="38" pageNumber="228">Frisea</emphasis>
|
||
</taxonomicName>
|
||
and 14 of them were resolved as polyphyletic on the chloroplast but not on the ITS trees. They were resolved in two or more well-supported clades together with individuals of species very different in vegetative, inflorescence, floral and fruit morphology, thus ruling out the introgression of nuclear genes. As an example,
|
||
<taxonomicName id="BDDC6CE02C554028FF83A20EFF206DC2" box="[103,211,891,914]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="38" pageNumber="228" phylum="Tracheophyta" rank="species" species="strictum">
|
||
<emphasis id="48A8CB712C554028FF83A20EFF206DC2" box="[103,211,891,914]" italics="true" pageId="38" pageNumber="228">T. strictum</emphasis>
|
||
</taxonomicName>
|
||
, a common and widespread species in the fynbos, is distinctive given its wand-plant architecture and other morphological features. It was resolved as monophyletic with strong support on the ITS tree (clade 36). In contrast, on the chloroplast trees these accessions were placed among several well-supported clades with species such as
|
||
<taxonomicName id="BDDC6CE02C554028FE4FA56EFDC76A62" box="[427,564,1051,1074]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="38" pageNumber="228" phylum="Tracheophyta" rank="species" species="pubescens">
|
||
<emphasis id="48A8CB712C554028FE4FA56EFDC76A62" box="[427,564,1051,1074]" italics="true" pageId="38" pageNumber="228">T. pubescens</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="BDDC6CE02C554028FDAEA56EFD146A62" box="[586,743,1051,1074]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="38" pageNumber="228" phylum="Tracheophyta" rank="species" species="pycnanthum">
|
||
<emphasis id="48A8CB712C554028FDAEA56EFD146A62" box="[586,743,1051,1074]" italics="true" pageId="38" pageNumber="228">T. pycnanthum</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="BDDC6CE02C554028FF83A54EFF016A02" baseAuthorityName="Zhigila & Muasya" baseAuthorityYear="2022" box="[103,242,1083,1106]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="38" pageNumber="228" phylum="Tracheophyta" rank="species" species="hispidulum">
|
||
<emphasis id="48A8CB712C554028FF83A54EFF016A02" box="[103,242,1083,1106]" italics="true" pageId="38" pageNumber="228">T. hispidulum</emphasis>
|
||
</taxonomicName>
|
||
, and
|
||
<taxonomicName id="BDDC6CE02C554028FEC8A54EFE476A02" box="[300,436,1083,1106]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="38" pageNumber="228" phylum="Tracheophyta" rank="species" species="prostratum">
|
||
<emphasis id="48A8CB712C554028FEC8A54EFE476A02" box="[300,436,1083,1106]" italics="true" pageId="38" pageNumber="228">T. prostratum</emphasis>
|
||
</taxonomicName>
|
||
, all morphologically quite different. If hybridization and successive backcrosses were involved, without convergent evolution of the nuclear ribosomal cistron, one would expect to find cases of ITS sequence polymorphisms with ribotypes of the same species resolved on different clades on the ITS trees. In the case of convergent evolution, it should have occurred in all the species and always homogenized towards the paternal progenitor because species are monophyletic on the ITS trees and clades are congruent with morphology.
|
||
</paragraph>
|
||
<paragraph id="7A6317632C554028FF70A42FFAA96F02" blockId="38.[103,751,187,1970]" lastBlockId="38.[789,1437,187,1682]" pageId="38" pageNumber="228">
|
||
<emphasis id="48A8CB712C554028FF70A42FFECF6BC2" bold="true" pageId="38" pageNumber="228">
|
||
Stochastic incomplete lineage sorting of ancestral polymorphisms.
|
||
<emphasis id="48A8CB712C554028FEC5A408FECF6BC2" bold="true" box="[289,316,1405,1426]" italics="true" pageId="38" pageNumber="228">—</emphasis>
|
||
</emphasis>
|
||
Incomplete lineage sorting might explain some incongruences between nuclear and chloroplast phylogenies, especially under high rates of diversification. This biological factor was considered by
|
||
<bibRefCitation id="1E4D6A922C554028FE79A4AEFD826BA2" author="Zhigila, D. A. & Verboom, G. A. & Muasya, A. M." box="[413,625,1499,1522]" pageId="38" pageNumber="228" pagination="100 - 123" refId="ref46295" refString="Zhigila, D. A., Verboom, G. A. & Muasya, A. M. 2020. An infrageneric classification of Thesium (Santalaceae) based on molecular phylogenetic data. Taxon 69: 100 - 123. https: // doi. org / 10.1002 / tax. 12202" type="journal article" year="2020">Zhigila & al. (2020)</bibRefCitation>
|
||
as the most probable explanation of the incongruences in
|
||
<taxonomicName id="BDDC6CE02C554028FDD4A48EFF7A6862" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="38" pageNumber="228" phylum="Tracheophyta" rank="subGenus" subGenus="Frisea">
|
||
<emphasis id="48A8CB712C554028FDD4A48EFD746842" box="[560,647,1531,1554]" italics="true" pageId="38" pageNumber="228">Thesium</emphasis>
|
||
subg.
