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<document id="70ACF308A04C9B0F90905F81779FEFEC" ID-CLB-Dataset="224103" ID-DOI="10.37520/fi.2022.016" ID-GBIF-Dataset="a360f5c7-16da-4c69-afbe-01e6ad032418" ID-ISSN="2533-4069" ID-Zenodo-Dep="7522801" IM.bibliography_approvedBy="juliana" IM.illustrations_approvedBy="juliana" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_approvedBy="juliana" IM.taxonomicNames_approvedBy="juliana" IM.treatments_approvedBy="juliana" checkinTime="1673382488781" checkinUser="felipe" docAuthor="Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel &amp; Kvaček, Jiří" docDate="2022" docId="03FD87F2FFEDFFFEFF14F998C6C6FCC4" docLanguage="en" docName="FossilImprint.78.2.341-424.pdf" docOrigin="Fossil Imprint 78 (2)" docSource="http://dx.doi.org/10.37520/fi.2022.016" docStyle="DocumentStyle:0EC54BF35E227CA4627F894F6BA30634.4:FossilImprint.2016-.journal_article" docStyleId="0EC54BF35E227CA4627F894F6BA30634" docStyleName="FossilImprint.2016-.journal_article" docStyleVersion="4" docTitle="Clavatipollenites" docType="treatment" docVersion="2" lastPageNumber="357" masterDocId="FFC4FF8AFFFDFFEEFFBFFFFFC378FFA6" masterDocTitle="The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms" masterLastPageNumber="424" masterPageNumber="341" pageNumber="357" updateTime="1728648832095" updateUser="juliana" zenodo-license-document="CC-BY-NC-ND-3.0">
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<mods:title id="2238D689F32AB2BD273D380ABB13AC6A">The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms</mods:title>
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<mods:namePart id="BA772D8031F3E98873041F4E6407D974">Friis, Else Marie</mods:namePart>
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<mods:namePart id="FD493E6C7011E53215E1BDCB8149EBE3">Pedersen, Kaj Raunsgaard</mods:namePart>
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<mods:namePart id="6B01B3263869D8FDAE976A99B26B0BE0">Mendes, Mário Miguel</mods:namePart>
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<subSubSection id="C34E656FFFEDFFFEFF14F998C178F906" pageId="16" pageNumber="357" type="nomenclature">
<paragraph id="8BEB36E4FFEDFFFEFF14F998C178F906" blockId="16.[171,725,1639,1696]" pageId="16" pageNumber="357">
<heading id="D0A38188FFEDFFFEFF14F998C178F906" centered="true" fontSize="8" level="2" pageId="16" pageNumber="357" reason="2">
<emphasis id="B920EAF6FFEDFFFEFF14F998C178F906" bold="true" pageId="16" pageNumber="357">
Stamens with monocolpate
<taxonomicName id="4C544D67FFEDFFFEFE5CF998C1DCF9D9" authority="(Walker and Walker 1984)" box="[483,676,1639,1663]" class="Magnoliopsida" family="Chloranthaceae" genus="Clavatipollenites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Chloranthales" pageId="16" pageNumber="357" phylum="Tracheophyta" rank="genus">
<emphasis id="B920EAF6FFEDFFFEFE5CF998C1DCF9D9" bold="true" box="[483,676,1639,1663]" italics="true" pageId="16" pageNumber="357">Clavatipollenites-</emphasis>
</taxonomicName>
<typeStatus id="54EF8846FFEDFFFEFD1BF997C1ADF926" box="[676,725,1640,1664]" pageId="16" pageNumber="357">type</typeStatus>
pollen sp. 1
</emphasis>
</heading>
</paragraph>
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<subSubSection id="C34E656FFFEDFFFEFED0F956C6C6FCC4" pageId="16" pageNumber="357" type="description">
<paragraph id="8BEB36E4FFEDFFFEFED0F956C16AF967" blockId="16.[367,530,1705,1729]" box="[367,530,1705,1729]" pageId="16" pageNumber="357">
