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<mods:title id="0830F4C06530050E2C90F7B035311DA9">A tough nutlet to crack: Resolving the phylogeny of Thesium (Thesiaceae), the largest genus in Santalales</mods:title>
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<mods:affiliation id="66B578B6F51EABD749B1E8E650615A70">&amp; Daniel L. Nickrent &amp; 28014 Madrid, Spain &amp; Harry Butler Institute, Murdoch University, 90 South Street, Murdoch 6150, Perth, Australia &amp; &amp; Daniel L. Nickrent &amp; Department of Geography and Environmental Studies, Stellenbosch University, Stellenbosch, South Africa</mods:affiliation>
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<mods:namePart id="B8C93C929A267F61B19DCE89595C69B5">Nickrent, Daniel L.</mods:namePart>
<mods:affiliation id="CBE9E9C60726FB21D7F245D2354599E6">Plant Biology Section, School of Integrative Plant Science, 332 Emerson Hall, College of Agriculture and Life Science,</mods:affiliation>
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5.
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clade:
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<subSubSection id="32C644E82C794002FBA7A649FDDE6A42" lastPageId="12" lastPageNumber="202" pageId="10" pageNumber="200" type="discussion">
<paragraph id="7A6317632C794002FBA7A649FE776C82" blockId="10.[789,1437,187,1970]" lastBlockId="12.[103,751,187,1714]" lastPageId="12" lastPageNumber="202" pageId="10" pageNumber="200">
<taxonomicName id="BDDC6CE02C794004FBA7A649FA6F6902" box="[1091,1436,1850,1874]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="subGenus" subGenus="Hagnothesium">
<emphasis id="48A8CB712C794004FBA7A649FB6E6902" bold="true" box="[1091,1181,1852,1874]" italics="true" pageId="10" pageNumber="200">Thesium</emphasis>
subg.
<emphasis id="48A8CB712C794004FB1BA649FA6F6902" bold="true" box="[1279,1436,1852,1874]" italics="true" pageId="10" pageNumber="200">Hagnothesium</emphasis>
</taxonomicName>
(BIPP 1).
<emphasis id="48A8CB712C794004FC67A628FC6D6922" bold="true" box="[899,926,1885,1906]" italics="true" pageId="10" pageNumber="200">—</emphasis>
To maintain a monophyletic
<taxonomicName id="BDDC6CE02C794004FB09A62EFC2E69C2" authority="(Forest &amp; Manning, 2013)" baseAuthorityName="Forest &amp; Manning" baseAuthorityYear="2013" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="genus">
<emphasis id="48A8CB712C794004FB09A62EFAB46922" box="[1261,1351,1883,1906]" italics="true" pageId="10" pageNumber="200">Thesium</emphasis>
(
<bibRefCitation id="1E4D6A922C794004FABDA62EFC2069C2" author="Forest, F. &amp; Manning, J. C." pageId="10" pageNumber="200" pagination="214 - 216" refId="ref42043" refString="Forest, F. &amp; Manning, J. C. 2013. The minor genera Kunkeliella and Thesidium included in Thesium. Bothalia 43: 214 - 216. https: // doi. org / 10.4102 / abc. v 43 i 2.98" type="journal article" year="2013">Forest &amp; Manning, 2013</bibRefCitation>
)
</taxonomicName>
, species previously classified in the genus
<taxonomicName id="BDDC6CE02C794004FCF1A6EEFBD869E2" authority="(Sonder, 1857)" baseAuthorityName="Sonder" baseAuthorityYear="1857" box="[789,1067,1947,1970]" class="Magnoliopsida" family="Thesiaceae" genus="Thesidium" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="genus">
<emphasis id="48A8CB712C794004FCF1A6EEFC7069E2" box="[789,899,1947,1970]" italics="true" pageId="10" pageNumber="200">Thesidium</emphasis>
(
<bibRefCitation id="1E4D6A922C794004FC77A6EEFBD169E2" author="Sonder, W." box="[915,1058,1947,1970]" pageId="10" pageNumber="200" pagination="353 - 365" refId="ref45473" refString="Sonder, W. 1857. Enumeratio Santalacearum in Africa australi extratropica crescentium quas Dr. Ecklon et C. Zeyher collegerunt. Flora 40: 353 - 365." type="journal article" year="1857">Sonder, 1857</bibRefCitation>
)
</taxonomicName>
were reclassified as
<taxonomicName id="BDDC6CE02C794002FAEFA6EEFEF26E82" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" lastPageId="12" lastPageNumber="202" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="section" section="Hagnothesium">
<emphasis id="48A8CB712C794004FAEFA6EEFA9669E2" box="[1291,1381,1947,1970]" italics="true" pageId="10" pageNumber="200">Thesium</emphasis>
sect.
<emphasis id="48A8CB712C7F4002FF83A1CEFEF26E82" box="[103,257,187,210]" italics="true" pageId="12" pageNumber="202">Hagnothesium</emphasis>
</taxonomicName>
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.DC.
<bibRefCitation id="1E4D6A922C7F4002FEB2A1CEFDC36E82" author="Zhigila, D. A. &amp; Verboom, G. A. &amp; Muasya, A. M." box="[342,560,187,210]" pageId="12" pageNumber="202" pagination="100 - 123" refId="ref46295" refString="Zhigila, D. A., Verboom, G. A. &amp; 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 &amp; al. (2020)</bibRefCitation>
. They later raised this section to the rank of subgenus (
<emphasis id="48A8CB712C7F4002FE12A1AEFDF66EA2" box="[502,517,219,242]" italics="true" pageId="12" pageNumber="202">
<collectionCode id="1CCD8FA62C7F4002FE12A1AEFDF66EA2" box="[502,517,219,242]" name="Tavera, Department of Geology and Geophysics" pageId="12" pageNumber="202">T</collectionCode>
</emphasis>
. subg.