|
||
<emphasis id="48A8CB712C554028FD23A48EFF7A6862" italics="true" pageId="38" pageNumber="228">Frisea</emphasis>
|
||
</taxonomicName>
|
||
. This hypothesis is supported by the fact that incongruences seem random and not geographically biased. However, it is difficult to use incomplete lineage sorting to explain the incongruence found by
|
||
<bibRefCitation id="1E4D6A922C554028FF18A70EFE3468C2" author="Zhigila, D. A. & Verboom, G. A. & Muasya, A. M." box="[252,455,1659,1682]" pageId="38" pageNumber="228" pagination="100 - 123" refId="ref46295" refString="Zhigila, D. A., Verboom, G. A. & Muasya, A. M. 2020. An infrageneric classification of Thesium (Santalaceae) based on molecular phylogenetic data. Taxon 69: 100 - 123. https: // doi. org / 10.1002 / tax. 12202" type="journal article" year="2020">Zhigila & al. (2020)</bibRefCitation>
|
||
of one individual of
|
||
<taxonomicName id="BDDC6CE02C554028FD7DA70EFD1C68C2" box="[665,751,1659,1682]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="38" pageNumber="228" phylum="Tracheophyta" rank="species" species="minus">
|
||
<emphasis id="48A8CB712C554028FD7DA70EFD1C68C2" box="[665,751,1659,1682]" italics="true" pageId="38" pageNumber="228">T. minus</emphasis>
|
||
</taxonomicName>
|
||
(subg.
|
||
<taxonomicName id="BDDC6CE02C554028FF57A7EEFEBA68E2" box="[179,329,1691,1714]" pageId="38" pageNumber="232" rank="section" section="Hagnothesium">
|
||
<emphasis id="48A8CB712C554028FF57A7EEFEBA68E2" box="[179,329,1691,1714]" italics="true" pageId="38" pageNumber="228">Hagnothesium</emphasis>
|
||
</taxonomicName>
|
||
) resolved in
|
||
<emphasis id="48A8CB712C554028FE00A7EEFE0168E2" box="[484,498,1691,1714]" italics="true" pageId="38" pageNumber="228">
|
||
<collectionCode id="1CCD8FA62C554028FE00A7EEFE0168E2" box="[484,498,1691,1714]" name="Tavera, Department of Geology and Geophysics" pageId="38" pageNumber="228">T</collectionCode>
|
||
</emphasis>
|
||
. subg.
|
||
<taxonomicName id="BDDC6CE02C554028FDAEA7EEFD7868E2" box="[586,651,1691,1714]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="38" pageNumber="228" phylum="Tracheophyta" rank="species" species="frisea">
|
||
<emphasis id="48A8CB712C554028FDAEA7EEFD7868E2" box="[586,651,1691,1714]" italics="true" pageId="38" pageNumber="228">Frisea</emphasis>
|
||
</taxonomicName>
|
||
on their concatenated chloroplast tree. It is more probable that the cpDNA in this individual was acquired after the diversification of the four subgenera. To our knowledge no cases have been documented in plants of maintenance of ancestral polymorphism long after speciation or at least with such a wide sequence diversity among individuals of the same species as we found in
|
||
<emphasis id="48A8CB712C554028FF83A60EFF8669C2" box="[103,117,1915,1938]" italics="true" pageId="38" pageNumber="228">
|
||
<collectionCode id="1CCD8FA62C554028FF83A60EFF8669C2" box="[103,117,1915,1938]" name="Tavera, Department of Geology and Geophysics" pageId="38" pageNumber="228">T</collectionCode>
|
||
</emphasis>
|
||
. subg.
