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<document ID-DOI="10.37520/fi.2021.016" ID-ISSN="2533-4069" ID-Zenodo-Dep="7167561" approvalRequired="48" approvalRequired_for_taxonomicNames="27" approvalRequired_for_textStreams="19" approvalRequired_for_treatments="2" checkinTime="1665260427164" checkinUser="felipe" docAuthor="Friis, Else Marie, Crane, Peter R. &amp; Pedersen, Kaj Raunsgaard" docDate="2021" docId="03F487DBFF8F246FFC54FA39FA86C40F" docLanguage="en" docName="FossilImprint.77.2.214-230.pdf" docOrigin="Fossil Imprint 77 (2)" docSource="http://dx.doi.org/10.37520/fi.2021.016" docStyle="DocumentStyle:0EC54BF35E227CA4627F894F6BA30634.2:FossilImprint.2016-.journal_article" docStyleId="0EC54BF35E227CA4627F894F6BA30634" docStyleName="FossilImprint.2016-.journal_article" docStyleVersion="2" docTitle="Miranthus elegans E. M. FRIIS, P. R. CRANE et K. R. PEDERSEN 2021, sp. nov." docType="treatment" docVersion="2" lastPageNumber="221" masterDocId="FFCDFFA3FF8D2468FFF2FF81FFF7CC3C" masterDocTitle="Early Flowers Of Primuloid Ericales From The Late Cretaceous Of Portugal And Their Ecological And Phytogeographic Implications" masterLastPageNumber="230" masterPageNumber="214" pageNumber="216" updateTime="1665273363288" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-NC-ND-3.0">
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<mods:titleInfo>
<mods:title>Early Flowers Of Primuloid Ericales From The Late Cretaceous Of Portugal And Their Ecological And Phytogeographic Implications</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Friis, Else Marie</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Crane, Peter R.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Pedersen, Kaj Raunsgaard</mods:namePart>
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<mods:title>Fossil Imprint</mods:title>
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<mods:date>2021</mods:date>
<mods:detail type="pubDate">
<mods:number>2021-12-24</mods:number>
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<mods:detail type="volume">
<mods:number>77</mods:number>
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<mods:detail type="issue">
<mods:number>2</mods:number>
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<mods:url>http://dx.doi.org/10.37520/fi.2021.016</mods:url>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">10.37520/fi.2021.016</mods:identifier>
<mods:identifier type="ISSN">2533-4069</mods:identifier>
<mods:identifier type="Zenodo-Dep">7167561</mods:identifier>
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<treatment LSID="urn:lsid:plazi:treatment:03F487DBFF8F246FFC54FA39FA86C40F" httpUri="http://treatment.plazi.org/id/03F487DBFF8F246FFC54FA39FA86C40F" lastPageId="7" lastPageNumber="221" pageId="2" pageNumber="216">
<subSubSection pageId="2" pageNumber="216" type="nomenclature">
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<taxonomicName authority="E.M.FRIIS, P.R.CRANE et K.R.PEDERSEN" authorityName="E. M. FRIIS, P. R. CRANE et K. R. PEDERSEN" authorityYear="2021" class="Magnoliopsida" family="Primulaceae" genus="Miranthus" kingdom="Plantae" order="Ericales" pageId="2" pageNumber="216" phylum="Tracheophyta" rank="species" species="elegans" status="sp. nov.">
<emphasis bold="true" box="[934,1141,1464,1488]" italics="true" pageId="2" pageNumber="216">Miranthus elegans</emphasis>
E.M.FRIIS, P.R.CRANE et K.R.PEDERSEN
</taxonomicName>
<taxonomicNameLabel box="[1216,1301,1497,1521]" pageId="2" pageNumber="216" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
<paragraph blockId="2.[1092,1232,1529,1553]" box="[1092,1232,1529,1553]" pageId="2" pageNumber="216">
<figureCitation box="[1092,1232,1529,1553]" captionStart-0="Text-fig" captionStart-1="Text-fig" captionStart-2="Text-fig" captionStart-3="Text-fig" captionStart-4="Text-fig" captionStart-5="Text-fig" captionStartId-0="3.[125,206,1774,1795]" captionStartId-1="4.[147,228,1735,1756]" captionStartId-2="5.[123,204,1939,1960]" captionStartId-3="7.[125,207,1562,1583]" captionStartId-4="8.[148,226,1363,1384]" captionStartId-5="9.[125,206,1251,1272]" captionTargetBox-0="[125,1440,151,1758]" captionTargetBox-1="[148,1463,151,1719]" captionTargetBox-2="[123,1438,152,1923]" captionTargetBox-3="[125,1439,151,1546]" captionTargetBox-4="[148,1461,144,1347]" captionTargetBox-5="[125,1440,151,1236]" captionTargetId-0="figure-2@3.[-26,1561,55,2300]" captionTargetId-1="figure-2@4.[-3,1584,55,2300]" captionTargetId-2="figure-2@5.[-28,1559,56,2301]" captionTargetId-3="figure-148@7.[-27,1560,47,2292]" captionTargetId-4="figure-291@8.[-5,1582,47,2292]" captionTargetId-5="figure-297@9.[-26,1561,55,2300]" captionTargetPageId-0="3" captionTargetPageId-1="4" captionTargetPageId-2="5" captionTargetPageId-3="7" captionTargetPageId-4="8" captionTargetPageId-5="9" captionText-0="Text-fig. 1. SEM images of flowers of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Flowers in lateral view showing elongated pedicel, narrowly triangular sepals and elongated protruding style (a); note the large openings in the floral tissue and pedicel (asterisks) interpreted as schizogenous secretory cavities. c: Flower in lateral view with portion of the calyx missing exposing the ovary wall and slightly raised nectariferous ring with probable stomata-like secretory structures (arrow). d: Flower in lateral view showing long pedicel and three of the five tepals; note the elongated narrowly triangular form of the sepals. e: Flower in oblique lateral view with portion of the calyx missing exposing the ovary and elongated style. f, g: Flowers in apical view showing the bases of five sepals (f) and apex of the five-parted ovary; note larger openings in the floral tissue (asterisk) interpreted as schizogenous secretory cavities. Specimens, Mira 100-S170155 (a, holotype), Mira 100-S153145 (b, c, g), Mira 100- S101267 (d), Mira 105-S100732 (e), Mira 100-S101268 (f). Scale bars = 600 µm (a–g)." captionText-1="Text-fig. 2. SEM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Flowers in oblique lateral view showing remains of calyx and slightly semi-inferior ovary with elongated apical style (a); note larger openings in the floral tissue (asterisk) interpreted as schizogenous secretory cavities and the stomata-like secretory structures on the upper portion of the ovary (arrows) that are interpreted as nectariferous (b). c: Detail of ovary surface showing secretory stomata-like structures (arrows). d: Flower in lateral view showing fragmentary calyx and broken slightly semi-inferior ovary with secretory stomata-like structures; note the point of attachment of the central placenta (pl). e: Cluster of seeds removed from the ovary in (d) showing reticulate surface. f: Outer (abaxial)surface of calyx lobe showing the slightly pointed papillae and scattered, fine trichomes (arrows). g: Triaperturate pollen grains from the ovary surface. Specimens, Mira 100-S153146 (a, b), Mira 100-S170155 (c), Mira 100-S101266 (d, e), Mira 105-S100732 (f), Mira 100-S170125 (g). Scale bars = 600 µm (a, b, d), 300 µm (f), 100 µm (c, e), 10 µm (g)." captionText-2="Text-fig. 3. SRXTM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Volume renderings of flower bud in two different lateral views showing long pedicel, distinct calyx (ca) with almost equiaxial epidermal cells and corolla (co) with nearly smooth surface. c–e: Transverse sections (c, orthoslice xy1500; d, orthoslice xy1760; e, orthoslice xy1850) through flower bud at levels below the anthers showing stamen filaments (yellow) opposite the corolla lobes (co) and smaller staminodes (orange)" captionText-3="Text-fig. 4. SRXTM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Transverse (a) and longitudinal (b) sections of flower showing the ovary wall (ow) and numerous ovules (ov) borne on the mushroom-shaped central globose placenta (pl, blue); placenta shown as a voltex rendering added to orthoslices; note the large openings in the floral tissue (asterisks) interpreted as schizogenous secretory cavities. c, d: Longitudinal (c) and transverse (d) sections of flower (c, volume rendering cut between orthoslice yz0970-1005; d, volume rendering cut between orthoslice xy0780-0820) showing semiinferior ovary (ow, ovary wall) with sepals inserted at the rim of the hypanthium, central column (cc) with mushroom-shaped globose placenta (pl) bearing numerous ovules (ov). Specimens, Mira 100-S153146 (a, b), Mira 100-S170155 (c, d, holotype). Scale bars = 600 µm (a–d)." captionText-4="Text-fig. 5. SRXTM imagesof Miranthuselegans gen. et sp. nov.;Mira locality, Portugal. a, b: Transverse sections offlower (a, orthoslice xy0665 close to the apex of placenta; b, orthoslice xy0800 in middle part of placenta) showing remains of calyx with distinct bundles (arrows), ovary wall (ow) and numerous ovules (ov) on the central mushroom-shaped globose placenta (pl) with central column (cc). c: Transverse section of flower (orthoslice xy0620) through perianth and ovary (ow) at a level above the placenta showing ovules (ov) and cellular preservation of the sepal bundles (arrows shown for one sepal); note abaxial surface of sepals with thick-walled epidermal cells, thick cuticle, and fine pointed verrucae. d: Longitudinal section of flower (orthoslice xz0500) showing perigynous position of calyx and semi-inferior ovary (ow, ovary wall) with a central placenta (pl), central column (cc) and numerous ovules (ov); note spiny verrucae on abaxial surface of calyx lobes. Specimens, Mira 100-S153145 (a, b), Mira 100-S170155 (c, d, holotype). Scale bars = 600 µm (a–d)." captionText-5="Text-fig. 6. SRXTM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a: Volume rendering of flower in apical view showing five narrow calyx lobes; note that the apical part of the five-angled style is sealed (arrow). b: Volume rendering of flower (cut at orthoslice xy0796) showing transverse sections of calyx lobes with three distinct bundles indicated by arrows in one of the lobes, ovary wall (ow) and central placenta (pl) bearing numerous ovules (ov). c, d: Transverse sections of style (c, orthoslice xy0153; d, orthoslice xy0222) showing five angled form, sealed near apex (c) and with prominent central canal further farther down (d). e: Volume rendering of flower (cut at orthoslice xz1024) in longitudinal section showing calyx, semi-inferior ovary with ovary wall (ow), and central placenta (pl) bearing numerous ovules (ov). Specimen, Mira 100-S153145 (a–d). Scale bars = 600 µm (a, b, e), 100 µm (c, d)." figureDoi-0="http://doi.org/10.5281/zenodo.7167563" figureDoi-1="http://doi.org/10.5281/zenodo.7167565" figureDoi-2="http://doi.org/10.5281/zenodo.7167567" figureDoi-3="http://doi.org/10.5281/zenodo.7167569" figureDoi-4="http://doi.org/10.5281/zenodo.7167571" figureDoi-5="http://doi.org/10.5281/zenodo.7167573" httpUri-0="https://zenodo.org/record/7167563/files/figure.png" httpUri-1="https://zenodo.org/record/7167565/files/figure.png" httpUri-2="https://zenodo.org/record/7167567/files/figure.png" httpUri-3="https://zenodo.org/record/7167569/files/figure.png" httpUri-4="https://zenodo.org/record/7167571/files/figure.png" httpUri-5="https://zenodo.org/record/7167573/files/figure.png" pageId="2" pageNumber="216">Text-figs 1–6</figureCitation>
</paragraph>
</subSubSection>
<subSubSection lastPageId="7" lastPageNumber="221" pageId="2" pageNumber="216" type="description">
<paragraph blockId="2.[843,1482,1577,1633]" pageId="2" pageNumber="216">H o l o t y p e. S170155 (Mira sample 100; figured Textfigs 1a, 4c, d, 5c, d).</paragraph>
<paragraph blockId="2.[843,1481,1657,1779]" pageId="2" pageNumber="216">P a r a t y p e s. S170106 (Mira sample 99), S101263, S101264, S101266–S101268, S153144–S153146, S154535– S154538, S170123, S170125, S266059 (Mira sample 100), S100732 (Mira sample 105).</paragraph>
<paragraph blockId="2.[843,1479,1802,1859]" box="[881,1479,1802,1827]" pageId="2" pageNumber="216">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 blockId="2.[843,1479,1802,1859]" box="[843,1169,1835,1859]" pageId="2" pageNumber="216">PFN002666 (for new species).</paragraph>
<paragraph blockId="2.[843,1481,1883,1939]" box="[881,1481,1883,1907]" pageId="2" pageNumber="216">R e p o s i t o r y. Palaeobotanical Collections, Swedish</paragraph>
<paragraph blockId="2.[843,1481,1883,1939]" box="[843,1409,1915,1939]" pageId="2" pageNumber="216">
<materialsCitation box="[843,1405,1915,1939]" collectionCode="S" country="Sweden" location="Museum of Natural History" pageId="2" pageNumber="216" specimenCount="1" stateProvince="Stockholm">
<location LSID="urn:lsid:plazi:treatment:03F487DBFF8F246FFC54FA39FA86C40F:8E826016FF8F246AFCB9F8FDFB84CBAF" box="[843,1139,1915,1939]" country="Sweden" name="Museum of Natural History" pageId="2" pageNumber="216" stateProvince="Stockholm">Museum of Natural History</location>
,
<collectingRegion box="[1150,1268,1915,1939]" country="Sweden" name="Stockholm" pageId="2" pageNumber="216">Stockholm</collectingRegion>
,
<collectingCountry box="[1279,1363,1915,1939]" name="Sweden" pageId="2" pageNumber="216">Sweden</collectingCountry>
(
<collectionCode box="[1379,1395,1915,1939]" country="Sweden" lsid="urn:lsid:biocol.org:col:15668" name="Department of Botany, Swedish Museum of Natural History" pageId="2" pageNumber="216" type="Herbarium">S</collectionCode>
)
</materialsCitation>
.
</paragraph>
<paragraph blockId="2.[843,1482,1963,2019]" pageId="2" pageNumber="216">E t y m o l o g y. From the elegant shape of the pedicellate flower.</paragraph>
<paragraph blockId="2.[843,1481,2042,2100]" lastBlockId="6.[129,767,151,208]" lastPageId="6" lastPageNumber="220" pageId="2" pageNumber="216">
<materialsCitation collectionCode="T" collectorName="Mira" latitude="40.416668" location="Aveiro" longLatPrecision="925" longitude="-8.7375" pageId="2" pageNumber="216" specimenCount="1" stateProvince="Aveiro">
<collectionCode box="[881,897,2044,2067]" name="Tavera, Department of Geology and Geophysics" pageId="2" pageNumber="216">T</collectionCode>
y p e l o c a l i t y.
<collectorName box="[1108,1160,2043,2067]" pageId="2" pageNumber="216">Mira</collectorName>
(
<geoCoordinate box="[1178,1291,2042,2067]" degrees="40" direction="north" minutes="25" orientation="latitude" pageId="2" pageNumber="216" precision="925" value="40.416668">40° 25ʹ N</geoCoordinate>
,
<geoCoordinate box="[1305,1470,2042,2067]" degrees="08" direction="west" minutes="44" orientation="longitude" pageId="2" pageNumber="216" precision="15" seconds="15" value="-8.7375">08° 44ʹ 15ʺ W</geoCoordinate>
), about
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south of
<collectingRegion box="[1071,1146,2076,2100]" country="Portugal" name="Aveiro" pageId="2" pageNumber="216">Aveiro</collectingRegion>
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,
<materialsCitation box="[1157,1473,2076,2100]" country="Portugal" location="Coimbra" pageId="2" pageNumber="216" specimenCount="1" stateProvince="Coimbra">
District of
<collectingRegion box="[1272,1368,2076,2100]" country="Portugal" name="Coimbra" pageId="2" pageNumber="216">Coimbra</collectingRegion>
,
<collectingCountry box="[1379,1473,2076,2100]" name="Portugal" pageId="2" pageNumber="216">Portugal</collectingCountry>
</materialsCitation>
.
<materialsCitation box="[166,764,151,176]" collectionCode="T" country="Portugal" location="Argilas de Vagos" pageId="6" pageNumber="220" specimenCount="1" stateProvince="Coimbra">
<collectionCode box="[166,182,152,175]" name="Tavera, Department of Geology and Geophysics" pageId="6" pageNumber="220">T</collectionCode>
y p e s t r a t u m a n d a g e. “
<location LSID="urn:lsid:plazi:treatment:03F487DBFF8F246FFC54FA39FA86C40F:8E826016FF8B246EFDDAFF16FD05CC8C" box="[552,754,151,176]" country="Portugal" name="Argilas de Vagos" pageId="6" pageNumber="220" stateProvince="Coimbra">Argilas de Vagos</location>
”
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,
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7167563" ID-Zenodo-Dep="7167563" httpUri="https://zenodo.org/record/7167563/files/figure.png" pageId="3" pageNumber="217" startId="3.[125,206,1774,1795]" targetBox="[125,1440,151,1758]" targetPageId="3">
<paragraph blockId="3.[125,1440,1774,2029]" pageId="3" pageNumber="217">
Text-fig. 1. SEM images of flowers of
<taxonomicName authority="Friis &amp; Crane &amp; Pedersen, 2021" authorityName="Friis &amp; Crane &amp; Pedersen" authorityYear="2021" box="[523,712,1774,1795]" class="Magnoliopsida" family="Primulaceae" genus="Miranthus" kingdom="Plantae" order="Ericales" pageId="3" pageNumber="217" phylum="Tracheophyta" rank="species" species="elegans" status="gen. et sp. nov.">
<emphasis bold="true" box="[523,712,1774,1795]" italics="true" pageId="3" pageNumber="217">Miranthus elegans</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[720,880,1774,1795]" pageId="3" pageNumber="217" rank="species">gen. et sp. nov.</taxonomicNameLabel>
; Mira locality, Portugal. a, b: Flowers in lateral view showing elongated pedicel, narrowly triangular sepals and elongated protruding style (a); note the large openings in the floral tissue and pedicel (asterisks) interpreted as schizogenous secretory cavities. c: Flower in lateral view with portion of the calyx missing exposing the ovary wall and slightly raised nectariferous ring with probable stomata-like secretory structures (arrow). d: Flower in lateral view showing long pedicel and three of the five tepals; note the elongated narrowly triangular form of the sepals. e: Flower in oblique lateral view with portion of the calyx missing exposing the ovary and elongated style. f, g: Flowers in apical view showing the bases of five sepals (f) and apex of the five-parted ovary; note larger openings in the floral tissue (asterisk) interpreted as schizogenous secretory cavities. Specimens, Mira 100-S170155 (a, holotype), Mira 100-S153145 (b, c, g), Mira 100- S101267 (d), Mira 105-S100732 (e), Mira 100-S101268 (f). Scale bars = 600 µm (a–g).
</paragraph>
</caption>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7167565" ID-Zenodo-Dep="7167565" httpUri="https://zenodo.org/record/7167565/files/figure.png" pageId="4" pageNumber="218" startId="4.[147,228,1735,1756]" targetBox="[148,1463,151,1719]" targetPageId="4">
<paragraph blockId="4.[147,1463,1735,1990]" pageId="4" pageNumber="218">
Text-fig. 2. SEM images of
<taxonomicName authority="Friis &amp; Crane &amp; Pedersen, 2021" authorityName="Friis &amp; Crane &amp; Pedersen" authorityYear="2021" box="[419,605,1735,1756]" class="Magnoliopsida" family="Primulaceae" genus="Miranthus" kingdom="Plantae" order="Ericales" pageId="4" pageNumber="218" phylum="Tracheophyta" rank="species" species="elegans" status="gen. et sp. nov.">
<emphasis bold="true" box="[419,605,1735,1756]" italics="true" pageId="4" pageNumber="218">Miranthus elegans</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[610,760,1735,1756]" pageId="4" pageNumber="218" rank="species">gen. et sp. nov.</taxonomicNameLabel>
; Mira locality, Portugal. a, b: Flowers in oblique lateral view showing remains of calyx and slightly semi-inferior ovary with elongated apical style (a); note larger openings in the floral tissue (asterisk) interpreted as schizogenous secretory cavities and the stomata-like secretory structures on the upper portion of the ovary (arrows) that are interpreted as nectariferous (b). c: Detail of ovary surface showing secretory stomata-like structures (arrows). d: Flower in lateral view showing fragmentary calyx and broken slightly semi-inferior ovary with secretory stomata-like structures; note the point of attachment of the central placenta (pl). e: Cluster of seeds removed from the ovary in (d) showing reticulate surface. f: Outer (abaxial) surface of calyx lobe showing the slightly pointed papillae and scattered, fine trichomes (arrows). g: Triaperturate pollen grains from the ovary surface. Specimens, Mira 100-S153146 (a, b), Mira 100-S170155 (c), Mira 100-S101266 (d, e), Mira 105-S100732 (f), Mira 100-S170125 (g). Scale bars = 600 µm (a, b, d), 300 µm (f), 100 µm (c, e), 10 µm (g).
</paragraph>
</caption>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7167567" ID-Zenodo-Dep="7167567" httpUri="https://zenodo.org/record/7167567/files/figure.png" pageId="5" pageNumber="219" startId="5.[123,204,1939,1960]" targetBox="[123,1438,152,1923]" targetPageId="5">
<paragraph blockId="5.[123,1442,1939,2048]" pageId="5" pageNumber="219">
Text-fig. 3. SRXTM images of
<taxonomicName authority="Friis &amp; Crane &amp; Pedersen, 2021" authorityName="Friis &amp; Crane &amp; Pedersen" authorityYear="2021" box="[438,626,1939,1960]" class="Magnoliopsida" family="Primulaceae" genus="Miranthus" kingdom="Plantae" order="Ericales" pageId="5" pageNumber="219" phylum="Tracheophyta" rank="species" species="elegans" status="gen. et sp. nov.">
<emphasis bold="true" box="[438,626,1939,1960]" italics="true" pageId="5" pageNumber="219">Miranthus elegans</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[633,788,1939,1960]" pageId="5" pageNumber="219" rank="species">gen. et sp. nov.</taxonomicNameLabel>
; Mira locality, Portugal. a, b: Volume renderings of flower bud in two different lateral views showing long pedicel, distinct calyx (ca) with almost equiaxial epidermal cells and corolla (co) with nearly smooth surface. c–e: Transverse sections (c, orthoslice xy1500; d, orthoslice xy1760; e, orthoslice xy1850) through flower bud at levels below the anthers showing stamen filaments (yellow) opposite the corolla lobes (co) and smaller staminodes (orange)
</paragraph>
</caption>
<paragraph blockId="6.[129,767,151,208]" box="[129,609,184,208]" pageId="6" pageNumber="220">Late Cretaceous (Campanian/Maastrichtian).</paragraph>
<paragraph blockId="6.[129,768,231,352]" pageId="6" pageNumber="220">D i a g n o s i s. As for the genus with the following addition. Epidermis of pedicel and outer (abaxial) epidermis of calyx composed of cells that each have a central pointed papilla. Trichomes scattered and thin.</paragraph>
<paragraph blockId="6.[128,767,375,464]" pageId="6" pageNumber="220">
D i m e n s i o n s. Length of flower without pedicel and corolla: 0.8–1.0 mm; diameter: 0.7–1.0 mm. Length of flower bud with corolla and without pedicel:
<quantity box="[609,691,440,464]" metricMagnitude="-3" metricUnit="m" metricValue="1.5" pageId="6" pageNumber="220" unit="mm" value="1.5">1.5 mm</quantity>
.
</paragraph>
<paragraph blockId="6.[128,768,487,1899]" pageId="6" pageNumber="220">
Description and comments on the s p e c i e s. The fossil flowers were briefly described and illustrated by
<bibRefCitation author="Friis, E. M. &amp; Pedersen, K. R. &amp; Crane, P. R." box="[269,439,552,576]" pageId="6" pageNumber="220" pagination="341 - 361" refId="ref11380" refString="Friis, E. M., Pedersen, K. R., Crane, P. R. (2010): Cretaceous diversification of angiosperms in the western part of the Iberian Peninsula. - Review of Palaeobotany and Palynology, 162: 341 - 361. https: // doi. org / 10.1016 / j. revpalbo. 2009.11.009" type="journal article" year="2010">Friis et al. (2010</bibRefCitation>
,
<bibRefCitation author="Friis, E. M. &amp; Crane, P. R. &amp; Pedersen, K. R." box="[447,503,552,576]" pageId="6" pageNumber="220" refId="ref10807" refString="Friis, E. M., Crane, P. R., Pedersen, K. R. (2011): Early flowers and angiosperm evolution. - Cambridge University Press, Cambridge, x + 585 pp. https: // doi. org / 10.1017 / CBO 9780511980206" type="book" year="2011">2011</bibRefCitation>
), but not formally named. More material has subsequently been discovered, including a small, informative flower bud. In most flowers the corolla and androecium have been shed (
<figureCitation box="[489,648,649,673]" captionStart="Text-fig" captionStartId="3.[125,206,1774,1795]" captionTargetBox="[125,1440,151,1758]" captionTargetId="figure-2@3.[-26,1561,55,2300]" captionTargetPageId="3" captionText="Text-fig. 1. SEM images of flowers of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Flowers in lateral view showing elongated pedicel, narrowly triangular sepals and elongated protruding style (a); note the large openings in the floral tissue and pedicel (asterisks) interpreted as schizogenous secretory cavities. c: Flower in lateral view with portion of the calyx missing exposing the ovary wall and slightly raised nectariferous ring with probable stomata-like secretory structures (arrow). d: Flower in lateral view showing long pedicel and three of the five tepals; note the elongated narrowly triangular form of the sepals. e: Flower in oblique lateral view with portion of the calyx missing exposing the ovary and elongated style. f, g: Flowers in apical view showing the bases of five sepals (f) and apex of the five-parted ovary; note larger openings in the floral tissue (asterisk) interpreted as schizogenous secretory cavities. Specimens, Mira 100-S170155 (a, holotype), Mira 100-S153145 (b, c, g), Mira 100- S101267 (d), Mira 105-S100732 (e), Mira 100-S101268 (f). Scale bars = 600 µm (a–g)." figureDoi="http://doi.org/10.5281/zenodo.7167563" httpUri="https://zenodo.org/record/7167563/files/figure.png" pageId="6" pageNumber="220">Text-figs 1a–g</figureCitation>
,
<figureCitation box="[660,714,649,673]" captionStart="Text-fig" captionStartId="4.[147,228,1735,1756]" captionTargetBox="[148,1463,151,1719]" captionTargetId="figure-2@4.[-3,1584,55,2300]" captionTargetPageId="4" captionText="Text-fig. 2. SEM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Flowers in oblique lateral view showing remains of calyx and slightly semi-inferior ovary with elongated apical style (a); note larger openings in the floral tissue (asterisk) interpreted as schizogenous secretory cavities and the stomata-like secretory structures on the upper portion of the ovary (arrows) that are interpreted as nectariferous (b). c: Detail of ovary surface showing secretory stomata-like structures (arrows). d: Flower in lateral view showing fragmentary calyx and broken slightly semi-inferior ovary with secretory stomata-like structures; note the point of attachment of the central placenta (pl). e: Cluster of seeds removed from the ovary in (d) showing reticulate surface. f: Outer (abaxial)surface of calyx lobe showing the slightly pointed papillae and scattered, fine trichomes (arrows). g: Triaperturate pollen grains from the ovary surface. Specimens, Mira 100-S153146 (a, b), Mira 100-S170155 (c), Mira 100-S101266 (d, e), Mira 105-S100732 (f), Mira 100-S170125 (g). Scale bars = 600 µm (a, b, d), 300 µm (f), 100 µm (c, e), 10 µm (g)." figureDoi="http://doi.org/10.5281/zenodo.7167565" httpUri="https://zenodo.org/record/7167565/files/figure.png" pageId="6" pageNumber="220">2a–d</figureCitation>
) and the flowers are probably fossilized at a post-anthetic stage. The flower bud has remains of corolla and stamens preserved (
<figureCitation box="[136,299,745,769]" captionStart="Text-fig" captionStartId="5.[123,204,1939,1960]" captionTargetBox="[123,1438,152,1923]" captionTargetId="figure-2@5.[-28,1559,56,2301]" captionTargetPageId="5" captionText="Text-fig. 3. SRXTM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Volume renderings of flower bud in two different lateral views showing long pedicel, distinct calyx (ca) with almost equiaxial epidermal cells and corolla (co) with nearly smooth surface. c–e: Transverse sections (c, orthoslice xy1500; d, orthoslice xy1760; e, orthoslice xy1850) through flower bud at levels below the anthers showing stamen filaments (yellow) opposite the corolla lobes (co) and smaller staminodes (orange)" figureDoi="http://doi.org/10.5281/zenodo.7167567" httpUri="https://zenodo.org/record/7167567/files/figure.png" pageId="6" pageNumber="220">Text-fig. 3a–f</figureCitation>
), but unfortunately, internal structures, including the ovary, ovules and parts of the androecium, are not well preserved. These internal structures were partially destroyed during fossilization and are only preserved on one side of the flower, which complicates interpretations of their organisation. The flower bud is assigned to the same species as the post-anthetic flowers based on features of the calyx. A second flower bud from the Mira mesofossil flora is described as a separate species,
<taxonomicName authorityName="Friis &amp; Crane &amp; Pedersen" authorityYear="2021" box="[489,694,1003,1027]" class="Magnoliopsida" family="Primulaceae" genus="Miranthus" kingdom="Plantae" order="Ericales" pageId="6" pageNumber="220" phylum="Tracheophyta" rank="species" species="kvacekii">
<emphasis box="[489,694,1003,1027]" italics="true" pageId="6" pageNumber="220">Miranthus kvacekii</emphasis>
</taxonomicName>
, based on epidermal differences from
<taxonomicName authorityName="Friis &amp; Crane &amp; Pedersen" authorityYear="2021" box="[458,657,1036,1060]" class="Magnoliopsida" family="Primulaceae" genus="Miranthus" kingdom="Plantae" order="Ericales" pageId="6" pageNumber="220" phylum="Tracheophyta" rank="species" species="elegans">
<emphasis box="[458,657,1036,1060]" italics="true" pageId="6" pageNumber="220">Miranthus elegans</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph blockId="6.[128,768,487,1899]" pageId="6" pageNumber="220">
The flowers are borne on a long, slender pedicel (Textfigs 1a–d, 2d, 3a, b) and lack bracts or bracteoles immediately below the calyx. The pedicel is up to about
<quantity box="[631,708,1133,1157]" metricMagnitude="-4" metricUnit="m" metricValue="9.0" pageId="6" pageNumber="220" unit="mm" value="0.9">0.9 mm</quantity>
long (
<figureCitation box="[136,263,1165,1189]" captionStart="Text-fig" captionStartId="3.[125,206,1774,1795]" captionTargetBox="[125,1440,151,1758]" captionTargetId="figure-2@3.[-26,1561,55,2300]" captionTargetPageId="3" captionText="Text-fig. 1. SEM images of flowers of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Flowers in lateral view showing elongated pedicel, narrowly triangular sepals and elongated protruding style (a); note the large openings in the floral tissue and pedicel (asterisks) interpreted as schizogenous secretory cavities. c: Flower in lateral view with portion of the calyx missing exposing the ovary wall and slightly raised nectariferous ring with probable stomata-like secretory structures (arrow). d: Flower in lateral view showing long pedicel and three of the five tepals; note the elongated narrowly triangular form of the sepals. e: Flower in oblique lateral view with portion of the calyx missing exposing the ovary and elongated style. f, g: Flowers in apical view showing the bases of five sepals (f) and apex of the five-parted ovary; note larger openings in the floral tissue (asterisk) interpreted as schizogenous secretory cavities. Specimens, Mira 100-S170155 (a, holotype), Mira 100-S153145 (b, c, g), Mira 100- S101267 (d), Mira 105-S100732 (e), Mira 100-S101268 (f). Scale bars = 600 µm (a–g)." figureDoi="http://doi.org/10.5281/zenodo.7167563" httpUri="https://zenodo.org/record/7167563/files/figure.png" pageId="6" pageNumber="220">Text-fig. 1d</figureCitation>
). All of the fossil flowers are isolated and there is no information on inflorescence structure, or how the pedicel may have been attached to the plant.
</paragraph>
<paragraph blockId="6.[128,768,487,1899]" lastBlockId="6.[843,1482,152,1899]" pageId="6" pageNumber="220">
The flowers are structurally bisexual, actinomorphic, pentamerous and isomerous, about 0.8–1.0 mm long, excluding the pedicel and corolla, and 0.7–1.0 mm in diameter. Including the corolla, but excluding the pedicel, the flower bud is about
<quantity box="[381,459,1391,1415]" metricMagnitude="-3" metricUnit="m" metricValue="1.5" pageId="6" pageNumber="220" unit="mm" value="1.5">1.5 mm</quantity>
long. The calyx is persistent and consists of five imbricate sepals in a single whorl that are fused at the base (
<figureCitation box="[365,524,1455,1479]" captionStart="Text-fig" captionStartId="3.[125,206,1774,1795]" captionTargetBox="[125,1440,151,1758]" captionTargetId="figure-2@3.[-26,1561,55,2300]" captionTargetPageId="3" captionText="Text-fig. 1. SEM images of flowers of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Flowers in lateral view showing elongated pedicel, narrowly triangular sepals and elongated protruding style (a); note the large openings in the floral tissue and pedicel (asterisks) interpreted as schizogenous secretory cavities. c: Flower in lateral view with portion of the calyx missing exposing the ovary wall and slightly raised nectariferous ring with probable stomata-like secretory structures (arrow). d: Flower in lateral view showing long pedicel and three of the five tepals; note the elongated narrowly triangular form of the sepals. e: Flower in oblique lateral view with portion of the calyx missing exposing the ovary and elongated style. f, g: Flowers in apical view showing the bases of five sepals (f) and apex of the five-parted ovary; note larger openings in the floral tissue (asterisk) interpreted as schizogenous secretory cavities. Specimens, Mira 100-S170155 (a, holotype), Mira 100-S153145 (b, c, g), Mira 100- S101267 (d), Mira 105-S100732 (e), Mira 100-S101268 (f). Scale bars = 600 µm (a–g)." figureDoi="http://doi.org/10.5281/zenodo.7167563" httpUri="https://zenodo.org/record/7167563/files/figure.png" pageId="6" pageNumber="220">Text-figs 1a–g</figureCitation>
,
<figureCitation box="[536,592,1455,1479]" captionStart="Text-fig" captionStartId="8.[148,226,1363,1384]" captionTargetBox="[148,1461,144,1347]" captionTargetId="figure-291@8.[-5,1582,47,2292]" captionTargetPageId="8" captionText="Text-fig. 5. SRXTM imagesof Miranthuselegans gen. et sp. nov.;Mira locality, Portugal. a, b: Transverse sections offlower (a, orthoslice xy0665 close to the apex of placenta; b, orthoslice xy0800 in middle part of placenta) showing remains of calyx with distinct bundles (arrows), ovary wall (ow) and numerous ovules (ov) on the central mushroom-shaped globose placenta (pl) with central column (cc). c: Transverse section of flower (orthoslice xy0620) through perianth and ovary (ow) at a level above the placenta showing ovules (ov) and cellular preservation of the sepal bundles (arrows shown for one sepal); note abaxial surface of sepals with thick-walled epidermal cells, thick cuticle, and fine pointed verrucae. d: Longitudinal section of flower (orthoslice xz0500) showing perigynous position of calyx and semi-inferior ovary (ow, ovary wall) with a central placenta (pl), central column (cc) and numerous ovules (ov); note spiny verrucae on abaxial surface of calyx lobes. Specimens, Mira 100-S153145 (a, b), Mira 100-S170155 (c, d, holotype). Scale bars = 600 µm (a–d)." figureDoi="http://doi.org/10.5281/zenodo.7167571" httpUri="https://zenodo.org/record/7167571/files/figure.png" pageId="6" pageNumber="220">5a, b</figureCitation>
). The sepals are narrowly triangular, with the free portion up to about
<quantity box="[689,767,1488,1512]" metricMagnitude="-4" metricUnit="m" metricValue="9.0" pageId="6" pageNumber="220" unit="mm" value="0.9">0.9 mm</quantity>
long, which tapers into a long tip (
<figureCitation box="[484,636,1520,1544]" captionStart="Text-fig" captionStartId="3.[125,206,1774,1795]" captionTargetBox="[125,1440,151,1758]" captionTargetId="figure-2@3.[-26,1561,55,2300]" captionTargetPageId="3" captionText="Text-fig. 1. SEM images of flowers of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Flowers in lateral view showing elongated pedicel, narrowly triangular sepals and elongated protruding style (a); note the large openings in the floral tissue and pedicel (asterisks) interpreted as schizogenous secretory cavities. c: Flower in lateral view with portion of the calyx missing exposing the ovary wall and slightly raised nectariferous ring with probable stomata-like secretory structures (arrow). d: Flower in lateral view showing long pedicel and three of the five tepals; note the elongated narrowly triangular form of the sepals. e: Flower in oblique lateral view with portion of the calyx missing exposing the ovary and elongated style. f, g: Flowers in apical view showing the bases of five sepals (f) and apex of the five-parted ovary; note larger openings in the floral tissue (asterisk) interpreted as schizogenous secretory cavities. Specimens, Mira 100-S170155 (a, holotype), Mira 100-S153145 (b, c, g), Mira 100- S101267 (d), Mira 105-S100732 (e), Mira 100-S101268 (f). Scale bars = 600 µm (a–g)." figureDoi="http://doi.org/10.5281/zenodo.7167563" httpUri="https://zenodo.org/record/7167563/files/figure.png" pageId="6" pageNumber="220">Text-fig. 1a–g</figureCitation>
). Each sepal is supplied by three distinct vascular bundles, each of which typically appears as a hollow space in transverse sections (
<figureCitation box="[135,271,1616,1641]" captionStart="Text-fig" captionStartId="8.[148,226,1363,1384]" captionTargetBox="[148,1461,144,1347]" captionTargetId="figure-291@8.[-5,1582,47,2292]" captionTargetPageId="8" captionText="Text-fig. 5. SRXTM imagesof Miranthuselegans gen. et sp. nov.;Mira locality, Portugal. a, b: Transverse sections offlower (a, orthoslice xy0665 close to the apex of placenta; b, orthoslice xy0800 in middle part of placenta) showing remains of calyx with distinct bundles (arrows), ovary wall (ow) and numerous ovules (ov) on the central mushroom-shaped globose placenta (pl) with central column (cc). c: Transverse section of flower (orthoslice xy0620) through perianth and ovary (ow) at a level above the placenta showing ovules (ov) and cellular preservation of the sepal bundles (arrows shown for one sepal); note abaxial surface of sepals with thick-walled epidermal cells, thick cuticle, and fine pointed verrucae. d: Longitudinal section of flower (orthoslice xz0500) showing perigynous position of calyx and semi-inferior ovary (ow, ovary wall) with a central placenta (pl), central column (cc) and numerous ovules (ov); note spiny verrucae on abaxial surface of calyx lobes. Specimens, Mira 100-S153145 (a, b), Mira 100-S170155 (c, d, holotype). Scale bars = 600 µm (a–d)." figureDoi="http://doi.org/10.5281/zenodo.7167571" httpUri="https://zenodo.org/record/7167571/files/figure.png" pageId="6" pageNumber="220">Text-figs 5a</figureCitation>
,
<figureCitation box="[287,347,1617,1641]" captionStart="Text-fig" captionStartId="9.[125,206,1251,1272]" captionTargetBox="[125,1440,151,1236]" captionTargetId="figure-297@9.[-26,1561,55,2300]" captionTargetPageId="9" captionText="Text-fig. 6. SRXTM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a: Volume rendering of flower in apical view showing five narrow calyx lobes; note that the apical part of the five-angled style is sealed (arrow). b: Volume rendering of flower (cut at orthoslice xy0796) showing transverse sections of calyx lobes with three distinct bundles indicated by arrows in one of the lobes, ovary wall (ow) and central placenta (pl) bearing numerous ovules (ov). c, d: Transverse sections of style (c, orthoslice xy0153; d, orthoslice xy0222) showing five angled form, sealed near apex (c) and with prominent central canal further farther down (d). e: Volume rendering of flower (cut at orthoslice xz1024) in longitudinal section showing calyx, semi-inferior ovary with ovary wall (ow), and central placenta (pl) bearing numerous ovules (ov). Specimen, Mira 100-S153145 (a–d). Scale bars = 600 µm (a, b, e), 100 µm (c, d)." figureDoi="http://doi.org/10.5281/zenodo.7167573" httpUri="https://zenodo.org/record/7167573/files/figure.png" pageId="6" pageNumber="220">6a, b</figureCitation>
). In some specimens, minor vascular bundles are seen close to the sepal margins. The epidermis of the pedicel, and the abaxial (outer) epidermis of the calyx, is covered by a thick cuticle. The epidermal cells are small, almost equiaxial, thick-walled and typically with a pointed papilla that gives the surface a faint verrucate to spiny appearance. Similar epidermal features are also observed on the outer surface of the young ovaries (
<figureCitation box="[564,725,1842,1867]" captionStart="Text-fig" captionStartId="3.[125,206,1774,1795]" captionTargetBox="[125,1440,151,1758]" captionTargetId="figure-2@3.[-26,1561,55,2300]" captionTargetPageId="3" captionText="Text-fig. 1. SEM images of flowers of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Flowers in lateral view showing elongated pedicel, narrowly triangular sepals and elongated protruding style (a); note the large openings in the floral tissue and pedicel (asterisks) interpreted as schizogenous secretory cavities. c: Flower in lateral view with portion of the calyx missing exposing the ovary wall and slightly raised nectariferous ring with probable stomata-like secretory structures (arrow). d: Flower in lateral view showing long pedicel and three of the five tepals; note the elongated narrowly triangular form of the sepals. e: Flower in oblique lateral view with portion of the calyx missing exposing the ovary and elongated style. f, g: Flowers in apical view showing the bases of five sepals (f) and apex of the five-parted ovary; note larger openings in the floral tissue (asterisk) interpreted as schizogenous secretory cavities. Specimens, Mira 100-S170155 (a, holotype), Mira 100-S153145 (b, c, g), Mira 100- S101267 (d), Mira 105-S100732 (e), Mira 100-S101268 (f). Scale bars = 600 µm (a–g)." figureDoi="http://doi.org/10.5281/zenodo.7167563" httpUri="https://zenodo.org/record/7167563/files/figure.png" pageId="6" pageNumber="220">Text-figs 1a–g</figureCitation>
,
<figureCitation box="[738,763,1842,1866]" captionStart="Text-fig" captionStartId="4.[147,228,1735,1756]" captionTargetBox="[148,1463,151,1719]" captionTargetId="figure-2@4.[-3,1584,55,2300]" captionTargetPageId="4" captionText="Text-fig. 2. SEM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Flowers in oblique lateral view showing remains of calyx and slightly semi-inferior ovary with elongated apical style (a); note larger openings in the floral tissue (asterisk) interpreted as schizogenous secretory cavities and the stomata-like secretory structures on the upper portion of the ovary (arrows) that are interpreted as nectariferous (b). c: Detail of ovary surface showing secretory stomata-like structures (arrows). d: Flower in lateral view showing fragmentary calyx and broken slightly semi-inferior ovary with secretory stomata-like structures; note the point of attachment of the central placenta (pl). e: Cluster of seeds removed from the ovary in (d) showing reticulate surface. f: Outer (abaxial)surface of calyx lobe showing the slightly pointed papillae and scattered, fine trichomes (arrows). g: Triaperturate pollen grains from the ovary surface. Specimens, Mira 100-S153146 (a, b), Mira 100-S170155 (c), Mira 100-S101266 (d, e), Mira 105-S100732 (f), Mira 100-S170125 (g). Scale bars = 600 µm (a, b, d), 300 µm (f), 100 µm (c, e), 10 µm (g)." figureDoi="http://doi.org/10.5281/zenodo.7167565" httpUri="https://zenodo.org/record/7167565/files/figure.png" pageId="6" pageNumber="220">2f</figureCitation>
,
<figureCitation box="[128,182,1875,1899]" captionStart="Text-fig" captionStartId="5.[123,204,1939,1960]" captionTargetBox="[123,1438,152,1923]" captionTargetId="figure-2@5.[-28,1559,56,2301]" captionTargetPageId="5" captionText="Text-fig. 3. SRXTM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Volume renderings of flower bud in two different lateral views showing long pedicel, distinct calyx (ca) with almost equiaxial epidermal cells and corolla (co) with nearly smooth surface. c–e: Transverse sections (c, orthoslice xy1500; d, orthoslice xy1760; e, orthoslice xy1850) through flower bud at levels below the anthers showing stamen filaments (yellow) opposite the corolla lobes (co) and smaller staminodes (orange)" figureDoi="http://doi.org/10.5281/zenodo.7167567" httpUri="https://zenodo.org/record/7167567/files/figure.png" pageId="6" pageNumber="220">3c–e</figureCitation>
,
<figureCitation box="[194,248,1875,1899]" captionStart="Text-fig" captionStartId="8.[148,226,1363,1384]" captionTargetBox="[148,1461,144,1347]" captionTargetId="figure-291@8.[-5,1582,47,2292]" captionTargetPageId="8" captionText="Text-fig. 5. SRXTM imagesof Miranthuselegans gen. et sp. nov.;Mira locality, Portugal. a, b: Transverse sections offlower (a, orthoslice xy0665 close to the apex of placenta; b, orthoslice xy0800 in middle part of placenta) showing remains of calyx with distinct bundles (arrows), ovary wall (ow) and numerous ovules (ov) on the central mushroom-shaped globose placenta (pl) with central column (cc). c: Transverse section of flower (orthoslice xy0620) through perianth and ovary (ow) at a level above the placenta showing ovules (ov) and cellular preservation of the sepal bundles (arrows shown for one sepal); note abaxial surface of sepals with thick-walled epidermal cells, thick cuticle, and fine pointed verrucae. d: Longitudinal section of flower (orthoslice xz0500) showing perigynous position of calyx and semi-inferior ovary (ow, ovary wall) with a central placenta (pl), central column (cc) and numerous ovules (ov); note spiny verrucae on abaxial surface of calyx lobes. Specimens, Mira 100-S153145 (a, b), Mira 100-S170155 (c, d, holotype). Scale bars = 600 µm (a–d)." figureDoi="http://doi.org/10.5281/zenodo.7167571" httpUri="https://zenodo.org/record/7167571/files/figure.png" pageId="6" pageNumber="220">5a–d</figureCitation>
,
<figureCitation box="[261,290,1875,1899]" captionStart="Text-fig" captionStartId="9.[125,206,1251,1272]" captionTargetBox="[125,1440,151,1236]" captionTargetId="figure-297@9.[-26,1561,55,2300]" captionTargetPageId="9" captionText="Text-fig. 6. SRXTM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a: Volume rendering of flower in apical view showing five narrow calyx lobes; note that the apical part of the five-angled style is sealed (arrow). b: Volume rendering of flower (cut at orthoslice xy0796) showing transverse sections of calyx lobes with three distinct bundles indicated by arrows in one of the lobes, ovary wall (ow) and central placenta (pl) bearing numerous ovules (ov). c, d: Transverse sections of style (c, orthoslice xy0153; d, orthoslice xy0222) showing five angled form, sealed near apex (c) and with prominent central canal further farther down (d). e: Volume rendering of flower (cut at orthoslice xz1024) in longitudinal section showing calyx, semi-inferior ovary with ovary wall (ow), and central placenta (pl) bearing numerous ovules (ov). Specimen, Mira 100-S153145 (a–d). Scale bars = 600 µm (a, b, e), 100 µm (c, d)." figureDoi="http://doi.org/10.5281/zenodo.7167573" httpUri="https://zenodo.org/record/7167573/files/figure.png" pageId="6" pageNumber="220">6a</figureCitation>
). Very slender trichomes occur scattered on the outer surface of the sepals (
<figureCitation box="[1181,1307,152,176]" captionStart="Text-fig" captionStartId="4.[147,228,1735,1756]" captionTargetBox="[148,1463,151,1719]" captionTargetId="figure-2@4.[-3,1584,55,2300]" captionTargetPageId="4" captionText="Text-fig. 2. SEM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Flowers in oblique lateral view showing remains of calyx and slightly semi-inferior ovary with elongated apical style (a); note larger openings in the floral tissue (asterisk) interpreted as schizogenous secretory cavities and the stomata-like secretory structures on the upper portion of the ovary (arrows) that are interpreted as nectariferous (b). c: Detail of ovary surface showing secretory stomata-like structures (arrows). d: Flower in lateral view showing fragmentary calyx and broken slightly semi-inferior ovary with secretory stomata-like structures; note the point of attachment of the central placenta (pl). e: Cluster of seeds removed from the ovary in (d) showing reticulate surface. f: Outer (abaxial)surface of calyx lobe showing the slightly pointed papillae and scattered, fine trichomes (arrows). g: Triaperturate pollen grains from the ovary surface. Specimens, Mira 100-S153146 (a, b), Mira 100-S170155 (c), Mira 100-S101266 (d, e), Mira 105-S100732 (f), Mira 100-S170125 (g). Scale bars = 600 µm (a, b, d), 300 µm (f), 100 µm (c, e), 10 µm (g)." figureDoi="http://doi.org/10.5281/zenodo.7167565" httpUri="https://zenodo.org/record/7167565/files/figure.png" pageId="6" pageNumber="220">Text-fig. 2f</figureCitation>
). The epidermal cells of the adaxial (inner) epidermis of the calyx lobes are small, equiaxial, thin-walled and covered with a thin cuticle (
<figureCitation box="[850,1005,249,274]" captionStart="Text-fig" captionStartId="5.[123,204,1939,1960]" captionTargetBox="[123,1438,152,1923]" captionTargetId="figure-2@5.[-28,1559,56,2301]" captionTargetPageId="5" captionText="Text-fig. 3. SRXTM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Volume renderings of flower bud in two different lateral views showing long pedicel, distinct calyx (ca) with almost equiaxial epidermal cells and corolla (co) with nearly smooth surface. c–e: Transverse sections (c, orthoslice xy1500; d, orthoslice xy1760; e, orthoslice xy1850) through flower bud at levels below the anthers showing stamen filaments (yellow) opposite the corolla lobes (co) and smaller staminodes (orange)" figureDoi="http://doi.org/10.5281/zenodo.7167567" httpUri="https://zenodo.org/record/7167567/files/figure.png" pageId="6" pageNumber="220">Text-fig. 3c–e</figureCitation>
). The calyx of the single flower bud is very similar in shape, size and epidermal features to the calyx post-anthetic flowers (
<figureCitation box="[1079,1230,314,339]" captionStart="Text-fig" captionStartId="5.[123,204,1939,1960]" captionTargetBox="[123,1438,152,1923]" captionTargetId="figure-2@5.[-28,1559,56,2301]" captionTargetPageId="5" captionText="Text-fig. 3. SRXTM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Volume renderings of flower bud in two different lateral views showing long pedicel, distinct calyx (ca) with almost equiaxial epidermal cells and corolla (co) with nearly smooth surface. c–e: Transverse sections (c, orthoslice xy1500; d, orthoslice xy1760; e, orthoslice xy1850) through flower bud at levels below the anthers showing stamen filaments (yellow) opposite the corolla lobes (co) and smaller staminodes (orange)" figureDoi="http://doi.org/10.5281/zenodo.7167567" httpUri="https://zenodo.org/record/7167567/files/figure.png" pageId="6" pageNumber="220">Text-fig. 3a, b</figureCitation>
) and the anatomy of all preserved organs is also identical.
</paragraph>
<paragraph blockId="6.[843,1482,152,1899]" pageId="6" pageNumber="220">
There is no trace of corolla and androecium in the supposed post-anthetic flowers. Corolla and androecium were most likely shed together after flowering, which would also be consistent with stamens that were fused to the corolla. The corolla is five-lobed, with imbricate lobes that are free for most of their length (
<figureCitation box="[1104,1258,542,566]" captionStart="Text-fig" captionStartId="5.[123,204,1939,1960]" captionTargetBox="[123,1438,152,1923]" captionTargetId="figure-2@5.[-28,1559,56,2301]" captionTargetPageId="5" captionText="Text-fig. 3. SRXTM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Volume renderings of flower bud in two different lateral views showing long pedicel, distinct calyx (ca) with almost equiaxial epidermal cells and corolla (co) with nearly smooth surface. c–e: Transverse sections (c, orthoslice xy1500; d, orthoslice xy1760; e, orthoslice xy1850) through flower bud at levels below the anthers showing stamen filaments (yellow) opposite the corolla lobes (co) and smaller staminodes (orange)" figureDoi="http://doi.org/10.5281/zenodo.7167567" httpUri="https://zenodo.org/record/7167567/files/figure.png" pageId="6" pageNumber="220">Text-fig. 3c–e</figureCitation>
). They appear united at the base, but the preservation does not allow a fully secure conclusion that they were sympetalous.
</paragraph>
<paragraph blockId="6.[843,1482,152,1899]" pageId="6" pageNumber="220">
The internal structures are preserved on one side of the flower bud, but based on the symmetry of the structures that remain, the androecium can be reconstructed as pentamerous and obhaplostemonous with five antepetalous stamens, which are seen near the base of the flower as stout filaments positioned in front of the petal lobes (
<figureCitation box="[1270,1424,802,826]" captionStart="Text-fig" captionStartId="5.[123,204,1939,1960]" captionTargetBox="[123,1438,152,1923]" captionTargetId="figure-2@5.[-28,1559,56,2301]" captionTargetPageId="5" captionText="Text-fig. 3. SRXTM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Volume renderings of flower bud in two different lateral views showing long pedicel, distinct calyx (ca) with almost equiaxial epidermal cells and corolla (co) with nearly smooth surface. c–e: Transverse sections (c, orthoslice xy1500; d, orthoslice xy1760; e, orthoslice xy1850) through flower bud at levels below the anthers showing stamen filaments (yellow) opposite the corolla lobes (co) and smaller staminodes (orange)" figureDoi="http://doi.org/10.5281/zenodo.7167567" httpUri="https://zenodo.org/record/7167567/files/figure.png" pageId="6" pageNumber="220">Text-fig. 3c–e</figureCitation>
) and near the bud apex as remains of thecae. Smaller structures, interpreted as staminodes, are seen alternating with the petals (
<figureCitation box="[850,1004,900,924]" captionStart="Text-fig" captionStartId="5.[123,204,1939,1960]" captionTargetBox="[123,1438,152,1923]" captionTargetId="figure-2@5.[-28,1559,56,2301]" captionTargetPageId="5" captionText="Text-fig. 3. SRXTM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Volume renderings of flower bud in two different lateral views showing long pedicel, distinct calyx (ca) with almost equiaxial epidermal cells and corolla (co) with nearly smooth surface. c–e: Transverse sections (c, orthoslice xy1500; d, orthoslice xy1760; e, orthoslice xy1850) through flower bud at levels below the anthers showing stamen filaments (yellow) opposite the corolla lobes (co) and smaller staminodes (orange)" figureDoi="http://doi.org/10.5281/zenodo.7167567" httpUri="https://zenodo.org/record/7167567/files/figure.png" pageId="6" pageNumber="220">Text-fig. 3c–e</figureCitation>
). Both stamens and staminodes appear to be attached to the corolla near the base (
<figureCitation box="[1265,1393,932,956]" captionStart="Text-fig" captionStartId="5.[123,204,1939,1960]" captionTargetBox="[123,1438,152,1923]" captionTargetId="figure-2@5.[-28,1559,56,2301]" captionTargetPageId="5" captionText="Text-fig. 3. SRXTM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Volume renderings of flower bud in two different lateral views showing long pedicel, distinct calyx (ca) with almost equiaxial epidermal cells and corolla (co) with nearly smooth surface. c–e: Transverse sections (c, orthoslice xy1500; d, orthoslice xy1760; e, orthoslice xy1850) through flower bud at levels below the anthers showing stamen filaments (yellow) opposite the corolla lobes (co) and smaller staminodes (orange)" figureDoi="http://doi.org/10.5281/zenodo.7167567" httpUri="https://zenodo.org/record/7167567/files/figure.png" pageId="6" pageNumber="220">Text-fig. 3f</figureCitation>
), which is also consistent with the absence of stamens and stamen bases in the post-anthetic flowers.
</paragraph>
<paragraph blockId="6.[843,1482,152,1899]" pageId="6" pageNumber="220">
Normapolles-type pollen, which is very common on the surface of other Mira mesofossils, is also present on
<taxonomicName authorityName="E. M. FRIIS, P. R. CRANE et K. R. PEDERSEN" authorityYear="2021" box="[843,954,1095,1119]" class="Magnoliopsida" family="Primulaceae" genus="Miranthus" kingdom="Plantae" order="Ericales" pageId="6" pageNumber="220" phylum="Tracheophyta" rank="genus">
<emphasis box="[843,954,1095,1119]" italics="true" pageId="6" pageNumber="220">Miranthus</emphasis>
</taxonomicName>
flowers. However, in addition, two flowers have another kind of pollen on their surface. In both cases, the pollen is about 16 µm long and triaperturate with very long colpi that reach almost to the poles (
<figureCitation captionStart="Text-fig" captionStartId="4.[147,228,1735,1756]" captionTargetBox="[148,1463,151,1719]" captionTargetId="figure-2@4.[-3,1584,55,2300]" captionTargetPageId="4" captionText="Text-fig. 2. SEM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Flowers in oblique lateral view showing remains of calyx and slightly semi-inferior ovary with elongated apical style (a); note larger openings in the floral tissue (asterisk) interpreted as schizogenous secretory cavities and the stomata-like secretory structures on the upper portion of the ovary (arrows) that are interpreted as nectariferous (b). c: Detail of ovary surface showing secretory stomata-like structures (arrows). d: Flower in lateral view showing fragmentary calyx and broken slightly semi-inferior ovary with secretory stomata-like structures; note the point of attachment of the central placenta (pl). e: Cluster of seeds removed from the ovary in (d) showing reticulate surface. f: Outer (abaxial)surface of calyx lobe showing the slightly pointed papillae and scattered, fine trichomes (arrows). g: Triaperturate pollen grains from the ovary surface. Specimens, Mira 100-S153146 (a, b), Mira 100-S170155 (c), Mira 100-S101266 (d, e), Mira 105-S100732 (f), Mira 100-S170125 (g). Scale bars = 600 µm (a, b, d), 300 µm (f), 100 µm (c, e), 10 µm (g)." figureDoi="http://doi.org/10.5281/zenodo.7167565" httpUri="https://zenodo.org/record/7167565/files/figure.png" pageId="6" pageNumber="220">
Text-fig.
<quantity box="[843,873,1225,1249]" metricMagnitude="-3" metricUnit="kg" metricValue="2.0" pageId="6" pageNumber="220" unit="g" value="2.0">2g</quantity>
</figureCitation>
). Unfortunately, the grains are folded and it cannot be established whether the grains are tricolpate or tricolporate. The tectum is foveolate.
</paragraph>
<paragraph blockId="6.[843,1482,152,1899]" pageId="6" pageNumber="220">
The ovary is semi-inferior and unilocular (
<figureCitation captionStart="Text-fig" captionStartId="4.[147,228,1735,1756]" captionTargetBox="[148,1463,151,1719]" captionTargetId="figure-2@4.[-3,1584,55,2300]" captionTargetPageId="4" captionText="Text-fig. 2. SEM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Flowers in oblique lateral view showing remains of calyx and slightly semi-inferior ovary with elongated apical style (a); note larger openings in the floral tissue (asterisk) interpreted as schizogenous secretory cavities and the stomata-like secretory structures on the upper portion of the ovary (arrows) that are interpreted as nectariferous (b). c: Detail of ovary surface showing secretory stomata-like structures (arrows). d: Flower in lateral view showing fragmentary calyx and broken slightly semi-inferior ovary with secretory stomata-like structures; note the point of attachment of the central placenta (pl). e: Cluster of seeds removed from the ovary in (d) showing reticulate surface. f: Outer (abaxial)surface of calyx lobe showing the slightly pointed papillae and scattered, fine trichomes (arrows). g: Triaperturate pollen grains from the ovary surface. Specimens, Mira 100-S153146 (a, b), Mira 100-S170155 (c), Mira 100-S101266 (d, e), Mira 105-S100732 (f), Mira 100-S170125 (g). Scale bars = 600 µm (a, b, d), 300 µm (f), 100 µm (c, e), 10 µm (g)." figureDoi="http://doi.org/10.5281/zenodo.7167565" httpUri="https://zenodo.org/record/7167565/files/figure.png" pageId="6" pageNumber="220">Text-figs 2d</figureCitation>
,
<figureCitation box="[886,941,1355,1379]" captionStart="Text-fig" captionStartId="7.[125,207,1562,1583]" captionTargetBox="[125,1439,151,1546]" captionTargetId="figure-148@7.[-27,1560,47,2292]" captionTargetPageId="7" captionText="Text-fig. 4. SRXTM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Transverse (a) and longitudinal (b) sections of flower showing the ovary wall (ow) and numerous ovules (ov) borne on the mushroom-shaped central globose placenta (pl, blue); placenta shown as a voltex rendering added to orthoslices; note the large openings in the floral tissue (asterisks) interpreted as schizogenous secretory cavities. c, d: Longitudinal (c) and transverse (d) sections of flower (c, volume rendering cut between orthoslice yz0970-1005; d, volume rendering cut between orthoslice xy0780-0820) showing semiinferior ovary (ow, ovary wall) with sepals inserted at the rim of the hypanthium, central column (cc) with mushroom-shaped globose placenta (pl) bearing numerous ovules (ov). Specimens, Mira 100-S153146 (a, b), Mira 100-S170155 (c, d, holotype). Scale bars = 600 µm (a–d)." figureDoi="http://doi.org/10.5281/zenodo.7167569" httpUri="https://zenodo.org/record/7167569/files/figure.png" pageId="6" pageNumber="220">4a–d</figureCitation>
,
<figureCitation box="[954,1011,1355,1379]" captionStart="Text-fig" captionStartId="8.[148,226,1363,1384]" captionTargetBox="[148,1461,144,1347]" captionTargetId="figure-291@8.[-5,1582,47,2292]" captionTargetPageId="8" captionText="Text-fig. 5. SRXTM imagesof Miranthuselegans gen. et sp. nov.;Mira locality, Portugal. a, b: Transverse sections offlower (a, orthoslice xy0665 close to the apex of placenta; b, orthoslice xy0800 in middle part of placenta) showing remains of calyx with distinct bundles (arrows), ovary wall (ow) and numerous ovules (ov) on the central mushroom-shaped globose placenta (pl) with central column (cc). c: Transverse section of flower (orthoslice xy0620) through perianth and ovary (ow) at a level above the placenta showing ovules (ov) and cellular preservation of the sepal bundles (arrows shown for one sepal); note abaxial surface of sepals with thick-walled epidermal cells, thick cuticle, and fine pointed verrucae. d: Longitudinal section of flower (orthoslice xz0500) showing perigynous position of calyx and semi-inferior ovary (ow, ovary wall) with a central placenta (pl), central column (cc) and numerous ovules (ov); note spiny verrucae on abaxial surface of calyx lobes. Specimens, Mira 100-S153145 (a, b), Mira 100-S170155 (c, d, holotype). Scale bars = 600 µm (a–d)." figureDoi="http://doi.org/10.5281/zenodo.7167571" httpUri="https://zenodo.org/record/7167571/files/figure.png" pageId="6" pageNumber="220">5a, b</figureCitation>
,
<figureCitation box="[1026,1112,1355,1379]" captionStart="Text-fig" captionStartId="9.[125,206,1251,1272]" captionTargetBox="[125,1440,151,1236]" captionTargetId="figure-297@9.[-26,1561,55,2300]" captionTargetPageId="9" captionText="Text-fig. 6. SRXTM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a: Volume rendering of flower in apical view showing five narrow calyx lobes; note that the apical part of the five-angled style is sealed (arrow). b: Volume rendering of flower (cut at orthoslice xy0796) showing transverse sections of calyx lobes with three distinct bundles indicated by arrows in one of the lobes, ovary wall (ow) and central placenta (pl) bearing numerous ovules (ov). c, d: Transverse sections of style (c, orthoslice xy0153; d, orthoslice xy0222) showing five angled form, sealed near apex (c) and with prominent central canal further farther down (d). e: Volume rendering of flower (cut at orthoslice xz1024) in longitudinal section showing calyx, semi-inferior ovary with ovary wall (ow), and central placenta (pl) bearing numerous ovules (ov). Specimen, Mira 100-S153145 (a–d). Scale bars = 600 µm (a, b, e), 100 µm (c, d)." figureDoi="http://doi.org/10.5281/zenodo.7167573" httpUri="https://zenodo.org/record/7167573/files/figure.png" pageId="6" pageNumber="220">6a, b, e</figureCitation>
). The apical portion of the ovary has a slightly protruding ring with stomata-like openings (
<figureCitation box="[850,982,1420,1444]" captionStart="Text-fig" captionStartId="3.[125,206,1774,1795]" captionTargetBox="[125,1440,151,1758]" captionTargetId="figure-2@3.[-26,1561,55,2300]" captionTargetPageId="3" captionText="Text-fig. 1. SEM images of flowers of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Flowers in lateral view showing elongated pedicel, narrowly triangular sepals and elongated protruding style (a); note the large openings in the floral tissue and pedicel (asterisks) interpreted as schizogenous secretory cavities. c: Flower in lateral view with portion of the calyx missing exposing the ovary wall and slightly raised nectariferous ring with probable stomata-like secretory structures (arrow). d: Flower in lateral view showing long pedicel and three of the five tepals; note the elongated narrowly triangular form of the sepals. e: Flower in oblique lateral view with portion of the calyx missing exposing the ovary and elongated style. f, g: Flowers in apical view showing the bases of five sepals (f) and apex of the five-parted ovary; note larger openings in the floral tissue (asterisk) interpreted as schizogenous secretory cavities. Specimens, Mira 100-S170155 (a, holotype), Mira 100-S153145 (b, c, g), Mira 100- S101267 (d), Mira 105-S100732 (e), Mira 100-S101268 (f). Scale bars = 600 µm (a–g)." figureDoi="http://doi.org/10.5281/zenodo.7167563" httpUri="https://zenodo.org/record/7167563/files/figure.png" pageId="6" pageNumber="220">Text-figs 1c</figureCitation>
,
<figureCitation box="[998,1055,1420,1444]" captionStart="Text-fig" captionStartId="4.[147,228,1735,1756]" captionTargetBox="[148,1463,151,1719]" captionTargetId="figure-2@4.[-3,1584,55,2300]" captionTargetPageId="4" captionText="Text-fig. 2. SEM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Flowers in oblique lateral view showing remains of calyx and slightly semi-inferior ovary with elongated apical style (a); note larger openings in the floral tissue (asterisk) interpreted as schizogenous secretory cavities and the stomata-like secretory structures on the upper portion of the ovary (arrows) that are interpreted as nectariferous (b). c: Detail of ovary surface showing secretory stomata-like structures (arrows). d: Flower in lateral view showing fragmentary calyx and broken slightly semi-inferior ovary with secretory stomata-like structures; note the point of attachment of the central placenta (pl). e: Cluster of seeds removed from the ovary in (d) showing reticulate surface. f: Outer (abaxial)surface of calyx lobe showing the slightly pointed papillae and scattered, fine trichomes (arrows). g: Triaperturate pollen grains from the ovary surface. Specimens, Mira 100-S153146 (a, b), Mira 100-S170155 (c), Mira 100-S101266 (d, e), Mira 105-S100732 (f), Mira 100-S170125 (g). Scale bars = 600 µm (a, b, d), 300 µm (f), 100 µm (c, e), 10 µm (g)." figureDoi="http://doi.org/10.5281/zenodo.7167565" httpUri="https://zenodo.org/record/7167565/files/figure.png" pageId="6" pageNumber="220">2b, c</figureCitation>
). We think it likely that this zone was nectariferous and that the openings were nectar secreting. In some specimens there appear to be remains of septa in the apical part of the ovary, but these may be invaginations of the ovary wall and they do not reach the centre of the ovary (
<figureCitation box="[922,1055,1582,1606]" captionStart="Text-fig" captionStartId="8.[148,226,1363,1384]" captionTargetBox="[148,1461,144,1347]" captionTargetId="figure-291@8.[-5,1582,47,2292]" captionTargetPageId="8" captionText="Text-fig. 5. SRXTM imagesof Miranthuselegans gen. et sp. nov.;Mira locality, Portugal. a, b: Transverse sections offlower (a, orthoslice xy0665 close to the apex of placenta; b, orthoslice xy0800 in middle part of placenta) showing remains of calyx with distinct bundles (arrows), ovary wall (ow) and numerous ovules (ov) on the central mushroom-shaped globose placenta (pl) with central column (cc). c: Transverse section of flower (orthoslice xy0620) through perianth and ovary (ow) at a level above the placenta showing ovules (ov) and cellular preservation of the sepal bundles (arrows shown for one sepal); note abaxial surface of sepals with thick-walled epidermal cells, thick cuticle, and fine pointed verrucae. d: Longitudinal section of flower (orthoslice xz0500) showing perigynous position of calyx and semi-inferior ovary (ow, ovary wall) with a central placenta (pl), central column (cc) and numerous ovules (ov); note spiny verrucae on abaxial surface of calyx lobes. Specimens, Mira 100-S153145 (a, b), Mira 100-S170155 (c, d, holotype). Scale bars = 600 µm (a–d)." figureDoi="http://doi.org/10.5281/zenodo.7167571" httpUri="https://zenodo.org/record/7167571/files/figure.png" pageId="6" pageNumber="220">Text-fig. 5a</figureCitation>
). The style is long, slender (
<figureCitation captionStart="Text-fig" captionStartId="3.[125,206,1774,1795]" captionTargetBox="[125,1440,151,1758]" captionTargetId="figure-2@3.[-26,1561,55,2300]" captionTargetPageId="3" captionText="Text-fig. 1. SEM images of flowers of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Flowers in lateral view showing elongated pedicel, narrowly triangular sepals and elongated protruding style (a); note the large openings in the floral tissue and pedicel (asterisks) interpreted as schizogenous secretory cavities. c: Flower in lateral view with portion of the calyx missing exposing the ovary wall and slightly raised nectariferous ring with probable stomata-like secretory structures (arrow). d: Flower in lateral view showing long pedicel and three of the five tepals; note the elongated narrowly triangular form of the sepals. e: Flower in oblique lateral view with portion of the calyx missing exposing the ovary and elongated style. f, g: Flowers in apical view showing the bases of five sepals (f) and apex of the five-parted ovary; note larger openings in the floral tissue (asterisk) interpreted as schizogenous secretory cavities. Specimens, Mira 100-S170155 (a, holotype), Mira 100-S153145 (b, c, g), Mira 100- S101267 (d), Mira 105-S100732 (e), Mira 100-S101268 (f). Scale bars = 600 µm (a–g)." figureDoi="http://doi.org/10.5281/zenodo.7167563" httpUri="https://zenodo.org/record/7167563/files/figure.png" pageId="6" pageNumber="220">Text-figs 1e</figureCitation>
,
<figureCitation box="[889,917,1615,1639]" captionStart="Text-fig" captionStartId="7.[125,207,1562,1583]" captionTargetBox="[125,1439,151,1546]" captionTargetId="figure-148@7.[-27,1560,47,2292]" captionTargetPageId="7" captionText="Text-fig. 4. SRXTM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Transverse (a) and longitudinal (b) sections of flower showing the ovary wall (ow) and numerous ovules (ov) borne on the mushroom-shaped central globose placenta (pl, blue); placenta shown as a voltex rendering added to orthoslices; note the large openings in the floral tissue (asterisks) interpreted as schizogenous secretory cavities. c, d: Longitudinal (c) and transverse (d) sections of flower (c, volume rendering cut between orthoslice yz0970-1005; d, volume rendering cut between orthoslice xy0780-0820) showing semiinferior ovary (ow, ovary wall) with sepals inserted at the rim of the hypanthium, central column (cc) with mushroom-shaped globose placenta (pl) bearing numerous ovules (ov). Specimens, Mira 100-S153146 (a, b), Mira 100-S170155 (c, d, holotype). Scale bars = 600 µm (a–d)." figureDoi="http://doi.org/10.5281/zenodo.7167569" httpUri="https://zenodo.org/record/7167569/files/figure.png" pageId="6" pageNumber="220">4c</figureCitation>
), and distinctly five-angled in cross-section with a vascular bundle that extends for the length of the style in each of the angles (
<figureCitation box="[1089,1248,1680,1704]" captionStart="Text-fig" captionStartId="9.[125,206,1251,1272]" captionTargetBox="[125,1440,151,1236]" captionTargetId="figure-297@9.[-26,1561,55,2300]" captionTargetPageId="9" captionText="Text-fig. 6. SRXTM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a: Volume rendering of flower in apical view showing five narrow calyx lobes; note that the apical part of the five-angled style is sealed (arrow). b: Volume rendering of flower (cut at orthoslice xy0796) showing transverse sections of calyx lobes with three distinct bundles indicated by arrows in one of the lobes, ovary wall (ow) and central placenta (pl) bearing numerous ovules (ov). c, d: Transverse sections of style (c, orthoslice xy0153; d, orthoslice xy0222) showing five angled form, sealed near apex (c) and with prominent central canal further farther down (d). e: Volume rendering of flower (cut at orthoslice xz1024) in longitudinal section showing calyx, semi-inferior ovary with ovary wall (ow), and central placenta (pl) bearing numerous ovules (ov). Specimen, Mira 100-S153145 (a–d). Scale bars = 600 µm (a, b, e), 100 µm (c, d)." figureDoi="http://doi.org/10.5281/zenodo.7167573" httpUri="https://zenodo.org/record/7167573/files/figure.png" pageId="6" pageNumber="220">Text-fig. 6c, d</figureCitation>
). The style is hollow and the stylar canal is wide along its full length (
<figureCitation captionStart="Text-fig" captionStartId="9.[125,206,1251,1272]" captionTargetBox="[125,1440,151,1236]" captionTargetId="figure-297@9.[-26,1561,55,2300]" captionTargetPageId="9" captionText="Text-fig. 6. SRXTM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a: Volume rendering of flower in apical view showing five narrow calyx lobes; note that the apical part of the five-angled style is sealed (arrow). b: Volume rendering of flower (cut at orthoslice xy0796) showing transverse sections of calyx lobes with three distinct bundles indicated by arrows in one of the lobes, ovary wall (ow) and central placenta (pl) bearing numerous ovules (ov). c, d: Transverse sections of style (c, orthoslice xy0153; d, orthoslice xy0222) showing five angled form, sealed near apex (c) and with prominent central canal further farther down (d). e: Volume rendering of flower (cut at orthoslice xz1024) in longitudinal section showing calyx, semi-inferior ovary with ovary wall (ow), and central placenta (pl) bearing numerous ovules (ov). Specimen, Mira 100-S153145 (a–d). Scale bars = 600 µm (a, b, e), 100 µm (c, d)." figureDoi="http://doi.org/10.5281/zenodo.7167573" httpUri="https://zenodo.org/record/7167573/files/figure.png" pageId="6" pageNumber="220">Text-fig. 6d</figureCitation>
), except near the tip where it is closed (
<figureCitation box="[1337,1469,1745,1769]" captionStart="Text-fig" captionStartId="9.[125,206,1251,1272]" captionTargetBox="[125,1440,151,1236]" captionTargetId="figure-297@9.[-26,1561,55,2300]" captionTargetPageId="9" captionText="Text-fig. 6. SRXTM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a: Volume rendering of flower in apical view showing five narrow calyx lobes; note that the apical part of the five-angled style is sealed (arrow). b: Volume rendering of flower (cut at orthoslice xy0796) showing transverse sections of calyx lobes with three distinct bundles indicated by arrows in one of the lobes, ovary wall (ow) and central placenta (pl) bearing numerous ovules (ov). c, d: Transverse sections of style (c, orthoslice xy0153; d, orthoslice xy0222) showing five angled form, sealed near apex (c) and with prominent central canal further farther down (d). e: Volume rendering of flower (cut at orthoslice xz1024) in longitudinal section showing calyx, semi-inferior ovary with ovary wall (ow), and central placenta (pl) bearing numerous ovules (ov). Specimen, Mira 100-S153145 (a–d). Scale bars = 600 µm (a, b, e), 100 µm (c, d)." figureDoi="http://doi.org/10.5281/zenodo.7167573" httpUri="https://zenodo.org/record/7167573/files/figure.png" pageId="6" pageNumber="220">Text-fig. 6c</figureCitation>
). In some specimens, the style is broken, but the specimens available give no indication of different style lengths and the flower is interpreted as homostylous. Placentation is free and central, with a dome-shaped, almost globose placenta,
</paragraph>
<paragraph blockId="6.[129,1482,1969,2048]" pageId="6" pageNumber="220">
<emphasis bold="true" pageId="6" pageNumber="220">alternating with the corolla lobes; note verrucate abaxial surface of calyx lobes (ca) and ovary wall (ow) partly distorted and displaced to one side. f: Longitudinal section (orthoslice yz0567) of flower bud showing ovary wall (ow) and insertion of calyx (ca), corolla (co) and stamens (st) on a hypanthium rim. Specimen, Mira 100-S266059 (a–f). Scale bars = 600 µm (a, b), 300 µm (c–f).</emphasis>
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7167569" ID-Zenodo-Dep="7167569" httpUri="https://zenodo.org/record/7167569/files/figure.png" pageId="7" pageNumber="221" startId="7.[125,207,1562,1583]" targetBox="[125,1439,151,1546]" targetPageId="7">
<paragraph blockId="7.[125,1440,1562,1788]" pageId="7" pageNumber="221">
Text-fig. 4. SRXTM images of
<taxonomicName authority="Friis &amp; Crane &amp; Pedersen, 2021" authorityName="Friis &amp; Crane &amp; Pedersen" authorityYear="2021" box="[443,632,1562,1583]" class="Magnoliopsida" family="Primulaceae" genus="Miranthus" kingdom="Plantae" order="Ericales" pageId="7" pageNumber="221" phylum="Tracheophyta" rank="species" species="elegans" status="gen. et sp. nov.">
<emphasis bold="true" box="[443,632,1562,1583]" italics="true" pageId="7" pageNumber="221">Miranthus elegans</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[639,797,1562,1583]" pageId="7" pageNumber="221" rank="species">gen. et sp. nov.</taxonomicNameLabel>
; Mira locality, Portugal. a, b: Transverse (a) and longitudinal (b) sections of flower showing the ovary wall (ow) and numerous ovules (ov) borne on the mushroom-shaped central globose placenta (pl, blue); placenta shown as a voltex rendering added to orthoslices; note the large openings in the floral tissue (asterisks) interpreted as schizogenous secretory cavities. c, d: Longitudinal (c) and transverse (d) sections of flower (c, volume rendering cut between orthoslice yz0970-1005; d, volume rendering cut between orthoslice xy0780-0820) showing semiinferior ovary (ow, ovary wall) with sepals inserted at the rim of the hypanthium, central column (cc) with mushroom-shaped globose placenta (pl) bearing numerous ovules (ov). Specimens, Mira 100-S153146 (a, b), Mira 100-S170155 (c, d, holotype). Scale bars = 600 µm (a–d).
</paragraph>
</caption>
<paragraph blockId="7.[106,746,1851,2100]" pageId="7" pageNumber="221">
borne on a central column (
<figureCitation box="[408,568,1851,1875]" captionStart="Text-fig" captionStartId="7.[125,207,1562,1583]" captionTargetBox="[125,1439,151,1546]" captionTargetId="figure-148@7.[-27,1560,47,2292]" captionTargetPageId="7" captionText="Text-fig. 4. SRXTM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Transverse (a) and longitudinal (b) sections of flower showing the ovary wall (ow) and numerous ovules (ov) borne on the mushroom-shaped central globose placenta (pl, blue); placenta shown as a voltex rendering added to orthoslices; note the large openings in the floral tissue (asterisks) interpreted as schizogenous secretory cavities. c, d: Longitudinal (c) and transverse (d) sections of flower (c, volume rendering cut between orthoslice yz0970-1005; d, volume rendering cut between orthoslice xy0780-0820) showing semiinferior ovary (ow, ovary wall) with sepals inserted at the rim of the hypanthium, central column (cc) with mushroom-shaped globose placenta (pl) bearing numerous ovules (ov). Specimens, Mira 100-S153146 (a, b), Mira 100-S170155 (c, d, holotype). Scale bars = 600 µm (a–d)." figureDoi="http://doi.org/10.5281/zenodo.7167569" httpUri="https://zenodo.org/record/7167569/files/figure.png" pageId="7" pageNumber="221">Text-figs 4a–d</figureCitation>
,
<figureCitation box="[581,635,1851,1875]" captionStart="Text-fig" captionStartId="8.[148,226,1363,1384]" captionTargetBox="[148,1461,144,1347]" captionTargetId="figure-291@8.[-5,1582,47,2292]" captionTargetPageId="8" captionText="Text-fig. 5. SRXTM imagesof Miranthuselegans gen. et sp. nov.;Mira locality, Portugal. a, b: Transverse sections offlower (a, orthoslice xy0665 close to the apex of placenta; b, orthoslice xy0800 in middle part of placenta) showing remains of calyx with distinct bundles (arrows), ovary wall (ow) and numerous ovules (ov) on the central mushroom-shaped globose placenta (pl) with central column (cc). c: Transverse section of flower (orthoslice xy0620) through perianth and ovary (ow) at a level above the placenta showing ovules (ov) and cellular preservation of the sepal bundles (arrows shown for one sepal); note abaxial surface of sepals with thick-walled epidermal cells, thick cuticle, and fine pointed verrucae. d: Longitudinal section of flower (orthoslice xz0500) showing perigynous position of calyx and semi-inferior ovary (ow, ovary wall) with a central placenta (pl), central column (cc) and numerous ovules (ov); note spiny verrucae on abaxial surface of calyx lobes. Specimens, Mira 100-S153145 (a, b), Mira 100-S170155 (c, d, holotype). Scale bars = 600 µm (a–d)." figureDoi="http://doi.org/10.5281/zenodo.7167571" httpUri="https://zenodo.org/record/7167571/files/figure.png" pageId="7" pageNumber="221">5a–b</figureCitation>
,
<figureCitation box="[648,731,1851,1876]" captionStart="Text-fig" captionStartId="9.[125,206,1251,1272]" captionTargetBox="[125,1440,151,1236]" captionTargetId="figure-297@9.[-26,1561,55,2300]" captionTargetPageId="9" captionText="Text-fig. 6. SRXTM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a: Volume rendering of flower in apical view showing five narrow calyx lobes; note that the apical part of the five-angled style is sealed (arrow). b: Volume rendering of flower (cut at orthoslice xy0796) showing transverse sections of calyx lobes with three distinct bundles indicated by arrows in one of the lobes, ovary wall (ow) and central placenta (pl) bearing numerous ovules (ov). c, d: Transverse sections of style (c, orthoslice xy0153; d, orthoslice xy0222) showing five angled form, sealed near apex (c) and with prominent central canal further farther down (d). e: Volume rendering of flower (cut at orthoslice xz1024) in longitudinal section showing calyx, semi-inferior ovary with ovary wall (ow), and central placenta (pl) bearing numerous ovules (ov). Specimen, Mira 100-S153145 (a–d). Scale bars = 600 µm (a, b, e), 100 µm (c, d)." figureDoi="http://doi.org/10.5281/zenodo.7167573" httpUri="https://zenodo.org/record/7167573/files/figure.png" pageId="7" pageNumber="221">6a, b, e</figureCitation>
). Numerous ovules are densely spaced on the placenta, but not immersed in its surface (
<figureCitation box="[407,537,1915,1940]" captionStart="Text-fig" captionStartId="4.[147,228,1735,1756]" captionTargetBox="[148,1463,151,1719]" captionTargetId="figure-2@4.[-3,1584,55,2300]" captionTargetPageId="4" captionText="Text-fig. 2. SEM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Flowers in oblique lateral view showing remains of calyx and slightly semi-inferior ovary with elongated apical style (a); note larger openings in the floral tissue (asterisk) interpreted as schizogenous secretory cavities and the stomata-like secretory structures on the upper portion of the ovary (arrows) that are interpreted as nectariferous (b). c: Detail of ovary surface showing secretory stomata-like structures (arrows). d: Flower in lateral view showing fragmentary calyx and broken slightly semi-inferior ovary with secretory stomata-like structures; note the point of attachment of the central placenta (pl). e: Cluster of seeds removed from the ovary in (d) showing reticulate surface. f: Outer (abaxial)surface of calyx lobe showing the slightly pointed papillae and scattered, fine trichomes (arrows). g: Triaperturate pollen grains from the ovary surface. Specimens, Mira 100-S153146 (a, b), Mira 100-S170155 (c), Mira 100-S101266 (d, e), Mira 105-S100732 (f), Mira 100-S170125 (g). Scale bars = 600 µm (a, b, d), 300 µm (f), 100 µm (c, e), 10 µm (g)." figureDoi="http://doi.org/10.5281/zenodo.7167565" httpUri="https://zenodo.org/record/7167565/files/figure.png" pageId="7" pageNumber="221">Text-figs 2e</figureCitation>
,
<figureCitation box="[548,602,1915,1939]" captionStart="Text-fig" captionStartId="7.[125,207,1562,1583]" captionTargetBox="[125,1439,151,1546]" captionTargetId="figure-148@7.[-27,1560,47,2292]" captionTargetPageId="7" captionText="Text-fig. 4. SRXTM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Transverse (a) and longitudinal (b) sections of flower showing the ovary wall (ow) and numerous ovules (ov) borne on the mushroom-shaped central globose placenta (pl, blue); placenta shown as a voltex rendering added to orthoslices; note the large openings in the floral tissue (asterisks) interpreted as schizogenous secretory cavities. c, d: Longitudinal (c) and transverse (d) sections of flower (c, volume rendering cut between orthoslice yz0970-1005; d, volume rendering cut between orthoslice xy0780-0820) showing semiinferior ovary (ow, ovary wall) with sepals inserted at the rim of the hypanthium, central column (cc) with mushroom-shaped globose placenta (pl) bearing numerous ovules (ov). Specimens, Mira 100-S153146 (a, b), Mira 100-S170155 (c, d, holotype). Scale bars = 600 µm (a–d)." figureDoi="http://doi.org/10.5281/zenodo.7167569" httpUri="https://zenodo.org/record/7167569/files/figure.png" pageId="7" pageNumber="221">4a–d</figureCitation>
,
<figureCitation box="[613,667,1915,1939]" captionStart="Text-fig" captionStartId="8.[148,226,1363,1384]" captionTargetBox="[148,1461,144,1347]" captionTargetId="figure-291@8.[-5,1582,47,2292]" captionTargetPageId="8" captionText="Text-fig. 5. SRXTM imagesof Miranthuselegans gen. et sp. nov.;Mira locality, Portugal. a, b: Transverse sections offlower (a, orthoslice xy0665 close to the apex of placenta; b, orthoslice xy0800 in middle part of placenta) showing remains of calyx with distinct bundles (arrows), ovary wall (ow) and numerous ovules (ov) on the central mushroom-shaped globose placenta (pl) with central column (cc). c: Transverse section of flower (orthoslice xy0620) through perianth and ovary (ow) at a level above the placenta showing ovules (ov) and cellular preservation of the sepal bundles (arrows shown for one sepal); note abaxial surface of sepals with thick-walled epidermal cells, thick cuticle, and fine pointed verrucae. d: Longitudinal section of flower (orthoslice xz0500) showing perigynous position of calyx and semi-inferior ovary (ow, ovary wall) with a central placenta (pl), central column (cc) and numerous ovules (ov); note spiny verrucae on abaxial surface of calyx lobes. Specimens, Mira 100-S153145 (a, b), Mira 100-S170155 (c, d, holotype). Scale bars = 600 µm (a–d)." figureDoi="http://doi.org/10.5281/zenodo.7167571" httpUri="https://zenodo.org/record/7167571/files/figure.png" pageId="7" pageNumber="221">5a–d</figureCitation>
,
<figureCitation box="[677,731,1915,1940]" captionStart="Text-fig" captionStartId="9.[125,206,1251,1272]" captionTargetBox="[125,1440,151,1236]" captionTargetId="figure-297@9.[-26,1561,55,2300]" captionTargetPageId="9" captionText="Text-fig. 6. SRXTM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a: Volume rendering of flower in apical view showing five narrow calyx lobes; note that the apical part of the five-angled style is sealed (arrow). b: Volume rendering of flower (cut at orthoslice xy0796) showing transverse sections of calyx lobes with three distinct bundles indicated by arrows in one of the lobes, ovary wall (ow) and central placenta (pl) bearing numerous ovules (ov). c, d: Transverse sections of style (c, orthoslice xy0153; d, orthoslice xy0222) showing five angled form, sealed near apex (c) and with prominent central canal further farther down (d). e: Volume rendering of flower (cut at orthoslice xz1024) in longitudinal section showing calyx, semi-inferior ovary with ovary wall (ow), and central placenta (pl) bearing numerous ovules (ov). Specimen, Mira 100-S153145 (a–d). Scale bars = 600 µm (a, b, e), 100 µm (c, d)." figureDoi="http://doi.org/10.5281/zenodo.7167573" httpUri="https://zenodo.org/record/7167573/files/figure.png" pageId="7" pageNumber="221">6b, e</figureCitation>
). Ovules are bitegmic and anatropous, about
<quantity box="[589,680,1948,1972]" metricMagnitude="-4" metricUnit="m" metricValue="1.4" pageId="7" pageNumber="221" unit="mm" value="0.14">0.14 mm</quantity>
long, with a reticulate seed surface (
<figureCitation box="[429,556,1979,2004]" captionStart="Text-fig" captionStartId="4.[147,228,1735,1756]" captionTargetBox="[148,1463,151,1719]" captionTargetId="figure-2@4.[-3,1584,55,2300]" captionTargetPageId="4" captionText="Text-fig. 2. SEM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Flowers in oblique lateral view showing remains of calyx and slightly semi-inferior ovary with elongated apical style (a); note larger openings in the floral tissue (asterisk) interpreted as schizogenous secretory cavities and the stomata-like secretory structures on the upper portion of the ovary (arrows) that are interpreted as nectariferous (b). c: Detail of ovary surface showing secretory stomata-like structures (arrows). d: Flower in lateral view showing fragmentary calyx and broken slightly semi-inferior ovary with secretory stomata-like structures; note the point of attachment of the central placenta (pl). e: Cluster of seeds removed from the ovary in (d) showing reticulate surface. f: Outer (abaxial)surface of calyx lobe showing the slightly pointed papillae and scattered, fine trichomes (arrows). g: Triaperturate pollen grains from the ovary surface. Specimens, Mira 100-S153146 (a, b), Mira 100-S170155 (c), Mira 100-S101266 (d, e), Mira 105-S100732 (f), Mira 100-S170125 (g). Scale bars = 600 µm (a, b, d), 300 µm (f), 100 µm (c, e), 10 µm (g)." figureDoi="http://doi.org/10.5281/zenodo.7167565" httpUri="https://zenodo.org/record/7167565/files/figure.png" pageId="7" pageNumber="221">Text-fig. 2e</figureCitation>
).
</paragraph>
<paragraph blockId="7.[106,746,1851,2100]" lastBlockId="7.[820,1459,1851,2099]" pageId="7" pageNumber="221">
Among the post-anthetic flowers, none has a fully developed fruit. However, some are clearly five-parted, have five valves and are interpreted as young capsules (
<figureCitation box="[652,872,1851,2099]" captionStart="Text-fig" captionStartId="3.[125,206,1774,1795]" captionTargetBox="[125,1440,151,1758]" captionTargetId="figure-2@3.[-26,1561,55,2300]" captionTargetPageId="3" captionText="Text-fig. 1. SEM images of flowers of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Flowers in lateral view showing elongated pedicel, narrowly triangular sepals and elongated protruding style (a); note the large openings in the floral tissue and pedicel (asterisks) interpreted as schizogenous secretory cavities. c: Flower in lateral view with portion of the calyx missing exposing the ovary wall and slightly raised nectariferous ring with probable stomata-like secretory structures (arrow). d: Flower in lateral view showing long pedicel and three of the five tepals; note the elongated narrowly triangular form of the sepals. e: Flower in oblique lateral view with portion of the calyx missing exposing the ovary and elongated style. f, g: Flowers in apical view showing the bases of five sepals (f) and apex of the five-parted ovary; note larger openings in the floral tissue (asterisk) interpreted as schizogenous secretory cavities. Specimens, Mira 100-S170155 (a, holotype), Mira 100-S153145 (b, c, g), Mira 100- S101267 (d), Mira 105-S100732 (e), Mira 100-S101268 (f). Scale bars = 600 µm (a–g)." figureDoi="http://doi.org/10.5281/zenodo.7167563" httpUri="https://zenodo.org/record/7167563/files/figure.png" pageId="7" pageNumber="221">Text-fig. 1e, f</figureCitation>
). In
<specimenCount box="[922,1069,1851,1875]" pageId="7" pageNumber="221" type="generic">one specimen</specimenCount>
the ovary is split open along one of the valves (
<figureCitation box="[945,1076,1883,1907]" captionStart="Text-fig" captionStartId="4.[147,228,1735,1756]" captionTargetBox="[148,1463,151,1719]" captionTargetId="figure-2@4.[-3,1584,55,2300]" captionTargetPageId="4" captionText="Text-fig. 2. SEM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Flowers in oblique lateral view showing remains of calyx and slightly semi-inferior ovary with elongated apical style (a); note larger openings in the floral tissue (asterisk) interpreted as schizogenous secretory cavities and the stomata-like secretory structures on the upper portion of the ovary (arrows) that are interpreted as nectariferous (b). c: Detail of ovary surface showing secretory stomata-like structures (arrows). d: Flower in lateral view showing fragmentary calyx and broken slightly semi-inferior ovary with secretory stomata-like structures; note the point of attachment of the central placenta (pl). e: Cluster of seeds removed from the ovary in (d) showing reticulate surface. f: Outer (abaxial)surface of calyx lobe showing the slightly pointed papillae and scattered, fine trichomes (arrows). g: Triaperturate pollen grains from the ovary surface. Specimens, Mira 100-S153146 (a, b), Mira 100-S170155 (c), Mira 100-S101266 (d, e), Mira 105-S100732 (f), Mira 100-S170125 (g). Scale bars = 600 µm (a, b, d), 300 µm (f), 100 µm (c, e), 10 µm (g)." figureDoi="http://doi.org/10.5281/zenodo.7167565" httpUri="https://zenodo.org/record/7167565/files/figure.png" pageId="7" pageNumber="221">Text-fig. 2d</figureCitation>
). The epidermal cells of the young capsules are small, equiaxial, and often with tiny papillate projections. Stomata occur scattered over the surface of the calyx lobes and pedicel together with smaller openings that are probably trichome bases. Larger openings are irregularly scattered and are interpreted as schizogenous secretory cavities that are sometimes burst (
<figureCitation box="[1182,1339,2075,2099]" captionStart="Text-fig" captionStartId="3.[125,206,1774,1795]" captionTargetBox="[125,1440,151,1758]" captionTargetId="figure-2@3.[-26,1561,55,2300]" captionTargetPageId="3" captionText="Text-fig. 1. SEM images of flowers of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Flowers in lateral view showing elongated pedicel, narrowly triangular sepals and elongated protruding style (a); note the large openings in the floral tissue and pedicel (asterisks) interpreted as schizogenous secretory cavities. c: Flower in lateral view with portion of the calyx missing exposing the ovary wall and slightly raised nectariferous ring with probable stomata-like secretory structures (arrow). d: Flower in lateral view showing long pedicel and three of the five tepals; note the elongated narrowly triangular form of the sepals. e: Flower in oblique lateral view with portion of the calyx missing exposing the ovary and elongated style. f, g: Flowers in apical view showing the bases of five sepals (f) and apex of the five-parted ovary; note larger openings in the floral tissue (asterisk) interpreted as schizogenous secretory cavities. Specimens, Mira 100-S170155 (a, holotype), Mira 100-S153145 (b, c, g), Mira 100- S101267 (d), Mira 105-S100732 (e), Mira 100-S101268 (f). Scale bars = 600 µm (a–g)." figureDoi="http://doi.org/10.5281/zenodo.7167563" httpUri="https://zenodo.org/record/7167563/files/figure.png" pageId="7" pageNumber="221">Text-figs 1b, c</figureCitation>
,
<figureCitation box="[1350,1380,2075,2099]" captionStart="Text-fig" captionStartId="7.[125,207,1562,1583]" captionTargetBox="[125,1439,151,1546]" captionTargetId="figure-148@7.[-27,1560,47,2292]" captionTargetPageId="7" captionText="Text-fig. 4. SRXTM images of Miranthus elegans gen. et sp. nov.; Mira locality, Portugal. a, b: Transverse (a) and longitudinal (b) sections of flower showing the ovary wall (ow) and numerous ovules (ov) borne on the mushroom-shaped central globose placenta (pl, blue); placenta shown as a voltex rendering added to orthoslices; note the large openings in the floral tissue (asterisks) interpreted as schizogenous secretory cavities. c, d: Longitudinal (c) and transverse (d) sections of flower (c, volume rendering cut between orthoslice yz0970-1005; d, volume rendering cut between orthoslice xy0780-0820) showing semiinferior ovary (ow, ovary wall) with sepals inserted at the rim of the hypanthium, central column (cc) with mushroom-shaped globose placenta (pl) bearing numerous ovules (ov). Specimens, Mira 100-S153146 (a, b), Mira 100-S170155 (c, d, holotype). Scale bars = 600 µm (a–d)." figureDoi="http://doi.org/10.5281/zenodo.7167569" httpUri="https://zenodo.org/record/7167569/files/figure.png" pageId="7" pageNumber="221">4b</figureCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>
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