<figureCitation id="136F2A61FFEDFFFEFED0F956C16AF967" box="[367,530,1705,1729]" captionStart="Text-fig" captionStartId="18.[147,226,1475,1496]" captionTargetBox="[148,1462,151,1459]" captionTargetId="figure-186@18.[145,1466,148,1461]" captionTargetPageId="18" captionText="Text-fig. 10. Scanning electron microscope (SEM) images of isolated “Stamen fragment with Clavatipollenites-type pollen sp. 1”; Catefica locality, Portugal. a) Fragment of tetrasporangiate stamen with pollen in situ; b) Detail from stamen fragment showing distal and proximal surfaces of in situ pollen grains and tiny orbicules on the inner surface of the anther wall (arrows); c) Pollen grains in distal view showing short colpi with irregular margins and aperture membrane with irregular verrucae; d) Detail of pollen wall showing the semitectate-reticulate tectum and long, scattered columellae supporting the narrow muri with finely verrucate supratectal ornamentation. Specimen, Catefica 50-S170387 (ad). Scale bars = 600 Μm (a), 20 Μm (b), 6 Μm (c), 1.5 Μm (d)." figureDoi="http://doi.org/10.5281/zenodo.7522821" httpUri="https://zenodo.org/record/7522821/files/figure.png" pageId="16" pageNumber="357">Text-fig. 10ad</figureCitation>
</paragraph>
<paragraph id="8BEB36E4FFEDFFFEFF19F926C66BFEF7" blockId="16.[129,768,1752,2100]" lastBlockId="16.[843,1483,152,337]" pageId="16" pageNumber="357">
Description and remarks. The material comprises two stamen fragments that have a long, narrow, elongated tetrasporangiate anther (
<figureCitation id="136F2A61FFEDFFFEFE52F8E6C10EF897" box="[493,630,1817,1841]" captionStart="Text-fig" captionStartId="18.[147,226,1475,1496]" captionTargetBox="[148,1462,151,1459]" captionTargetId="figure-186@18.[145,1466,148,1461]" captionTargetPageId="18" captionText="Text-fig. 10. Scanning electron microscope (SEM) images of isolated “Stamen fragment with Clavatipollenites-type pollen sp. 1”; Catefica locality, Portugal. a) Fragment of tetrasporangiate stamen with pollen in situ; b) Detail from stamen fragment showing distal and proximal surfaces of in situ pollen grains and tiny orbicules on the inner surface of the anther wall (arrows); c) Pollen grains in distal view showing short colpi with irregular margins and aperture membrane with irregular verrucae; d) Detail of pollen wall showing the semitectate-reticulate tectum and long, scattered columellae supporting the narrow muri with finely verrucate supratectal ornamentation. Specimen, Catefica 50-S170387 (ad). Scale bars = 600 Μm (a), 20 Μm (b), 6 Μm (c), 1.5 Μm (d)." figureDoi="http://doi.org/10.5281/zenodo.7522821" httpUri="https://zenodo.org/record/7522821/files/figure.png" pageId="16" pageNumber="357">Text-fig. 10a</figureCitation>
) with in situ
<taxonomicName id="4C544D67FFEDFFFEFF3EF8C6C24BF8F7" baseAuthorityName="Walker and Walker" baseAuthorityYear="1984" box="[129,307,1849,1873]" class="Magnoliopsida" family="Chloranthaceae" genus="Clavatipollenites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Chloranthales" pageId="16" pageNumber="357" phylum="Tracheophyta" rank="genus">
<emphasis id="B920EAF6FFEDFFFEFF3EF8C6C24BF8F7" box="[129,307,1849,1873]" italics="true" pageId="16" pageNumber="357">Clavatipollenites</emphasis>
</taxonomicName>
-
<typeStatus id="54EF8846FFEDFFFEFE86F8C5C210F8F4" box="[313,360,1850,1874]" pageId="16" pageNumber="357">type</typeStatus>
pollen (
<figureCitation id="136F2A61FFEDFFFEFE77F8C6C10BF8F7" box="[456,627,1849,1873]" captionStart="Text-fig" captionStartId="18.[147,226,1475,1496]" captionTargetBox="[148,1462,151,1459]" captionTargetId="figure-186@18.[145,1466,148,1461]" captionTargetPageId="18" captionText="Text-fig. 10. Scanning electron microscope (SEM) images of isolated “Stamen fragment with Clavatipollenites-type pollen sp. 1”; Catefica locality, Portugal. a) Fragment of tetrasporangiate stamen with pollen in situ; b) Detail from stamen fragment showing distal and proximal surfaces of in situ pollen grains and tiny orbicules on the inner surface of the anther wall (arrows); c) Pollen grains in distal view showing short colpi with irregular margins and aperture membrane with irregular verrucae; d) Detail of pollen wall showing the semitectate-reticulate tectum and long, scattered columellae supporting the narrow muri with finely verrucate supratectal ornamentation. Specimen, Catefica 50-S170387 (ad). Scale bars = 600 Μm (a), 20 Μm (b), 6 Μm (c), 1.5 Μm (d)." figureDoi="http://doi.org/10.5281/zenodo.7522821" httpUri="https://zenodo.org/record/7522821/files/figure.png" pageId="16" pageNumber="357">Text-fig. 10bd</figureCitation>
). The anther is about
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long and about
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broad. Pollen is monoaperturate, almost circular in equatorial outline and about 1720 µm in diameter. The aperture is about 12 µm long, with an irregular, indistinctly delimited margin and a colpus membrane covered by irregular verrucae that grade into the non-apertural reticulum (
<figureCitation id="136F2A61FFEDFFFEFE61F803C1FBF7B2" box="[478,643,2044,2068]" captionStart="Text-fig" captionStartId="18.[147,226,1475,1496]" captionTargetBox="[148,1462,151,1459]" captionTargetId="figure-186@18.[145,1466,148,1461]" captionTargetPageId="18" captionText="Text-fig. 10. Scanning electron microscope (SEM) images of isolated “Stamen fragment with Clavatipollenites-type pollen sp. 1”; Catefica locality, Portugal. a) Fragment of tetrasporangiate stamen with pollen in situ; b) Detail from stamen fragment showing distal and proximal surfaces of in situ pollen grains and tiny orbicules on the inner surface of the anther wall (arrows); c) Pollen grains in distal view showing short colpi with irregular margins and aperture membrane with irregular verrucae; d) Detail of pollen wall showing the semitectate-reticulate tectum and long, scattered columellae supporting the narrow muri with finely verrucate supratectal ornamentation. Specimen, Catefica 50-S170387 (ad). Scale bars = 600 Μm (a), 20 Μm (b), 6 Μm (c), 1.5 Μm (d)." figureDoi="http://doi.org/10.5281/zenodo.7522821" httpUri="https://zenodo.org/record/7522821/files/figure.png" pageId="16" pageNumber="357">Text-fig. 10b, c</figureCitation>
). The exine is semitectate-reticulate with a homobrochate reticulum (
<figureCitation id="136F2A61FFEDFFFEFCEDFF67C080FF16" box="[850,1016,152,176]" captionStart="Text-fig" captionStartId="18.[147,226,1475,1496]" captionTargetBox="[148,1462,151,1459]" captionTargetId="figure-186@18.[145,1466,148,1461]" captionTargetPageId="18" captionText="Text-fig. 10. Scanning electron microscope (SEM) images of isolated “Stamen fragment with Clavatipollenites-type pollen sp. 1”; Catefica locality, Portugal. a) Fragment of tetrasporangiate stamen with pollen in situ; b) Detail from stamen fragment showing distal and proximal surfaces of in situ pollen grains and tiny orbicules on the inner surface of the anther wall (arrows); c) Pollen grains in distal view showing short colpi with irregular margins and aperture membrane with irregular verrucae; d) Detail of pollen wall showing the semitectate-reticulate tectum and long, scattered columellae supporting the narrow muri with finely verrucate supratectal ornamentation. Specimen, Catefica 50-S170387 (ad). Scale bars = 600 Μm (a), 20 Μm (b), 6 Μm (c), 1.5 Μm (d)." figureDoi="http://doi.org/10.5281/zenodo.7522821" httpUri="https://zenodo.org/record/7522821/files/figure.png" pageId="16" pageNumber="357">Text-fig. 10bd</figureCitation>
). Lumina are irregular, up to about 1.4 µm in diameter. Muri are narrow, 0.20.4 µm wide, and ornamented with minute verrucae arranged in a single longitudinal row. Columellae are long, about 0.6 µm, and widely spaced (Textfig. 10d). Orbicules observed on the inner surface of the anther wall are tiny, spherical and with fine spines.
</paragraph>
<paragraph id="8BEB36E4FFEDFFFEFCCEFE97C64AFDC7" blockId="16.[843,1482,360,866]" pageId="16" pageNumber="357">
<collectionCode id="ED45AE21FFEDFFFEFCCEFE97C0FCFE26" box="[881,900,360,384]" country="USA" lsid="urn:lsid:biocol.org:col:15406" name="Harvard University - Arnold Arboretum" pageId="16" pageNumber="357" type="Herbarium">Affinity</collectionCode>
and other occurrences. The general shape of the pollen grains, as well as their semitectate reticulate pollen wall, indistinctly delimited aperture, and aperture membrane covered by irregular verrucae, is closely similar to dispersed pollen assigned to the extinct pollen genus
<taxonomicName id="4C544D67FFEDFFFEFC68FDF7C7F5FD86" baseAuthorityName="Walker and Walker" baseAuthorityYear="1984" box="[983,1165,520,544]" class="Magnoliopsida" family="Chloranthaceae" genus="Clavatipollenites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Chloranthales" pageId="16" pageNumber="357" phylum="Tracheophyta" rank="genus">
<emphasis id="B920EAF6FFEDFFFEFC68FDF7C7F5FD86" box="[983,1165,520,544]" italics="true" pageId="16" pageNumber="357">Clavatipollenites</emphasis>
</taxonomicName>
and pollen of extant
<taxonomicName id="4C544D67FFEDFFFEFAD6FDF6C71FFDE7" authority="(Walker and Walker 1984)" baseAuthorityName="Walker and Walker" baseAuthorityYear="1984" class="Magnoliopsida" family="Chloranthaceae" genus="Ascarina" kingdom="Plantae" order="Chloranthales" pageId="16" pageNumber="357" phylum="Tracheophyta" rank="genus">
<emphasis id="B920EAF6FFEDFFFEFAD6FDF6C6B1FD86" box="[1385,1481,521,544]" italics="true" pageId="16" pageNumber="357">Ascarina</emphasis>
(
<bibRefCitation id="EFC54B15FFEDFFFEFCEDFDD6C727FDE7" author="Walker, J. W. &amp; Walker, A. G." box="[850,1119,553,577]" pageId="16" pageNumber="357" pagination="464 - 521" refId="ref60968" refString="Walker, J. W., Walker, A. G. (1984): Ultrastructure of Lower Cretaceous angiosperm pollen and the origin and early evolution of flowering plants. - Annals of the Missouri Botanical Garden, 71: 464 - 521. https: // doi. org / 10.2307 / 2399035" type="journal article" year="1984">Walker and Walker 1984</bibRefCitation>
)
</taxonomicName>
. Based on these similarities these fossils are included here in the
<taxonomicName id="4C544D67FFEDFFFEFB28FDB6C657FDC7" authorityName="R. BR. ex SIMS" authorityYear="1821" box="[1175,1327,585,609]" class="Magnoliopsida" higherTaxonomySource="GBIF" kingdom="Plantae" order="Chloranthales" pageId="16" pageNumber="357" phylum="Tracheophyta" rank="order">Chloranthales</taxonomicName>
.
</paragraph>
<paragraph id="8BEB36E4FFEDFFFEFCCEFD96C6C6FCC4" blockId="16.[843,1482,360,866]" pageId="16" pageNumber="357">
The four staminate structures described here from Catefica that have monocolpate
<taxonomicName id="4C544D67FFEDFFFEFB29FD76C634FD07" baseAuthorityName="Walker and Walker" baseAuthorityYear="1984" box="[1174,1356,649,673]" class="Magnoliopsida" family="Chloranthaceae" genus="Clavatipollenites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Chloranthales" pageId="16" pageNumber="357" phylum="Tracheophyta" rank="genus">
<emphasis id="B920EAF6FFEDFFFEFB29FD76C634FD07" box="[1174,1356,649,673]" italics="true" pageId="16" pageNumber="357">Clavatipollenites</emphasis>
</taxonomicName>
-
<typeStatus id="54EF8846FFEDFFFEFAECFD75C6FBFD04" box="[1363,1411,650,674]" pageId="16" pageNumber="357">type</typeStatus>
pollen differ from each other in size and shape of anthers and in details of the pollen wall (
<tableCitation id="C6D6035FFFEDFFFEFBE1FD36C7D8FD47" box="[1118,1184,713,737]" captionStart="Table 1" captionStartId="5.[106,162,152,173]" captionTargetId="graphics-427@5.[106,1462,246,1415]" captionTargetPageId="5" captionText="Table 1. Chloranthoid taxa recognized in the Catefica mesofossil flora based on inflorescences and flowers as well as isolated fruits, seeds and stamens with pollen characters added for those taxa where pollen are known." httpUri="http://table.plazi.org/id/DF2B666CFFF8FFEBFFD5FF67C09DFF6C" pageId="16" pageNumber="357" tableUuid="DF2B666CFFF8FFEBFFD5FF67C09DFF6C">Tab. 1</tableCitation>
). Similar
<emphasis id="B920EAF6FFEDFFFEFAB4FD36C001FCA4" italics="true" pageId="16" pageNumber="357">Clavatipollenitestype</emphasis>
pollen grains are also present in the palynological preparations from Catefica.
<taxonomicName id="4C544D67FFEDFFFEFB33FCF6C63AFC87" baseAuthorityName="Walker and Walker" baseAuthorityYear="1984" box="[1164,1346,777,801]" class="Magnoliopsida" family="Chloranthaceae" genus="Clavatipollenites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Chloranthales" pageId="16" pageNumber="357" phylum="Tracheophyta" rank="genus">
<emphasis id="B920EAF6FFEDFFFEFB33FCF6C63AFC87" box="[1164,1346,777,801]" italics="true" pageId="16" pageNumber="357">Clavatipollenites</emphasis>
</taxonomicName>
-
<typeStatus id="54EF8846FFEDFFFEFAF6FCF5C601FC84" box="[1353,1401,778,802]" pageId="16" pageNumber="357">type</typeStatus>
pollen observed on
<taxonomicName id="4C544D67FFEDFFFEFC5CFCD5C67AFCE7" authorityName="E. M. FRIIS, G. W. GRIMM, M. M. MENDES et K. R. PEDERSEN" authorityYear="2015" box="[995,1282,810,834]" class="Magnoliopsida" family="Chloranthaceae" genus="Canrightiopsis" higherTaxonomySource="GBIF" kingdom="Plantae" order="Chloranthales" pageId="16" pageNumber="357" phylum="Tracheophyta" rank="species" species="crassitesta">
<emphasis id="B920EAF6FFEDFFFEFC5CFCD5C67AFCE7" box="[995,1282,810,834]" italics="true" pageId="16" pageNumber="357">Canrightiopsis crassitesta</emphasis>
</taxonomicName>
from Catefica is much smaller and with a more open reticulum (see above).
</paragraph>
</subSubSection>
</treatment>
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