568 lines
80 KiB
XML
568 lines
80 KiB
XML
<document ID-DOI="10.1073/pnas.2013442118" ID-PMC="PMC7980368" ID-PubMed="33836571" ID-Zenodo-Dep="4596604" approvalRequired="21" approvalRequired_for_document="1" approvalRequired_for_textStreams="19" approvalRequired_for_treatments="1" checkinTime="1615465050340" checkinUser="donat" docAuthor="Wang, Jun, Hilton, Jason, Pfefferkorn, Hermann W., Wang, Shijun, Zhang, Yi, Bek, Jiri, Pšenička, Josef, Seyfullah, Leyla J. & Dilcher, David" docDate="2021" docId="987687A30E44FFFF87E3FA5F6D78FE50" docLanguage="en" docName="pnas.e2013442118.full.pdf" docOrigin="Proceedings of the National Academy of Sciences 118 (11)" docSource="http://dx.doi.org/10.1073/pnas.2013442118" docTitle="Paratingia wuhaia J. Wang 2021, sp. nov." docType="treatment" docVersion="4" lastPageNumber="4" masterDocId="644FFFDB0E45FFFC8769FFAD6909FF9E" masterDocTitle="Ancient noeggerathialean reveals the seed plant sister group diversified alongside the primary seed plant radiation" masterLastPageNumber="7" masterPageNumber="1" pageNumber="2" updateTime="1668143302897" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0" zenodo-license-treatments="CC0-1.0">
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<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:titleInfo>
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<mods:title>Ancient noeggerathialean reveals the seed plant sister group diversified alongside the primary seed plant radiation</mods:title>
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</mods:titleInfo>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Wang, Jun</mods:namePart>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Hilton, Jason</mods:namePart>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Pfefferkorn, Hermann W.</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Wang, Shijun</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Zhang, Yi</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Bek, Jiri</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Pšenička, Josef</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Seyfullah, Leyla J.</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Dilcher, David</mods:namePart>
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<mods:typeOfResource>text</mods:typeOfResource>
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<mods:title>Proceedings of the National Academy of Sciences</mods:title>
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<mods:part>
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<mods:date>2021</mods:date>
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<mods:detail type="pubDate">
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<mods:number>2021-03-08</mods:number>
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<mods:detail type="volume">
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<mods:number>118</mods:number>
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<mods:detail type="issue">
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<mods:number>11</mods:number>
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<mods:extent unit="page">
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<mods:start>1</mods:start>
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<mods:url>http://dx.doi.org/10.1073/pnas.2013442118</mods:url>
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<mods:classification>journal article</mods:classification>
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<mods:identifier type="DOI">10.1073/pnas.2013442118</mods:identifier>
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<mods:identifier type="PMC">PMC7980368</mods:identifier>
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<mods:identifier type="PubMed">33836571</mods:identifier>
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<mods:identifier type="Zenodo-Dep">4596604</mods:identifier>
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<treatment LSID="urn:lsid:plazi:treatment:987687A30E44FFFF87E3FA5F6D78FE50" httpUri="http://treatment.plazi.org/id/987687A30E44FFFF87E3FA5F6D78FE50" lastPageId="3" lastPageNumber="4" pageId="1" pageNumber="2">
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<subSubSection box="[138,654,1522,1544]" pageId="1" pageNumber="2" type="nomenclature">
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<paragraph blockId="1.[138,769,1163,1943]" box="[138,654,1522,1544]" pageId="1" pageNumber="2">
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Species—
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<taxonomicName authority="J. Wang et al." authorityName="J. Wang" authorityYear="2021" box="[237,568,1522,1544]" class="Noeggerathiopsida" family="Paratingia" kingdom="Plantae" order="Noeggerathiales" pageId="1" pageNumber="2" rank="species" species="wuhaia" status="sp. nov.">
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<emphasis box="[237,416,1522,1544]" italics="true" pageId="1" pageNumber="2">Paratingia wuhaia</emphasis>
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J. Wang et al.
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</taxonomicName>
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<taxonomicNameLabel box="[575,654,1523,1544]" pageId="1" pageNumber="2" rank="species">sp. nov.</taxonomicNameLabel>
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</paragraph>
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</subSubSection>
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<subSubSection pageId="1" pageNumber="2" type="etymology">
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<paragraph blockId="1.[138,769,1163,1943]" pageId="1" pageNumber="2">Etymology—The specific epithet refers to Wuhai city to which the Wuda Coalfield belongs.</paragraph>
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</subSubSection>
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<subSubSection pageId="1" pageNumber="2" type="diagnosis">
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<paragraph blockId="1.[138,769,1163,1943]" lastBlockId="1.[828,1458,141,787]" pageId="1" pageNumber="2">
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Species diagnosis—Monopodial stem with once-pinnate compound leaves and pseudostrobili forming a crown. Stem eustele with pith; cauline primary xylem strand adaxially curved; secondary xylem pycnoxylic; leaf traces tangentially elongated. Rachis of leaf and pseudostrobilus with a reversed Ω- shaped vascular bundle with protoxylem at lateral margin. Pinnules with radiate venation, arranged in four rows, with two rows of large, lanceolate pinnules on lower surface, two rows of small, oblong pinnules on upper surface of the axis. Pseudostrobilus heterosporous; sporophylls whorled, proximally diskshaped or dissected into segments, distally upturned and dissected. Sporangia attached adaxially and proximally to sporophyll in single ring. Megasporangia in vertical row bearing single functional megaspore. Microspores of the
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<emphasis box="[828,954,194,216]" italics="true" pageId="1" pageNumber="2">Calamospora</emphasis>
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<typeStatus box="[961,1005,194,215]" pageId="1" pageNumber="2">type</typeStatus>
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.
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</paragraph>
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</subSubSection>
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<subSubSection pageId="1" pageNumber="2" type="materials_examined">
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<paragraph blockId="1.[828,1458,141,787]" box="[828,1397,233,256]" pageId="1" pageNumber="2">
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<materialsCitation box="[828,1397,233,256]" collectionCode="PB" pageId="1" pageNumber="2" specimenCode="PB22126" specimenCount="1" typeStatus="holotype-">
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<typeStatus box="[828,946,233,256]" pageId="1" pageNumber="2">Holotype—</typeStatus>
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<specimenCode box="[946,1036,233,255]" pageId="1" pageNumber="2">PB22126</specimenCode>
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(
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<figureCitation box="[1051,1127,233,255]" captionStart="Fig" captionStartId="2.[102,130,1656,1673]" captionTargetBox="[174,1375,147,1630]" captionTargetId="figure-1@2.[172,1377,145,1632]" captionTargetPageId="2" captionText="Fig. 1. P. wuhaia J. Wang et al. sp. nov. from the early Permian Taiyuan Formation of Wuda Coalfield, Inner Mongolia. (A) Holotype with an entire crown consisting of pseudostrobili and leaves. (B) Once-pinnate compound leaf with both large and small pinnules visible. Reprinted with permission from ref. 15. (C) Cross-section of a crown illustrating pseudostrobili around the stem. (D) Cross-section of pseudostrobilus with microsporangia around the axis with bilateral, inversed Ω-shaped vascular bundle. (E) Cross-section of a leaf rachis showing the same form of vascular bundle as that of pseudostrobili axes. (F–H) Partial cross, radial, and tangential sections of the stem showing the secondary xylem (wood). (I) Tangential section of pseudostrobilus showing sporangial arrangement with single line of megasporangia along with the axis (middle-right pseudostrobilus of the fragmental crown illustrated SI Appendix, Fig. S8; specimen PB22132; magnified in SI Appendix, Fig. S21C.) (J) Radial section of pseudostrobilus showing adaxial sporangia and axis lacking nodes (upmost-left pseudostrobilus of the fragmental crown in SI Appendix, Fig. S8; magnified in SI Appendix, Fig. S21A). (K) Tangential section showing adaxial sporangia and a single line of megasporangia along with the axis (SI Appendix, Fig. S7; specimen PB22131, second up-right pseudostrobilus). (L) Tangential section through same specimen as K showing megasporangial arrangement (magnified in SI Appendix, Figs. S21B and S23A). (M) Detail of the middle part of L showing the megasporangia and microsporangia. (N) Single spore macerated from the holotype. (Scale bars: A, 10 cm; B, 3 cm; C–E, 1 cm; F, 100 μm; G and H, 200 μm; I–L, 5 mm; M, 2 mm; N, 10 μm.)" figureDoi="http://doi.org/10.5281/zenodo.4596608" httpUri="https://zenodo.org/record/4596608/files/figure.png" pageId="1" pageNumber="2">
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Fig. 1
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<emphasis box="[1111,1127,233,255]" italics="true" pageId="1" pageNumber="2">A</emphasis>
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</figureCitation>
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and
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<emphasis box="[1181,1303,233,255]" italics="true" pageId="1" pageNumber="2">SI Appendix</emphasis>
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,
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<figureCitation box="[1317,1388,233,255]" captionStart="Fig" captionStartId="3.[804,832,1876,1893]" captionTargetBox="[805,1457,953,1852]" captionTargetId="figure-907@3.[805,1457,953,1852]" captionTargetPageId="3" captionText="Fig. 2. Reconstruction of the aerial parts of P. wuhaia J. Wang et al. sp.nov. from the early Permian of China (based on the holotype). Image credit: Yugao Ren and Sijia Tang (artists)." figureDoi="http://doi.org/10.5281/zenodo.4596612" httpUri="https://zenodo.org/record/4596612/files/figure.png" pageId="1" pageNumber="2">Fig. S2</figureCitation>
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)
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</materialsCitation>
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</paragraph>
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<paragraph blockId="1.[828,1458,141,787]" box="[828,1399,273,296]" pageId="1" pageNumber="2">
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<materialsCitation box="[828,1394,273,296]" collectionCode="PB" pageId="1" pageNumber="2" specimenCode="PB22127-22138" specimenCount="1" typeStatus="paratypes-">
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<typeStatus box="[828,950,274,296]" pageId="1" pageNumber="2">Paratypes—</typeStatus>
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<specimenCode box="[950,1107,273,295]" pageId="1" pageNumber="2">PB22127-22138</specimenCode>
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(
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<emphasis box="[1124,1246,273,295]" italics="true" pageId="1" pageNumber="2">SI Appendix</emphasis>
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||
,
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<figureCitation box="[1260,1360,273,296]" captionStart="Fig" captionStartId="4.[102,130,1148,1165]" captionTargetBox="[348,1560,143,1123]" captionTargetId="figure-354@4.[346,1202,142,1124]" captionTargetPageId="4" captionText="Fig. 3. Diagrammatic reconstruction of the key features of P. wuhaia J. Wang et al. (A) Stem bears pseudostrobili and once-pinnate compound leaves; pseudostrobili are attached to the fertile zone, above which the leaves are shown from the abaxial side, and below which the leaves are shown from the adaxial side. (B) A pseudostrobilus shown its longitudinal surface. (C) Cross-section of a pseudostrobilus showing both megasporangia and microsporangia, the bilateral symmetrical Ω-shaped vascular bundle and sporophylls; noting the relative position of the presumed two megasporangia and the orientation of the opening of the vascular bundle. (D) Partial longitudinal section through the central axis of the pseudostrobilus and the symmetric axis of the vascular bundle, approximately corresponding to the dashed line in C; (E) partial of the leaf showing the arrangement of the small and big pinnules, seen from the abaxial (Upper) and adaxial (Lower) sides, respectively; (F) a pair of small and big pinnules showing the radiate venation; (G and H) cross-section of the stem showing the anatomy; H is the magnification of a part (within the dashed line) of the stem in G. CT, cortex; LT, leaf trace; MEG, megasporangium; MIG, microsporangium; PD, periderm; PT, pith; PX, primary xylem; SP, sporophyll; SX, secondary xylem; VB, vascular bundle." figureDoi="http://doi.org/10.5281/zenodo.4596614" httpUri="https://zenodo.org/record/4596614/files/figure.png" pageId="1" pageNumber="2">Figs S3‒S</figureCitation>
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14)
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</materialsCitation>
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.
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</paragraph>
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<paragraph blockId="1.[828,1458,141,787]" box="[828,1455,313,335]" pageId="1" pageNumber="2">
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<materialsCitation box="[828,1455,313,335]" collectionCode="Nanjing Institute of Geology and Paleontology, Chinese Academy of Sciences" country="China" location="Wuda Coalfield, Wuhai" pageId="1" pageNumber="2" specimenCount="1" stateProvince="Inner Mongolia" typeStatus="type">
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<typeStatus box="[828,877,314,335]" pageId="1" pageNumber="2">Type</typeStatus>
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locality—
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<location LSID="urn:lsid:plazi:treatment:987687A30E44FFFF87E3FA5F6D78FE50:1500606E0E44FFFD84B9FE946DB5FED1" box="[976,1212,313,335]" country="China" name="Wuda Coalfield, Wuhai" pageId="1" pageNumber="2" stateProvince="Inner Mongolia">Wuda Coalfield, Wuhai</location>
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,
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<collectingRegion box="[1223,1380,313,335]" country="China" name="Nei Mongol" pageId="1" pageNumber="2">Inner Mongolia</collectingRegion>
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,
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<collectingCountry box="[1390,1451,313,335]" name="China" pageId="1" pageNumber="2">China</collectingCountry>
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.
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</materialsCitation>
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</paragraph>
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<paragraph blockId="1.[828,1458,141,787]" box="[828,1318,353,375]" pageId="1" pageNumber="2">
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<geologicalTimeScale box="[828,1318,353,375]" pageId="1" pageNumber="2">Horizon—Uppermost part, Taiyuan Formation.</geologicalTimeScale>
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</paragraph>
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<paragraph blockId="1.[828,1458,141,787]" pageId="1" pageNumber="2">Stratigraphic age—298.34 ± 0.09 Ma, Asselian, Cisuralian, Permian.</paragraph>
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<paragraph blockId="1.[828,1458,141,787]" pageId="1" pageNumber="2">
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Repository—All specimens are deposited at
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<collectionCode pageId="1" pageNumber="2">Nanjing Institute of Geology and Paleontology, Chinese Academy of Sciences</collectionCode>
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.
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</paragraph>
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</subSubSection>
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<subSubSection lastPageId="3" lastPageNumber="4" pageId="1" pageNumber="2" type="discussion">
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<paragraph blockId="1.[828,1458,141,787]" pageId="1" pageNumber="2">
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Remarks—Following Wang et al. (
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<bibRefCitation author="S. J. Wang" box="[1191,1205,526,548]" journalOrPublisher="Am. J. Bot." pageId="1" pageNumber="2" pagination="127 - 149" part="104" refId="ref6909" refString="9. S. J. Wang et al., Anatomically preserved " strobili " and leaves from the Permian of China (Dorsalistachyaceae, fam. nov.) broaden knowledge of Noeggerathiales and constrain their possible taxonomic affinities. Am. J. Bot. 104, 127 - 149 (2017)." title="Anatomically preserved " strobili " and leaves from the Permian of China (Dorsalistachyaceae, fam. nov.) broaden knowledge of Noeggerathiales and constrain their possible taxonomic affinities" type="journal article" year="2017">9</bibRefCitation>
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), we consider the noeggerathialean fertile shoot to be a pseudostrobilus as despite its radial exterior appearance, it is distinctly bilateral in organization and represents a complex, three-dimensional sporophyll. We use quotation marks to denote that what would traditionally be interpreted as sporophylls on the pseudostrobilus axis represent pinnule homologs and are not in fact sporophylls in the traditional sense. Detailed descriptions, measurement, and further illustrations are provided in the supporting information.
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</paragraph>
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<paragraph blockId="1.[804,1458,845,1943]" pageId="1" pageNumber="2">
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<emphasis bold="true" box="[804,917,846,866]" pageId="1" pageNumber="2">Description.</emphasis>
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The
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<typeStatus box="[968,1056,845,867]" pageId="1" pageNumber="2">holotype</typeStatus>
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is
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<quantity box="[1084,1156,845,867]" metricMagnitude="0" metricUnit="m" metricValue="1.6" pageId="1" pageNumber="2" unit="cm" value="160.0">160 cm</quantity>
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long and
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<quantity box="[1256,1316,845,867]" metricMagnitude="-1" metricUnit="m" metricValue="6.0" pageId="1" pageNumber="2" unit="cm" value="60.0">60 cm</quantity>
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||
wide (
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<figureCitation box="[1382,1457,845,867]" captionStart="Fig" captionStartId="2.[102,130,1656,1673]" captionTargetBox="[174,1375,147,1630]" captionTargetId="figure-1@2.[172,1377,145,1632]" captionTargetPageId="2" captionText="Fig. 1. P. wuhaia J. Wang et al. sp. nov. from the early Permian Taiyuan Formation of Wuda Coalfield, Inner Mongolia. (A) Holotype with an entire crown consisting of pseudostrobili and leaves. (B) Once-pinnate compound leaf with both large and small pinnules visible. Reprinted with permission from ref. 15. (C) Cross-section of a crown illustrating pseudostrobili around the stem. (D) Cross-section of pseudostrobilus with microsporangia around the axis with bilateral, inversed Ω-shaped vascular bundle. (E) Cross-section of a leaf rachis showing the same form of vascular bundle as that of pseudostrobili axes. (F–H) Partial cross, radial, and tangential sections of the stem showing the secondary xylem (wood). (I) Tangential section of pseudostrobilus showing sporangial arrangement with single line of megasporangia along with the axis (middle-right pseudostrobilus of the fragmental crown illustrated SI Appendix, Fig. S8; specimen PB22132; magnified in SI Appendix, Fig. S21C.) (J) Radial section of pseudostrobilus showing adaxial sporangia and axis lacking nodes (upmost-left pseudostrobilus of the fragmental crown in SI Appendix, Fig. S8; magnified in SI Appendix, Fig. S21A). (K) Tangential section showing adaxial sporangia and a single line of megasporangia along with the axis (SI Appendix, Fig. S7; specimen PB22131, second up-right pseudostrobilus). (L) Tangential section through same specimen as K showing megasporangial arrangement (magnified in SI Appendix, Figs. S21B and S23A). (M) Detail of the middle part of L showing the megasporangia and microsporangia. (N) Single spore macerated from the holotype. (Scale bars: A, 10 cm; B, 3 cm; C–E, 1 cm; F, 100 μm; G and H, 200 μm; I–L, 5 mm; M, 2 mm; N, 10 μm.)" figureDoi="http://doi.org/10.5281/zenodo.4596608" httpUri="https://zenodo.org/record/4596608/files/figure.png" pageId="1" pageNumber="2">
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Fig. 1
|
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<emphasis box="[1441,1457,845,867]" italics="true" pageId="1" pageNumber="2">A</emphasis>
|
||
</figureCitation>
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||
and
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||
<emphasis box="[847,966,872,894]" italics="true" pageId="1" pageNumber="2">SI Appendix</emphasis>
|
||
,
|
||
<figureCitation box="[978,1046,872,894]" captionStart="Fig" captionStartId="3.[804,832,1876,1893]" captionTargetBox="[805,1457,953,1852]" captionTargetId="figure-907@3.[805,1457,953,1852]" captionTargetPageId="3" captionText="Fig. 2. Reconstruction of the aerial parts of P. wuhaia J. Wang et al. sp.nov. from the early Permian of China (based on the holotype). Image credit: Yugao Ren and Sijia Tang (artists)." figureDoi="http://doi.org/10.5281/zenodo.4596612" httpUri="https://zenodo.org/record/4596612/files/figure.png" pageId="1" pageNumber="2">Fig. S2</figureCitation>
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) and comprises a stem with apical crown of helically attached megaphyllous leaves and pseudostrobili. The crown has three parts: a 13-cm-long basal region bearing leaves, a middle 23-cm-long fertile zone, and apically a 6-cm-long region bearing more leaves. Below the crown, the stem is naked, extending for>
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<quantity box="[968,1034,1010,1032]" metricMagnitude="-1" metricUnit="m" metricValue="3.2" pageId="1" pageNumber="2" unit="cm" value="32.0">32 cm</quantity>
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, tapers little, and is ∼
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<quantity box="[1273,1325,1010,1032]" metricMagnitude="-2" metricUnit="m" metricValue="5.0" pageId="1" pageNumber="2" unit="cm" value="5.0">5 cm</quantity>
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wide with a smooth exterior, lacking evidence of abscission structures. More than 60 once-pinnate, megaphyllous leaves are attached to the stem above and below the fertile zone, with typical leaf rachises being 10.4 mm wide and <
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<quantity box="[1090,1154,1121,1143]" metricMagnitude="-1" metricUnit="m" metricValue="5.0" pageId="1" pageNumber="2" unit="cm" value="50.0">50 cm</quantity>
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long. Most pinnules are not fully discernible and are either matted together, obscured by other parts of the plant overlying them, or remain embedded within the rock. A
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<typeStatus box="[998,1086,1204,1225]" pageId="1" pageNumber="2">paratype</typeStatus>
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(
|
||
<figureCitation box="[1102,1177,1204,1226]" captionStart="Fig" captionStartId="2.[102,130,1656,1673]" captionTargetBox="[174,1375,147,1630]" captionTargetId="figure-1@2.[172,1377,145,1632]" captionTargetPageId="2" captionText="Fig. 1. P. wuhaia J. Wang et al. sp. nov. from the early Permian Taiyuan Formation of Wuda Coalfield, Inner Mongolia. (A) Holotype with an entire crown consisting of pseudostrobili and leaves. (B) Once-pinnate compound leaf with both large and small pinnules visible. Reprinted with permission from ref. 15. (C) Cross-section of a crown illustrating pseudostrobili around the stem. (D) Cross-section of pseudostrobilus with microsporangia around the axis with bilateral, inversed Ω-shaped vascular bundle. (E) Cross-section of a leaf rachis showing the same form of vascular bundle as that of pseudostrobili axes. (F–H) Partial cross, radial, and tangential sections of the stem showing the secondary xylem (wood). (I) Tangential section of pseudostrobilus showing sporangial arrangement with single line of megasporangia along with the axis (middle-right pseudostrobilus of the fragmental crown illustrated SI Appendix, Fig. S8; specimen PB22132; magnified in SI Appendix, Fig. S21C.) (J) Radial section of pseudostrobilus showing adaxial sporangia and axis lacking nodes (upmost-left pseudostrobilus of the fragmental crown in SI Appendix, Fig. S8; magnified in SI Appendix, Fig. S21A). (K) Tangential section showing adaxial sporangia and a single line of megasporangia along with the axis (SI Appendix, Fig. S7; specimen PB22131, second up-right pseudostrobilus). (L) Tangential section through same specimen as K showing megasporangial arrangement (magnified in SI Appendix, Figs. S21B and S23A). (M) Detail of the middle part of L showing the megasporangia and microsporangia. (N) Single spore macerated from the holotype. (Scale bars: A, 10 cm; B, 3 cm; C–E, 1 cm; F, 100 μm; G and H, 200 μm; I–L, 5 mm; M, 2 mm; N, 10 μm.)" figureDoi="http://doi.org/10.5281/zenodo.4596608" httpUri="https://zenodo.org/record/4596608/files/figure.png" pageId="1" pageNumber="2">
|
||
Fig. 1
|
||
<emphasis box="[1162,1177,1205,1226]" italics="true" pageId="1" pageNumber="2">B</emphasis>
|
||
</figureCitation>
|
||
and
|
||
<emphasis box="[1230,1351,1204,1226]" italics="true" pageId="1" pageNumber="2">SI Appendix</emphasis>
|
||
, Fig. S12) demonstrates clearly lanceolate, large pinnules on the lower surface of the leaf, and oval, small pinnules on the upper surface, both with radiate venation.
|
||
</paragraph>
|
||
<paragraph blockId="1.[804,1458,845,1943]" pageId="1" pageNumber="2">
|
||
Anatomically, the stem has an eustele with a large pith (
|
||
<figureCitation box="[1383,1457,1314,1336]" captionStart="Fig" captionStartId="2.[102,130,1656,1673]" captionTargetBox="[174,1375,147,1630]" captionTargetId="figure-1@2.[172,1377,145,1632]" captionTargetPageId="2" captionText="Fig. 1. P. wuhaia J. Wang et al. sp. nov. from the early Permian Taiyuan Formation of Wuda Coalfield, Inner Mongolia. (A) Holotype with an entire crown consisting of pseudostrobili and leaves. (B) Once-pinnate compound leaf with both large and small pinnules visible. Reprinted with permission from ref. 15. (C) Cross-section of a crown illustrating pseudostrobili around the stem. (D) Cross-section of pseudostrobilus with microsporangia around the axis with bilateral, inversed Ω-shaped vascular bundle. (E) Cross-section of a leaf rachis showing the same form of vascular bundle as that of pseudostrobili axes. (F–H) Partial cross, radial, and tangential sections of the stem showing the secondary xylem (wood). (I) Tangential section of pseudostrobilus showing sporangial arrangement with single line of megasporangia along with the axis (middle-right pseudostrobilus of the fragmental crown illustrated SI Appendix, Fig. S8; specimen PB22132; magnified in SI Appendix, Fig. S21C.) (J) Radial section of pseudostrobilus showing adaxial sporangia and axis lacking nodes (upmost-left pseudostrobilus of the fragmental crown in SI Appendix, Fig. S8; magnified in SI Appendix, Fig. S21A). (K) Tangential section showing adaxial sporangia and a single line of megasporangia along with the axis (SI Appendix, Fig. S7; specimen PB22131, second up-right pseudostrobilus). (L) Tangential section through same specimen as K showing megasporangial arrangement (magnified in SI Appendix, Figs. S21B and S23A). (M) Detail of the middle part of L showing the megasporangia and microsporangia. (N) Single spore macerated from the holotype. (Scale bars: A, 10 cm; B, 3 cm; C–E, 1 cm; F, 100 μm; G and H, 200 μm; I–L, 5 mm; M, 2 mm; N, 10 μm.)" figureDoi="http://doi.org/10.5281/zenodo.4596608" httpUri="https://zenodo.org/record/4596608/files/figure.png" pageId="1" pageNumber="2">
|
||
Fig. 1
|
||
<emphasis box="[1440,1457,1314,1336]" italics="true" pageId="1" pageNumber="2">C</emphasis>
|
||
</figureCitation>
|
||
and
|
||
<emphasis box="[853,975,1341,1363]" italics="true" pageId="1" pageNumber="2">SI Appendix</emphasis>
|
||
, Fig. S15); cauline primary xylem strands are adaxially curved (
|
||
<figureCitation box="[970,1039,1369,1391]" captionStart="Fig" captionStartId="2.[102,130,1656,1673]" captionTargetBox="[174,1375,147,1630]" captionTargetId="figure-1@2.[172,1377,145,1632]" captionTargetPageId="2" captionText="Fig. 1. P. wuhaia J. Wang et al. sp. nov. from the early Permian Taiyuan Formation of Wuda Coalfield, Inner Mongolia. (A) Holotype with an entire crown consisting of pseudostrobili and leaves. (B) Once-pinnate compound leaf with both large and small pinnules visible. Reprinted with permission from ref. 15. (C) Cross-section of a crown illustrating pseudostrobili around the stem. (D) Cross-section of pseudostrobilus with microsporangia around the axis with bilateral, inversed Ω-shaped vascular bundle. (E) Cross-section of a leaf rachis showing the same form of vascular bundle as that of pseudostrobili axes. (F–H) Partial cross, radial, and tangential sections of the stem showing the secondary xylem (wood). (I) Tangential section of pseudostrobilus showing sporangial arrangement with single line of megasporangia along with the axis (middle-right pseudostrobilus of the fragmental crown illustrated SI Appendix, Fig. S8; specimen PB22132; magnified in SI Appendix, Fig. S21C.) (J) Radial section of pseudostrobilus showing adaxial sporangia and axis lacking nodes (upmost-left pseudostrobilus of the fragmental crown in SI Appendix, Fig. S8; magnified in SI Appendix, Fig. S21A). (K) Tangential section showing adaxial sporangia and a single line of megasporangia along with the axis (SI Appendix, Fig. S7; specimen PB22131, second up-right pseudostrobilus). (L) Tangential section through same specimen as K showing megasporangial arrangement (magnified in SI Appendix, Figs. S21B and S23A). (M) Detail of the middle part of L showing the megasporangia and microsporangia. (N) Single spore macerated from the holotype. (Scale bars: A, 10 cm; B, 3 cm; C–E, 1 cm; F, 100 μm; G and H, 200 μm; I–L, 5 mm; M, 2 mm; N, 10 μm.)" figureDoi="http://doi.org/10.5281/zenodo.4596608" httpUri="https://zenodo.org/record/4596608/files/figure.png" pageId="1" pageNumber="2">
|
||
Fig. 1
|
||
<emphasis box="[1025,1039,1370,1391]" italics="true" pageId="1" pageNumber="2">F</emphasis>
|
||
</figureCitation>
|
||
and
|
||
<emphasis box="[1085,1200,1369,1391]" italics="true" pageId="1" pageNumber="2">SI Appendix</emphasis>
|
||
, Fig. S16
|
||
<emphasis box="[1293,1337,1369,1391]" italics="true" pageId="1" pageNumber="2">A–C</emphasis>
|
||
). Secondary xylem is pycnoxylic with uniseriate rays, which are mostly short/low and vary remarkably in size from one to seven cells high (
|
||
<figureCitation box="[1381,1457,1424,1446]" captionStart="Fig" captionStartId="2.[102,130,1656,1673]" captionTargetBox="[174,1375,147,1630]" captionTargetId="figure-1@2.[172,1377,145,1632]" captionTargetPageId="2" captionText="Fig. 1. P. wuhaia J. Wang et al. sp. nov. from the early Permian Taiyuan Formation of Wuda Coalfield, Inner Mongolia. (A) Holotype with an entire crown consisting of pseudostrobili and leaves. (B) Once-pinnate compound leaf with both large and small pinnules visible. Reprinted with permission from ref. 15. (C) Cross-section of a crown illustrating pseudostrobili around the stem. (D) Cross-section of pseudostrobilus with microsporangia around the axis with bilateral, inversed Ω-shaped vascular bundle. (E) Cross-section of a leaf rachis showing the same form of vascular bundle as that of pseudostrobili axes. (F–H) Partial cross, radial, and tangential sections of the stem showing the secondary xylem (wood). (I) Tangential section of pseudostrobilus showing sporangial arrangement with single line of megasporangia along with the axis (middle-right pseudostrobilus of the fragmental crown illustrated SI Appendix, Fig. S8; specimen PB22132; magnified in SI Appendix, Fig. S21C.) (J) Radial section of pseudostrobilus showing adaxial sporangia and axis lacking nodes (upmost-left pseudostrobilus of the fragmental crown in SI Appendix, Fig. S8; magnified in SI Appendix, Fig. S21A). (K) Tangential section showing adaxial sporangia and a single line of megasporangia along with the axis (SI Appendix, Fig. S7; specimen PB22131, second up-right pseudostrobilus). (L) Tangential section through same specimen as K showing megasporangial arrangement (magnified in SI Appendix, Figs. S21B and S23A). (M) Detail of the middle part of L showing the megasporangia and microsporangia. (N) Single spore macerated from the holotype. (Scale bars: A, 10 cm; B, 3 cm; C–E, 1 cm; F, 100 μm; G and H, 200 μm; I–L, 5 mm; M, 2 mm; N, 10 μm.)" figureDoi="http://doi.org/10.5281/zenodo.4596608" httpUri="https://zenodo.org/record/4596608/files/figure.png" pageId="1" pageNumber="2">
|
||
Fig. 1
|
||
<emphasis box="[1439,1457,1425,1446]" italics="true" pageId="1" pageNumber="2">H</emphasis>
|
||
</figureCitation>
|
||
and
|
||
<emphasis box="[847,962,1452,1474]" italics="true" pageId="1" pageNumber="2">SI Appendix</emphasis>
|
||
, Fig. S17
|
||
<emphasis box="[1060,1077,1452,1474]" italics="true" pageId="1" pageNumber="2">C</emphasis>
|
||
and
|
||
<emphasis box="[1126,1143,1453,1474]" italics="true" pageId="1" pageNumber="2">D</emphasis>
|
||
). Secondary tracheid walls have scalariform thickenings (
|
||
<figureCitation box="[1045,1122,1480,1502]" captionStart="Fig" captionStartId="2.[102,130,1656,1673]" captionTargetBox="[174,1375,147,1630]" captionTargetId="figure-1@2.[172,1377,145,1632]" captionTargetPageId="2" captionText="Fig. 1. P. wuhaia J. Wang et al. sp. nov. from the early Permian Taiyuan Formation of Wuda Coalfield, Inner Mongolia. (A) Holotype with an entire crown consisting of pseudostrobili and leaves. (B) Once-pinnate compound leaf with both large and small pinnules visible. Reprinted with permission from ref. 15. (C) Cross-section of a crown illustrating pseudostrobili around the stem. (D) Cross-section of pseudostrobilus with microsporangia around the axis with bilateral, inversed Ω-shaped vascular bundle. (E) Cross-section of a leaf rachis showing the same form of vascular bundle as that of pseudostrobili axes. (F–H) Partial cross, radial, and tangential sections of the stem showing the secondary xylem (wood). (I) Tangential section of pseudostrobilus showing sporangial arrangement with single line of megasporangia along with the axis (middle-right pseudostrobilus of the fragmental crown illustrated SI Appendix, Fig. S8; specimen PB22132; magnified in SI Appendix, Fig. S21C.) (J) Radial section of pseudostrobilus showing adaxial sporangia and axis lacking nodes (upmost-left pseudostrobilus of the fragmental crown in SI Appendix, Fig. S8; magnified in SI Appendix, Fig. S21A). (K) Tangential section showing adaxial sporangia and a single line of megasporangia along with the axis (SI Appendix, Fig. S7; specimen PB22131, second up-right pseudostrobilus). (L) Tangential section through same specimen as K showing megasporangial arrangement (magnified in SI Appendix, Figs. S21B and S23A). (M) Detail of the middle part of L showing the megasporangia and microsporangia. (N) Single spore macerated from the holotype. (Scale bars: A, 10 cm; B, 3 cm; C–E, 1 cm; F, 100 μm; G and H, 200 μm; I–L, 5 mm; M, 2 mm; N, 10 μm.)" figureDoi="http://doi.org/10.5281/zenodo.4596608" httpUri="https://zenodo.org/record/4596608/files/figure.png" pageId="1" pageNumber="2">
|
||
Fig. 1
|
||
<emphasis box="[1103,1122,1480,1502]" italics="true" pageId="1" pageNumber="2">G</emphasis>
|
||
</figureCitation>
|
||
and
|
||
<emphasis box="[1175,1293,1480,1502]" italics="true" pageId="1" pageNumber="2">SI Appendix</emphasis>
|
||
, Fig. S17
|
||
<emphasis box="[1396,1412,1480,1502]" italics="true" pageId="1" pageNumber="2">A</emphasis>
|
||
and
|
||
<emphasis box="[804,819,1508,1529]" italics="true" pageId="1" pageNumber="2">B</emphasis>
|
||
), and the cortex has collenchyma in its outer part. Leaf traces are tangentially elongated.