|
||
<taxonomicName id="BDDC6CE02C554028FF2DA60EFEF969C2" box="[201,266,1915,1938]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="38" pageNumber="228" phylum="Tracheophyta" rank="species" species="frisea">
|
||
<emphasis id="48A8CB712C554028FF2DA60EFEF969C2" box="[201,266,1915,1938]" italics="true" pageId="38" pageNumber="228">Frisea</emphasis>
|
||
</taxonomicName>
|
||
. In this work we have detected polyphyletic cpDNA haplotypes among individuals of different populations of the same species, but it remains to be studied if it occurs also within the same population. Although incomplete lineage sorting in
|
||
<emphasis id="48A8CB712C554028FCBAA18EFC9F6F42" box="[862,876,251,274]" italics="true" pageId="38" pageNumber="228">
|
||
<collectionCode id="1CCD8FA62C554028FCBAA18EFC9F6F42" box="[862,876,251,274]" name="Tavera, Department of Geology and Geophysics" pageId="38" pageNumber="228">T</collectionCode>
|
||
</emphasis>
|
||
. subg.
|
||
<taxonomicName id="BDDC6CE02C554028FC59A18EFC0D6F42" box="[957,1022,251,274]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="38" pageNumber="228" phylum="Tracheophyta" rank="species" species="frisea">
|
||
<emphasis id="48A8CB712C554028FC59A18EFC0D6F42" box="[957,1022,251,274]" italics="true" pageId="38" pageNumber="228">Frisea</emphasis>
|
||
</taxonomicName>
|
||
or even deep reticulation might be involved, it alone cannot explain the incongruences found in
|
||
<emphasis id="48A8CB712C554028FCF1A04EFCD06F02" box="[789,803,315,338]" italics="true" pageId="38" pageNumber="228">
|
||
<collectionCode id="1CCD8FA62C554028FCF1A04EFCD06F02" box="[789,803,315,338]" name="Tavera, Department of Geology and Geophysics" pageId="38" pageNumber="228">T</collectionCode>
|
||
</emphasis>
|
||
. subg.
|
||
<taxonomicName id="BDDC6CE02C554028FC8AA04EFC5C6F02" box="[878,943,315,338]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="38" pageNumber="228" phylum="Tracheophyta" rank="species" species="frisea">
|
||
<emphasis id="48A8CB712C554028FC8AA04EFC5C6F02" box="[878,943,315,338]" italics="true" pageId="38" pageNumber="228">Frisea</emphasis>
|
||
</taxonomicName>
|
||
by
|
||
<bibRefCitation id="1E4D6A922C554028FC32A04EFB556F02" author="Zhigila, D. A. & Verboom, G. A. & Muasya, A. M." box="[982,1190,315,338]" pageId="38" pageNumber="228" pagination="100 - 123" refId="ref46295" refString="Zhigila, D. A., Verboom, G. A. & Muasya, A. M. 2020. An infrageneric classification of Thesium (Santalaceae) based on molecular phylogenetic data. Taxon 69: 100 - 123. https: // doi. org / 10.1002 / tax. 12202" type="journal article" year="2020">Zhigila & al. (2020)</bibRefCitation>
|
||
and in this work.
|
||
</paragraph>
|
||
<paragraph id="7A6317632C554028FCA6A02EFB1068C2" blockId="38.[789,1437,187,1682]" pageId="38" pageNumber="228">
|
||
<emphasis id="48A8CB712C554028FCA6A02EFB7B6F22" bold="true" box="[834,1160,347,371]" pageId="38" pageNumber="228">
|
||
Horizontal gene transfer.
|
||
<emphasis id="48A8CB712C554028FB89A028FB7B6F22" bold="true" box="[1133,1160,349,370]" italics="true" pageId="38" pageNumber="228">—</emphasis>
|
||
</emphasis>
|
||
Horizontal gene transfer (HGT) is recognized to have contributed to the genome plasticity and adaptive evolution of prokaryotes and unicellular eukaryotes, and played an important role in the origin and evolution of green plants (
|
||
<bibRefCitation id="1E4D6A922C554028FC1AA0AEFB436FA2" author="Chen, R. & Huangfu, L. & Lu, Y. & Fang, H. & Xu, Y. & Li, P. & Zhou, Y. & Xu, C. & Huang, J. & Yang, Z." box="[1022,1200,475,498]" pageId="38" pageNumber="228" refId="ref41441" refString="Chen, R., Huangfu, L., Lu, Y., Fang, H., Xu, Y., Li, P., Zhou, Y., Xu, C., Huang, J. & Yang, Z. 2021. Adaptive innovation of green plants by horizontal gene transfer. Biotechnol. Advances 46: 107671. https: // doi. org / 10.1016 / j. biotechadv. 2020.107671" type="journal volume" year="2021">Chen & al., 2021</bibRefCitation>
|
||
). HGT has been documented to be frequent in several groups of parasitic plants (
|
||
<bibRefCitation id="1E4D6A922C554028FCF8A36EFC2B6C62" author="Davis, C. C. & Xi, Z." box="[796,984,539,562]" pageId="38" pageNumber="228" pagination="14 - 19" refId="ref41749" refString="Davis, C. C. & Xi, Z. 2015. Horizontal gene transfer in parasitic plants. Curr. Opin. Pl. Biol. 26: 14 - 19. https: // doi. org / 10.1016 / j. pbi. 2015.05. 008" type="journal article" year="2015">Davis & Xi, 2015</bibRefCitation>
|
||
). Normally HGT involves particular genes but these transfers can be large, such as the acquisition of entire mitochondrial genomes from three green algae, one moss and fragments of other angiosperms in
|
||
<taxonomicName id="BDDC6CE02C554028FB01A30EFAA76CC2" box="[1253,1364,635,658]" class="Magnoliopsida" family="Amborellaceae" genus="Amborella" kingdom="Plantae" order="Amborellales" pageId="38" pageNumber="228" phylum="Tracheophyta" rank="genus">
|
||
<emphasis id="48A8CB712C554028FB01A30EFAA76CC2" box="[1253,1364,635,658]" italics="true" pageId="38" pageNumber="228">Amborella</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation id="1E4D6A922C554028FA8EA30EFC616CE2" author="Rice, D. W. & Alverson, A. J. & Richardson, A. O. & Young, G. J. & Sanchez-Puerta, M. V. & Munzinger, J. & Barry, K. & Boore, J. L. & Zhang, Y. & DePamphilis, C. W. & Knox, E. B. & Palmer, J. D." pageId="38" pageNumber="228" pagination="1468 - 1473" refId="ref45038" refString="Rice, D. W., Alverson, A. J., Richardson, A. O., Young, G. J., Sanchez-Puerta, M. V., Munzinger, J., Barry, K., Boore, J. L., Zhang, Y., DePamphilis, C. W., Knox, E. B. & Palmer, J. D. 2013. Horizontal transfer of entire genomes via mitochondrial fusion in the angiosperm Amborella. Science 342 (6165): 1468 - 1473. https: // doi. org / 10.1126 / science. 1246275" type="journal article" year="2013">Rice & al., 2013</bibRefCitation>
|
||
). Although HGT of mitochondrial genes is frequent, especially in parasitic plants, it only rarely affects plastid genomes (
|
||
<bibRefCitation id="1E4D6A922C554028FC52A3AEFB566CA2" author="Sanchez-Puerta, M. V." box="[950,1189,731,754]" pageId="38" pageNumber="228" pagination="317 - 323" refId="ref45427" refString="Sanchez-Puerta, M. V. 2014. Involvement of plastid, mitochondrial and nuclear genomes in plant-to-plant horizontal gene transfer. Acta Soc. Bot. Poloniae 83: 317 - 323. https: // doi. org / 10.5586 / asbp. 2014.041" type="book chapter" year="2014">Sánchez-Puerta, 2014</bibRefCitation>
|
||
). We sequenced two regions of the chloroplast genome from different locations. The topologies recovered with each of them are largely congruent (see, e.g., suppl. Figs. S8, S11) and similar results were obtained by
|
||
<bibRefCitation id="1E4D6A922C554028FC7FA22EFB856D22" author="Zhigila, D. A. & Verboom, G. A. & Muasya, A. M." box="[923,1142,859,882]" pageId="38" pageNumber="228" pagination="100 - 123" refId="ref46295" refString="Zhigila, D. A., Verboom, G. A. & Muasya, A. M. 2020. An infrageneric classification of Thesium (Santalaceae) based on molecular phylogenetic data. Taxon 69: 100 - 123. https: // doi. org / 10.1002 / tax. 12202" type="journal article" year="2020">Zhigila & al. (2020)</bibRefCitation>
|
||
comparing their
|
||
<emphasis id="48A8CB712C554028FAD4A22EFA9D6D22" box="[1328,1390,859,882]" italics="true" pageId="38" pageNumber="228">trnLF</emphasis>
|
||
and
|
||
<emphasis id="48A8CB712C554028FCF1A20EFCBC6DC2" box="[789,847,891,914]" italics="true" pageId="38" pageNumber="228">matK</emphasis>
|
||
phylogenies. This suggests that whole plastomes, and not just fragments, have been transmitted horizontally between species. Chloroplast capture in plants is typically associated with reticulation events. However, asexual chloroplast capture in the absence of nuclear introgression has been demonstrated to occur through natural grafts (
|
||
<bibRefCitation id="1E4D6A922C554028FB45A56EFA7E6A62" author="Stegemann, S. & Keuthe, M. & Greiner, S. & Bock, R." box="[1185,1421,1051,1074]" pageId="38" pageNumber="228" pagination="2434 - 2438" refId="ref45534" refString="Stegemann, S., Keuthe, M., Greiner, S. & Bock, R. 2012. Horizontal transfer of chloroplast genomes between plant species. Proc. Natl. Acad. Sci. U. S. A. 109 (7): 2434 - 2438. https: // doi. org / 10.1073 / pnas. 1114076109" type="journal article" year="2012">Stegemann & al., 2012</bibRefCitation>
|
||
). Plastids can dedifferentiate into motile ameboid organelles and move into neighboring cells through connective pores after cell wall disintegration (
|
||
<bibRefCitation id="1E4D6A922C554028FC04A50EFB6F6AC2" author="Hertle, A. P. & Haberl, B. & Bock, R." box="[992,1180,1147,1170]" pageId="38" pageNumber="228" refId="ref42710" refString="Hertle, A. P., Haberl, B. & Bock, R. 2021. Horizontal genome transfer by cell-to-cell travel of whole organelles. Sci. Advances 7: eabd 8215. https: // doi. org / 10.1126 / sciadv. abd 8215" type="journal volume" year="2021">Hertle & al., 2021</bibRefCitation>
|
||
). In parasitic plants, intimate connections between cells of the host and parasite are formed during haustoria development, which mechanistically resemble grafting. In
|
||
<taxonomicName id="BDDC6CE02C554028FBE1A5AEFBAE6AA2" authorityName="García & Mucina & Nickrent" authorityYear="2024" box="[1029,1117,1243,1266]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="38" pageNumber="228" phylum="Tracheophyta" rank="genus">
|
||
<emphasis id="48A8CB712C554028FBE1A5AEFBAE6AA2" box="[1029,1117,1243,1266]" italics="true" pageId="38" pageNumber="228">Thesium</emphasis>
|
||
</taxonomicName>
|
||
, autoparasitism through selfhaustoria and connections between different individuals are known to be common (
|
||
<bibRefCitation id="1E4D6A922C554028FBE6A46EFAC36B62" author="Melnyk, C. W. & Meyerowitz, E. M." box="[1026,1328,1307,1330]" pageId="38" pageNumber="228" pagination="183 - R 188" refId="ref43658" refString="Melnyk, C. W. & Meyerowitz, E. M. 2015. Plant grafting. Curr. Biol. 25: R 183 - R 188. https: // doi. org / 10.1016 / j. cub. 2015.01.029" type="journal article" year="2015">Melnyk & Meyerowitz, 2015</bibRefCitation>
|
||
).
|
||
<taxonomicName id="BDDC6CE02C554028FAA1A46EFC6A6B02" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="38" pageNumber="228" phylum="Tracheophyta" rank="subGenus" subGenus="Frisea">
|
||
<emphasis id="48A8CB712C554028FAA1A46EFA6E6B62" box="[1349,1437,1307,1330]" italics="true" pageId="38" pageNumber="228">Thesium</emphasis>
|
||
subg.
|
||
<emphasis id="48A8CB712C554028FCB3A44EFC6A6B02" box="[855,921,1339,1362]" italics="true" pageId="38" pageNumber="228">Frisea</emphasis>
|
||
</taxonomicName>
|
||
is greatly diversified, and several species commonly grow in the same area in close proximity to each other. This might allow the formation of haustorial connections among individuals of different species, thus in theory HGT of whole plastids could occur between them. Although this is proposed here as a hypothesis, experimentation could yield significant insights to help explain the observations.
|
||
<taxonomicName id="BDDC6CE02C554028FAE6A48EFAA96842" authorityName="García & Mucina & Nickrent" authorityYear="2024" box="[1282,1370,1531,1554]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="38" pageNumber="228" phylum="Tracheophyta" rank="genus">
|
||
<emphasis id="48A8CB712C554028FAE6A48EFAA96842" box="[1282,1370,1531,1554]" italics="true" pageId="38" pageNumber="228">Thesium</emphasis>
|
||
</taxonomicName>
|
||
offers an attractive group in which to investigate whether this phenomenon is actually taking place and whether it explains the unprecedented level of incongruence documented here between nuclear and chloroplast gene trees.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |