<document id="1776AF4C43B89EA169BFD780E41D6BCE" ID-DOI="10.2478/if-2018-0010" ID-ISSN="2533-4069" ID-Zenodo-Dep="5383379" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1630296376118" checkinUser="diego" docAuthor="Friis, Else Marie, Crane, Peter R. &amp; Pedersen, Kaj Raunsgaard" docDate="2018" docId="1A4787F5FFB2E119FCA166E8FD92BB79" docLanguage="en" docName="10_Friis_135158.pdf" docOrigin="Fossil Imprint 74 (1 - 2)" docSource="http://dx.doi.org/10.2478/if-2018-0010" docStyle="DocumentStyle:0EC54BF35E227CA4627F894F6BA30634.4:FossilImprint.2016-.journal_article" docStyleId="0EC54BF35E227CA4627F894F6BA30634" docStyleName="FossilImprint.2016-.journal_article" docStyleVersion="4" docTitle="Gastonispermum portugallicum Friis &amp; Crane &amp; Pedersen 2018, gen. et sp. nov." docType="treatment" docVersion="3" lastPageNumber="140" masterDocId="E67EFF8DFFB1E11CFFCF6751FFA3BC0C" masterDocTitle="Extinct Taxa Of Exotestal Seeds Close To Austrobaileyales And Nymphaeales From The Early Cretaceous Of Portugal" masterLastPageNumber="158" masterPageNumber="135" pageNumber="138" updateTime="1699219021208" updateUser="plazi" zenodo-license-document="CC-BY-NC-ND-3.0">
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<mods:title id="C543168647817190226D18162B7D2C4E">Extinct Taxa Of Exotestal Seeds Close To Austrobaileyales And Nymphaeales From The Early Cretaceous Of Portugal</mods:title>
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<mods:namePart id="386933FEACC83A8D59B7866A97F49572">Friis, Else Marie</mods:namePart>
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<mods:namePart id="586A4FE2426EC212A5F2E690560A86C1">Crane, Peter R.</mods:namePart>
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<mods:title id="EB08A62231732F654E47B99B5DDB2782">Fossil Imprint</mods:title>
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<mods:date id="6A0A49363767BC5D4DFFF041A0CCE7C6">2018</mods:date>
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<paragraph id="925136E3FFB2E11FFCA166E8FADABDDD" blockId="3.[878,1401,440,465]" box="[878,1401,440,465]" pageId="3" pageNumber="138">
<heading id="C919818FFFB2E11FFCA166E8FADABDDD" box="[878,1401,440,465]" centered="true" fontSize="8" level="2" pageId="3" pageNumber="138" reason="2">
<emphasis id="A09AEAF1FFB2E11FFCA166E8FADABDDD" bold="true" box="[878,1401,440,465]" pageId="3" pageNumber="138">
<taxonomicName id="55EE4D60FFB2E11FFCA166E8FB6EBDDC" authority="Friis &amp; Crane &amp; Pedersen, 2018" authorityName="Friis &amp; Crane &amp; Pedersen" authorityYear="2018" box="[878,1229,440,465]" class="Magnoliopsida" family="Chloranthaceae" genus="Gastonispermum" kingdom="Plantae" order="Chloranthales" pageId="3" pageNumber="138" phylum="Tracheophyta" rank="species" species="portugallicum" status="gen. et sp. nov.">
<emphasis id="A09AEAF1FFB2E11FFCA166E8FB6EBDDC" bold="true" box="[878,1229,440,465]" italics="true" pageId="3" pageNumber="138">Gastonispermum portugallicum</emphasis>
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<taxonomicNameLabel id="BBA9578AFFB2E11FFB1C66E8FADABDDD" box="[1235,1401,441,465]" pageId="3" pageNumber="138" rank="species">gen. et sp. nov.</taxonomicNameLabel>
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<paragraph id="925136E3FFB2E11FFBE26688FB18BDFD" blockId="3.[1069,1211,473,497]" box="[1069,1211,473,497]" pageId="3" pageNumber="138">
<figureCitation id="0AD52A66FFB2E11FFBE26688FB18BDFD" box="[1069,1211,473,497]" captionStart-0="Text-fig" captionStart-1="Text-fig" captionStartId-0="2.[147,228,1713,1734]" captionStartId-1="3.[125,205,1667,1688]" captionTargetBox-0="[147,1462,151,1686]" captionTargetId-0="figure-2@2.[147,1463,151,1686]" captionTargetPageId-0="2" captionText-0="Text-fig. 1. Gastonispermum portugallicum gen. et sp. nov. seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, volume renderings). Note remains of mounting media on several seeds (¤). a) Seed in oblique view showing seed shape, the slightly raised raphal ridge and the position of hilum (hi) and micropyle (mi) on the raphal side of the seed (S170218). b, c) Seeds in lateral view (b, S170234; c, S175095). d–f) Holotype (S174820); seed in lateral view (d) and cut volume rendering (e, f) through the median plane of the seed showing palisade-shaped sclerenchyma cells of exotesta and remains of embryo (emb) and surrounding nutritive tissue (e, cut between yz0440-0530; f, cut between slices yz440-480). g) Hilum (hi) and micropyle (mi) of seed in (1a) showing the Y-shaped micropylar slit in the outer integument. h) Cut volume rendering through the median plane of the seed (cut at yz0492) showing seed coat mainly composed of palisade-shaped cells of the exotesta (S174435). i) Seed surface showing the raised outlines of the undulate anticlinal walls of the exotestal cells (S175045). Scale bars = 500 µm (a–e); 250 µm (g); 125 µm (f, i)." captionText-1="Text-fig. 2. Gastonispermum portugallicum gen. et sp. nov. seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, orthoslices). a) Longitudinal orthoslice (yz0745) through the median plane of the seed (holotype, S174820) showing taller exotestal cells on raphal side, well preserved embryo (em) and partly preserved nutritive tissue. b) Longitudinal orthoslice (xz1355) through the micropylar region perpendicular to the median plane showing the exotestal tissue surrounding the micropyle (mi) forming a Y-shaped micropylar slit (S105218). c) Longitudinal orthoslice (yz0564) through the hilum (hi) and micropyle (mi) in the median plane showing the gradual shortening of the exotestal cells above the micropyle and palisade-shaped sclerenchyma cells lining the micropylar slit (S170234). d) Longitudinal orthoslice (1322) through apical part of seed in the median plane (holotype, S174820) showing embryo with two rudimentary cotyledons (asterisks); note the tiny cells of embryo containing structures we interpret as the fossilized remains of nuclei. e) Longitudinal orthoslice (yz1170) through the apical part of seed in the median plane showing tiny embryo with two rudimentary cotyledons (asterisks) and remains of nutritive tissue with scattered granules that probably representing the remains of protein or lipid bodies; nutritive tissue immediately surrounding embryo is probably endosperm (endo) (S174821). f) Longitudinal orthoslice (xz0590) through apical part of seed perpendicular to the median plane showing the empty space formerly occupied by the embryo, surrounded by cellular nutritive tissue (S174430). g) Transverse orthoslice (xy0969) through the middle part of a seed showing palisade-shaped exotestal cells and remains of mesotesta and tegmen (S175044). Scale bars = 500 µm (a); 250 µm (c–e, g); 125 µm (b, f)." figureDoi-0="http://doi.org/10.5281/zenodo.5383381" httpUri-0="https://zenodo.org/record/5383381/files/figure.png" pageId="3" pageNumber="138">Text-figs 1–2</figureCitation>
</paragraph>
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<paragraph id="925136E3FFB2E11FFC956558FAC7BE4C" blockId="3.[820,1459,520,577]" pageId="3" pageNumber="138">
H o l o t y p e. Designated here, S174820 (Famalicão sample 025; illustrated here in 
<figureCitation id="0AD52A66FFB2E11FFBB26579FAB5BE4C" box="[1149,1302,552,576]" captionStart="Text-fig" captionStartId="2.[147,228,1713,1734]" captionTargetBox="[147,1462,151,1686]" captionTargetId="figure-2@2.[147,1463,151,1686]" captionTargetPageId="2" captionText="Text-fig. 1. Gastonispermum portugallicum gen. et sp. nov. seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, volume renderings). Note remains of mounting media on several seeds (¤). a) Seed in oblique view showing seed shape, the slightly raised raphal ridge and the position of hilum (hi) and micropyle (mi) on the raphal side of the seed (S170218). b, c) Seeds in lateral view (b, S170234; c, S175095). d–f) Holotype (S174820); seed in lateral view (d) and cut volume rendering (e, f) through the median plane of the seed showing palisade-shaped sclerenchyma cells of exotesta and remains of embryo (emb) and surrounding nutritive tissue (e, cut between yz0440-0530; f, cut between slices yz440-480). g) Hilum (hi) and micropyle (mi) of seed in (1a) showing the Y-shaped micropylar slit in the outer integument. h) Cut volume rendering through the median plane of the seed (cut at yz0492) showing seed coat mainly composed of palisade-shaped cells of the exotesta (S174435). i) Seed surface showing the raised outlines of the undulate anticlinal walls of the exotestal cells (S175045). Scale bars = 500 µm (a–e); 250 µm (g); 125 µm (f, i)." figureDoi="http://doi.org/10.5281/zenodo.5383381" httpUri="https://zenodo.org/record/5383381/files/figure.png" pageId="3" pageNumber="138">Text-figs 1d–f</figureCitation>
, 
<figureCitation id="0AD52A66FFB2E11FFAEE6578FAF4BE4C" box="[1313,1367,552,577]" captionStart="Text-fig" captionStartId="3.[125,205,1667,1688]" captionText="Text-fig. 2. Gastonispermum portugallicum gen. et sp. nov. seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, orthoslices). a) Longitudinal orthoslice (yz0745) through the median plane of the seed (holotype, S174820) showing taller exotestal cells on raphal side, well preserved embryo (em) and partly preserved nutritive tissue. b) Longitudinal orthoslice (xz1355) through the micropylar region perpendicular to the median plane showing the exotestal tissue surrounding the micropyle (mi) forming a Y-shaped micropylar slit (S105218). c) Longitudinal orthoslice (yz0564) through the hilum (hi) and micropyle (mi) in the median plane showing the gradual shortening of the exotestal cells above the micropyle and palisade-shaped sclerenchyma cells lining the micropylar slit (S170234). d) Longitudinal orthoslice (1322) through apical part of seed in the median plane (holotype, S174820) showing embryo with two rudimentary cotyledons (asterisks); note the tiny cells of embryo containing structures we interpret as the fossilized remains of nuclei. e) Longitudinal orthoslice (yz1170) through the apical part of seed in the median plane showing tiny embryo with two rudimentary cotyledons (asterisks) and remains of nutritive tissue with scattered granules that probably representing the remains of protein or lipid bodies; nutritive tissue immediately surrounding embryo is probably endosperm (endo) (S174821). f) Longitudinal orthoslice (xz0590) through apical part of seed perpendicular to the median plane showing the empty space formerly occupied by the embryo, surrounded by cellular nutritive tissue (S174430). g) Transverse orthoslice (xy0969) through the middle part of a seed showing palisade-shaped exotestal cells and remains of mesotesta and tegmen (S175044). Scale bars = 500 µm (a); 250 µm (c–e, g); 125 µm (b, f)." pageId="3" pageNumber="138">2a, d</figureCitation>
).
</paragraph>
<paragraph id="925136E3FFB2E11FFC956509FA13BE63" blockId="3.[820,1456,600,656]" box="[858,1456,600,624]" pageId="3" pageNumber="138">P l a n t F o s s i l N a m e s R e g i s t r y N u m b e r.</paragraph>
<paragraph id="925136E3FFB2E11FFCFB6529FBD8BE9C" blockId="3.[820,1456,600,656]" box="[820,1147,632,656]" pageId="3" pageNumber="138">PFN000089 (for new species).</paragraph>
<paragraph id="925136E3FFB2E11FFC9565F9FB14BF2C" blockId="3.[820,1459,679,800]" pageId="3" pageNumber="138">P a r a t y p e s. Designated here, S105218, S105220, S170234, S174343, S174430, S174435, S174819, S174821, S175044, S175045, S175082, S175084, S175095, S175100 – S175104 (Famalicão sample 025).</paragraph>
<paragraph id="925136E3FFB2E11FFC956466FB94BF83" blockId="3.[820,1459,823,911]" pageId="3" pageNumber="138">
R e p o s i t o r y. Palaeobotanical Collections, Department of Palaeobiology, the Swedish Museum of Natural History, 
<collectingRegion id="502AF801FFB2E11FFCFB6426FC09BF83" box="[820,938,887,911]" country="Sweden" name="Stockholm" pageId="3" pageNumber="138">Stockholm</collectingRegion>
, 
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(S).
</paragraph>
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E t y m o l o g y. The species is named for 
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where the fossils were collected.
</paragraph>
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<materialsCitation id="22863CBEFFB2E11FFC9564A6FA10B802" box="[858,1459,1013,1039]" collectionCode="T" country="Portugal" latitude="39.704445" location="Famalicao" pageId="3" pageNumber="138" specimenCount="1">
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y p e l o c a l i t y. 
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, 
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(
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;
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<geoCoordinate id="F7DA5024FFB2E11FFCFB6344FC13B822" box="[820,944,1045,1070]" degrees="8" direction="west" minutes="46" orientation="longitude" pageId="3" pageNumber="138" precision="15" seconds="12" value="-8.7699995">8°46′12″W</geoCoordinate>
).
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<paragraph id="925136E3FFB2E11FFC956317FC3BB892" blockId="3.[820,1459,1093,1182]" pageId="3" pageNumber="138">T y p e s t r a t u m a n d a g e. Below the Figueira da Foz Formation; Early Cretaceous (late Aptian – early Albian or older).</paragraph>
<paragraph id="925136E3FFB2E11FFC9563E7FB14B8C2" blockId="3.[858,1207,1205,1230]" box="[858,1207,1205,1230]" pageId="3" pageNumber="138">D i a g n o s i s. As for the genus.</paragraph>
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D i m e n s i o n s. Length of seeds: 
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; width of seeds: 
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.
</paragraph>
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O t h e r s p e c i m e n s. S175108 (
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de Água sample
</paragraph>
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330); S175110 (
<collectingRegion id="502AF801FFB2E11FFC106205FBB3B960" box="[991,1040,1364,1388]" country="Switzerland" name="Valais" pageId="3" pageNumber="138">Vale</collectingRegion>
de Água sample 333).
</paragraph>
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D e s c r i p t i o n a n d r e m a r k s. The species is based on about 230 isolated seeds of which 
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were studied using 
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(
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, 
<specimenCode id="C2489E98FFB2E11FFB2F6292FAE2B9D7" box="[1248,1345,1475,1499]" collectionCode="S" country="Sweden" lsid="urn:lsid:biocol.org:col:15668" name="Department of Botany, Swedish Museum of Natural History" pageId="3" pageNumber="138" type="Herbarium">S105220</specimenCode>
, 
<specimenCode id="C2489E98FFB2E11FFA826295FA0CB9D0" box="[1357,1455,1476,1500]" collectionCode="S" country="Sweden" lsid="urn:lsid:biocol.org:col:15668" name="Department of Botany, Swedish Museum of Natural History" pageId="3" pageNumber="138" type="Herbarium">S170234</specimenCode>
, 
<specimenCode id="C2489E98FFB2E11FFCFB62B5FC35B9F0" box="[820,918,1508,1532]" collectionCode="S" country="Sweden" lsid="urn:lsid:biocol.org:col:15668" name="Department of Botany, Swedish Museum of Natural History" pageId="3" pageNumber="138" type="Herbarium">S174343</specimenCode>
, 
<specimenCode id="C2489E98FFB2E11FFC6F62B5FBA2B9F0" box="[928,1025,1508,1532]" collectionCode="S" country="Sweden" lsid="urn:lsid:biocol.org:col:15668" name="Department of Botany, Swedish Museum of Natural History" pageId="3" pageNumber="138" type="Herbarium">S174430</specimenCode>
, 
<specimenCode id="C2489E98FFB2E11FFBC462B5FBCFB9F0" box="[1035,1132,1508,1532]" collectionCode="S" country="Sweden" lsid="urn:lsid:biocol.org:col:15668" name="Department of Botany, Swedish Museum of Natural History" pageId="3" pageNumber="138" type="Herbarium">S174435</specimenCode>
, 
<specimenCode id="C2489E98FFB2E11FFBB862B5FB7BB9F0" box="[1143,1240,1508,1532]" collectionCode="S" country="Sweden" lsid="urn:lsid:biocol.org:col:15668" name="Department of Botany, Swedish Museum of Natural History" pageId="3" pageNumber="138" type="Herbarium">S174819</specimenCode>
, 
<specimenCode id="C2489E98FFB2E11FFB2D62B5FAE0B9F0" box="[1250,1347,1508,1532]" collectionCode="S" country="Sweden" lsid="urn:lsid:biocol.org:col:15668" name="Department of Botany, Swedish Museum of Natural History" pageId="3" pageNumber="138" type="Herbarium">S174820</specimenCode>
, 
<specimenCode id="C2489E98FFB2E11FFA8262B5FA0CB9F0" box="[1357,1455,1508,1532]" collectionCode="S" country="Sweden" lsid="urn:lsid:biocol.org:col:15668" name="Department of Botany, Swedish Museum of Natural History" pageId="3" pageNumber="138" type="Herbarium">S174821</specimenCode>
, 
<specimenCode id="C2489E98FFB2E11FFCFB6155FC36BA10" box="[820,917,1540,1564]" collectionCode="S" country="Sweden" lsid="urn:lsid:biocol.org:col:15668" name="Department of Botany, Swedish Museum of Natural History" pageId="3" pageNumber="138" type="Herbarium">S175044</specimenCode>
, 
<specimenCode id="C2489E98FFB2E11FFC6E6155FBA1BA10" box="[929,1026,1540,1564]" collectionCode="S" country="Sweden" lsid="urn:lsid:biocol.org:col:15668" name="Department of Botany, Swedish Museum of Natural History" pageId="3" pageNumber="138" type="Herbarium">S175045</specimenCode>
, 
<specimenCode id="C2489E98FFB2E11FFBC26155FBCCBA10" box="[1037,1135,1540,1564]" collectionCode="S" country="Sweden" lsid="urn:lsid:biocol.org:col:15668" name="Department of Botany, Swedish Museum of Natural History" pageId="3" pageNumber="138" type="Herbarium">S175082</specimenCode>
, 
<specimenCode id="C2489E98FFB2E11FFBB56155FB78BA10" box="[1146,1243,1540,1564]" collectionCode="S" country="Sweden" lsid="urn:lsid:biocol.org:col:15668" name="Department of Botany, Swedish Museum of Natural History" pageId="3" pageNumber="138" type="Herbarium">S175084</specimenCode>
, 
<specimenCode id="C2489E98FFB2E11FFB286155FAE9BA10" box="[1255,1354,1540,1564]" collectionCode="S" country="Sweden" lsid="urn:lsid:biocol.org:col:15668" name="Department of Botany, Swedish Museum of Natural History" pageId="3" pageNumber="138" type="Herbarium">S175095</specimenCode>
). There is no information on the fruits in which the seeds were borne. The seeds are small, anatropous, bitegmic and exotestal with bilateral symmetry. They are elliptical in lateral view (
<figureCitation id="0AD52A66FFB5E118FF486F47FEE3B422" box="[135,320,2070,2094]" captionStart="Text-fig" captionStartId="2.[147,228,1713,1734]" captionTargetBox="[147,1462,151,1686]" captionTargetId="figure-2@2.[147,1463,151,1686]" captionTargetPageId="2" captionText="Text-fig. 1. Gastonispermum portugallicum gen. et sp. nov. seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, volume renderings). Note remains of mounting media on several seeds (¤). a) Seed in oblique view showing seed shape, the slightly raised raphal ridge and the position of hilum (hi) and micropyle (mi) on the raphal side of the seed (S170218). b, c) Seeds in lateral view (b, S170234; c, S175095). d–f) Holotype (S174820); seed in lateral view (d) and cut volume rendering (e, f) through the median plane of the seed showing palisade-shaped sclerenchyma cells of exotesta and remains of embryo (emb) and surrounding nutritive tissue (e, cut between yz0440-0530; f, cut between slices yz440-480). g) Hilum (hi) and micropyle (mi) of seed in (1a) showing the Y-shaped micropylar slit in the outer integument. h) Cut volume rendering through the median plane of the seed (cut at yz0492) showing seed coat mainly composed of palisade-shaped cells of the exotesta (S174435). i) Seed surface showing the raised outlines of the undulate anticlinal walls of the exotestal cells (S175045). Scale bars = 500 µm (a–e); 250 µm (g); 125 µm (f, i)." figureDoi="http://doi.org/10.5281/zenodo.5383381" httpUri="https://zenodo.org/record/5383381/files/figure.png" pageId="4" pageNumber="139">Text-figs 1a–e, h</figureCitation>
, 
<figureCitation id="0AD52A66FFB5E118FE826F47FEC4B422" box="[333,359,2070,2094]" captionStart="Text-fig" captionStartId="3.[125,205,1667,1688]" captionText="Text-fig. 2. Gastonispermum portugallicum gen. et sp. nov. seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, orthoslices). a) Longitudinal orthoslice (yz0745) through the median plane of the seed (holotype, S174820) showing taller exotestal cells on raphal side, well preserved embryo (em) and partly preserved nutritive tissue. b) Longitudinal orthoslice (xz1355) through the micropylar region perpendicular to the median plane showing the exotestal tissue surrounding the micropyle (mi) forming a Y-shaped micropylar slit (S105218). c) Longitudinal orthoslice (yz0564) through the hilum (hi) and micropyle (mi) in the median plane showing the gradual shortening of the exotestal cells above the micropyle and palisade-shaped sclerenchyma cells lining the micropylar slit (S170234). d) Longitudinal orthoslice (1322) through apical part of seed in the median plane (holotype, S174820) showing embryo with two rudimentary cotyledons (asterisks); note the tiny cells of embryo containing structures we interpret as the fossilized remains of nuclei. e) Longitudinal orthoslice (yz1170) through the apical part of seed in the median plane showing tiny embryo with two rudimentary cotyledons (asterisks) and remains of nutritive tissue with scattered granules that probably representing the remains of protein or lipid bodies; nutritive tissue immediately surrounding embryo is probably endosperm (endo) (S174821). f) Longitudinal orthoslice (xz0590) through apical part of seed perpendicular to the median plane showing the empty space formerly occupied by the embryo, surrounded by cellular nutritive tissue (S174430). g) Transverse orthoslice (xy0969) through the middle part of a seed showing palisade-shaped exotestal cells and remains of mesotesta and tegmen (S175044). Scale bars = 500 µm (a); 250 µm (c–e, g); 125 µm (b, f)." pageId="4" pageNumber="139">2a</figureCitation>
) and also in transverse section (Textfig. 
<quantity id="55169B06FFB5E118FCBA6087FC30BBE2" box="[885,915,2006,2030]" metricMagnitude="-3" metricUnit="kg" metricValue="2.0" pageId="4" pageNumber="139" unit="g" value="2.0">2g</quantity>
). The seed surface is dull, almost smooth, but with a jigsaw puzzle-shaped pattern formed from the slightly raised undulate anticlinal walls of the exotesta cells
</materialsCitation>
.
</paragraph>
<caption id="C691666BFFB2E11FFFB261D2FBC5B43F" pageId="3" pageNumber="138" startId="3.[125,205,1667,1688]">
<paragraph id="925136E3FFB2E11FFFB261D2FBC5B43F" blockId="3.[125,1441,1667,2099]" pageId="3" pageNumber="138">
Text-fig. 2. 
<taxonomicName id="55EE4D60FFB2E11FFF3861D2FD90BA94" authority="Friis &amp; Crane &amp; Pedersen, 2018" authorityName="Friis &amp; Crane &amp; Pedersen" authorityYear="2018" box="[247,563,1667,1688]" class="Magnoliopsida" family="Chloranthaceae" genus="Gastonispermum" kingdom="Plantae" order="Chloranthales" pageId="3" pageNumber="138" phylum="Tracheophyta" rank="species" species="portugallicum" status="gen. et sp. nov.">
<emphasis id="A09AEAF1FFB2E11FFF3861D2FD90BA94" bold="true" box="[247,563,1667,1688]" italics="true" pageId="3" pageNumber="138">Gastonispermum portugallicum</emphasis>
</taxonomicName>
<taxonomicNameLabel id="BBA9578AFFB2E11FFDF161D2FD7CBA94" box="[574,735,1667,1688]" pageId="3" pageNumber="138" rank="species">gen. et sp. nov.</taxonomicNameLabel>
seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, orthoslices). a) Longitudinal orthoslice (yz0745) through the median plane of the seed (holotype, S174820) showing taller exotestal cells on raphal side, well preserved embryo (em) and partly preserved nutritive tissue. b) Longitudinal orthoslice (xz1355) through the micropylar region perpendicular to the median plane showing the exotestal tissue surrounding the micropyle (mi) forming a Y-shaped micropylar slit (S105218). c) Longitudinal orthoslice (yz0564) through the hilum (hi) and micropyle (mi) in the median plane showing the gradual shortening of the exotestal cells above the micropyle and palisade-shaped sclerenchyma cells lining the micropylar slit (S170234). d) Longitudinal orthoslice (1322) through apical part of seed in the median plane (holotype, S174820) showing embryo with two rudimentary cotyledons (asterisks); note the tiny cells of embryo containing structures we interpret as the fossilized remains of nuclei. e) Longitudinal orthoslice (yz1170) through the apical part of seed in the median plane showing tiny embryo with two rudimentary cotyledons (asterisks) and remains of nutritive tissue with scattered granules that probably representing the remains of protein or lipid bodies; nutritive tissue immediately surrounding embryo is probably endosperm (endo) (S174821). f) Longitudinal orthoslice (xz0590) through apical part of seed perpendicular to the median plane showing the empty space formerly occupied by the embryo, surrounded by cellular nutritive tissue (S174430). g) Transverse orthoslice (xy0969) through the middle part of a seed showing palisade-shaped exotestal cells and remains of mesotesta and tegmen (S175044). Scale bars = 500 µm (a); 250 µm (c–e, g); 125 µm (b, f).
</paragraph>
</caption>
<paragraph id="925136E3FFB4E119FF5F67C6FEF7BDFD" blockId="5.[106,746,151,1909]" pageId="5" pageNumber="140">
Micropyle and hilum are very close together and displaced towards the raphal side of the seed (
<figureCitation id="0AD52A66FFB4E119FD9167E9FFD9BCFC" captionStart="Text-fig" captionStartId="2.[147,228,1713,1734]" captionTargetBox="[147,1462,151,1686]" captionTargetId="figure-2@2.[147,1463,151,1686]" captionTargetPageId="2" captionText="Text-fig. 1. Gastonispermum portugallicum gen. et sp. nov. seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, volume renderings). Note remains of mounting media on several seeds (¤). a) Seed in oblique view showing seed shape, the slightly raised raphal ridge and the position of hilum (hi) and micropyle (mi) on the raphal side of the seed (S170218). b, c) Seeds in lateral view (b, S170234; c, S175095). d–f) Holotype (S174820); seed in lateral view (d) and cut volume rendering (e, f) through the median plane of the seed showing palisade-shaped sclerenchyma cells of exotesta and remains of embryo (emb) and surrounding nutritive tissue (e, cut between yz0440-0530; f, cut between slices yz440-480). g) Hilum (hi) and micropyle (mi) of seed in (1a) showing the Y-shaped micropylar slit in the outer integument. h) Cut volume rendering through the median plane of the seed (cut at yz0492) showing seed coat mainly composed of palisade-shaped cells of the exotesta (S174435). i) Seed surface showing the raised outlines of the undulate anticlinal walls of the exotestal cells (S175045). Scale bars = 500 µm (a–e); 250 µm (g); 125 µm (f, i)." figureDoi="http://doi.org/10.5281/zenodo.5383381" httpUri="https://zenodo.org/record/5383381/files/figure.png" pageId="5" pageNumber="140">Text-fig. 1a– d</figureCitation>
). The hilum is small, circular in outline and slightly raised without a hilar rim (
<figureCitation id="0AD52A66FFB4E119FE8667A9FE68BD1C" box="[329,459,248,272]" captionStart="Text-fig" captionStartId="2.[147,228,1713,1734]" captionTargetBox="[147,1462,151,1686]" captionTargetId="figure-2@2.[147,1463,151,1686]" captionTargetPageId="2" captionText="Text-fig. 1. Gastonispermum portugallicum gen. et sp. nov. seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, volume renderings). Note remains of mounting media on several seeds (¤). a) Seed in oblique view showing seed shape, the slightly raised raphal ridge and the position of hilum (hi) and micropyle (mi) on the raphal side of the seed (S170218). b, c) Seeds in lateral view (b, S170234; c, S175095). d–f) Holotype (S174820); seed in lateral view (d) and cut volume rendering (e, f) through the median plane of the seed showing palisade-shaped sclerenchyma cells of exotesta and remains of embryo (emb) and surrounding nutritive tissue (e, cut between yz0440-0530; f, cut between slices yz440-480). g) Hilum (hi) and micropyle (mi) of seed in (1a) showing the Y-shaped micropylar slit in the outer integument. h) Cut volume rendering through the median plane of the seed (cut at yz0492) showing seed coat mainly composed of palisade-shaped cells of the exotesta (S174435). i) Seed surface showing the raised outlines of the undulate anticlinal walls of the exotestal cells (S175045). Scale bars = 500 µm (a–e); 250 µm (g); 125 µm (f, i)." figureDoi="http://doi.org/10.5281/zenodo.5383381" httpUri="https://zenodo.org/record/5383381/files/figure.png" pageId="5" pageNumber="140">
Text-fig. 
<quantity id="55169B06FFB4E119FE6067A9FE68BD1C" box="[431,459,248,272]" metricMagnitude="-3" metricUnit="kg" metricValue="1.0" pageId="5" pageNumber="140" unit="g" value="1.0">1g</quantity>
</figureCitation>
). The course of the raphe is distinct on the seed surface and seen as a slightly raised, rounded ridge that extends from hilum to the chalazal end of the seed (
<figureCitation id="0AD52A66FFB4E119FF006609FECABD7C" box="[207,361,344,368]" captionStart="Text-fig" captionStartId="2.[147,228,1713,1734]" captionTargetBox="[147,1462,151,1686]" captionTargetId="figure-2@2.[147,1463,151,1686]" captionTargetPageId="2" captionText="Text-fig. 1. Gastonispermum portugallicum gen. et sp. nov. seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, volume renderings). Note remains of mounting media on several seeds (¤). a) Seed in oblique view showing seed shape, the slightly raised raphal ridge and the position of hilum (hi) and micropyle (mi) on the raphal side of the seed (S170218). b, c) Seeds in lateral view (b, S170234; c, S175095). d–f) Holotype (S174820); seed in lateral view (d) and cut volume rendering (e, f) through the median plane of the seed showing palisade-shaped sclerenchyma cells of exotesta and remains of embryo (emb) and surrounding nutritive tissue (e, cut between yz0440-0530; f, cut between slices yz440-480). g) Hilum (hi) and micropyle (mi) of seed in (1a) showing the Y-shaped micropylar slit in the outer integument. h) Cut volume rendering through the median plane of the seed (cut at yz0492) showing seed coat mainly composed of palisade-shaped cells of the exotesta (S174435). i) Seed surface showing the raised outlines of the undulate anticlinal walls of the exotestal cells (S175045). Scale bars = 500 µm (a–e); 250 µm (g); 125 µm (f, i)." figureDoi="http://doi.org/10.5281/zenodo.5383381" httpUri="https://zenodo.org/record/5383381/files/figure.png" pageId="5" pageNumber="140">Text-fig. 1a, b</figureCitation>
). The micropyle is formed from the inner integument. The position of the micropyle is marked by a branched, Y-shaped slit in the testa (
<figureCitation id="0AD52A66FFB4E119FDE366C9FD12BDBC" box="[556,689,408,432]" captionStart="Text-fig" captionStartId="2.[147,228,1713,1734]" captionTargetBox="[147,1462,151,1686]" captionTargetId="figure-2@2.[147,1463,151,1686]" captionTargetPageId="2" captionText="Text-fig. 1. Gastonispermum portugallicum gen. et sp. nov. seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, volume renderings). Note remains of mounting media on several seeds (¤). a) Seed in oblique view showing seed shape, the slightly raised raphal ridge and the position of hilum (hi) and micropyle (mi) on the raphal side of the seed (S170218). b, c) Seeds in lateral view (b, S170234; c, S175095). d–f) Holotype (S174820); seed in lateral view (d) and cut volume rendering (e, f) through the median plane of the seed showing palisade-shaped sclerenchyma cells of exotesta and remains of embryo (emb) and surrounding nutritive tissue (e, cut between yz0440-0530; f, cut between slices yz440-480). g) Hilum (hi) and micropyle (mi) of seed in (1a) showing the Y-shaped micropylar slit in the outer integument. h) Cut volume rendering through the median plane of the seed (cut at yz0492) showing seed coat mainly composed of palisade-shaped cells of the exotesta (S174435). i) Seed surface showing the raised outlines of the undulate anticlinal walls of the exotestal cells (S175045). Scale bars = 500 µm (a–e); 250 µm (g); 125 µm (f, i)." figureDoi="http://doi.org/10.5281/zenodo.5383381" httpUri="https://zenodo.org/record/5383381/files/figure.png" pageId="5" pageNumber="140">
Text-figs 
<quantity id="55169B06FFB4E119FD5A66C9FD12BDBC" box="[661,689,408,432]" metricMagnitude="-3" metricUnit="kg" metricValue="1.0" pageId="5" pageNumber="140" unit="g" value="1.0">1g</quantity>
</figureCitation>
, 
<figureCitation id="0AD52A66FFB4E119FD7166C9FD7FBDBC" box="[702,732,408,432]" captionStart="Text-fig" captionStartId="3.[125,205,1667,1688]" captionText="Text-fig. 2. Gastonispermum portugallicum gen. et sp. nov. seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, orthoslices). a) Longitudinal orthoslice (yz0745) through the median plane of the seed (holotype, S174820) showing taller exotestal cells on raphal side, well preserved embryo (em) and partly preserved nutritive tissue. b) Longitudinal orthoslice (xz1355) through the micropylar region perpendicular to the median plane showing the exotestal tissue surrounding the micropyle (mi) forming a Y-shaped micropylar slit (S105218). c) Longitudinal orthoslice (yz0564) through the hilum (hi) and micropyle (mi) in the median plane showing the gradual shortening of the exotestal cells above the micropyle and palisade-shaped sclerenchyma cells lining the micropylar slit (S170234). d) Longitudinal orthoslice (1322) through apical part of seed in the median plane (holotype, S174820) showing embryo with two rudimentary cotyledons (asterisks); note the tiny cells of embryo containing structures we interpret as the fossilized remains of nuclei. e) Longitudinal orthoslice (yz1170) through the apical part of seed in the median plane showing tiny embryo with two rudimentary cotyledons (asterisks) and remains of nutritive tissue with scattered granules that probably representing the remains of protein or lipid bodies; nutritive tissue immediately surrounding embryo is probably endosperm (endo) (S174821). f) Longitudinal orthoslice (xz0590) through apical part of seed perpendicular to the median plane showing the empty space formerly occupied by the embryo, surrounded by cellular nutritive tissue (S174430). g) Transverse orthoslice (xy0969) through the middle part of a seed showing palisade-shaped exotestal cells and remains of mesotesta and tegmen (S175044). Scale bars = 500 µm (a); 250 µm (c–e, g); 125 µm (b, f)." pageId="5" pageNumber="140">2b</figureCitation>
). Internally the micropylar slit is lined by testal cells on all sides (
<figureCitation id="0AD52A66FFB4E119FF626688FEE4BDFD" box="[173,327,473,497]" captionStart="Text-fig" captionStartId="3.[125,205,1667,1688]" captionText="Text-fig. 2. Gastonispermum portugallicum gen. et sp. nov. seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, orthoslices). a) Longitudinal orthoslice (yz0745) through the median plane of the seed (holotype, S174820) showing taller exotestal cells on raphal side, well preserved embryo (em) and partly preserved nutritive tissue. b) Longitudinal orthoslice (xz1355) through the micropylar region perpendicular to the median plane showing the exotestal tissue surrounding the micropyle (mi) forming a Y-shaped micropylar slit (S105218). c) Longitudinal orthoslice (yz0564) through the hilum (hi) and micropyle (mi) in the median plane showing the gradual shortening of the exotestal cells above the micropyle and palisade-shaped sclerenchyma cells lining the micropylar slit (S170234). d) Longitudinal orthoslice (1322) through apical part of seed in the median plane (holotype, S174820) showing embryo with two rudimentary cotyledons (asterisks); note the tiny cells of embryo containing structures we interpret as the fossilized remains of nuclei. e) Longitudinal orthoslice (yz1170) through the apical part of seed in the median plane showing tiny embryo with two rudimentary cotyledons (asterisks) and remains of nutritive tissue with scattered granules that probably representing the remains of protein or lipid bodies; nutritive tissue immediately surrounding embryo is probably endosperm (endo) (S174821). f) Longitudinal orthoslice (xz0590) through apical part of seed perpendicular to the median plane showing the empty space formerly occupied by the embryo, surrounded by cellular nutritive tissue (S174430). g) Transverse orthoslice (xy0969) through the middle part of a seed showing palisade-shaped exotestal cells and remains of mesotesta and tegmen (S175044). Scale bars = 500 µm (a); 250 µm (c–e, g); 125 µm (b, f)." pageId="5" pageNumber="140">Text-fig. 2a, b</figureCitation>
).
</paragraph>
<paragraph id="925136E3FFB4E119FF5F66A8FD0CB8BF" blockId="5.[106,746,151,1909]" pageId="5" pageNumber="140">
The seed coat is composed of a thick exotesta, a thinner mesotesta/endotesta, and a thin tegmen. The exotesta consists of a single layer of tall, columnar sclerenchyma that vary in height from about 55 µm laterally and on the antiraphal sides of the seed, to about 80 µm over the raphe (
<figureCitation id="0AD52A66FFB4E119FD8A6528FD47BE9D" box="[581,740,633,657]" captionStart="Text-fig" captionStartId="2.[147,228,1713,1734]" captionTargetBox="[147,1462,151,1686]" captionTargetId="figure-2@2.[147,1463,151,1686]" captionTargetPageId="2" captionText="Text-fig. 1. Gastonispermum portugallicum gen. et sp. nov. seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, volume renderings). Note remains of mounting media on several seeds (¤). a) Seed in oblique view showing seed shape, the slightly raised raphal ridge and the position of hilum (hi) and micropyle (mi) on the raphal side of the seed (S170218). b, c) Seeds in lateral view (b, S170234; c, S175095). d–f) Holotype (S174820); seed in lateral view (d) and cut volume rendering (e, f) through the median plane of the seed showing palisade-shaped sclerenchyma cells of exotesta and remains of embryo (emb) and surrounding nutritive tissue (e, cut between yz0440-0530; f, cut between slices yz440-480). g) Hilum (hi) and micropyle (mi) of seed in (1a) showing the Y-shaped micropylar slit in the outer integument. h) Cut volume rendering through the median plane of the seed (cut at yz0492) showing seed coat mainly composed of palisade-shaped cells of the exotesta (S174435). i) Seed surface showing the raised outlines of the undulate anticlinal walls of the exotestal cells (S175045). Scale bars = 500 µm (a–e); 250 µm (g); 125 µm (f, i)." figureDoi="http://doi.org/10.5281/zenodo.5383381" httpUri="https://zenodo.org/record/5383381/files/figure.png" pageId="5" pageNumber="140">Text-figs 1e, h</figureCitation>
, 
<figureCitation id="0AD52A66FFB4E119FFA565C8FF02BEBD" box="[106,161,665,689]" captionStart="Text-fig" captionStartId="3.[125,205,1667,1688]" captionText="Text-fig. 2. Gastonispermum portugallicum gen. et sp. nov. seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, orthoslices). a) Longitudinal orthoslice (yz0745) through the median plane of the seed (holotype, S174820) showing taller exotestal cells on raphal side, well preserved embryo (em) and partly preserved nutritive tissue. b) Longitudinal orthoslice (xz1355) through the micropylar region perpendicular to the median plane showing the exotestal tissue surrounding the micropyle (mi) forming a Y-shaped micropylar slit (S105218). c) Longitudinal orthoslice (yz0564) through the hilum (hi) and micropyle (mi) in the median plane showing the gradual shortening of the exotestal cells above the micropyle and palisade-shaped sclerenchyma cells lining the micropylar slit (S170234). d) Longitudinal orthoslice (1322) through apical part of seed in the median plane (holotype, S174820) showing embryo with two rudimentary cotyledons (asterisks); note the tiny cells of embryo containing structures we interpret as the fossilized remains of nuclei. e) Longitudinal orthoslice (yz1170) through the apical part of seed in the median plane showing tiny embryo with two rudimentary cotyledons (asterisks) and remains of nutritive tissue with scattered granules that probably representing the remains of protein or lipid bodies; nutritive tissue immediately surrounding embryo is probably endosperm (endo) (S174821). f) Longitudinal orthoslice (xz0590) through apical part of seed perpendicular to the median plane showing the empty space formerly occupied by the embryo, surrounded by cellular nutritive tissue (S174430). g) Transverse orthoslice (xy0969) through the middle part of a seed showing palisade-shaped exotestal cells and remains of mesotesta and tegmen (S175044). Scale bars = 500 µm (a); 250 µm (c–e, g); 125 µm (b, f)." pageId="5" pageNumber="140">2a, g</figureCitation>
). In the micropylar region the columnar sclerenchyma cells are shorter and become much shorter towards the micropylar slit (
<figureCitation id="0AD52A66FFB4E119FEDE6588FE1CBEFD" box="[273,447,729,754]" captionStart="Text-fig" captionStartId="2.[147,228,1713,1734]" captionTargetBox="[147,1462,151,1686]" captionTargetId="figure-2@2.[147,1463,151,1686]" captionTargetPageId="2" captionText="Text-fig. 1. Gastonispermum portugallicum gen. et sp. nov. seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, volume renderings). Note remains of mounting media on several seeds (¤). a) Seed in oblique view showing seed shape, the slightly raised raphal ridge and the position of hilum (hi) and micropyle (mi) on the raphal side of the seed (S170218). b, c) Seeds in lateral view (b, S170234; c, S175095). d–f) Holotype (S174820); seed in lateral view (d) and cut volume rendering (e, f) through the median plane of the seed showing palisade-shaped sclerenchyma cells of exotesta and remains of embryo (emb) and surrounding nutritive tissue (e, cut between yz0440-0530; f, cut between slices yz440-480). g) Hilum (hi) and micropyle (mi) of seed in (1a) showing the Y-shaped micropylar slit in the outer integument. h) Cut volume rendering through the median plane of the seed (cut at yz0492) showing seed coat mainly composed of palisade-shaped cells of the exotesta (S174435). i) Seed surface showing the raised outlines of the undulate anticlinal walls of the exotestal cells (S175045). Scale bars = 500 µm (a–e); 250 µm (g); 125 µm (f, i)." figureDoi="http://doi.org/10.5281/zenodo.5383381" httpUri="https://zenodo.org/record/5383381/files/figure.png" pageId="5" pageNumber="140">Text-figs 1e, f, h</figureCitation>
, 
<figureCitation id="0AD52A66FFB4E119FE06658BFDB0BEFE" box="[457,531,730,754]" captionStart="Text-fig" captionStartId="3.[125,205,1667,1688]" captionText="Text-fig. 2. Gastonispermum portugallicum gen. et sp. nov. seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, orthoslices). a) Longitudinal orthoslice (yz0745) through the median plane of the seed (holotype, S174820) showing taller exotestal cells on raphal side, well preserved embryo (em) and partly preserved nutritive tissue. b) Longitudinal orthoslice (xz1355) through the micropylar region perpendicular to the median plane showing the exotestal tissue surrounding the micropyle (mi) forming a Y-shaped micropylar slit (S105218). c) Longitudinal orthoslice (yz0564) through the hilum (hi) and micropyle (mi) in the median plane showing the gradual shortening of the exotestal cells above the micropyle and palisade-shaped sclerenchyma cells lining the micropylar slit (S170234). d) Longitudinal orthoslice (1322) through apical part of seed in the median plane (holotype, S174820) showing embryo with two rudimentary cotyledons (asterisks); note the tiny cells of embryo containing structures we interpret as the fossilized remains of nuclei. e) Longitudinal orthoslice (yz1170) through the apical part of seed in the median plane showing tiny embryo with two rudimentary cotyledons (asterisks) and remains of nutritive tissue with scattered granules that probably representing the remains of protein or lipid bodies; nutritive tissue immediately surrounding embryo is probably endosperm (endo) (S174821). f) Longitudinal orthoslice (xz0590) through apical part of seed perpendicular to the median plane showing the empty space formerly occupied by the embryo, surrounded by cellular nutritive tissue (S174430). g) Transverse orthoslice (xy0969) through the middle part of a seed showing palisade-shaped exotestal cells and remains of mesotesta and tegmen (S175044). Scale bars = 500 µm (a); 250 µm (c–e, g); 125 µm (b, f)." pageId="5" pageNumber="140">2a, c, e</figureCitation>
). The exotestal cells are arranged in indistinct longitudinal rows. The anticlinal walls of the exotestal cells are thickened, slightly raised on the surface, and of almost even thickness from the outside to the inside, resulting in an almost straight lumen (
<figureCitation id="0AD52A66FFB4E119FD47640BFF25BF9E" captionStart="Text-fig" captionStartId="2.[147,228,1713,1734]" captionTargetBox="[147,1462,151,1686]" captionTargetId="figure-2@2.[147,1463,151,1686]" captionTargetPageId="2" captionText="Text-fig. 1. Gastonispermum portugallicum gen. et sp. nov. seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, volume renderings). Note remains of mounting media on several seeds (¤). a) Seed in oblique view showing seed shape, the slightly raised raphal ridge and the position of hilum (hi) and micropyle (mi) on the raphal side of the seed (S170218). b, c) Seeds in lateral view (b, S170234; c, S175095). d–f) Holotype (S174820); seed in lateral view (d) and cut volume rendering (e, f) through the median plane of the seed showing palisade-shaped sclerenchyma cells of exotesta and remains of embryo (emb) and surrounding nutritive tissue (e, cut between yz0440-0530; f, cut between slices yz440-480). g) Hilum (hi) and micropyle (mi) of seed in (1a) showing the Y-shaped micropylar slit in the outer integument. h) Cut volume rendering through the median plane of the seed (cut at yz0492) showing seed coat mainly composed of palisade-shaped cells of the exotesta (S174435). i) Seed surface showing the raised outlines of the undulate anticlinal walls of the exotestal cells (S175045). Scale bars = 500 µm (a–e); 250 µm (g); 125 µm (f, i)." figureDoi="http://doi.org/10.5281/zenodo.5383381" httpUri="https://zenodo.org/record/5383381/files/figure.png" pageId="5" pageNumber="140">Text-figs 1h</figureCitation>
, 
<figureCitation id="0AD52A66FFB4E119FF5C642BFF69BF9E" box="[147,202,890,914]" captionStart="Text-fig" captionStartId="3.[125,205,1667,1688]" captionText="Text-fig. 2. Gastonispermum portugallicum gen. et sp. nov. seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, orthoslices). a) Longitudinal orthoslice (yz0745) through the median plane of the seed (holotype, S174820) showing taller exotestal cells on raphal side, well preserved embryo (em) and partly preserved nutritive tissue. b) Longitudinal orthoslice (xz1355) through the micropylar region perpendicular to the median plane showing the exotestal tissue surrounding the micropyle (mi) forming a Y-shaped micropylar slit (S105218). c) Longitudinal orthoslice (yz0564) through the hilum (hi) and micropyle (mi) in the median plane showing the gradual shortening of the exotestal cells above the micropyle and palisade-shaped sclerenchyma cells lining the micropylar slit (S170234). d) Longitudinal orthoslice (1322) through apical part of seed in the median plane (holotype, S174820) showing embryo with two rudimentary cotyledons (asterisks); note the tiny cells of embryo containing structures we interpret as the fossilized remains of nuclei. e) Longitudinal orthoslice (yz1170) through the apical part of seed in the median plane showing tiny embryo with two rudimentary cotyledons (asterisks) and remains of nutritive tissue with scattered granules that probably representing the remains of protein or lipid bodies; nutritive tissue immediately surrounding embryo is probably endosperm (endo) (S174821). f) Longitudinal orthoslice (xz0590) through apical part of seed perpendicular to the median plane showing the empty space formerly occupied by the embryo, surrounded by cellular nutritive tissue (S174430). g) Transverse orthoslice (xy0969) through the middle part of a seed showing palisade-shaped exotestal cells and remains of mesotesta and tegmen (S175044). Scale bars = 500 µm (a); 250 µm (c–e, g); 125 µm (b, f)." pageId="5" pageNumber="140">2a, g</figureCitation>
). Towards the outside and inside they are strongly undulate resulting in stellate-undulate facets and a jigsaw puzzle-like pattern on the surface with rounded, undivided lobes (
<figureCitation id="0AD52A66FFB4E119FF60648BFE84BFFE" box="[175,295,986,1010]" captionStart="Text-fig" captionStartId="2.[147,228,1713,1734]" captionTargetBox="[147,1462,151,1686]" captionTargetId="figure-2@2.[147,1463,151,1686]" captionTargetPageId="2" captionText="Text-fig. 1. Gastonispermum portugallicum gen. et sp. nov. seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, volume renderings). Note remains of mounting media on several seeds (¤). a) Seed in oblique view showing seed shape, the slightly raised raphal ridge and the position of hilum (hi) and micropyle (mi) on the raphal side of the seed (S170218). b, c) Seeds in lateral view (b, S170234; c, S175095). d–f) Holotype (S174820); seed in lateral view (d) and cut volume rendering (e, f) through the median plane of the seed showing palisade-shaped sclerenchyma cells of exotesta and remains of embryo (emb) and surrounding nutritive tissue (e, cut between yz0440-0530; f, cut between slices yz440-480). g) Hilum (hi) and micropyle (mi) of seed in (1a) showing the Y-shaped micropylar slit in the outer integument. h) Cut volume rendering through the median plane of the seed (cut at yz0492) showing seed coat mainly composed of palisade-shaped cells of the exotesta (S174435). i) Seed surface showing the raised outlines of the undulate anticlinal walls of the exotestal cells (S175045). Scale bars = 500 µm (a–e); 250 µm (g); 125 µm (f, i)." figureDoi="http://doi.org/10.5281/zenodo.5383381" httpUri="https://zenodo.org/record/5383381/files/figure.png" pageId="5" pageNumber="140">Text-fig. 1i</figureCitation>
). Exotestal cells are also undulate over the raphe. Exotesta cells around the micropylar slit are slender and tall (
<figureCitation id="0AD52A66FFB4E119FF03634BFEEDB83F" box="[204,334,1050,1075]" captionStart="Text-fig" captionStartId="3.[125,205,1667,1688]" captionText="Text-fig. 2. Gastonispermum portugallicum gen. et sp. nov. seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, orthoslices). a) Longitudinal orthoslice (yz0745) through the median plane of the seed (holotype, S174820) showing taller exotestal cells on raphal side, well preserved embryo (em) and partly preserved nutritive tissue. b) Longitudinal orthoslice (xz1355) through the micropylar region perpendicular to the median plane showing the exotestal tissue surrounding the micropyle (mi) forming a Y-shaped micropylar slit (S105218). c) Longitudinal orthoslice (yz0564) through the hilum (hi) and micropyle (mi) in the median plane showing the gradual shortening of the exotestal cells above the micropyle and palisade-shaped sclerenchyma cells lining the micropylar slit (S170234). d) Longitudinal orthoslice (1322) through apical part of seed in the median plane (holotype, S174820) showing embryo with two rudimentary cotyledons (asterisks); note the tiny cells of embryo containing structures we interpret as the fossilized remains of nuclei. e) Longitudinal orthoslice (yz1170) through the apical part of seed in the median plane showing tiny embryo with two rudimentary cotyledons (asterisks) and remains of nutritive tissue with scattered granules that probably representing the remains of protein or lipid bodies; nutritive tissue immediately surrounding embryo is probably endosperm (endo) (S174821). f) Longitudinal orthoslice (xz0590) through apical part of seed perpendicular to the median plane showing the empty space formerly occupied by the embryo, surrounded by cellular nutritive tissue (S174430). g) Transverse orthoslice (xy0969) through the middle part of a seed showing palisade-shaped exotestal cells and remains of mesotesta and tegmen (S175044). Scale bars = 500 µm (a); 250 µm (c–e, g); 125 µm (b, f)." pageId="5" pageNumber="140">Text-fig. 2c</figureCitation>
). The mesotesta/endotesta consists of two or three layers of collapsed parenchyma cells with thick cell walls (
<figureCitation id="0AD52A66FFB4E119FF2E630AFEC0B87F" box="[225,355,1115,1139]" captionStart="Text-fig" captionStartId="3.[125,205,1667,1688]" captionText="Text-fig. 2. Gastonispermum portugallicum gen. et sp. nov. seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, orthoslices). a) Longitudinal orthoslice (yz0745) through the median plane of the seed (holotype, S174820) showing taller exotestal cells on raphal side, well preserved embryo (em) and partly preserved nutritive tissue. b) Longitudinal orthoslice (xz1355) through the micropylar region perpendicular to the median plane showing the exotestal tissue surrounding the micropyle (mi) forming a Y-shaped micropylar slit (S105218). c) Longitudinal orthoslice (yz0564) through the hilum (hi) and micropyle (mi) in the median plane showing the gradual shortening of the exotestal cells above the micropyle and palisade-shaped sclerenchyma cells lining the micropylar slit (S170234). d) Longitudinal orthoslice (1322) through apical part of seed in the median plane (holotype, S174820) showing embryo with two rudimentary cotyledons (asterisks); note the tiny cells of embryo containing structures we interpret as the fossilized remains of nuclei. e) Longitudinal orthoslice (yz1170) through the apical part of seed in the median plane showing tiny embryo with two rudimentary cotyledons (asterisks) and remains of nutritive tissue with scattered granules that probably representing the remains of protein or lipid bodies; nutritive tissue immediately surrounding embryo is probably endosperm (endo) (S174821). f) Longitudinal orthoslice (xz0590) through apical part of seed perpendicular to the median plane showing the empty space formerly occupied by the embryo, surrounded by cellular nutritive tissue (S174430). g) Transverse orthoslice (xy0969) through the middle part of a seed showing palisade-shaped exotestal cells and remains of mesotesta and tegmen (S175044). Scale bars = 500 µm (a); 250 µm (c–e, g); 125 µm (b, f)." pageId="5" pageNumber="140">Text-fig. 2a</figureCitation>
). The tegmen is thin and composed of cells that are collapsed over most of the seed, but more robust and slightly thickened in the micropylar region.
</paragraph>
<paragraph id="925136E3FFB4E119FF5F63EAFD92BB79" blockId="5.[106,746,151,1909]" pageId="5" pageNumber="140">
Nutritive tissue is partly preserved in five of the specimens that were studied using SRXTM. Two of these specimens (S174820, S174821) also had a complete embryo preserved. In both cases the embryo is tiny, about 130 µm long and 130 µm broad, with two rudimentary cotyledons and an embryo to seed (E: S) ratio of about 0.02. The embryo is composed of cells that are much smaller than the surrounding nutritive tissue. Each cell contains a dense, central structure that may be the fossilized remains of a nucleus (
<figureCitation id="0AD52A66FFB4E119FDC662EDFD01B9D8" box="[521,674,1468,1492]" captionStart="Text-fig" captionStartId="2.[147,228,1713,1734]" captionTargetBox="[147,1462,151,1686]" captionTargetId="figure-2@2.[147,1463,151,1686]" captionTargetPageId="2" captionText="Text-fig. 1. Gastonispermum portugallicum gen. et sp. nov. seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, volume renderings). Note remains of mounting media on several seeds (¤). a) Seed in oblique view showing seed shape, the slightly raised raphal ridge and the position of hilum (hi) and micropyle (mi) on the raphal side of the seed (S170218). b, c) Seeds in lateral view (b, S170234; c, S175095). d–f) Holotype (S174820); seed in lateral view (d) and cut volume rendering (e, f) through the median plane of the seed showing palisade-shaped sclerenchyma cells of exotesta and remains of embryo (emb) and surrounding nutritive tissue (e, cut between yz0440-0530; f, cut between slices yz440-480). g) Hilum (hi) and micropyle (mi) of seed in (1a) showing the Y-shaped micropylar slit in the outer integument. h) Cut volume rendering through the median plane of the seed (cut at yz0492) showing seed coat mainly composed of palisade-shaped cells of the exotesta (S174435). i) Seed surface showing the raised outlines of the undulate anticlinal walls of the exotestal cells (S175045). Scale bars = 500 µm (a–e); 250 µm (g); 125 µm (f, i)." figureDoi="http://doi.org/10.5281/zenodo.5383381" httpUri="https://zenodo.org/record/5383381/files/figure.png" pageId="5" pageNumber="140">Text-figs 1e, f</figureCitation>
, 
<figureCitation id="0AD52A66FFB4E119FD6162EDFFDBB9F8" captionStart="Text-fig" captionStartId="3.[125,205,1667,1688]" captionText="Text-fig. 2. Gastonispermum portugallicum gen. et sp. nov. seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, orthoslices). a) Longitudinal orthoslice (yz0745) through the median plane of the seed (holotype, S174820) showing taller exotestal cells on raphal side, well preserved embryo (em) and partly preserved nutritive tissue. b) Longitudinal orthoslice (xz1355) through the micropylar region perpendicular to the median plane showing the exotestal tissue surrounding the micropyle (mi) forming a Y-shaped micropylar slit (S105218). c) Longitudinal orthoslice (yz0564) through the hilum (hi) and micropyle (mi) in the median plane showing the gradual shortening of the exotestal cells above the micropyle and palisade-shaped sclerenchyma cells lining the micropylar slit (S170234). d) Longitudinal orthoslice (1322) through apical part of seed in the median plane (holotype, S174820) showing embryo with two rudimentary cotyledons (asterisks); note the tiny cells of embryo containing structures we interpret as the fossilized remains of nuclei. e) Longitudinal orthoslice (yz1170) through the apical part of seed in the median plane showing tiny embryo with two rudimentary cotyledons (asterisks) and remains of nutritive tissue with scattered granules that probably representing the remains of protein or lipid bodies; nutritive tissue immediately surrounding embryo is probably endosperm (endo) (S174821). f) Longitudinal orthoslice (xz0590) through apical part of seed perpendicular to the median plane showing the empty space formerly occupied by the embryo, surrounded by cellular nutritive tissue (S174430). g) Transverse orthoslice (xy0969) through the middle part of a seed showing palisade-shaped exotestal cells and remains of mesotesta and tegmen (S175044). Scale bars = 500 µm (a); 250 µm (c–e, g); 125 µm (b, f)." pageId="5" pageNumber="140">2a, d, e</figureCitation>
). The nutritive tissue is cellular, consisting of isodiametric and thin-walled cells, about 40 µm in diameter (
<figureCitation id="0AD52A66FFB4E119FD4362ADFF1EBA38" captionStart="Text-fig" captionStartId="3.[125,205,1667,1688]" captionText="Text-fig. 2. Gastonispermum portugallicum gen. et sp. nov. seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, orthoslices). a) Longitudinal orthoslice (yz0745) through the median plane of the seed (holotype, S174820) showing taller exotestal cells on raphal side, well preserved embryo (em) and partly preserved nutritive tissue. b) Longitudinal orthoslice (xz1355) through the micropylar region perpendicular to the median plane showing the exotestal tissue surrounding the micropyle (mi) forming a Y-shaped micropylar slit (S105218). c) Longitudinal orthoslice (yz0564) through the hilum (hi) and micropyle (mi) in the median plane showing the gradual shortening of the exotestal cells above the micropyle and palisade-shaped sclerenchyma cells lining the micropylar slit (S170234). d) Longitudinal orthoslice (1322) through apical part of seed in the median plane (holotype, S174820) showing embryo with two rudimentary cotyledons (asterisks); note the tiny cells of embryo containing structures we interpret as the fossilized remains of nuclei. e) Longitudinal orthoslice (yz1170) through the apical part of seed in the median plane showing tiny embryo with two rudimentary cotyledons (asterisks) and remains of nutritive tissue with scattered granules that probably representing the remains of protein or lipid bodies; nutritive tissue immediately surrounding embryo is probably endosperm (endo) (S174821). f) Longitudinal orthoslice (xz0590) through apical part of seed perpendicular to the median plane showing the empty space formerly occupied by the embryo, surrounded by cellular nutritive tissue (S174430). g) Transverse orthoslice (xy0969) through the middle part of a seed showing palisade-shaped exotestal cells and remains of mesotesta and tegmen (S175044). Scale bars = 500 µm (a); 250 µm (c–e, g); 125 µm (b, f)." pageId="5" pageNumber="140">Text-fig. 2a, e, f</figureCitation>
). Cell walls are slightly undulating or straight. In some specimens the cells of the nutritive tissue are empty (
<figureCitation id="0AD52A66FFB4E119FFBE610DFF4FBA78" box="[113,236,1628,1652]" captionStart="Text-fig" captionStartId="3.[125,205,1667,1688]" captionText="Text-fig. 2. Gastonispermum portugallicum gen. et sp. nov. seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, orthoslices). a) Longitudinal orthoslice (yz0745) through the median plane of the seed (holotype, S174820) showing taller exotestal cells on raphal side, well preserved embryo (em) and partly preserved nutritive tissue. b) Longitudinal orthoslice (xz1355) through the micropylar region perpendicular to the median plane showing the exotestal tissue surrounding the micropyle (mi) forming a Y-shaped micropylar slit (S105218). c) Longitudinal orthoslice (yz0564) through the hilum (hi) and micropyle (mi) in the median plane showing the gradual shortening of the exotestal cells above the micropyle and palisade-shaped sclerenchyma cells lining the micropylar slit (S170234). d) Longitudinal orthoslice (1322) through apical part of seed in the median plane (holotype, S174820) showing embryo with two rudimentary cotyledons (asterisks); note the tiny cells of embryo containing structures we interpret as the fossilized remains of nuclei. e) Longitudinal orthoslice (yz1170) through the apical part of seed in the median plane showing tiny embryo with two rudimentary cotyledons (asterisks) and remains of nutritive tissue with scattered granules that probably representing the remains of protein or lipid bodies; nutritive tissue immediately surrounding embryo is probably endosperm (endo) (S174821). f) Longitudinal orthoslice (xz0590) through apical part of seed perpendicular to the median plane showing the empty space formerly occupied by the embryo, surrounded by cellular nutritive tissue (S174430). g) Transverse orthoslice (xy0969) through the middle part of a seed showing palisade-shaped exotestal cells and remains of mesotesta and tegmen (S175044). Scale bars = 500 µm (a); 250 µm (c–e, g); 125 µm (b, f)." pageId="5" pageNumber="140">Text-fig. 2f</figureCitation>
), but in other cases these cells are filled by small granules. These granules are often clustered close to the cell walls (
<figureCitation id="0AD52A66FFB4E119FF7F61CCFE8DBAB9" box="[176,302,1693,1717]" captionStart="Text-fig" captionStartId="3.[125,205,1667,1688]" captionText="Text-fig. 2. Gastonispermum portugallicum gen. et sp. nov. seeds from the Early Cretaceous Famalicão locality (sample 025), Portugal; Synchrotron radiation X-ray tomographic microscopy (SRXTM, orthoslices). a) Longitudinal orthoslice (yz0745) through the median plane of the seed (holotype, S174820) showing taller exotestal cells on raphal side, well preserved embryo (em) and partly preserved nutritive tissue. b) Longitudinal orthoslice (xz1355) through the micropylar region perpendicular to the median plane showing the exotestal tissue surrounding the micropyle (mi) forming a Y-shaped micropylar slit (S105218). c) Longitudinal orthoslice (yz0564) through the hilum (hi) and micropyle (mi) in the median plane showing the gradual shortening of the exotestal cells above the micropyle and palisade-shaped sclerenchyma cells lining the micropylar slit (S170234). d) Longitudinal orthoslice (1322) through apical part of seed in the median plane (holotype, S174820) showing embryo with two rudimentary cotyledons (asterisks); note the tiny cells of embryo containing structures we interpret as the fossilized remains of nuclei. e) Longitudinal orthoslice (yz1170) through the apical part of seed in the median plane showing tiny embryo with two rudimentary cotyledons (asterisks) and remains of nutritive tissue with scattered granules that probably representing the remains of protein or lipid bodies; nutritive tissue immediately surrounding embryo is probably endosperm (endo) (S174821). f) Longitudinal orthoslice (xz0590) through apical part of seed perpendicular to the median plane showing the empty space formerly occupied by the embryo, surrounded by cellular nutritive tissue (S174430). g) Transverse orthoslice (xy0969) through the middle part of a seed showing palisade-shaped exotestal cells and remains of mesotesta and tegmen (S175044). Scale bars = 500 µm (a); 250 µm (c–e, g); 125 µm (b, f)." pageId="5" pageNumber="140">Text-fig. 2e</figureCitation>
) and we interpret them to be the remains of protein or lipid bodies. We interpret the nutritive tissue immediately surrounding the embryo as endosperm (Textfig. 2e), but the continuity of this inner tissue with the bulk of the nutritive tissue in the outer parts of the seed is not clear, and thus the possibility that the outer tissues may be perisperm cannot be definitively excluded.
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