<taxonomicName id="BDDC6CE02C7F4002FDB0A1AEFD1D6EA2" box="[596,750,219,242]" pageId="12" pageNumber="232" rank="section" section="Hagnothesium">
<emphasis id="48A8CB712C7F4002FDB0A1AEFD1D6EA2" box="[596,750,219,242]" italics="true" pageId="12" pageNumber="202">Hagnothesium</emphasis>
</taxonomicName>
(
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.DC.) Zhigila &amp; al.). These species are CFR endemics and include annual to shrubby plants with mostly tetramerous flowers and a dioecious sexual condition. Some species, including
<taxonomicName id="BDDC6CE02C7F4002FF22A02EFE856F22" box="[198,374,347,370]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="12" pageNumber="202" phylum="Tracheophyta" rank="species" species="microcarpum">
<emphasis id="48A8CB712C7F4002FF22A02EFE856F22" box="[198,374,347,370]" italics="true" pageId="12" pageNumber="202">T. microcarpum</emphasis>
</taxonomicName>
<collectionCode id="1CCD8FA62C7F4002FE61A02EFE696F22" box="[389,410,347,370]" country="USA" lsid="urn:lsid:biocol.org:col:15406" name="Harvard University - Arnold Arboretum" pageId="12" pageNumber="202" type="Herbarium">A</collectionCode>
.DC,
<taxonomicName id="BDDC6CE02C7F4002FE3DA02EFDB26F22" box="[473,577,347,370]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="12" pageNumber="202" phylum="Tracheophyta" rank="species" species="fragile">
<emphasis id="48A8CB712C7F4002FE3DA02EFDB26F22" box="[473,577,347,370]" italics="true" pageId="12" pageNumber="202">T. fragile</emphasis>
</taxonomicName>
(
<figureCitation id="E2E70BE62C7F4002FDB3A02EFD436F22" box="[599,688,347,370]" captionStart="Fig" captionStartId="11.[684,716,1893,1914]" captionTargetBox="[152,1484,187,1858]" captionTargetId="figure-0@11.[152,1484,187,1859]" captionTargetPageId="11" captionText="Fig. 3. Caption on next page." figureDoi="http://doi.org/10.5281/zenodo.14047590" httpUri="https://zenodo.org/record/14047590/files/figure.png" pageId="12" pageNumber="202">Fig. 3C</figureCitation>
) and
<taxonomicName id="BDDC6CE02C7F4002FF83A00EFE956FC2" authority="Zhigila" authorityName="Zhigila" box="[103,358,379,402]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="12" pageNumber="202" phylum="Tracheophyta" rank="species" species="quartzicolum">
<emphasis id="48A8CB712C7F4002FF83A00EFEE36FC2" box="[103,272,379,402]" italics="true" pageId="12" pageNumber="202">T. quartzicolum</emphasis>
Zhigila
</taxonomicName>
&amp; al. have morphologically similar male and female plants (monomorphic) whereas
<taxonomicName id="BDDC6CE02C7F4002FD94A0EEFF616F82" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="12" pageNumber="202" phylum="Tracheophyta" rank="species" species="fruticulosum">
<emphasis id="48A8CB712C7F4002FD94A0EEFF616F82" italics="true" pageId="12" pageNumber="202">T. fruticulosum</emphasis>
</taxonomicName>
(
<collectionCode id="1CCD8FA62C7F4002FF7BA0CEFF476F82" box="[159,180,443,466]" country="USA" lsid="urn:lsid:biocol.org:col:15406" name="Harvard University - Arnold Arboretum" pageId="12" pageNumber="202" type="Herbarium">A</collectionCode>
.
<collectionCode id="1CCD8FA62C7F4002FF53A0CEFF206F82" box="[183,211,443,466]" country="Austria" lsid="urn:lsid:biocol.org:col:15588" name="Naturhistorisches Museum Wien" pageId="12" pageNumber="202" type="Herbarium">W</collectionCode>
.Hill) J.
<collectionCode id="1CCD8FA62C7F4002FEFAA0CEFEC36F82" box="[286,304,443,466]" country="Denmark" name="University of Copenhagen" pageId="12" pageNumber="202" type="Herbarium">C</collectionCode>
.Manning &amp; F.Forest,
<taxonomicName id="BDDC6CE02C7F4002FDF9A0CEFD866F82" box="[541,629,443,466]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="12" pageNumber="202" phylum="Tracheophyta" rank="species" species="minus">
<emphasis id="48A8CB712C7F4002FDF9A0CEFD866F82" box="[541,629,443,466]" italics="true" pageId="12" pageNumber="202">T. minus</emphasis>
</taxonomicName>
(
<collectionCode id="1CCD8FA62C7F4002FD67A0CEFD6B6F82" box="[643,664,443,466]" country="USA" lsid="urn:lsid:biocol.org:col:15406" name="Harvard University - Arnold Arboretum" pageId="12" pageNumber="202" type="Herbarium">A</collectionCode>
.
<collectionCode id="1CCD8FA62C7F4002FD7FA0CEFD446F82" box="[667,695,443,466]" country="Austria" lsid="urn:lsid:biocol.org:col:15588" name="Naturhistorisches Museum Wien" pageId="12" pageNumber="202" type="Herbarium">W</collectionCode>
.Hill) J.
<collectionCode id="1CCD8FA62C7F4002FF91A0AEFF746FA2" box="[117,135,475,498]" country="Denmark" name="University of Copenhagen" pageId="12" pageNumber="202" type="Herbarium">C</collectionCode>
.Manning &amp; F.Forest,
<taxonomicName id="BDDC6CE02C7F4002FE99A0AEFDE66FA2" box="[381,533,475,498]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="12" pageNumber="202" phylum="Tracheophyta" rank="species" species="strigulosum">
<emphasis id="48A8CB712C7F4002FE99A0AEFDE66FA2" box="[381,533,475,498]" italics="true" pageId="12" pageNumber="202">T. strigulosum</emphasis>
</taxonomicName>
<collectionCode id="1CCD8FA62C7F4002FDFAA0AEFDC06FA2" box="[542,563,475,498]" country="USA" lsid="urn:lsid:biocol.org:col:15406" name="Harvard University - Arnold Arboretum" pageId="12" pageNumber="202" type="Herbarium">A</collectionCode>
.DC. (=
<taxonomicName id="BDDC6CE02C7F4002FD68A0AEFEFF6C42" authority="(Sond.) Zhigila" authorityName="Zhigila" baseAuthorityName="Sond." class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="12" pageNumber="202" phylum="Tracheophyta" rank="species" species="hirtum">
<emphasis id="48A8CB712C7F4002FD68A0AEFD1C6FA2" box="[652,751,475,498]" italics="true" pageId="12" pageNumber="202">T. hirtum</emphasis>
(Sond.) Zhigila
</taxonomicName>
&amp; al., nom. illeg.),
<taxonomicName id="BDDC6CE02C7F4002FE09A08EFD236C42" authority="Zhigila" authorityName="Zhigila" box="[493,720,507,530]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="12" pageNumber="202" phylum="Tracheophyta" rank="species" species="longicaule">
<emphasis id="48A8CB712C7F4002FE09A08EFD896C42" box="[493,634,507,530]" italics="true" pageId="12" pageNumber="202">T. longicaule</emphasis>
Zhigila
</taxonomicName>
&amp; al. and
<taxonomicName id="BDDC6CE02C7F4002FF5EA36EFE836C62" box="[186,368,539,562]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="12" pageNumber="202" phylum="Tracheophyta" rank="species" species="leptostachyum">
<emphasis id="48A8CB712C7F4002FF5EA36EFE836C62" box="[186,368,539,562]" italics="true" pageId="12" pageNumber="202">T. leptostachyum</emphasis>
</taxonomicName>
<collectionCode id="1CCD8FA62C7F4002FE9DA36EFE7D6C62" box="[377,398,539,562]" country="USA" lsid="urn:lsid:biocol.org:col:15406" name="Harvard University - Arnold Arboretum" pageId="12" pageNumber="202" type="Herbarium">A</collectionCode>
.DC. are dimorphic. In the latter case, male plants usually have small herbaceous bracts and female plants have conspicuous leafy bracts. Although we only sampled five species, the ITS phylogeny (
<figureCitation id="E2E70BE62C7F4002FDD6A30EFD746CC2" box="[562,647,635,658]" captionStart="Fig" captionStartId="5.[151,183,1831,1852]" captionTargetBox="[300,1336,189,1795]" captionTargetId="graphics-0@5.[300,1001,198,1789]" captionTargetPageId="5" captionText="Fig. 2. Phylogenetic analysis of Thesium using nuclear ribosomal internal transcribed spacer (ITS) sequences. This tree, rooted with Osyridicarpos and Lacomucinaea (outgroups) gives Bayesian posterior probabilities at the nodes. Clades discussed in the text are numbered. Geographical sources for the sampled taxa are abbreviated as follows: AFN = Northern Africa, AFS = Southern Africa, AFT = Tropical Africa, ASC = Central Asia, ASE = Eastern Asia, ASN = Northern Asia, ASS = Southern Asia, ASW = Western Asia, AUS = Australia, CIS = Canary Islands, EUR = Europe, MAD = Madagascar, SAM = South America, and USA = United States of America." figureDoi="http://doi.org/10.5281/zenodo.14047588" httpUri="https://zenodo.org/record/14047588/files/figure.png" pageId="12" pageNumber="202">Fig. 2A</figureCitation>
) resolves two highly supported clades separating sexually dimorphic and monomorphic species.
</paragraph>
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<emphasis id="48A8CB712C784005FD48A610FD14692A" bold="true" box="[684,743,1893,1914]" pageId="11" pageNumber="201">Fig. 3.</emphasis>
Caption on next page.
</paragraph>
</caption>
<paragraph id="7A6317632C7F4002FF70A3AEFD986D82" blockId="12.[103,751,187,1714]" pageId="12" pageNumber="202">
The nuclear and chloroplast phylogenies suggest that
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<emphasis id="48A8CB712C7F4002FD5BA3AEFF6B6D42" italics="true" pageId="12" pageNumber="202">Thesium</emphasis>
subg.
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</taxonomicName>
is an early-divergent lineage that, in spite of being endemic to the CFR, is more closely related to Eurasian, North African, and
<collectingRegion id="B818D9812C7F4002FE7CA24EFDC06D02" box="[408,563,827,850]" country="Spain" name="Canarias" pageId="12" pageNumber="202">Canary Islands</collectingRegion>
lineages (
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</emphasis>
. subg.
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<emphasis id="48A8CB712C7F4002FF83A22EFF4C6D22" box="[103,191,859,882]" italics="true" pageId="12" pageNumber="202">Thesium</emphasis>
</taxonomicName>
). According to our chloroplast phylogeny it is sister to
<emphasis id="48A8CB712C7F4002FF83A20EFF866DC2" box="[103,117,891,914]" italics="true" pageId="12" pageNumber="202">
<collectionCode id="1CCD8FA62C7F4002FF83A20EFF866DC2" box="[103,117,891,914]" name="Tavera, Department of Geology and Geophysics" pageId="12" pageNumber="202">T</collectionCode>
</emphasis>
. subg.
<taxonomicName id="BDDC6CE02C7F4002FF26A20EFEE96DC2" baseAuthorityName="Forest &amp; Manning" baseAuthorityYear="2013" box="[194,282,891,914]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="12" pageNumber="202" phylum="Tracheophyta" rank="genus">
<emphasis id="48A8CB712C7F4002FF26A20EFEE96DC2" box="[194,282,891,914]" italics="true" pageId="12" pageNumber="202">Thesium</emphasis>
</taxonomicName>
, however this relationship is not resolved by ITS. Instead,
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<collectionCode id="1CCD8FA62C7F4002FF10A2EEFEF06DE2" box="[244,259,923,946]" name="Tavera, Department of Geology and Geophysics" pageId="12" pageNumber="202">T</collectionCode>
</emphasis>
. subg.
<taxonomicName id="BDDC6CE02C7F4002FEABA2EEFE146DE2" box="[335,487,923,946]" pageId="12" pageNumber="232" rank="section" section="Hagnothesium">
<emphasis id="48A8CB712C7F4002FEABA2EEFE146DE2" box="[335,487,923,946]" italics="true" pageId="12" pageNumber="202">Hagnothesium</emphasis>
</taxonomicName>
is nested within
<emphasis id="48A8CB712C7F4002FD7EA2EEFD5A6DE2" box="[666,681,923,946]" italics="true" pageId="12" pageNumber="202">
<collectionCode id="1CCD8FA62C7F4002FD7EA2EEFD5A6DE2" box="[666,681,923,946]" name="Tavera, Department of Geology and Geophysics" pageId="12" pageNumber="202">T</collectionCode>
</emphasis>
. subg.
<taxonomicName id="BDDC6CE02C7F4002FF83A2CEFF4C6D82" baseAuthorityName="Forest &amp; Manning" baseAuthorityYear="2013" box="[103,191,955,978]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="12" pageNumber="202" phylum="Tracheophyta" rank="genus">
<emphasis id="48A8CB712C7F4002FF83A2CEFF4C6D82" box="[103,191,955,978]" italics="true" pageId="12" pageNumber="202">Thesium</emphasis>
</taxonomicName>
(clade 2), making the latter paraphyletic.
</paragraph>
<paragraph id="7A6317632C7F4002FF70A2AEFDDE6A42" blockId="12.[103,751,187,1714]" pageId="12" pageNumber="202">
Unsampled species include
<taxonomicName id="BDDC6CE02C7F4002FE2CA2AEFD8C6DA2" box="[456,639,987,1010]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="12" pageNumber="202" phylum="Tracheophyta" rank="species" species="leptostachyum">
<emphasis id="48A8CB712C7F4002FE2CA2AEFD8C6DA2" box="[456,639,987,1010]" italics="true" pageId="12" pageNumber="202">T. leptostachyum</emphasis>
</taxonomicName>
,
<taxonomicName id="BDDC6CE02C7F4002FD75A2AEFF536A42" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="12" pageNumber="202" phylum="Tracheophyta" rank="species" species="longicaule">
<emphasis id="48A8CB712C7F4002FD75A2AEFF536A42" italics="true" pageId="12" pageNumber="202">T. longicaule</emphasis>
</taxonomicName>
,
<taxonomicName id="BDDC6CE02C7F4002FF4AA28EFEA76A42" box="[174,340,1019,1042]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="12" pageNumber="202" phylum="Tracheophyta" rank="species" species="quartzicolum">
<emphasis id="48A8CB712C7F4002FF4AA28EFEA76A42" box="[174,340,1019,1042]" italics="true" pageId="12" pageNumber="202">T. quartzicolum</emphasis>
</taxonomicName>
, and
<taxonomicName id="BDDC6CE02C7F4002FE6BA28EFDD56A42" box="[399,550,1019,1042]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="12" pageNumber="202" phylum="Tracheophyta" rank="species" species="strigulosum">
<emphasis id="48A8CB712C7F4002FE6BA28EFDD56A42" box="[399,550,1019,1042]" italics="true" pageId="12" pageNumber="202">T. strigulosum</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
</emphasis>
</treatment>
</document>

186
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4.
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clade (BIPP 1).
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<paragraph id="7A6317632C794004FB77A388FB0668C2" blockId="10.[789,1437,187,1970]" pageId="10" pageNumber="200">
<emphasis id="48A8CB712C794004FB77A388FB5D6D42" bold="true" box="[1171,1198,765,786]" italics="true" pageId="10" pageNumber="200">—</emphasis>
This clade (
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), composed of endemics to the
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archipelago, was formerly placed in the genus
<taxonomicName id="BDDC6CE02C794004FB66A24EFA6E6D02" authority="(Stearn, 1972)" authorityName="J. C. Manning &amp; F. Forest" baseAuthorityName="Stearn" baseAuthorityYear="1972" box="[1154,1437,827,850]" class="Magnoliopsida" family="Thesiaceae" genus="Kunkeliella" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="genus">
<emphasis id="48A8CB712C794004FB66A24EFB086D02" box="[1154,1275,827,850]" italics="true" pageId="10" pageNumber="200">Kunkeliella</emphasis>
(
<bibRefCitation id="1E4D6A922C794004FAEFA24EFA606D02" author="Stearn, W. T." box="[1291,1427,827,850]" pageId="10" pageNumber="200" pagination="11 - 26" refId="ref45503" refString="Stearn, W. T. 1972. Kunkeliella, a new genus of Santalaceae in the Canary Islands. Cuad. Bot. Canaria 16: 11 - 26." type="journal article" year="1972">Stearn, 1972</bibRefCitation>
)
</taxonomicName>
and members were reclassified as
<taxonomicName id="BDDC6CE02C794004FB6DA22EFA6F6D22" box="[1161,1436,859,882]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="section" section="Kunkeliella">
<emphasis id="48A8CB712C794004FB6DA22EFB106D22" box="[1161,1251,859,882]" italics="true" pageId="10" pageNumber="200">Thesium</emphasis>
sect.
<emphasis id="48A8CB712C794004FAC7A22EFA6F6D22" box="[1315,1436,859,882]" italics="true" pageId="10" pageNumber="200">Kunkeliella</emphasis>
</taxonomicName>
by
<bibRefCitation id="1E4D6A922C794004FCD0A20EFBB06DC2" author="Forest, F. &amp; Manning, J. C." box="[820,1091,891,914]" pageId="10" pageNumber="200" pagination="214 - 216" refId="ref42043" refString="Forest, F. &amp; Manning, J. C. 2013. The minor genera Kunkeliella and Thesidium included in Thesium. Bothalia 43: 214 - 216. https: // doi. org / 10.4102 / abc. v 43 i 2.98" type="journal article" year="2013">Forest &amp; Manning (2013)</bibRefCitation>
. Two species were sampled here:
<taxonomicName id="BDDC6CE02C794004FCF1A2EEFA6F6DE2" authority="(Kammer) J. C. Manning &amp; F. Forest" authorityName="J. C. Manning &amp; F. Forest" baseAuthorityName="Kammer" box="[789,1436,923,946]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="species" species="subsucculentum">
<emphasis id="48A8CB712C794004FCF1A2EEFC156DE2" box="[789,998,923,946]" italics="true" pageId="10" pageNumber="200">T. subsucculentum</emphasis>
(Kämmer) J.C.Manning &amp; F.Forest
</taxonomicName>
(
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) from the north coast of Tenerife (
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,
<collectingCountry id="02CB57F32C794004FCF1A2AEFCA06DA2" box="[789,851,987,1010]" name="Spain" pageId="10" pageNumber="200">Spain</collectingCountry>
) and
<taxonomicName id="BDDC6CE02C794004FC69A2AEFBEF6DA2" box="[909,1052,987,1010]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="species" species="retamoides">
<emphasis id="48A8CB712C794004FC69A2AEFBEF6DA2" box="[909,1052,987,1010]" italics="true" pageId="10" pageNumber="200">T. retamoides</emphasis>
</taxonomicName>
(
<collectionCode id="1CCD8FA62C794004FBCEA2AEFBCD6DA2" box="[1066,1086,987,1010]" country="USA" lsid="urn:lsid:biocol.org:col:15406" name="Harvard University - Arnold Arboretum" pageId="10" pageNumber="200" type="Herbarium">A</collectionCode>
.Santos) J.
<collectionCode id="1CCD8FA62C794004FB4CA2AEFB496DA2" box="[1192,1210,987,1010]" country="Denmark" name="University of Copenhagen" pageId="10" pageNumber="200" type="Herbarium">C</collectionCode>
.Manning &amp; F.Forest from the southern part of the same island. The three ITS clones of
<taxonomicName id="BDDC6CE02C794004FC62A56EFBBD6A62" box="[902,1102,1051,1074]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="species" species="subsucculentum">
<emphasis id="48A8CB712C794004FC62A56EFBBD6A62" box="[902,1102,1051,1074]" italics="true" pageId="10" pageNumber="200">T. subsucculentum</emphasis>
</taxonomicName>
(
<collectionCode id="1CCD8FA62C794004FB86A56EFB8B6A62" box="[1122,1144,1051,1074]" country="USA" lsid="urn:lsid:biocol.org:col:15406" name="Harvard University - Arnold Arboretum" pageId="10" pageNumber="200" type="Herbarium">A</collectionCode>
, B,
<collectionCode id="1CCD8FA62C794004FB48A56EFB4C6A62" box="[1196,1215,1051,1074]" country="Denmark" name="University of Copenhagen" pageId="10" pageNumber="200" type="Herbarium">C</collectionCode>
) and two clones of
<taxonomicName id="BDDC6CE02C794004FCF1A54EFC506A02" box="[789,931,1083,1106]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="species" species="retamoides">
<emphasis id="48A8CB712C794004FCF1A54EFC506A02" box="[789,931,1083,1106]" italics="true" pageId="10" pageNumber="200">T. retamoides</emphasis>
</taxonomicName>
(
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, B) were not monophyletic based on species. However,
<bibRefCitation id="1E4D6A922C794004FC60A52EFB0F6A22" author="Rodriguez-Rodriguez, P. &amp; Fernandez de Castro, A. G. &amp; Perez de Paz, P. L. &amp; Curbelo, L. &amp; Palomares, A. &amp; Mesa, R. &amp; Acevedo, A. &amp; Sosa, P. A." box="[900,1276,1115,1138]" pageId="10" pageNumber="200" 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., &amp; 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 &amp; al. (2022)</bibRefCitation>
found enough genetic variation based on microsatellite markers to consider them different species, a conclusion also supported by morphology. On Tenerife, all the species have stems covered with an indument of tiny hairs, whereas
<taxonomicName id="BDDC6CE02C794004FB61A5AEFC026B42" authority="(Stearn) J. C. Manning &amp; F. Forest" authorityName="J. C. Manning &amp; F. Forest" baseAuthorityName="Stearn" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="species" species="canariense">
<emphasis id="48A8CB712C794004FB61A5AEFAE16AA2" box="[1157,1298,1243,1266]" italics="true" pageId="10" pageNumber="200">T. canariense</emphasis>
(Stearn) J.C. Manning &amp; F.Forest
</taxonomicName>
, endemic to Gran Canaria Island, is glabrous. Otherwise, the flowers, fruits and general habit of these plants are similar except for the smaller flowers of
<taxonomicName id="BDDC6CE02C794004FCF1A42EFC516B22" box="[789,930,1371,1394]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="species" species="canariense">
<emphasis id="48A8CB712C794004FCF1A42EFC516B22" box="[789,930,1371,1394]" italics="true" pageId="10" pageNumber="200">T. canariense</emphasis>
</taxonomicName>
. That species is the only one in the genus known to be gynodioecious, i.e., with bisexual and female flowers on separate plants (
<bibRefCitation id="1E4D6A922C794004FC5AA4EEFB326BE2" author="Perez de Paz, J. &amp; Ortega, C. &amp; Febles, R. &amp; Garcia-Medina, J. &amp; Nogales, I." box="[958,1217,1435,1458]" pageId="10" pageNumber="200" refId="ref44567" refString="Perez de Paz, J., Ortega, C., Febles, R., Garcia-Medina, J. &amp; Nogales, I. 2015. Kunkeliella canariensis, nuevo caso de" type="book" year="2015">Pérez de Paz &amp; al., 2015</bibRefCitation>
).
<bibRefCitation id="1E4D6A922C794004FB33A4EEFA9B6BE2" author="Stearn, W. T." box="[1239,1384,1435,1458]" pageId="10" pageNumber="200" pagination="11 - 26" refId="ref45503" refString="Stearn, W. T. 1972. Kunkeliella, a new genus of Santalaceae in the Canary Islands. Cuad. Bot. Canaria 16: 11 - 26." type="journal article" year="1972">Stearn (1972)</bibRefCitation>
considered this a genus separate from
<taxonomicName id="BDDC6CE02C794004FB9DA4CEFB206B82" box="[1145,1235,1467,1490]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="genus">
<emphasis id="48A8CB712C794004FB9DA4CEFB206B82" box="[1145,1235,1467,1490]" italics="true" pageId="10" pageNumber="200">Thesium</emphasis>
</taxonomicName>
based mainly upon its possession of isopolar pollen and drupaceous fruits. Isopolar pollen is of questionably diagnostic value as it has been found in several South African species of
<taxonomicName id="BDDC6CE02C794004FB35A76EFAD86862" box="[1233,1323,1563,1586]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="genus">
<emphasis id="48A8CB712C794004FB35A76EFAD86862" box="[1233,1323,1563,1586]" italics="true" pageId="10" pageNumber="200">Thesium</emphasis>
</taxonomicName>
plus intermediates between heteropolar and isopolar pollen
<typeStatus id="A567A9C12C794004FADFA74EFA806802" box="[1339,1395,1595,1618]" pageId="10" pageNumber="200">types</typeStatus>
are known (
<bibRefCitation id="1E4D6A922C794004FC8FA72EFBA66822" author="Lobreau-Callen, D." box="[875,1109,1627,1650]" pageId="10" pageNumber="200" pagination="23 - 30" refId="ref43303" refString="Lobreau-Callen, D. 1980. Variations morphologiques due pollen du genre Thesium (Santalaceae). Mem. Mus. Natl. Hist. Nat., Ser. B, Bot. 27: 23 - 30." type="journal article" year="1980">Lobreau-Callen, 1980</bibRefCitation>
). Fleshy fruits are also present in
<taxonomicName id="BDDC6CE02C794004FCD5A70EFC6368C2" box="[817,912,1659,1682]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="species" species="fragile">
<emphasis id="48A8CB712C794004FCD5A70EFC6368C2" box="[817,912,1659,1682]" italics="true" pageId="10" pageNumber="200">T. fragile</emphasis>
</taxonomicName>
L.f.,
<taxonomicName id="BDDC6CE02C794004FC2EA70EFBB068C2" box="[970,1091,1659,1682]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="species" species="triflorum">
<emphasis id="48A8CB712C794004FC2EA70EFBB068C2" box="[970,1091,1659,1682]" italics="true" pageId="10" pageNumber="200">T. triflorum</emphasis>
</taxonomicName>
and
<taxonomicName id="BDDC6CE02C794004FB9CA70EFB1D68C2" box="[1144,1262,1659,1682]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="species" species="radicans">
<emphasis id="48A8CB712C794004FB9CA70EFB1D68C2" box="[1144,1262,1659,1682]" italics="true" pageId="10" pageNumber="200">T. radicans</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="7A6317632C794004FCA6A7EEFC816962" blockId="10.[789,1437,187,1970]" pageId="10" pageNumber="200">
Unsampled species include
<taxonomicName id="BDDC6CE02C794004FBBAA7EEFAD968E2" box="[1118,1322,1691,1714]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="species" species="canariense">
<emphasis id="48A8CB712C794004FBBAA7EEFAD968E2" box="[1118,1322,1691,1714]" italics="true" pageId="10" pageNumber="200">Thesium canariense</emphasis>
</taxonomicName>
from Gran Canaria and
<taxonomicName id="BDDC6CE02C794004FC70A7CEFB716882" authority="Svent." authorityName="Svent." box="[916,1154,1723,1746]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="species" species="psilotocladum">
<emphasis id="48A8CB712C794004FC70A7CEFBC86882" box="[916,1083,1723,1746]" italics="true" pageId="10" pageNumber="200">T. psilotocladum</emphasis>
Svent.
</taxonomicName>
from Tenerife (extinct). Recently, a new species,
<taxonomicName id="BDDC6CE02C794004FC1BA7AEFACD68A2" authority="P. Perez &amp; P. Sosa" authorityName="P. Perez &amp; P. Sosa" box="[1023,1342,1755,1778]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="species" species="palmense">
<emphasis id="48A8CB712C794004FC1BA7AEFB8868A2" box="[1023,1147,1755,1778]" italics="true" pageId="10" pageNumber="200">T. palmense</emphasis>
P.Pérez &amp; P.Sosa
</taxonomicName>
, was described from La Palma, closely related to
<taxonomicName id="BDDC6CE02C794004FB31A78EFA926942" box="[1237,1377,1787,1810]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="species" species="retamoides">
<emphasis id="48A8CB712C794004FB31A78EFA926942" box="[1237,1377,1787,1810]" italics="true" pageId="10" pageNumber="200">T. retamoides</emphasis>
</taxonomicName>
from Tenerife.
</paragraph>
</subSubSection>
</emphasis>
</treatment>
</document>

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@ -0,0 +1,115 @@
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<emphasis id="48A8CB712C794004FF70A34EFDD96C02" bold="true" box="[148,554,571,595]" pageId="10" pageNumber="200">
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0.
<taxonomicName id="BDDC6CE02C794004FF50A34EFEA26C03" ID-CoL="59W6" box="[180,337,571,595]" class="Magnoliopsida" family="Thesiaceae" genus="Lacomucinaea" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="genus">Lacomucinaea</taxonomicName>
clade (BIPP 1).
</paragraph>
</subSubSection>
<subSubSection id="32C644E82C794004FDEBA348FF466A82" pageId="10" pageNumber="200" type="discussion">
<paragraph id="7A6317632C794004FDEBA348FF466A82" blockId="10.[103,751,186,1970]" pageId="10" pageNumber="200">
<emphasis id="48A8CB712C794004FDEBA348FDD96C02" box="[527,554,573,594]" italics="true" pageId="10" pageNumber="200">—</emphasis>
<taxonomicName id="BDDC6CE02C794004FDD0A34EFF526C22" class="Magnoliopsida" family="Thesiaceae" genus="Lacomucinaea" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="species" species="lineata">Lacomucinaea lineata</taxonomicName>
(L.f.) Nickrent &amp; M.
<collectionCode id="1CCD8FA62C794004FE7CA32EFE5F6C22" box="[408,428,603,626]" country="USA" lsid="urn:lsid:biocol.org:col:15406" name="Harvard University - Arnold Arboretum" pageId="10" pageNumber="200" type="Herbarium">A</collectionCode>
.García (
<figureCitation id="E2E70BE62C794004FDEAA32EFD966C22" box="[526,613,603,626]" captionStart="Fig" captionStartId="11.[684,716,1893,1914]" captionTargetBox="[152,1484,187,1858]" captionTargetId="figure-0@11.[152,1484,187,1859]" captionTargetPageId="11" captionText="Fig. 3. Caption on next page." figureDoi="http://doi.org/10.5281/zenodo.14047590" httpUri="https://zenodo.org/record/14047590/files/figure.png" pageId="10" pageNumber="200">Fig. 3A</figureCitation>
), previously known as
<taxonomicName id="BDDC6CE02C794004FF35A30EFE786CC2" box="[209,395,635,658]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="species" species="lineatum">
<emphasis id="48A8CB712C794004FF35A30EFE786CC2" box="[209,395,635,658]" italics="true" pageId="10" pageNumber="200">Thesium lineatum</emphasis>
</taxonomicName>
, was classified as a distinct genus by
<bibRefCitation id="1E4D6A922C794004FF63A3EEFE606CE2" author="Nickrent, D. L. &amp; Garcia, M. A." box="[135,403,667,690]" pageId="10" pageNumber="200" pagination="173 - 184" refId="ref44227" refString="Nickrent, D. L. &amp; 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 &amp; García (2015)</bibRefCitation>
based on molecular and morphological evidence. This genus is sister to
<taxonomicName id="BDDC6CE02C794004FDE5A3CEFD666C82" box="[513,661,699,722]" class="Magnoliopsida" family="Thesiaceae" genus="Osyridicarpos" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="genus">
<emphasis id="48A8CB712C794004FDE5A3CEFD666C82" box="[513,661,699,722]" italics="true" pageId="10" pageNumber="200">Osyridicarpos</emphasis>
</taxonomicName>
and that clade is sister
<taxonomicName id="BDDC6CE02C794004FF19A3AEFEA66CA2" box="[253,341,731,754]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="genus">
<emphasis id="48A8CB712C794004FF19A3AEFEA66CA2" box="[253,341,731,754]" italics="true" pageId="10" pageNumber="200">Thesium</emphasis>
</taxonomicName>
, relationships with the highest support from both ITS and the chloroplast genes (
<figureCitation id="E2E70BE62C794004FDC0A38EFD856D42" box="[548,630,763,786]" captionStart="Fig" captionStartId="5.[151,183,1831,1852]" captionTargetBox="[300,1336,189,1795]" captionTargetId="graphics-0@5.[300,1001,198,1789]" captionTargetPageId="5" captionText="Fig. 2. Phylogenetic analysis of Thesium using nuclear ribosomal internal transcribed spacer (ITS) sequences. This tree, rooted with Osyridicarpos and Lacomucinaea (outgroups) gives Bayesian posterior probabilities at the nodes. Clades discussed in the text are numbered. Geographical sources for the sampled taxa are abbreviated as follows: AFN = Northern Africa, AFS = Southern Africa, AFT = Tropical Africa, ASC = Central Asia, ASE = Eastern Asia, ASN = Northern Asia, ASS = Southern Asia, ASW = Western Asia, AUS = Australia, CIS = Canary Islands, EUR = Europe, MAD = Madagascar, SAM = South America, and USA = United States of America." figureDoi="http://doi.org/10.5281/zenodo.14047588" httpUri="https://zenodo.org/record/14047588/files/figure.png" pageId="10" pageNumber="200">Fig. 2A</figureCitation>
). One may ask why
<taxonomicName id="BDDC6CE02C794004FF5BA26EFEAB6D62" box="[191,344,795,818]" class="Magnoliopsida" family="Thesiaceae" genus="Lacomucinaea" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="genus">
<emphasis id="48A8CB712C794004FF5BA26EFEAB6D62" box="[191,344,795,818]" italics="true" pageId="10" pageNumber="200">Lacomucinaea</emphasis>
</taxonomicName>
was not included within
<taxonomicName id="BDDC6CE02C794004FDB0A26EFD1B6D62" box="[596,744,795,818]" class="Magnoliopsida" family="Thesiaceae" genus="Osyridicarpos" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="genus">
<emphasis id="48A8CB712C794004FDB0A26EFD1B6D62" box="[596,744,795,818]" italics="true" pageId="10" pageNumber="200">Osyridicarpos</emphasis>
</taxonomicName>
. These two genera share several morphological features, e.g., primary phloem fiber bundles near the stem surface, bisexual flowers, lack of a swollen pedicel, and fleshy fruits with smooth exocarps. They differ in other characters such as habit (shrub vs. scrambling subshrub), leaf morphology, leaf persistence, inflorescence structure, presence of external floral gland, style length and ribbing on ovary. Four species have been described in
<taxonomicName id="BDDC6CE02C794004FF65A56EFEE66A62" box="[129,277,1051,1074]" class="Magnoliopsida" family="Thesiaceae" genus="Osyridicarpos" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="genus">
<emphasis id="48A8CB712C794004FF65A56EFEE66A62" box="[129,277,1051,1074]" italics="true" pageId="10" pageNumber="200">Osyridicarpos</emphasis>
</taxonomicName>
although it is frequently treated as monotypic (
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<emphasis id="48A8CB712C794004FF8BA54EFEDA6A02" box="[111,297,1083,1106]" italics="true" pageId="10" pageNumber="200">O. schimperianus</emphasis>
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). Further studies might prove that several species should be recognized in the genus, all of them very different from
<taxonomicName id="BDDC6CE02C794004FF04A50EFE8F6AC2" box="[224,380,1147,1170]" class="Magnoliopsida" family="Thesiaceae" genus="Lacomucinaea" kingdom="Plantae" order="Santalales" pageId="10" pageNumber="200" phylum="Tracheophyta" rank="genus">
<emphasis id="48A8CB712C794004FF04A50EFE8F6AC2" box="[224,380,1147,1170]" italics="true" pageId="10" pageNumber="200">Lacomucinaea</emphasis>
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. Thus, the preponderance of molecular and morphological evidence justifies recognition of two genera.
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1.
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clade (BIPP 1).
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<emphasis id="48A8CB712C794004FE59A5A8FE2B6AA2" bold="true" box="[445,472,1245,1266]" italics="true" pageId="10" pageNumber="200">—</emphasis>
This clade (
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) contains all species now classified as
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<emphasis id="48A8CB712C794004FE37A58EFDDE6B42" box="[467,557,1275,1298]" italics="true" pageId="10" pageNumber="200">Thesium</emphasis>
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and includes taxa formerly placed in
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<emphasis id="48A8CB712C794004FED6A46EFE1B6B62" box="[306,488,1307,1330]" italics="true" pageId="10" pageNumber="200">Austroamericium</emphasis>
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,
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<emphasis id="48A8CB712C794004FE12A46EFD666B62" box="[502,661,1307,1330]" italics="true" pageId="10" pageNumber="200">Chrysothesium</emphasis>
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,
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<emphasis id="48A8CB712C794004FD47A46EFF6E6B02" italics="true" pageId="10" pageNumber="200">Kunkeliella</emphasis>
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, and
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<emphasis id="48A8CB712C794004FF3FA44EFEBA6B02" box="[219,329,1339,1362]" italics="true" pageId="10" pageNumber="200">Thesidium</emphasis>
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. The monophyly of
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<emphasis id="48A8CB712C794004FDCCA44EFD716B02" box="[552,642,1339,1362]" italics="true" pageId="10" pageNumber="200">Thesium</emphasis>
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including these genera and excluding
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<emphasis id="48A8CB712C794004FE79A42EFDCA6B22" box="[413,569,1371,1394]" italics="true" pageId="10" pageNumber="200">Lacomucinaea</emphasis>
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is also resolved with chloroplast markers with the highest support
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6.
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clade (BIPP 1).
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<emphasis id="48A8CB712C7F4002FE09A568FDFB6A62" bold="true" box="[493,520,1053,1074]" italics="true" pageId="12" pageNumber="202">—</emphasis>
In our sampling this clade contains a single species,
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<emphasis id="48A8CB712C7F4002FE0AA54EFD1D6A02" box="[494,750,1083,1106]" italics="true" pageId="12" pageNumber="202">Thesium mauritanicum</emphasis>
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), from North Africa (
<collectingCountry id="02CB57F32C7F4002FE41A52EFDFA6A22" box="[421,521,1115,1138]" name="Morocco" pageId="12" pageNumber="202">Morocco</collectingCountry>
,
<collectingCountry id="02CB57F32C7F4002FDF0A52EFD9B6A22" box="[532,616,1115,1138]" name="Algeria" pageId="12" pageNumber="202">Algeria</collectingCountry>
). Its position on the phylogenetic tree (
<figureCitation id="E2E70BE62C7F4002FE96A50EFE376AC2" box="[370,452,1147,1170]" captionStart="Fig" captionStartId="5.[151,183,1831,1852]" captionTargetBox="[300,1336,189,1795]" captionTargetId="graphics-0@5.[300,1001,198,1789]" captionTargetPageId="5" captionText="Fig. 2. Phylogenetic analysis of Thesium using nuclear ribosomal internal transcribed spacer (ITS) sequences. This tree, rooted with Osyridicarpos and Lacomucinaea (outgroups) gives Bayesian posterior probabilities at the nodes. Clades discussed in the text are numbered. Geographical sources for the sampled taxa are abbreviated as follows: AFN = Northern Africa, AFS = Southern Africa, AFT = Tropical Africa, ASC = Central Asia, ASE = Eastern Asia, ASN = Northern Asia, ASS = Southern Asia, ASW = Western Asia, AUS = Australia, CIS = Canary Islands, EUR = Europe, MAD = Madagascar, SAM = South America, and USA = United States of America." figureDoi="http://doi.org/10.5281/zenodo.14047588" httpUri="https://zenodo.org/record/14047588/files/figure.png" pageId="12" pageNumber="202">Fig. 2A</figureCitation>
) indicates that it is phylogenetically distinct from the African clade (26). When
<bibRefCitation id="1E4D6A922C7F4002FD45A5EEFF1A6A82" author="Battandier, M. J. A." pageId="12" pageNumber="202" pagination="385 - 393" refId="ref41110" refString="Battandier, M. J. A. 1889. Note sur quelques plantes d' Algerie rares ou nouvelles (1). Bull. Soc. Bot. France 35: 385 - 393. https: // doi. org / 10.1080 / 00378941.1888.10830389" type="journal article" year="1889">Battandier (1889)</bibRefCitation>
described
<taxonomicName id="BDDC6CE02C7F4002FE95A5CEFDD86A82" box="[369,555,1211,1234]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="12" pageNumber="202" phylum="Tracheophyta" rank="species" species="mauritanicum">
<emphasis id="48A8CB712C7F4002FE95A5CEFDD86A82" box="[369,555,1211,1234]" italics="true" pageId="12" pageNumber="202">T. mauritanicum</emphasis>
</taxonomicName>
he suggested a relationship with
<taxonomicName id="BDDC6CE02C7F4002FED9A5AEFDF66AA2" authority="Zucc." authorityName="Zucc." box="[317,517,1243,1266]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="12" pageNumber="202" phylum="Tracheophyta" rank="species" species="bergeri">
<emphasis id="48A8CB712C7F4002FED9A5AEFE406AA2" box="[317,435,1243,1266]" italics="true" pageId="12" pageNumber="202">T. bergeri</emphasis>
Zucc.
</taxonomicName>
(and its synonym
<taxonomicName id="BDDC6CE02C7F4002FF83A58EFE646B42" authority="Boiss. &amp; Spruner" authorityName="Boiss. &amp; Spruner" box="[103,407,1275,1298]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="12" pageNumber="202" phylum="Tracheophyta" rank="species" species="graecum">
<emphasis id="48A8CB712C7F4002FF83A58EFF2F6B42" box="[103,220,1275,1298]" italics="true" pageId="12" pageNumber="202">T. graecum</emphasis>
Boiss. &amp; Spruner
</taxonomicName>
) from
<collectionCode id="1CCD8FA62C7F4002FE38A58EFE066B42" box="[476,501,1275,1298]" country="Austria" lsid="urn:lsid:biocol.org:col:15588" name="Naturhistorisches Museum Wien" pageId="12" pageNumber="202" type="Herbarium">W</collectionCode>
Asia and
<collectingCountry id="02CB57F32C7F4002FDB8A58EFD596B42" box="[604,682,1275,1298]" name="Greece" pageId="12" pageNumber="202">Greece</collectingCountry>
. However, ITS and chloroplast sequences show that species as part of clade 9. The flowers of
<taxonomicName id="BDDC6CE02C7F4002FE6CA44EFDCB6B02" box="[392,568,1339,1362]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="12" pageNumber="202" phylum="Tracheophyta" rank="species" species="mauritanicum">
<emphasis id="48A8CB712C7F4002FE6CA44EFDCB6B02" box="[392,568,1339,1362]" italics="true" pageId="12" pageNumber="202">T. mauritanicum</emphasis>
</taxonomicName>
differ little from Eurasian
<taxonomicName id="BDDC6CE02C7F4002FF2DA42EFE866B22" box="[201,373,1371,1394]" pageId="12" pageNumber="202">
<emphasis id="48A8CB712C7F4002FF2DA42EFED06B22" box="[201,291,1371,1394]" italics="true" pageId="12" pageNumber="202">Thesium</emphasis>
species
</taxonomicName>
but dense pubescence on the stems, leaves, bracts, abaxial surface of the petals and fruits are uncommon characters in the genus. This clade corresponds to
<emphasis id="48A8CB712C7F4002FF83A4CEFF856B82" box="[103,118,1467,1490]" italics="true" pageId="12" pageNumber="202">
<collectionCode id="1CCD8FA62C7F4002FF83A4CEFF856B82" box="[103,118,1467,1490]" name="Tavera, Department of Geology and Geophysics" pageId="12" pageNumber="202">T</collectionCode>
</emphasis>
. ser.
<taxonomicName id="BDDC6CE02C7F4002FF57A4CEFE5A6B82" authority="Hendrych" authorityName="Hendrych" box="[179,425,1467,1490]" class="Gastropoda" family="Clausiliidae" genus="Mauritanica" kingdom="Animalia" order="Stylommatophora" pageId="12" pageNumber="202" phylum="Mollusca" rank="genus">
<emphasis id="48A8CB712C7F4002FF57A4CEFEC46B82" box="[179,311,1467,1490]" italics="true" pageId="12" pageNumber="202">Mauritanica</emphasis>
Hendrych
</taxonomicName>
that also included the Libyan endemic
<taxonomicName id="BDDC6CE02C7F4002FF2DA4AEFE246BA2" authority="Pamp. It" authorityName="Pamp. It" box="[201,471,1499,1522]" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="12" pageNumber="202" phylum="Tracheophyta" rank="species" species="erythronicum">
<emphasis id="48A8CB712C7F4002FF2DA4AEFE816BA2" box="[201,370,1499,1522]" italics="true" pageId="12" pageNumber="202">T. erythronicum</emphasis>
Pamp. It
</taxonomicName>
shares with
<taxonomicName id="BDDC6CE02C7F4002FD85A4AEFF606842" class="Magnoliopsida" family="Thesiaceae" genus="Thesium" kingdom="Plantae" order="Santalales" pageId="12" pageNumber="202" phylum="Tracheophyta" rank="species" species="mauritanicum">
<emphasis id="48A8CB712C7F4002FD85A4AEFF606842" italics="true" pageId="12" pageNumber="202">T. mauritanicum</emphasis>
</taxonomicName>
a subshrub habit and reticulate fruits; however, the two species differ in several vegetative and inflorescence features suggesting that it might be related to other Paleoboreal species, possibly in clades 8 or 9.
</paragraph>
</subSubSection>
</emphasis>
</treatment>
</document>