|
||
</paragraph>
|
||
<paragraph blockId="1.[804,1458,845,1943]" pageId="1" pageNumber="2">
|
||
Approximately 8 to 10 pseudostrobili are attached to the circumference of the fertile zone and about 20 more longitudinally attached along the stem (
|
||
<figureCitation box="[1067,1143,1617,1639]" captionStart="Fig" captionStartId="2.[102,130,1656,1673]" captionTargetBox="[174,1375,147,1630]" captionTargetId="figure-1@2.[172,1377,145,1632]" captionTargetPageId="2" captionText="Fig. 1. P. wuhaia J. Wang et al. sp. nov. from the early Permian Taiyuan Formation of Wuda Coalfield, Inner Mongolia. (A) Holotype with an entire crown consisting of pseudostrobili and leaves. (B) Once-pinnate compound leaf with both large and small pinnules visible. Reprinted with permission from ref. 15. (C) Cross-section of a crown illustrating pseudostrobili around the stem. (D) Cross-section of pseudostrobilus with microsporangia around the axis with bilateral, inversed Ω-shaped vascular bundle. (E) Cross-section of a leaf rachis showing the same form of vascular bundle as that of pseudostrobili axes. (F–H) Partial cross, radial, and tangential sections of the stem showing the secondary xylem (wood). (I) Tangential section of pseudostrobilus showing sporangial arrangement with single line of megasporangia along with the axis (middle-right pseudostrobilus of the fragmental crown illustrated SI Appendix, Fig. S8; specimen PB22132; magnified in SI Appendix, Fig. S21C.) (J) Radial section of pseudostrobilus showing adaxial sporangia and axis lacking nodes (upmost-left pseudostrobilus of the fragmental crown in SI Appendix, Fig. S8; magnified in SI Appendix, Fig. S21A). (K) Tangential section showing adaxial sporangia and a single line of megasporangia along with the axis (SI Appendix, Fig. S7; specimen PB22131, second up-right pseudostrobilus). (L) Tangential section through same specimen as K showing megasporangial arrangement (magnified in SI Appendix, Figs. S21B and S23A). (M) Detail of the middle part of L showing the megasporangia and microsporangia. (N) Single spore macerated from the holotype. (Scale bars: A, 10 cm; B, 3 cm; C–E, 1 cm; F, 100 μm; G and H, 200 μm; I–L, 5 mm; M, 2 mm; N, 10 μm.)" figureDoi="http://doi.org/10.5281/zenodo.4596608" httpUri="https://zenodo.org/record/4596608/files/figure.png" pageId="1" pageNumber="2">
|
||
Fig. 1
|
||
<emphasis box="[1127,1143,1617,1639]" italics="true" pageId="1" pageNumber="2">A</emphasis>
|
||
</figureCitation>
|
||
and
|
||
<emphasis box="[1198,1320,1617,1639]" italics="true" pageId="1" pageNumber="2">SI Appendix</emphasis>
|
||
,
|
||
<figureCitation box="[1334,1405,1617,1639]" captionStart="Fig" captionStartId="3.[804,832,1876,1893]" captionTargetBox="[805,1457,953,1852]" captionTargetId="figure-907@3.[805,1457,953,1852]" captionTargetPageId="3" captionText="Fig. 2. Reconstruction of the aerial parts of P. wuhaia J. Wang et al. sp.nov. from the early Permian of China (based on the holotype). Image credit: Yugao Ren and Sijia Tang (artists)." figureDoi="http://doi.org/10.5281/zenodo.4596612" httpUri="https://zenodo.org/record/4596612/files/figure.png" pageId="1" pageNumber="2">Fig. S2</figureCitation>
|
||
); we estimate an entire fertile zone would comprise 160 to 200 pseudostrobili. Pseudostrobili are elongated with parallel sides and bluntly rounded apices, and are attached more or less at right angles to the stem with one-third to one-quarter of their width without a stalk. They are
|
||
<quantity box="[1140,1276,1755,1777]" metricMagnitude="-2" metricUnit="m" metricValue="5.75" metricValueMax="6.5" metricValueMin="5.0" pageId="1" pageNumber="2" unit="mm" value="57.5" valueMax="65.0" valueMin="50.0">50 to 65 mm</quantity>
|
||
long and
|
||
<quantity box="[1385,1457,1755,1777]" metricMagnitude="-2" metricUnit="m" metricValue="1.3" pageId="1" pageNumber="2" unit="mm" value="13.0">13 mm</quantity>
|
||
wide (
|
||
<figureCitation box="[866,942,1783,1805]" captionStart="Fig" captionStartId="2.[102,130,1656,1673]" captionTargetBox="[174,1375,147,1630]" captionTargetId="figure-1@2.[172,1377,145,1632]" captionTargetPageId="2" captionText="Fig. 1. P. wuhaia J. Wang et al. sp. nov. from the early Permian Taiyuan Formation of Wuda Coalfield, Inner Mongolia. (A) Holotype with an entire crown consisting of pseudostrobili and leaves. (B) Once-pinnate compound leaf with both large and small pinnules visible. Reprinted with permission from ref. 15. (C) Cross-section of a crown illustrating pseudostrobili around the stem. (D) Cross-section of pseudostrobilus with microsporangia around the axis with bilateral, inversed Ω-shaped vascular bundle. (E) Cross-section of a leaf rachis showing the same form of vascular bundle as that of pseudostrobili axes. (F–H) Partial cross, radial, and tangential sections of the stem showing the secondary xylem (wood). (I) Tangential section of pseudostrobilus showing sporangial arrangement with single line of megasporangia along with the axis (middle-right pseudostrobilus of the fragmental crown illustrated SI Appendix, Fig. S8; specimen PB22132; magnified in SI Appendix, Fig. S21C.) (J) Radial section of pseudostrobilus showing adaxial sporangia and axis lacking nodes (upmost-left pseudostrobilus of the fragmental crown in SI Appendix, Fig. S8; magnified in SI Appendix, Fig. S21A). (K) Tangential section showing adaxial sporangia and a single line of megasporangia along with the axis (SI Appendix, Fig. S7; specimen PB22131, second up-right pseudostrobilus). (L) Tangential section through same specimen as K showing megasporangial arrangement (magnified in SI Appendix, Figs. S21B and S23A). (M) Detail of the middle part of L showing the megasporangia and microsporangia. (N) Single spore macerated from the holotype. (Scale bars: A, 10 cm; B, 3 cm; C–E, 1 cm; F, 100 μm; G and H, 200 μm; I–L, 5 mm; M, 2 mm; N, 10 μm.)" figureDoi="http://doi.org/10.5281/zenodo.4596608" httpUri="https://zenodo.org/record/4596608/files/figure.png" pageId="1" pageNumber="2">
|
||
Fig. 1
|
||
<emphasis box="[926,942,1783,1805]" italics="true" pageId="1" pageNumber="2">A</emphasis>
|
||
</figureCitation>
|
||
and
|
||
<emphasis box="[996,1117,1783,1805]" italics="true" pageId="1" pageNumber="2">SI Appendix</emphasis>
|
||
,
|
||
<figureCitation box="[1131,1237,1783,1805]" captionStart="Fig" captionStartId="4.[102,130,1148,1165]" captionTargetBox="[348,1560,143,1123]" captionTargetId="figure-354@4.[346,1202,142,1124]" captionTargetPageId="4" captionText="Fig. 3. Diagrammatic reconstruction of the key features of P. wuhaia J. Wang et al. (A) Stem bears pseudostrobili and once-pinnate compound leaves; pseudostrobili are attached to the fertile zone, above which the leaves are shown from the abaxial side, and below which the leaves are shown from the adaxial side. (B) A pseudostrobilus shown its longitudinal surface. (C) Cross-section of a pseudostrobilus showing both megasporangia and microsporangia, the bilateral symmetrical Ω-shaped vascular bundle and sporophylls; noting the relative position of the presumed two megasporangia and the orientation of the opening of the vascular bundle. (D) Partial longitudinal section through the central axis of the pseudostrobilus and the symmetric axis of the vascular bundle, approximately corresponding to the dashed line in C; (E) partial of the leaf showing the arrangement of the small and big pinnules, seen from the abaxial (Upper) and adaxial (Lower) sides, respectively; (F) a pair of small and big pinnules showing the radiate venation; (G and H) cross-section of the stem showing the anatomy; H is the magnification of a part (within the dashed line) of the stem in G. CT, cortex; LT, leaf trace; MEG, megasporangium; MIG, microsporangium; PD, periderm; PT, pith; PX, primary xylem; SP, sporophyll; SX, secondary xylem; VB, vascular bundle." figureDoi="http://doi.org/10.5281/zenodo.4596614" httpUri="https://zenodo.org/record/4596614/files/figure.png" pageId="1" pageNumber="2">Figs. S3–S</figureCitation>
|
||
8).
|
||
</paragraph>
|
||
<paragraph blockId="1.[804,1458,845,1943]" lastBlockId="3.[114,769,141,1199]" lastPageId="3" lastPageNumber="4" pageId="1" pageNumber="2">
|
||
Both the leaf (
|
||
<figureCitation box="[980,1056,1810,1832]" captionStart="Fig" captionStartId="2.[102,130,1656,1673]" captionTargetBox="[174,1375,147,1630]" captionTargetId="figure-1@2.[172,1377,145,1632]" captionTargetPageId="2" captionText="Fig. 1. P. wuhaia J. Wang et al. sp. nov. from the early Permian Taiyuan Formation of Wuda Coalfield, Inner Mongolia. (A) Holotype with an entire crown consisting of pseudostrobili and leaves. (B) Once-pinnate compound leaf with both large and small pinnules visible. Reprinted with permission from ref. 15. (C) Cross-section of a crown illustrating pseudostrobili around the stem. (D) Cross-section of pseudostrobilus with microsporangia around the axis with bilateral, inversed Ω-shaped vascular bundle. (E) Cross-section of a leaf rachis showing the same form of vascular bundle as that of pseudostrobili axes. (F–H) Partial cross, radial, and tangential sections of the stem showing the secondary xylem (wood). (I) Tangential section of pseudostrobilus showing sporangial arrangement with single line of megasporangia along with the axis (middle-right pseudostrobilus of the fragmental crown illustrated SI Appendix, Fig. S8; specimen PB22132; magnified in SI Appendix, Fig. S21C.) (J) Radial section of pseudostrobilus showing adaxial sporangia and axis lacking nodes (upmost-left pseudostrobilus of the fragmental crown in SI Appendix, Fig. S8; magnified in SI Appendix, Fig. S21A). (K) Tangential section showing adaxial sporangia and a single line of megasporangia along with the axis (SI Appendix, Fig. S7; specimen PB22131, second up-right pseudostrobilus). (L) Tangential section through same specimen as K showing megasporangial arrangement (magnified in SI Appendix, Figs. S21B and S23A). (M) Detail of the middle part of L showing the megasporangia and microsporangia. (N) Single spore macerated from the holotype. (Scale bars: A, 10 cm; B, 3 cm; C–E, 1 cm; F, 100 μm; G and H, 200 μm; I–L, 5 mm; M, 2 mm; N, 10 μm.)" figureDoi="http://doi.org/10.5281/zenodo.4596608" httpUri="https://zenodo.org/record/4596608/files/figure.png" pageId="1" pageNumber="2">
|
||
Fig. 1
|
||
<emphasis box="[1041,1056,1811,1832]" italics="true" pageId="1" pageNumber="2">B</emphasis>
|
||
</figureCitation>
|
||
and
|
||
<emphasis box="[1111,1233,1810,1832]" italics="true" pageId="1" pageNumber="2">SI Appendix</emphasis>
|
||
, Figs. S15 and S19
|
||
<emphasis box="[1441,1457,1810,1832]" italics="true" pageId="1" pageNumber="2">A</emphasis>
|
||
and
|
||
<emphasis box="[857,872,1839,1860]" italics="true" pageId="1" pageNumber="2">B</emphasis>
|
||
) and pseudostrobili (
|
||
<figureCitation box="[1115,1200,1838,1860]" captionStart="Fig" captionStartId="2.[102,130,1656,1673]" captionTargetBox="[174,1375,147,1630]" captionTargetId="figure-1@2.[172,1377,145,1632]" captionTargetPageId="2" captionText="Fig. 1. P. wuhaia J. Wang et al. sp. nov. from the early Permian Taiyuan Formation of Wuda Coalfield, Inner Mongolia. (A) Holotype with an entire crown consisting of pseudostrobili and leaves. (B) Once-pinnate compound leaf with both large and small pinnules visible. Reprinted with permission from ref. 15. (C) Cross-section of a crown illustrating pseudostrobili around the stem. (D) Cross-section of pseudostrobilus with microsporangia around the axis with bilateral, inversed Ω-shaped vascular bundle. (E) Cross-section of a leaf rachis showing the same form of vascular bundle as that of pseudostrobili axes. (F–H) Partial cross, radial, and tangential sections of the stem showing the secondary xylem (wood). (I) Tangential section of pseudostrobilus showing sporangial arrangement with single line of megasporangia along with the axis (middle-right pseudostrobilus of the fragmental crown illustrated SI Appendix, Fig. S8; specimen PB22132; magnified in SI Appendix, Fig. S21C.) (J) Radial section of pseudostrobilus showing adaxial sporangia and axis lacking nodes (upmost-left pseudostrobilus of the fragmental crown in SI Appendix, Fig. S8; magnified in SI Appendix, Fig. S21A). (K) Tangential section showing adaxial sporangia and a single line of megasporangia along with the axis (SI Appendix, Fig. S7; specimen PB22131, second up-right pseudostrobilus). (L) Tangential section through same specimen as K showing megasporangial arrangement (magnified in SI Appendix, Figs. S21B and S23A). (M) Detail of the middle part of L showing the megasporangia and microsporangia. (N) Single spore macerated from the holotype. (Scale bars: A, 10 cm; B, 3 cm; C–E, 1 cm; F, 100 μm; G and H, 200 μm; I–L, 5 mm; M, 2 mm; N, 10 μm.)" figureDoi="http://doi.org/10.5281/zenodo.4596608" httpUri="https://zenodo.org/record/4596608/files/figure.png" pageId="1" pageNumber="2">
|
||
Fig. 1
|
||
<emphasis box="[1182,1200,1839,1860]" italics="true" pageId="1" pageNumber="2">D</emphasis>
|
||
</figureCitation>
|
||
and
|
||
<emphasis box="[1268,1397,1838,1860]" italics="true" pageId="1" pageNumber="2">SI Appendix</emphasis>
|
||
, Fig. S22
|
||
<emphasis box="[851,896,1865,1888]" italics="true" pageId="1" pageNumber="2">A–C</emphasis>
|
||
) rachises have a reversed Ω- shaped vascular bundle. Protoxylem strands occur at the lateral margins of the bundle, while pinnule traces and sporophyll (i.e., pinnule homolog) traces are given off from the lateral ends of the vascular bundles of the rachis and “strobilus” axis, respectively (
|
||
<emphasis box="[588,708,168,190]" italics="true" pageId="3" pageNumber="4">SI Appendix</emphasis>
|
||
, Figs. S19
|
||
<emphasis box="[151,167,196,217]" italics="true" pageId="3" pageNumber="4">E</emphasis>
|
||
, S20
|
||
<emphasis box="[226,272,195,217]" italics="true" pageId="3" pageNumber="4">A‒C</emphasis>
|
||
, and S22
|
||
<emphasis box="[368,385,195,217]" italics="true" pageId="3" pageNumber="4">C</emphasis>
|
||
).
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.4596608" ID-Zenodo-Dep="4596608" httpUri="https://zenodo.org/record/4596608/files/figure.png" pageId="2" pageNumber="3" startId="2.[102,130,1656,1673]" targetBox="[174,1375,147,1630]" targetPageId="2">
|
||
<paragraph blockId="2.[102,1447,1656,1936]" pageId="2" pageNumber="3">
|
||
<emphasis bold="true" box="[102,154,1656,1673]" pageId="2" pageNumber="3">Fig. 1.</emphasis>
|
||
<taxonomicName authority="J. Wang et al." authorityName="J. Wang" authorityYear="2021" box="[173,383,1656,1673]" genus="Paratingia" pageId="2" pageNumber="3" rank="species" species="wuhaia" status="sp. nov.">
|
||
<emphasis box="[173,183,1656,1673]" italics="true" pageId="2" pageNumber="3">P</emphasis>
|
||
.
|
||
<emphasis box="[194,258,1656,1673]" italics="true" pageId="2" pageNumber="3">wuhaia</emphasis>
|
||
J. Wang et al.
|
||
</taxonomicName>
|
||
<taxonomicNameLabel box="[387,455,1656,1673]" pageId="2" pageNumber="3" rank="species">sp. nov.</taxonomicNameLabel>
|
||
from the early Permian Taiyuan Formation of Wuda Coalfield, Inner Mongolia. (
|
||
<emphasis box="[1154,1167,1656,1673]" italics="true" pageId="2" pageNumber="3">A</emphasis>
|
||
) Holotype with an entire crown consisting of pseudostrobili and leaves. (
|
||
<emphasis box="[452,463,1680,1697]" italics="true" pageId="2" pageNumber="3">B</emphasis>
|
||
) Once-pinnate compound leaf with both large and small pinnules visible. Reprinted with permission from ref. 15. (
|
||
<emphasis box="[108,119,1704,1721]" italics="true" pageId="2" pageNumber="3">C</emphasis>
|
||
) Cross-section of a crown illustrating pseudostrobili around the stem. (
|
||
<emphasis box="[742,755,1704,1721]" italics="true" pageId="2" pageNumber="3">D</emphasis>
|
||
) Cross-section of pseudostrobilus with microsporangia around the axis with bilateral, inversed Ω- shaped vascular bundle. (
|
||
<emphasis box="[485,495,1728,1745]" italics="true" pageId="2" pageNumber="3">E</emphasis>
|
||
) Cross-section of a leaf rachis showing the same form of vascular bundle as that of pseudostrobili axes. (
|
||
<emphasis box="[1407,1439,1728,1745]" italics="true" pageId="2" pageNumber="3">F–H</emphasis>
|
||
) Partial cross, radial, and tangential sections of the stem showing the secondary xylem (wood). (
|
||
<emphasis box="[930,935,1751,1768]" italics="true" pageId="2" pageNumber="3">I</emphasis>
|
||
) Tangential section of pseudostrobilus showing sporangial arrangement with single line of megasporangia along with the axis (middle-right pseudostrobilus of the fragmental crown illustrated
|
||
<emphasis box="[1270,1373,1775,1792]" italics="true" pageId="2" pageNumber="3">SI Appendix</emphasis>
|
||
, Fig. S8; specimen PB22132; magnified in
|
||
<emphasis box="[386,489,1799,1816]" italics="true" pageId="2" pageNumber="3">SI Appendix</emphasis>
|
||
, Fig. S21
|
||
<emphasis box="[565,576,1799,1816]" italics="true" pageId="2" pageNumber="3">C</emphasis>
|
||
.) (
|
||
<emphasis box="[599,606,1799,1816]" italics="true" pageId="2" pageNumber="3">J</emphasis>
|
||
) Radial section of pseudostrobilus showing adaxial sporangia and axis lacking nodes (upmost-left pseudostrobilus of the fragmental crown in
|
||
<emphasis box="[478,580,1823,1840]" italics="true" pageId="2" pageNumber="3">SI Appendix</emphasis>
|
||
, Fig. S8; magnified in
|
||
<emphasis box="[767,869,1823,1840]" italics="true" pageId="2" pageNumber="3">SI Appendix</emphasis>
|
||
, Fig. S21
|
||
<emphasis box="[944,957,1823,1840]" italics="true" pageId="2" pageNumber="3">A</emphasis>
|
||
). (
|
||
<emphasis box="[979,991,1823,1840]" italics="true" pageId="2" pageNumber="3">K</emphasis>
|
||
) Tangential section showing adaxial sporangia and a single line of megasporangia along with the axis (
|
||
<emphasis box="[538,641,1847,1864]" italics="true" pageId="2" pageNumber="3">SI Appendix</emphasis>
|
||
, Fig. S7; specimen PB22131, second up-right pseudostrobilus). (
|
||
<emphasis box="[1189,1198,1847,1864]" italics="true" pageId="2" pageNumber="3">L</emphasis>
|
||
) Tangential section through same specimen as
|
||
<emphasis box="[260,272,1871,1888]" italics="true" pageId="2" pageNumber="3">K</emphasis>
|
||
showing megasporangial arrangement (magnified in
|
||
<emphasis box="[740,843,1871,1888]" italics="true" pageId="2" pageNumber="3">SI Appendix</emphasis>
|
||
, Figs. S21
|
||
<emphasis box="[927,938,1871,1888]" italics="true" pageId="2" pageNumber="3">B</emphasis>
|
||
and S23
|
||
<emphasis box="[1012,1025,1871,1888]" italics="true" pageId="2" pageNumber="3">A</emphasis>
|
||
). (
|
||
<emphasis box="[1048,1065,1871,1888]" italics="true" pageId="2" pageNumber="3">M</emphasis>
|
||
) Detail of the middle part of
|
||
<emphasis box="[1324,1333,1871,1888]" italics="true" pageId="2" pageNumber="3">L</emphasis>
|
||
showing the megasporangia and microsporangia. (
|
||
<emphasis box="[428,441,1895,1912]" italics="true" pageId="2" pageNumber="3">N</emphasis>
|
||
) Single spore macerated from the holotype. (Scale bars:
|
||
<emphasis box="[927,940,1895,1912]" italics="true" pageId="2" pageNumber="3">A</emphasis>
|
||
, 10 cm;
|
||
<emphasis box="[1011,1022,1895,1912]" italics="true" pageId="2" pageNumber="3">B</emphasis>
|
||
, 3 cm;
|
||
<emphasis box="[1082,1113,1895,1912]" italics="true" pageId="2" pageNumber="3">C–E</emphasis>
|
||
, 1 cm;
|
||
<emphasis box="[1173,1182,1895,1912]" italics="true" pageId="2" pageNumber="3">F</emphasis>
|
||
, 100 μm;
|
||
<emphasis box="[1265,1278,1895,1912]" italics="true" pageId="2" pageNumber="3">G</emphasis>
|
||
and
|
||
<emphasis box="[1321,1334,1895,1912]" italics="true" pageId="2" pageNumber="3">H</emphasis>
|
||
, 200 μm;
|
||
<emphasis box="[1417,1440,1895,1912]" italics="true" pageId="2" pageNumber="3">I–L</emphasis>
|
||
, 5 mm;
|
||
<emphasis box="[163,180,1919,1936]" italics="true" pageId="2" pageNumber="3">M</emphasis>
|
||
, 2 mm;
|
||
<emphasis box="[252,265,1919,1936]" italics="true" pageId="2" pageNumber="3">N</emphasis>
|
||
, 10 μm.)
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="3.[114,769,141,1199]" pageId="3" pageNumber="4">
|
||
Many pseudostrobili show four to five rows of sporangia with rows arranged parallel to the rachis (
|
||
<figureCitation box="[499,647,250,272]" captionStart="Fig" captionStartId="2.[102,130,1656,1673]" captionTargetBox="[174,1375,147,1630]" captionTargetId="figure-1@2.[172,1377,145,1632]" captionTargetPageId="2" captionText="Fig. 1. P. wuhaia J. Wang et al. sp. nov. from the early Permian Taiyuan Formation of Wuda Coalfield, Inner Mongolia. (A) Holotype with an entire crown consisting of pseudostrobili and leaves. (B) Once-pinnate compound leaf with both large and small pinnules visible. Reprinted with permission from ref. 15. (C) Cross-section of a crown illustrating pseudostrobili around the stem. (D) Cross-section of pseudostrobilus with microsporangia around the axis with bilateral, inversed Ω-shaped vascular bundle. (E) Cross-section of a leaf rachis showing the same form of vascular bundle as that of pseudostrobili axes. (F–H) Partial cross, radial, and tangential sections of the stem showing the secondary xylem (wood). (I) Tangential section of pseudostrobilus showing sporangial arrangement with single line of megasporangia along with the axis (middle-right pseudostrobilus of the fragmental crown illustrated SI Appendix, Fig. S8; specimen PB22132; magnified in SI Appendix, Fig. S21C.) (J) Radial section of pseudostrobilus showing adaxial sporangia and axis lacking nodes (upmost-left pseudostrobilus of the fragmental crown in SI Appendix, Fig. S8; magnified in SI Appendix, Fig. S21A). (K) Tangential section showing adaxial sporangia and a single line of megasporangia along with the axis (SI Appendix, Fig. S7; specimen PB22131, second up-right pseudostrobilus). (L) Tangential section through same specimen as K showing megasporangial arrangement (magnified in SI Appendix, Figs. S21B and S23A). (M) Detail of the middle part of L showing the megasporangia and microsporangia. (N) Single spore macerated from the holotype. (Scale bars: A, 10 cm; B, 3 cm; C–E, 1 cm; F, 100 μm; G and H, 200 μm; I–L, 5 mm; M, 2 mm; N, 10 μm.)" figureDoi="http://doi.org/10.5281/zenodo.4596608" httpUri="https://zenodo.org/record/4596608/files/figure.png" pageId="3" pageNumber="4">
|
||
Fig. 1
|
||
<emphasis box="[568,577,250,271]" italics="true" pageId="3" pageNumber="4">I</emphasis>
|
||
and
|
||
<emphasis box="[631,647,250,271]" italics="true" pageId="3" pageNumber="4">K</emphasis>
|
||
</figureCitation>
|
||
and
|
||
<emphasis italics="true" pageId="3" pageNumber="4">SI Appendix</emphasis>
|
||
, Figs. S5‒S8). As the specimens have been affected by postmortem compression (
|
||
<emphasis box="[395,520,304,326]" italics="true" pageId="3" pageNumber="4">SI Appendix</emphasis>
|
||
, Figs. S5‒S8), we consider that there must have been a row of sporangia on either side that was laterally flattened; we estimate that a complete pseudostrobilus comprised 8 or 10 sporangia per whorl. Sporophylls are proximally fused laterally to adjacent sporophylls, while distally they are free with a small heel and upturned apex (
|
||
<emphasis italics="true" pageId="3" pageNumber="4">SI Appendix</emphasis>
|
||
, Figs. S21
|
||
<emphasis box="[314,332,469,490]" italics="true" pageId="3" pageNumber="4">D</emphasis>
|
||
and S23
|
||
<emphasis box="[424,440,469,490]" italics="true" pageId="3" pageNumber="4">E</emphasis>
|
||
). There are 32 to 35 sporangia per row in the pseudostrobili depending on the length of the pseudostrobili, producing five sporangia per centimeter’ s length. Sporangia are sessile, attached to the sporophylls adaxial surface (
|
||
<figureCitation box="[122,228,577,599]" captionStart="Fig" captionStartId="2.[102,130,1656,1673]" captionTargetBox="[174,1375,147,1630]" captionTargetId="figure-1@2.[172,1377,145,1632]" captionTargetPageId="2" captionText="Fig. 1. P. wuhaia J. Wang et al. sp. nov. from the early Permian Taiyuan Formation of Wuda Coalfield, Inner Mongolia. (A) Holotype with an entire crown consisting of pseudostrobili and leaves. (B) Once-pinnate compound leaf with both large and small pinnules visible. Reprinted with permission from ref. 15. (C) Cross-section of a crown illustrating pseudostrobili around the stem. (D) Cross-section of pseudostrobilus with microsporangia around the axis with bilateral, inversed Ω-shaped vascular bundle. (E) Cross-section of a leaf rachis showing the same form of vascular bundle as that of pseudostrobili axes. (F–H) Partial cross, radial, and tangential sections of the stem showing the secondary xylem (wood). (I) Tangential section of pseudostrobilus showing sporangial arrangement with single line of megasporangia along with the axis (middle-right pseudostrobilus of the fragmental crown illustrated SI Appendix, Fig. S8; specimen PB22132; magnified in SI Appendix, Fig. S21C.) (J) Radial section of pseudostrobilus showing adaxial sporangia and axis lacking nodes (upmost-left pseudostrobilus of the fragmental crown in SI Appendix, Fig. S8; magnified in SI Appendix, Fig. S21A). (K) Tangential section showing adaxial sporangia and a single line of megasporangia along with the axis (SI Appendix, Fig. S7; specimen PB22131, second up-right pseudostrobilus). (L) Tangential section through same specimen as K showing megasporangial arrangement (magnified in SI Appendix, Figs. S21B and S23A). (M) Detail of the middle part of L showing the megasporangia and microsporangia. (N) Single spore macerated from the holotype. (Scale bars: A, 10 cm; B, 3 cm; C–E, 1 cm; F, 100 μm; G and H, 200 μm; I–L, 5 mm; M, 2 mm; N, 10 μm.)" figureDoi="http://doi.org/10.5281/zenodo.4596608" httpUri="https://zenodo.org/record/4596608/files/figure.png" pageId="3" pageNumber="4">
|
||
Fig. 1
|
||
<emphasis box="[186,228,577,599]" italics="true" pageId="3" pageNumber="4">I–M</emphasis>
|
||
</figureCitation>
|
||
). Megasporangia occur in a single vertical row on the pseudostrobilus axis (
|
||
<figureCitation box="[336,455,604,627]" captionStart="Fig" captionStartId="2.[102,130,1656,1673]" captionTargetBox="[174,1375,147,1630]" captionTargetId="figure-1@2.[172,1377,145,1632]" captionTargetPageId="2" captionText="Fig. 1. P. wuhaia J. Wang et al. sp. nov. from the early Permian Taiyuan Formation of Wuda Coalfield, Inner Mongolia. (A) Holotype with an entire crown consisting of pseudostrobili and leaves. (B) Once-pinnate compound leaf with both large and small pinnules visible. Reprinted with permission from ref. 15. (C) Cross-section of a crown illustrating pseudostrobili around the stem. (D) Cross-section of pseudostrobilus with microsporangia around the axis with bilateral, inversed Ω-shaped vascular bundle. (E) Cross-section of a leaf rachis showing the same form of vascular bundle as that of pseudostrobili axes. (F–H) Partial cross, radial, and tangential sections of the stem showing the secondary xylem (wood). (I) Tangential section of pseudostrobilus showing sporangial arrangement with single line of megasporangia along with the axis (middle-right pseudostrobilus of the fragmental crown illustrated SI Appendix, Fig. S8; specimen PB22132; magnified in SI Appendix, Fig. S21C.) (J) Radial section of pseudostrobilus showing adaxial sporangia and axis lacking nodes (upmost-left pseudostrobilus of the fragmental crown in SI Appendix, Fig. S8; magnified in SI Appendix, Fig. S21A). (K) Tangential section showing adaxial sporangia and a single line of megasporangia along with the axis (SI Appendix, Fig. S7; specimen PB22131, second up-right pseudostrobilus). (L) Tangential section through same specimen as K showing megasporangial arrangement (magnified in SI Appendix, Figs. S21B and S23A). (M) Detail of the middle part of L showing the megasporangia and microsporangia. (N) Single spore macerated from the holotype. (Scale bars: A, 10 cm; B, 3 cm; C–E, 1 cm; F, 100 μm; G and H, 200 μm; I–L, 5 mm; M, 2 mm; N, 10 μm.)" figureDoi="http://doi.org/10.5281/zenodo.4596608" httpUri="https://zenodo.org/record/4596608/files/figure.png" pageId="3" pageNumber="4">
|
||
Fig. 1
|
||
<emphasis box="[407,455,605,627]" italics="true" pageId="3" pageNumber="4">K–M</emphasis>
|
||
</figureCitation>
|
||
and
|
||
<emphasis box="[511,634,604,626]" italics="true" pageId="3" pageNumber="4">SI Appendix</emphasis>
|
||
, Figs. S21
|
||
<emphasis box="[752,767,605,626]" italics="true" pageId="3" pageNumber="4">B</emphasis>
|
||
and
|
||
<emphasis box="[160,174,633,654]" italics="true" pageId="3" pageNumber="4">F</emphasis>
|
||
and S23
|
||
<emphasis box="[265,281,632,654]" italics="true" pageId="3" pageNumber="4">A</emphasis>
|
||
) and, although poorly preserved, have a single functional megaspore. As pseudostrobili are born horizontally from the stem, the vertical row of megaspores would have been organized horizontally in life. Due to preservation limitations, we cannot determine where in life the megasporangial row was positioned, but we suspect it was at the bottom of the pseudostrobilus and caused pooling of the hormone auxin under gravity to trigger megasporangial expression (
|
||
<bibRefCitation author="S. J. Wang" box="[502,517,823,845]" journalOrPublisher="Am. J. Bot." pageId="3" pageNumber="4" pagination="127 - 149" part="104" refId="ref6909" refString="9. S. J. Wang et al., Anatomically preserved " strobili " and leaves from the Permian of China (Dorsalistachyaceae, fam. nov.) broaden knowledge of Noeggerathiales and constrain their possible taxonomic affinities. Am. J. Bot. 104, 127 - 149 (2017)." title="Anatomically preserved " strobili " and leaves from the Permian of China (Dorsalistachyaceae, fam. nov.) broaden knowledge of Noeggerathiales and constrain their possible taxonomic affinities" type="journal article" year="2017">9</bibRefCitation>
|
||
). Microspores are of the
|
||
<emphasis box="[114,245,850,872]" italics="true" pageId="3" pageNumber="4">Calamospora</emphasis>
|
||
<typeStatus box="[259,304,851,872]" pageId="3" pageNumber="4">type</typeStatus>
|
||
; trilete circular, subcircular to oval, 30 to 40 μm in diameter, with straight rays of the trilete mark forming a labrum extending 3/4 or the whole radius a laevigate exine that forms irregular folds usually parallel to the margin (
|
||
<figureCitation box="[646,723,932,954]" captionStart="Fig" captionStartId="2.[102,130,1656,1673]" captionTargetBox="[174,1375,147,1630]" captionTargetId="figure-1@2.[172,1377,145,1632]" captionTargetPageId="2" captionText="Fig. 1. P. wuhaia J. Wang et al. sp. nov. from the early Permian Taiyuan Formation of Wuda Coalfield, Inner Mongolia. (A) Holotype with an entire crown consisting of pseudostrobili and leaves. (B) Once-pinnate compound leaf with both large and small pinnules visible. Reprinted with permission from ref. 15. (C) Cross-section of a crown illustrating pseudostrobili around the stem. (D) Cross-section of pseudostrobilus with microsporangia around the axis with bilateral, inversed Ω-shaped vascular bundle. (E) Cross-section of a leaf rachis showing the same form of vascular bundle as that of pseudostrobili axes. (F–H) Partial cross, radial, and tangential sections of the stem showing the secondary xylem (wood). (I) Tangential section of pseudostrobilus showing sporangial arrangement with single line of megasporangia along with the axis (middle-right pseudostrobilus of the fragmental crown illustrated SI Appendix, Fig. S8; specimen PB22132; magnified in SI Appendix, Fig. S21C.) (J) Radial section of pseudostrobilus showing adaxial sporangia and axis lacking nodes (upmost-left pseudostrobilus of the fragmental crown in SI Appendix, Fig. S8; magnified in SI Appendix, Fig. S21A). (K) Tangential section showing adaxial sporangia and a single line of megasporangia along with the axis (SI Appendix, Fig. S7; specimen PB22131, second up-right pseudostrobilus). (L) Tangential section through same specimen as K showing megasporangial arrangement (magnified in SI Appendix, Figs. S21B and S23A). (M) Detail of the middle part of L showing the megasporangia and microsporangia. (N) Single spore macerated from the holotype. (Scale bars: A, 10 cm; B, 3 cm; C–E, 1 cm; F, 100 μm; G and H, 200 μm; I–L, 5 mm; M, 2 mm; N, 10 μm.)" figureDoi="http://doi.org/10.5281/zenodo.4596608" httpUri="https://zenodo.org/record/4596608/files/figure.png" pageId="3" pageNumber="4">
|
||
Fig. 1
|
||
<emphasis box="[705,723,933,954]" italics="true" pageId="3" pageNumber="4">N</emphasis>
|
||
</figureCitation>
|
||
and
|
||
<emphasis box="[114,236,959,981]" italics="true" pageId="3" pageNumber="4">SI Appendix</emphasis>
|
||
, Fig. S23
|
||
<emphasis box="[333,351,960,981]" italics="true" pageId="3" pageNumber="4">D</emphasis>
|
||
).
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="3" pageNumber="4" type="description">
|
||
<paragraph blockId="3.[114,769,141,1199]" pageId="3" pageNumber="4">
|
||
Detailed descriptions, measurement, and further illustrations are provided in
|
||
<emphasis box="[279,615,1014,1036]" italics="true" pageId="3" pageNumber="4">SI Appendix, Detailed Description</emphasis>
|
||
.
|
||
</paragraph>
|
||
<paragraph blockId="3.[114,769,141,1199]" pageId="3" pageNumber="4">
|
||
<taxonomicName authority="J. Wang et al." authorityName="J. Wang" box="[138,392,1041,1063]" genus="Paratingia" pageId="3" pageNumber="4" rank="species" species="wuhaia">
|
||
<emphasis box="[138,152,1042,1063]" italics="true" pageId="3" pageNumber="4">P</emphasis>
|
||
.
|
||
<emphasis box="[167,238,1041,1063]" italics="true" pageId="3" pageNumber="4">wuhaia</emphasis>
|
||
J. Wang et al.
|
||
</taxonomicName>
|
||
is reconstructed as a small, slender, unbranched palm-like tree in
|
||
<figureCitation box="[413,472,1068,1090]" captionStart="Fig" captionStartId="3.[804,832,1876,1893]" captionTargetBox="[805,1457,953,1852]" captionTargetId="figure-907@3.[805,1457,953,1852]" captionTargetPageId="3" captionText="Fig. 2. Reconstruction of the aerial parts of P. wuhaia J. Wang et al. sp.nov. from the early Permian of China (based on the holotype). Image credit: Yugao Ren and Sijia Tang (artists)." figureDoi="http://doi.org/10.5281/zenodo.4596612" httpUri="https://zenodo.org/record/4596612/files/figure.png" pageId="3" pageNumber="4">Fig. 2</figureCitation>
|
||
, two to many meters high. Its major characteristics are illustrated by a diagrammatic reconstruction as shown in
|
||
<figureCitation box="[336,396,1123,1145]" captionStart="Fig" captionStartId="4.[102,130,1148,1165]" captionTargetBox="[348,1560,143,1123]" captionTargetId="figure-354@4.[346,1202,142,1124]" captionTargetPageId="4" captionText="Fig. 3. Diagrammatic reconstruction of the key features of P. wuhaia J. Wang et al. (A) Stem bears pseudostrobili and once-pinnate compound leaves; pseudostrobili are attached to the fertile zone, above which the leaves are shown from the abaxial side, and below which the leaves are shown from the adaxial side. (B) A pseudostrobilus shown its longitudinal surface. (C) Cross-section of a pseudostrobilus showing both megasporangia and microsporangia, the bilateral symmetrical Ω-shaped vascular bundle and sporophylls; noting the relative position of the presumed two megasporangia and the orientation of the opening of the vascular bundle. (D) Partial longitudinal section through the central axis of the pseudostrobilus and the symmetric axis of the vascular bundle, approximately corresponding to the dashed line in C; (E) partial of the leaf showing the arrangement of the small and big pinnules, seen from the abaxial (Upper) and adaxial (Lower) sides, respectively; (F) a pair of small and big pinnules showing the radiate venation; (G and H) cross-section of the stem showing the anatomy; H is the magnification of a part (within the dashed line) of the stem in G. CT, cortex; LT, leaf trace; MEG, megasporangium; MIG, microsporangium; PD, periderm; PT, pith; PX, primary xylem; SP, sporophyll; SX, secondary xylem; VB, vascular bundle." figureDoi="http://doi.org/10.5281/zenodo.4596614" httpUri="https://zenodo.org/record/4596614/files/figure.png" pageId="3" pageNumber="4">Fig. 3</figureCitation>
|
||
. The details of its stem anatomy are consistent with that of progymnosperms and early seed plants (
|
||
<emphasis italics="true" pageId="3" pageNumber="4">SI Appendix</emphasis>
|
||
, Figs. S16–S18).
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="3" pageNumber="4" type="discussion">
|
||
<paragraph blockId="3.[114,769,1238,1943]" pageId="3" pageNumber="4">
|
||
<emphasis bold="true" box="[114,241,1240,1260]" pageId="3" pageNumber="4">Comparisons.</emphasis>
|
||
Following Wang et al. (
|
||
<bibRefCitation author="S. J. Wang" box="[500,514,1238,1260]" journalOrPublisher="Am. J. Bot." pageId="3" pageNumber="4" pagination="127 - 149" part="104" refId="ref6909" refString="9. S. J. Wang et al., Anatomically preserved " strobili " and leaves from the Permian of China (Dorsalistachyaceae, fam. nov.) broaden knowledge of Noeggerathiales and constrain their possible taxonomic affinities. Am. J. Bot. 104, 127 - 149 (2017)." title="Anatomically preserved " strobili " and leaves from the Permian of China (Dorsalistachyaceae, fam. nov.) broaden knowledge of Noeggerathiales and constrain their possible taxonomic affinities" type="journal article" year="2017">9</bibRefCitation>
|
||
), we consider the noeggerathialean fertile shoot to be a pseudostrobilus because despite its radial exterior appearance, it is distinctly bilateral in organization and represents a complex, three-dimensional “sporophyll” (
|
||
<emphasis box="[123,246,1348,1370]" italics="true" pageId="3" pageNumber="4">SI Appendix</emphasis>
|
||
,
|
||
<emphasis box="[260,382,1348,1370]" italics="true" pageId="3" pageNumber="4">Terminology</emphasis>
|
||
). We use quotation marks to denote what would traditionally be interpreted as sporophylls on the pseudostrobilus axis that represent pinnule homologs and are not in fact sporophylls in the traditional sense.
|
||
</paragraph>
|
||
<paragraph blockId="3.[114,769,1238,1943]" lastBlockId="3.[804,1458,141,462]" pageId="3" pageNumber="4">
|
||
<taxonomicName authority="J. Wang et al." authorityName="J. Wang" box="[138,397,1457,1479]" genus="Paratingia" pageId="3" pageNumber="4" rank="species" species="wuhaia">
|
||
<emphasis box="[138,152,1457,1478]" italics="true" pageId="3" pageNumber="4">P</emphasis>
|
||
.
|
||
<emphasis box="[168,239,1457,1479]" italics="true" pageId="3" pageNumber="4">wuhaia</emphasis>
|
||
J. Wang et al.
|
||
</taxonomicName>
|
||
is most similar to the contemporaneous species
|
||
<taxonomicName box="[265,384,1484,1506]" genus="Paratingia" pageId="3" pageNumber="4" rank="species" species="wudensis">
|
||
<emphasis box="[265,384,1484,1506]" italics="true" pageId="3" pageNumber="4">P. wudensis</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation author="J. Wang & H. W. Pfefferkorn & J. Bek" box="[402,428,1484,1506]" journalOrPublisher="China. Am. J. Bot." pageId="3" pageNumber="4" pagination="1676 - 1689" part="96" refId="ref7308" refString="17. J. Wang, H. W. Pfefferkorn, J. Bek, Paratingia wudensis sp. nov., a whole noeggerathialean plant preserved in an earliest Permian air fall tuff in Inner Mongolia, China. Am. J. Bot. 96, 1676 - 1689 (2009)." title="Paratingia wudensis sp. nov., a whole noeggerathialean plant preserved in an earliest Permian air fall tuff in Inner Mongolia" type="journal article" year="2009">17</bibRefCitation>
|
||
) also from the Wuda Coalfield. Within the coalfield, the two species grew in different areas, with
|
||
<taxonomicName box="[114,221,1538,1560]" genus="Paratingia" pageId="3" pageNumber="4" rank="species" species="wuhaia">
|
||
<emphasis box="[114,221,1538,1560]" italics="true" pageId="3" pageNumber="4">P. wuhaia</emphasis>
|
||
</taxonomicName>
|
||
dominating central and northern parts while
|
||
<taxonomicName genus="Paratingia" pageId="3" pageNumber="4" rank="species" species="wudensis">
|
||
<emphasis italics="true" pageId="3" pageNumber="4">P. wudensis</emphasis>
|
||
</taxonomicName>
|
||
occurs in low density only in the southern parts.
|
||
<taxonomicName genus="Paratingia" pageId="3" pageNumber="4" rank="species" species="wudensis">
|
||
<emphasis italics="true" pageId="3" pageNumber="4">P. wudensis</emphasis>
|
||
</taxonomicName>
|
||
is distinguished from
|
||
<taxonomicName box="[424,521,1593,1615]" genus="Paratingia" pageId="3" pageNumber="4" rank="species" species="wuhaia">
|
||
<emphasis box="[424,521,1593,1615]" italics="true" pageId="3" pageNumber="4">P. wuhaia</emphasis>
|
||
</taxonomicName>
|
||
in forming smaller trees with comparatively larger pseudostrobili, and possessing needlelike small pinnules that are very different from the small, oval pinules in
|
||
<taxonomicName box="[224,324,1675,1697]" genus="Paratingia" pageId="3" pageNumber="4" rank="species" species="wuhaia">
|
||
<emphasis box="[224,324,1675,1697]" italics="true" pageId="3" pageNumber="4">P. wuhaia</emphasis>
|
||
</taxonomicName>
|
||
(see
|
||
<emphasis box="[384,506,1675,1697]" italics="true" pageId="3" pageNumber="4">SI Appendix</emphasis>
|
||
,
|
||
<emphasis box="[521,762,1675,1697]" italics="true" pageId="3" pageNumber="4">Systematic Comparisons</emphasis>
|
||
, for further details). Furthermore
|
||
<taxonomicName box="[455,570,1702,1724]" genus="Paratingia" pageId="3" pageNumber="4" rank="species" species="wudensis">
|
||
<emphasis box="[455,570,1702,1724]" italics="true" pageId="3" pageNumber="4">P. wudensis</emphasis>
|
||
</taxonomicName>
|
||
has more than one megaspore per megasporangium (
|
||
<bibRefCitation author="J. Wang & H. W. Pfefferkorn & J. Bek" box="[473,499,1729,1751]" journalOrPublisher="China. Am. J. Bot." pageId="3" pageNumber="4" pagination="1676 - 1689" part="96" refId="ref7308" refString="17. J. Wang, H. W. Pfefferkorn, J. Bek, Paratingia wudensis sp. nov., a whole noeggerathialean plant preserved in an earliest Permian air fall tuff in Inner Mongolia, China. Am. J. Bot. 96, 1676 - 1689 (2009)." title="Paratingia wudensis sp. nov., a whole noeggerathialean plant preserved in an earliest Permian air fall tuff in Inner Mongolia" type="journal article" year="2009">17</bibRefCitation>
|
||
) unlike the single megaspores in
|
||
<taxonomicName box="[223,327,1757,1779]" genus="Paratingia" pageId="3" pageNumber="4" rank="species" species="wuhaia">
|
||
<emphasis box="[223,327,1757,1779]" italics="true" pageId="3" pageNumber="4">P. wuhaia</emphasis>
|
||
</taxonomicName>
|
||
megasporangia. The fertile shoots of
|
||
<taxonomicName genus="Paratingia" pageId="3" pageNumber="4" rank="species" species="wuhaia">
|
||
<emphasis italics="true" pageId="3" pageNumber="4">P. wuhaia</emphasis>
|
||
</taxonomicName>
|
||
are similar to the late Permian heterosporous noeggerathialean pseudostrobilus
|
||
<emphasis box="[384,729,1811,1833]" italics="true" pageId="3" pageNumber="4">Dorsalistachya quadrisegmentorum</emphasis>
|
||
(
|
||
<bibRefCitation author="S. J. Wang" box="[746,760,1811,1833]" journalOrPublisher="Am. J. Bot." pageId="3" pageNumber="4" pagination="127 - 149" part="104" refId="ref6909" refString="9. S. J. Wang et al., Anatomically preserved " strobili " and leaves from the Permian of China (Dorsalistachyaceae, fam. nov.) broaden knowledge of Noeggerathiales and constrain their possible taxonomic affinities. Am. J. Bot. 104, 127 - 149 (2017)." title="Anatomically preserved " strobili " and leaves from the Permian of China (Dorsalistachyaceae, fam. nov.) broaden knowledge of Noeggerathiales and constrain their possible taxonomic affinities" type="journal article" year="2017">9</bibRefCitation>
|
||
) that is associated with
|
||
<emphasis box="[353,620,1839,1861]" italics="true" pageId="3" pageNumber="4">
|
||
<taxonomicName authorityName="R.Zeiller" authorityYear="1894" box="[353,486,1839,1861]" class="Progymnospermopsida" genus="Plagiozamites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Noeggerathiales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="genus">Plagiozamites</taxonomicName>
|
||
oblongifolius
|
||
</emphasis>
|
||
megaphyllous leaves; its leaves and pseudostrobili also have Ω- shaped bundles but lack secondary xylem.
|
||
<emphasis box="[383,526,1893,1915]" italics="true" pageId="3" pageNumber="4">Dorsalistachya</emphasis>
|
||
is further distinguished from
|
||
<taxonomicName box="[172,272,1921,1942]" genus="Paratingia" pageId="3" pageNumber="4" rank="genus">
|
||
<emphasis box="[172,272,1921,1942]" italics="true" pageId="3" pageNumber="4">Paratingia</emphasis>
|
||
</taxonomicName>
|
||
in having winged monomegasporangia in short, vertical rows (
|
||
<bibRefCitation author="S. J. Wang" box="[950,963,141,163]" journalOrPublisher="Am. J. Bot." pageId="3" pageNumber="4" pagination="127 - 149" part="104" refId="ref6909" refString="9. S. J. Wang et al., Anatomically preserved " strobili " and leaves from the Permian of China (Dorsalistachyaceae, fam. nov.) broaden knowledge of Noeggerathiales and constrain their possible taxonomic affinities. Am. J. Bot. 104, 127 - 149 (2017)." title="Anatomically preserved " strobili " and leaves from the Permian of China (Dorsalistachyaceae, fam. nov.) broaden knowledge of Noeggerathiales and constrain their possible taxonomic affinities" type="journal article" year="2017">9</bibRefCitation>
|
||
) and possessing more complexly divided sporophylls. Adaxially curved and C-shaped primary xylem strands at the innermost part of the xylem cylinder in
|
||
<taxonomicName box="[1185,1279,196,217]" genus="Paratingia" pageId="3" pageNumber="4" rank="genus">
|
||
<emphasis box="[1185,1279,196,217]" italics="true" pageId="3" pageNumber="4">Paratingia</emphasis>
|
||
</taxonomicName>
|
||
’s stem differ from those in gymnosperms and
|
||
<taxonomicName authorityName="J.W.Dawson" authorityYear="1871" box="[1065,1192,222,244]" class="Progymnospermopsida" family="Archaeopteridaceae" genus="Archaeopteris" higherTaxonomySource="GBIF" kingdom="Plantae" order="Archaeopteridales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="genus">
|
||
<emphasis box="[1065,1192,222,244]" italics="true" pageId="3" pageNumber="4">Archaeopteris</emphasis>
|
||
</taxonomicName>
|
||
(progymnosperm) in which strands are isodiametric in cross-section. Broad, arc-shaped leaf traces in the innermost part of the xylem cylinder of the progymnosperm
|
||
<taxonomicName box="[939,1030,305,326]" class="Progymnospermopsida" family="Protopityaceae" genus="Protopitys" higherTaxonomySource="GBIF" kingdom="Plantae" order="Protopityales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="genus">
|
||
<emphasis box="[939,1030,305,326]" italics="true" pageId="3" pageNumber="4">Protopitys</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation author="J. Walton & On Protopitys" box="[1049,1074,304,326]" journalOrPublisher="Trans. R. Soc. Edinb." pageId="3" pageNumber="4" pagination="333 - 340" part="63" refId="ref7364" refString="18. J. Walton, On Protopitys (Goppert): With a description of a fertile specimen Protopitys scotica sp. nov. from the Calciferous Sandstone Series of Dunbartonshire. Trans. R. Soc. Edinb. 63, 333 - 340 (1957)." title="(Goppert): With a description of a fertile specimen Protopitys scotica sp. nov. from the Calciferous Sandstone Series of Dunbartonshire" type="book" year="1957">18</bibRefCitation>
|
||
) are similar to the c-shaped primary xylem strands in
|
||
<taxonomicName box="[975,1070,332,353]" genus="Paratingia" pageId="3" pageNumber="4" rank="genus">
|
||
<emphasis box="[975,1070,332,353]" italics="true" pageId="3" pageNumber="4">Paratingia</emphasis>
|
||
</taxonomicName>
|
||
stems, but
|
||
<taxonomicName box="[1188,1279,332,353]" class="Progymnospermopsida" family="Protopityaceae" genus="Protopitys" higherTaxonomySource="GBIF" kingdom="Plantae" order="Protopityales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="genus">
|
||
<emphasis box="[1188,1279,332,353]" italics="true" pageId="3" pageNumber="4">Protopitys</emphasis>
|
||
</taxonomicName>
|
||
stems bear large, distichously arranged fronds with bilateral symmetry and plannated branching bearing terminal sporangia in fertile trusses that lack associated leaves or sporopylls (
|
||
<bibRefCitation author="R. M. Bateman & W. A. DiMichele" box="[1111,1122,413,435]" journalOrPublisher="Biol. Rev. Camb. Philos. Soc." pageId="3" pageNumber="4" pagination="345 - 417" part="69" refId="ref6576" refString="1. R. M. Bateman, W. A. DiMichele, Heterospory: The most iterative key innovation in the evolutionary history of the plant kingdom. Biol. Rev. Camb. Philos. Soc. 69, 345 - 417 (1994)." title="Heterospory: The most iterative key innovation in the evolutionary history of the plant kingdom" type="journal article" year="1994">1</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="S. P. Stubblefield & G. W. Rothwell" box="[1133,1146,413,435]" journalOrPublisher="Am. J. Bot." pageId="3" pageNumber="4" pagination="1415 - 1428" part="76" refId="ref6748" refString="5. S. P. Stubblefield, G. W. Rothwell, Cecropsis luculentum gen. et spec. nov.: Evidence for heterosporous progymnosperms in the Upper Pennsylvanian of North America. Am. J. Bot. 76, 1415 - 1428 (1989)." title="Cecropsis luculentum gen. et spec. nov.: Evidence for heterosporous progymnosperms in the Upper Pennsylvanian of North America" type="journal article" year="1989">5</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="J. Walton & On Protopitys" box="[1155,1179,413,435]" journalOrPublisher="Trans. R. Soc. Edinb." pageId="3" pageNumber="4" pagination="333 - 340" part="63" refId="ref7364" refString="18. J. Walton, On Protopitys (Goppert): With a description of a fertile specimen Protopitys scotica sp. nov. from the Calciferous Sandstone Series of Dunbartonshire. Trans. R. Soc. Edinb. 63, 333 - 340 (1957)." title="(Goppert): With a description of a fertile specimen Protopitys scotica sp. nov. from the Calciferous Sandstone Series of Dunbartonshire" type="book" year="1957">18</bibRefCitation>
|
||
) (see
|
||
<emphasis box="[1238,1352,413,435]" italics="true" pageId="3" pageNumber="4">SI Appendix</emphasis>
|
||
, Tables S1 and S2 for detailed comparisons).
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |