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<document id="75A59F95E3386D7C786DE2753A1459A1" ID-CLB-Dataset="300551" ID-DOI="10.5281/zenodo.13173975" ID-GBIF-Dataset="925f71cb-fbf8-439b-81c2-96067818d7e2" ID-ISSN="1732-2421" ID-Zenodo-Dep="13173975" IM.bibliography_approvedBy="felipe" IM.illustrations_approvedBy="juliana" IM.materialsCitations_approvedBy="juliana" IM.metadata_approvedBy="juliana" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="juliana" checkinTime="1721350226465" checkinUser="felipe" docAuthor="V. Naugolnykh, Serge" docDate="2002" docId="03EECE3DCF6BFFC28922FBAEFD5DF916" docLanguage="en" docName="ActaPalaeontolPol.47.3.469-478.pdf" docOrigin="Acta Palaeontologica Polonica 47 (3)" docStyle="DocumentStyle:93F5927C23D90727DE855278A6F17B37.1:ActaPalaeontolPol.2002-2009.journal_article" docStyleId="93F5927C23D90727DE855278A6F17B37" docStyleName="ActaPalaeontolPol.2002-2009.journal_article" docStyleVersion="1" docTitle="Todites lobulatus Naugolnykh, 2002, sp. nov." docType="treatment" docVersion="3" lastPageNumber="473" masterDocId="FFD7B645CF6AFFC68944FF8CFF9DFFE5" masterDocTitle="A new species of Todites (Pteridophyta) with in situ spores from the Upper Permian of Pechora Cis-Urals (Russia)" masterLastPageNumber="478" masterPageNumber="469" pageNumber="470" updateTime="1722619861464" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0">
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<mods:title id="B49C6CD19FB8B2997BDB069801EBFCCD">A new species of Todites (Pteridophyta) with in situ spores from the Upper Permian of Pechora Cis-Urals (Russia)</mods:title>
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<mods:name id="C7DF3B20A78F7B48B4428CFB0C6923B5" type="personal">
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<mods:namePart id="2572452CA381722F42496415D77651B4">V. Naugolnykh, Serge</mods:namePart>
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<mods:title id="EFC1C8249396A587BB147AD4142B4664">Acta Palaeontologica Polonica</mods:title>
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<mods:part id="A92B6DCC84C636D4434309FB7B6AFA32">
<mods:date id="FC63B74BC395485E5065FF558EFD3862">2002</mods:date>
<mods:detail id="9FC9F31141629E638DBD642B1E1AAD0C" type="pubDate">
<mods:number id="F38FF15C7E56057DE7DEE18040293CE1">2002-12-31</mods:number>
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<mods:number id="D80ADD4D0D9FAD32062B03E7F401EB86">47</mods:number>
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<mods:classification id="133CEC612D8DD880483017604759EF4D">journal article</mods:classification>
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<mods:identifier id="4F6EBE243B8C684FEC3B78C5A734BC7A" type="DOI">10.5281/zenodo.13173975</mods:identifier>
<mods:identifier id="2EAEA9B4AF455293173D0F3B7FC7342E" type="GBIF-Dataset">925f71cb-fbf8-439b-81c2-96067818d7e2</mods:identifier>
<mods:identifier id="F6936876D2DF840188925BA79FF3E500" type="ISSN">1732-2421</mods:identifier>
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<treatment id="03EECE3DCF6BFFC28922FBAEFD5DF916" ID-DOI="http://doi.org/10.5281/zenodo.13173979" ID-Zenodo-Dep="13173979" LSID="urn:lsid:plazi:treatment:03EECE3DCF6BFFC28922FBAEFD5DF916" httpUri="http://treatment.plazi.org/id/03EECE3DCF6BFFC28922FBAEFD5DF916" lastPageId="4" lastPageNumber="473" pageId="1" pageNumber="470">
<subSubSection id="C35D2CA0CF6BFFC78922FBAEFE37FBDA" box="[102,426,1058,1087]" pageId="1" pageNumber="470" type="nomenclature">
<paragraph id="8BF87F2BCF6BFFC78922FBAEFE37FBDA" blockId="1.[102,466,950,1122]" box="[102,426,1058,1087]" pageId="1" pageNumber="470">
<heading id="D0B0C847CF6BFFC78922FBAEFE37FBDA" box="[102,426,1058,1087]" fontSize="36" level="2" pageId="1" pageNumber="470" reason="7">
<taxonomicName id="4C4704A8CF6BFFC78922FBAEFEDDFBDA" authorityName="V.Naugolnykh" authorityYear="2002" box="[102,320,1058,1087]" class="Polypodiopsida" family="Osmundaceae" genus="Todites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Osmundales" pageId="1" pageNumber="470" phylum="Tracheophyta" rank="species" species="lobulatus" status="sp. nov.">
<emphasis id="B933A339CF6BFFC78922FBAEFEDDFBDA" box="[102,320,1058,1087]" italics="true" pageId="1" pageNumber="470">Todites lobulatus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2001E42CF6BFFC7880CFBAFFE37FBDA" box="[328,426,1059,1087]" pageId="1" pageNumber="470" rank="species">sp. nov.</taxonomicNameLabel>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C35D2CA0CF6BFFC78922FBC0FF5BFB87" box="[102,198,1100,1122]" pageId="1" pageNumber="470" type="description">
<paragraph id="8BF87F2BCF6BFFC78922FBC0FF5BFB87" blockId="1.[102,466,950,1122]" box="[102,198,1100,1122]" pageId="1" pageNumber="470">
<figureCitation id="137C63AECF6BFFC78922FBC0FF5FFB87" box="[102,194,1100,1122]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="A" captionStart-3="Fig" captionStart-4="Fig" captionStartId-0="2.[102,138,1630,1649]" captionStartId-1="3.[102,137,1288,1307]" captionStartId-2="4.[475,496,614,640]" captionStartId-3="5.[102,137,1856,1875]" captionStartId-4="6.[102,137,1751,1770]" captionTargetBox-0="[170,1417,226,1607]" captionTargetBox-1="[102,1266,232,1364]" captionTargetBox-3="[287,1295,227,1834]" captionTargetBox-4="[258,1327,226,1726]" captionTargetId-0="figure-155@2.[170,1417,226,1607]" captionTargetId-1="figure-343@3.[316,1272,226,1361]" captionTargetId-3="figure-9@5.[240,1348,227,1832]" captionTargetId-4="figure-70@6.[258,1330,226,1726]" captionTargetPageId-0="2" captionTargetPageId-1="3" captionTargetPageId-3="5" captionTargetPageId-4="6" captionText-0="Fig.3. Todites lobulatus sp.nov.Intinskian Formation, Ufimian, Upper Permian.Holotype GIN 4846/102. A, B.General morphology of fertile pinna, bef − ore (A) and after (B) preparation; A, × 1; B, × 4. C.Two prepared sporangia; left sporangium shows a group of thick−walled cells near the stomium (dark de − pression); × 18. D. The spore of Osmundacidites type, preserved in situ, under SEM, × 800." captionText-1="Fig.4. Todites lobulatus sp.nov.Intinskian Formation, Ufimian, Upper Permian.Holotype GIN 4846/102.Gen − eral morphology of a fertile pinna. Scale bar 0.5 cm." captionText-2="A Fig.5. Todites lobulatus sp.nov. Intinskian Formation, Ufimian, Upper Permian.Holotype GIN 4846/102.Sporangium with open stomium and thick−walled cells located near its right side, photo− graph (A) and line drawing of the same sporangium portion (B). Scale bar 20 mm. B" captionText-3="Fig.6. Todites lobulatus sp.nov.Intinskian Formation, Ufimian, Upper Permian.Macromorphology and in situ spores, based on holotype GIN 4846/102. A, B.Fertile pinnule reconstruction, upper (adaxial, A) and lower (abaxial, B) view, numerous sporangia attached to lower side of the pinnule surface (B). C.Individual sporangium with thick−walled cells on the top area. D.Granulate microstructure of the spore surface. E.General view on the isolated spore. F, G.Spore mass extracted from the sporangia, note different sizes (relatively small spores 1 and 2 and big spores 3 and 4 of F) and different outlines of the spores (round spores 2 and ovoid 1, 3 and 4 of F; round spore 1, rounded subtriangular spore 2 and ovoid spore 3 of G).Scale bars: A, B, 5 mm; C, 0.1 mm; E, F, G, 50 µm; D, 5 µm." captionText-4="Fig.7. Todites lobulatus sp.nov.Intinskian Formation, Ufimian, Upper Permian. In situ spores extracted from the holotype GIN 4846/102. A.Spore mass repeating the initial round form of sporangium, × 50. B.Several spores of different size and form, × 200. C.Spore with clearly seen laesura (left part of the photograph), × 200. D. Spore mass of ovoid form from relatively elongated sporangium, × 50. E. Spore mass in detail, × 200. F. Detail of D, × 100." figureDoi-0="http://doi.org/10.5281/zenodo.13173983" figureDoi-1="http://doi.org/10.5281/zenodo.13173985" figureDoi-3="http://doi.org/10.5281/zenodo.13173989" figureDoi-4="http://doi.org/10.5281/zenodo.13173991" httpUri-0="https://zenodo.org/record/13173983/files/figure.png" httpUri-1="https://zenodo.org/record/13173985/files/figure.png" httpUri-3="https://zenodo.org/record/13173989/files/figure.png" httpUri-4="https://zenodo.org/record/13173991/files/figure.png" pageId="1" pageNumber="470">Figs. 3–7</figureCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="C35D2CA0CF6BFFC78922FBFFFDF9FAEF" pageId="1" pageNumber="470" type="materials_examined">
<paragraph id="8BF87F2BCF6BFFC78922FBFFFEC0FB6C" blockId="1.[102,775,1139,1554]" box="[102,349,1139,1161]" pageId="1" pageNumber="470">
<materialsCitation id="3B2F7576CF6BFFC78922FBFFFEC0FB6C" box="[102,349,1139,1161]" collectionCode="GIN" country="Russia" county="Kozhim-Rudnik station" location="Kozhim River" municipality="Kozhim-Rudnik" pageId="1" pageNumber="470" specimenCode="GIN 4846/102" specimenCount="1" typeStatus="holotype">
<emphasis id="B933A339CF6BFFC78922FBFFFF23FB6C" box="[102,190,1139,1161]" italics="true" pageId="1" pageNumber="470">
<typeStatus id="54FCC189CF6BFFC78922FBFFFF23FB6C" box="[102,190,1139,1161]" pageId="1" pageNumber="470" type="holotype">Holotype</typeStatus>
</emphasis>
:
<specimenCode id="DBE1D750CF6BFFC7898EFBFFFEC5FB6C" box="[202,344,1139,1161]" pageId="1" pageNumber="470">GIN 4846/102</specimenCode>
.
</materialsCitation>
</paragraph>
<paragraph id="8BF87F2BCF6BFFC78922FB19FC9EFB4E" blockId="1.[102,775,1139,1554]" box="[102,771,1173,1195]" pageId="1" pageNumber="470">
<emphasis id="B933A339CF6BFFC78922FB19FF41FB4E" box="[102,220,1173,1195]" italics="true" pageId="1" pageNumber="470">Type horizon</emphasis>
:
<materialsCitation id="3B2F7576CF6BFFC789ADFB19FC9EFB4E" box="[233,771,1173,1195]" country="Russia" county="Kozhim-Rudnik station" location="Bed 103" municipality="Ufimian" pageId="1" pageNumber="470" specimenCount="1" typeStatus="holotype">
<location id="8E9829F0CF6BFFC789ADFB19FEA4FB4E" LSID="urn:lsid:plazi:treatment:03EECE3DCF6BFFC28922FBAEFD5DF916:8E9829F0CF6BFFC789ADFB19FEA4FB4E" box="[233,313,1173,1195]" country="Russia" county="Kozhim-Rudnik station" municipality="Ufimian" name="Bed 103" pageId="1" pageNumber="470">Bed 103</location>
,
<location id="8E9829F0CF6BFFC78806FB19FD98FB4E" LSID="urn:lsid:plazi:treatment:03EECE3DCF6BFFC28922FBAEFD5DF916:8E9829F0CF6BFFC78806FB19FD98FB4E" box="[322,517,1173,1195]" country="Russia" county="Kozhim-Rudnik station" municipality="Ufimian" name="Intinskian Formation" pageId="1" pageNumber="470">Intinskian Formation</location>
,
<location id="8E9829F0CF6BFFC78B4AFB19FD06FB4E" LSID="urn:lsid:plazi:treatment:03EECE3DCF6BFFC28922FBAEFD5DF916:8E9829F0CF6BFFC78B4AFB19FD06FB4E" box="[526,667,1173,1195]" country="Russia" county="Kozhim-Rudnik station" municipality="Ufimian" name="Upper Permian" pageId="1" pageNumber="470">Upper Permian</location>
(
<collectingMunicipality id="6B9CE551CF6BFFC78BE2FB19FD65FB4E" box="[678,760,1173,1195]" pageId="1" pageNumber="470">Ufimian</collectingMunicipality>
).
</materialsCitation>
</paragraph>
<paragraph id="8BF87F2BCF6BFFC78922FB3BFD76FB0D" blockId="1.[102,775,1139,1554]" pageId="1" pageNumber="470">
<materialsCitation id="3B2F7576CF6BFFC78922FB3BFD76FB0D" country="Russia" county="Kozhim-Rudnik station" location="Kozhim River" municipality="Kozhim-Rudnik" pageId="1" pageNumber="470" specimenCount="1" typeStatus="holotype">
<emphasis id="B933A339CF6BFFC78922FB3BFF7CFB29" box="[102,225,1206,1228]" italics="true" pageId="1" pageNumber="470">Type locality</emphasis>
: Left bank of
<location id="8E9829F0CF6BFFC78828FB3AFE6AFB29" LSID="urn:lsid:plazi:treatment:03EECE3DCF6BFFC28922FBAEFD5DF916:8E9829F0CF6BFFC78828FB3AFE6AFB29" box="[364,503,1206,1228]" country="Russia" county="Kozhim-Rudnik station" municipality="Kozhim-Rudnik" name="Kozhim River" pageId="1" pageNumber="470">Kozhim River</location>
,
<quantity id="4CBFD2CECF6BFFC78B45FB3AFDAFFB29" box="[513,562,1206,1228]" metricMagnitude="3" metricUnit="m" metricValue="4.0" pageId="1" pageNumber="470" unit="km" value="4.0">
<locationDeviation id="9A4A6609CF6BFFC78B45FB3AFDAFFB29" box="[513,562,1206,1228]" metricMagnitude="3" metricUnit="m" metricValue="4.0" pageId="1" pageNumber="470" unit="km" value="4.0">4 km</locationDeviation>
</quantity>
downstream from the bridge near
<collectingMunicipality id="6B9CE551CF6BFFC7899CFB5EFEE9FB0D" box="[216,372,1234,1256]" pageId="1" pageNumber="470">Kozhim−Rudnik</collectingMunicipality>
station, the
<location id="8E9829F0CF6BFFC788ADFB5EFD06FB0D" LSID="urn:lsid:plazi:treatment:03EECE3DCF6BFFC28922FBAEFD5DF916:8E9829F0CF6BFFC788ADFB5EFD06FB0D" box="[489,667,1234,1256]" country="Russia" county="Kozhim-Rudnik station" municipality="Kozhim-Rudnik" name="Pechora Cis-Urals" pageId="1" pageNumber="470">Pechora Cis−Urals</location>
,
<collectingCountry id="F3503FBBCF6BFFC78BE0FB5EFD7AFB0D" box="[676,743,1234,1256]" name="Russia" pageId="1" pageNumber="470">Russia</collectingCountry>
.
</materialsCitation>
</paragraph>
<paragraph id="8BF87F2BCF6BFFC78922FB79FDF9FAEF" blockId="1.[102,775,1139,1554]" box="[102,612,1268,1290]" pageId="1" pageNumber="470">
<emphasis id="B933A339CF6BFFC78922FB79FEDEFAEF" box="[102,323,1268,1290]" italics="true" pageId="1" pageNumber="470">Derivation of the name</emphasis>
: From Latin
<taxonomicName id="4C4704A8CF6BFFC78887FB78FD81FAEF" authorityName="V.Naugolnykh" authorityYear="2002" box="[451,540,1268,1290]" class="Polypodiopsida" family="Osmundaceae" genus="Todites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Osmundales" pageId="1" pageNumber="470" phylum="Tracheophyta" rank="species" species="lobulatus">
<emphasis id="B933A339CF6BFFC78887FB78FD81FAEF" box="[451,540,1268,1290]" italics="true" pageId="1" pageNumber="470">lobulatus</emphasis>
</taxonomicName>
, lobed.
</paragraph>
</subSubSection>
<subSubSection id="C35D2CA0CF6BFFC78922FA96FE08F9BF" pageId="1" pageNumber="470" type="diagnosis">
<paragraph id="8BF87F2BCF6BFFC78922FA96FF29F9F7" blockId="1.[102,775,1139,1554]" pageId="1" pageNumber="470">
<emphasis id="B933A339CF6BFFC78922FA96FF45FAD4" box="[102,216,1306,1329]" italics="true" pageId="1" pageNumber="470">Diagnosis</emphasis>
.—Ferns with tripinnate fronds. Pinnules deeply dissected with well developed lobes.Pinnule apex round. Sporangia free, located on abaxial leaf surface.Sporangia round or ovoid, small (
<quantity id="4CBFD2CECF6BFFC7882AFAF5FE5AFA74" box="[366,455,1401,1425]" metricMagnitude="-4" metricUnit="m" metricValue="5.0" pageId="1" pageNumber="470" unit="mm" value="0.5">0.5 mm</quantity>
in diameter), with group of thick−walled cells on sporangia top or side.Spores circular, laesura trilete, radii simple, straight, spore surface sculptured by granulate scabrae.Size of the spores ranges from 20 to 45 µm.
</paragraph>
<paragraph id="8BF87F2BCF6BFFC78922F9AEFE08F9BF" blockId="1.[102,774,1570,1626]" pageId="1" pageNumber="470">
<emphasis id="B933A339CF6BFFC78922F9AEFF55F9DF" box="[102,200,1570,1594]" italics="true" pageId="1" pageNumber="470">Material</emphasis>
.—Two representative pinnae of last order belonging to one individual plant.
</paragraph>
</subSubSection>
<caption id="DF382FA3CF6BFFC78A68F9DBFA90F98F" ID-DOI="http://doi.org/10.5281/zenodo.13173981" ID-Zenodo-Dep="13173981" box="[812,1293,1623,1643]" httpUri="https://zenodo.org/record/13173981/files/figure.png" pageId="1" pageNumber="470" startId="1.[812,845,1623,1642]" targetBox="[816,1485,226,1598]" targetPageId="1" targetType="figure">
<paragraph id="8BF87F2BCF6BFFC78A68F9DBFA90F98F" blockId="1.[812,1293,1623,1643]" box="[812,1293,1623,1643]" pageId="1" pageNumber="470">Fig. 2. Stratigraphical column of the locality studied.</paragraph>
</caption>
<subSubSection id="C35D2CA0CF6BFFC58922F9E7FA8CFA32" lastPageId="3" lastPageNumber="472" pageId="1" pageNumber="470" type="description">
<paragraph id="8BF87F2BCF6BFFC78922F9E7FD61F8C7" blockId="1.[102,775,1642,2018]" pageId="1" pageNumber="470">
<emphasis id="B933A339CF6BFFC78922F9E7FF77F967" box="[102,234,1643,1666]" italics="true" pageId="1" pageNumber="470">Description</emphasis>
.—The single representative fragment of a tripinnate frond with two preserved pinnae of the last order is shown on
<figureCitation id="137C63AECF6BFFC78999F926FED3F927" box="[221,334,1706,1730]" captionStart="Fig" captionStartId="2.[102,138,1630,1649]" captionTargetBox="[170,1417,226,1607]" captionTargetId="figure-155@2.[170,1417,226,1607]" captionTargetPageId="2" captionText="Fig.3. Todites lobulatus sp.nov.Intinskian Formation, Ufimian, Upper Permian.Holotype GIN 4846/102. A, B.General morphology of fertile pinna, bef − ore (A) and after (B) preparation; A, × 1; B, × 4. C.Two prepared sporangia; left sporangium shows a group of thick−walled cells near the stomium (dark de − pression); × 18. D. The spore of Osmundacidites type, preserved in situ, under SEM, × 800." figureDoi="http://doi.org/10.5281/zenodo.13173983" httpUri="https://zenodo.org/record/13173983/files/figure.png" pageId="1" pageNumber="470">Fig.3A–D</figureCitation>
and
<figureCitation id="137C63AECF6BFFC788CCF926FE5FF927" box="[392,450,1706,1730]" captionStart="Fig" captionStartId="3.[102,137,1288,1307]" captionTargetBox="[102,1266,232,1364]" captionTargetId="figure-343@3.[316,1272,226,1361]" captionTargetPageId="3" captionText="Fig.4. Todites lobulatus sp.nov.Intinskian Formation, Ufimian, Upper Permian.Holotype GIN 4846/102.Gen − eral morphology of a fertile pinna. Scale bar 0.5 cm." figureDoi="http://doi.org/10.5281/zenodo.13173985" httpUri="https://zenodo.org/record/13173985/files/figure.png" pageId="1" pageNumber="470">Fig.4</figureCitation>
.The preserved part of the most complete pinna is
<quantity id="4CBFD2CECF6BFFC78828F946FE4DF907" box="[364,464,1738,1762]" metricMagnitude="-2" metricUnit="m" metricValue="3.15" pageId="1" pageNumber="470" unit="mm" value="31.5">31.5 mm</quantity>
long and
<quantity id="4CBFD2CECF6BFFC78B78F946FD0CF907" box="[572,657,1738,1762]" metricMagnitude="-3" metricUnit="m" metricValue="2.8" pageId="1" pageNumber="470" unit="mm" value="2.8">2.8 mm</quantity>
wide. The pinna rachis is straight in the apical part, but slightly curved (undulating) in its proximal part.The rachis width is
<quantity id="4CBFD2CECF6BFFC78BE2F886FD65F8C7" box="[678,760,1802,1826]" metricMagnitude="-3" metricUnit="m" metricValue="1.5" pageId="1" pageNumber="470" unit="mm" value="1.5">1.5 mm</quantity>
.
</paragraph>
<paragraph id="8BF87F2BCF6BFFC78922F8A6FF45F807" blockId="1.[102,775,1642,2018]" pageId="1" pageNumber="470">
The narrow wing is present in the apical part of the pinna. One can see eight pinnules with lobed margins.The pinnules are attached to the pinna rachis in regularly alternate order. Pinnule length varies from 13.3 up to
<quantity id="4CBFD2CECF6BFFC78B6CF806FD0EF847" box="[552,659,1930,1954]" metricMagnitude="-2" metricUnit="m" metricValue="1.58" pageId="1" pageNumber="470" unit="mm" value="15.8">15.8 mm</quantity>
. Maximal width of the pinnules is
<quantity id="4CBFD2CECF6BFFC788CFF826FE4FF827" box="[395,466,1962,1986]" metricMagnitude="-3" metricUnit="m" metricValue="8.0" pageId="1" pageNumber="470" unit="mm" value="8.0">8 mm</quantity>
.Pinnule outlines are sub − triangular.
</paragraph>
<paragraph id="8BF87F2BCF6BFFC78A16F925FBDBF8C4" blockId="1.[812,1485,1705,2018]" pageId="1" pageNumber="470">
The frond surface is exposed on its adaxial side without sporangia.Since the leaf lamina is considerably eroded, the numerous sporangia attached to abaxial frond surface are clearly seen (
<figureCitation id="137C63AECF6BFFC78A84F885FBA4F8C4" box="[960,1081,1801,1825]" captionStart="Fig" captionStartId="2.[102,138,1630,1649]" captionTargetBox="[170,1417,226,1607]" captionTargetId="figure-155@2.[170,1417,226,1607]" captionTargetPageId="2" captionText="Fig.3. Todites lobulatus sp.nov.Intinskian Formation, Ufimian, Upper Permian.Holotype GIN 4846/102. A, B.General morphology of fertile pinna, bef − ore (A) and after (B) preparation; A, × 1; B, × 4. C.Two prepared sporangia; left sporangium shows a group of thick−walled cells near the stomium (dark de − pression); × 18. D. The spore of Osmundacidites type, preserved in situ, under SEM, × 800." figureDoi="http://doi.org/10.5281/zenodo.13173983" httpUri="https://zenodo.org/record/13173983/files/figure.png" pageId="1" pageNumber="470">Fig. 3B, C</figureCitation>
).
</paragraph>
<paragraph id="8BF87F2BCF6BFFC48A16F8A6FDDDF8BA" blockId="1.[812,1485,1705,2018]" lastBlockId="2.[102,775,1766,2019]" lastPageId="2" lastPageNumber="471" pageId="1" pageNumber="470">The pinnules are dissected into well developed lobes in 1/2 of pinnule width in their proximal parts.The apical parts of the pinnules are not so deeply dissected.The pinnate vena − tion is simple.One well developed robust midvein enters each pinnule.Lateral veins enter each pinnule lobe.Lateral veins in the apical part of the pinnule are simple; those of the proximal parts of the pinnules are rarely pinnately branched. Lateral veins originate from the midvein in alternating order, but sometimes almost opposite.The pinnule apex is round, the pinnule base is narrow, sphenopteroid.</paragraph>
<caption id="DF382FA3CF68FFC48922F9D2FC3BF94C" ID-DOI="http://doi.org/10.5281/zenodo.13173983" ID-Zenodo-Dep="13173983" httpUri="https://zenodo.org/record/13173983/files/figure.png" pageId="2" pageNumber="471" startId="2.[102,138,1630,1649]" targetBox="[170,1417,226,1607]" targetPageId="2" targetType="figure">
<paragraph id="8BF87F2BCF68FFC48922F9D2FC3BF94C" blockId="2.[102,1485,1630,1706]" pageId="2" pageNumber="471">
Fig.3.
<taxonomicName id="4C4704A8CF68FFC489E0F9D2FEA3F994" authorityName="V.Naugolnykh" authorityYear="2002" box="[164,318,1630,1649]" class="Polypodiopsida" family="Osmundaceae" genus="Todites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Osmundales" pageId="2" pageNumber="471" phylum="Tracheophyta" rank="species" species="lobulatus" status="sp. nov.">
<emphasis id="B933A339CF68FFC489E0F9D2FEA3F994" box="[164,318,1630,1649]" italics="true" pageId="2" pageNumber="471">Todites lobulatus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2001E42CF68FFC48807F9D2FE1DF994" box="[323,384,1630,1649]" pageId="2" pageNumber="471" rank="species">sp.nov.</taxonomicNameLabel>
Intinskian Formation, Ufimian, Upper Permian.Holotype GIN 4846/102.
<emphasis id="B933A339CF68FFC48D60F9D2FBA9F994" bold="true" box="[1060,1076,1630,1649]" pageId="2" pageNumber="471">A</emphasis>
,
<emphasis id="B933A339CF68FFC48D7BF9D2FBD3F994" bold="true" box="[1087,1102,1630,1649]" pageId="2" pageNumber="471">B</emphasis>
.General morphology of fertile pinna, bef − ore (
<emphasis id="B933A339CF68FFC489CAF9F6FF03F968" bold="true" box="[142,158,1658,1677]" pageId="2" pageNumber="471">A</emphasis>
) and after (
<emphasis id="B933A339CF68FFC48841F9F6FE89F968" bold="true" box="[261,276,1658,1677]" pageId="2" pageNumber="471">B</emphasis>
) preparation;
<emphasis id="B933A339CF68FFC488D5F9F6FE3CF968" bold="true" box="[401,417,1658,1677]" pageId="2" pageNumber="471">A</emphasis>
, × 1;
<emphasis id="B933A339CF68FFC48896F9F6FE7CF968" bold="true" box="[466,481,1658,1677]" pageId="2" pageNumber="471">B</emphasis>
, × 4.
<emphasis id="B933A339CF68FFC48B55F9F6FDBCF968" bold="true" box="[529,545,1658,1677]" pageId="2" pageNumber="471">C</emphasis>
.Two prepared sporangia; left sporangium shows a group of thick−walled cells near the stomium (dark de − pression); × 18.
<emphasis id="B933A339CF68FFC489BDF91AFE94F94C" bold="true" box="[249,265,1686,1705]" pageId="2" pageNumber="471">D</emphasis>
. The spore of
<taxonomicName id="4C4704A8CF68FFC488C9F91AFD82F94C" authorityName="Couper" authorityYear="1953" box="[397,543,1686,1705]" class="Polypodiopsida" genus="Osmundacidites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Polypodiales" pageId="2" pageNumber="471" phylum="Tracheophyta" rank="genus">
<emphasis id="B933A339CF68FFC488C9F91AFD82F94C" box="[397,543,1686,1705]" italics="true" pageId="2" pageNumber="471">Osmundacidites</emphasis>
</taxonomicName>
type, preserved
<emphasis id="B933A339CF68FFC48BF1F91BFD70F94C" box="[693,749,1687,1705]" italics="true" pageId="2" pageNumber="471">in situ</emphasis>
, under SEM, × 800.
</paragraph>
</caption>
<paragraph id="8BF87F2BCF68FFC489CFF8E5FA55F8DA" blockId="2.[102,775,1766,2019]" lastBlockId="2.[812,1485,1766,2018]" pageId="2" pageNumber="471">
The abaxial surface of the leaf lamina is covered by sporangia and one pinnule bears up to three hundred sporangia.Sporangia are so closely spaced, that they appear to form a mass, in which single sporangia are very difficult to observe.Individual sporangia are small (about
<quantity id="4CBFD2CECF68FFC48C74F96AFAE2F91B" box="[1328,1407,1766,1790]" metricMagnitude="-4" metricUnit="m" metricValue="5.0" pageId="2" pageNumber="471" unit="mm" value="0.5">0.5 mm</quantity>
in di − ameter), round or ovoid, and bear small stalks basally (sporangiophore), by which they are attached to the leaf lamina.
</paragraph>
<paragraph id="8BF87F2BCF68FFC48A16F8C5FAEAF807" blockId="2.[812,1485,1766,2018]" pageId="2" pageNumber="471">
One can see several well preserved sporangia, each showing a group of specialized cells with thicker walls located near the sporangial stomium (
<figureCitation id="137C63AECF68FFC48D3FF806FB4BF847" box="[1147,1238,1930,1954]" captionStart="Fig" captionStartId="2.[102,138,1630,1649]" captionTargetBox="[170,1417,226,1607]" captionTargetId="figure-155@2.[170,1417,226,1607]" captionTargetPageId="2" captionText="Fig.3. Todites lobulatus sp.nov.Intinskian Formation, Ufimian, Upper Permian.Holotype GIN 4846/102. A, B.General morphology of fertile pinna, bef − ore (A) and after (B) preparation; A, × 1; B, × 4. C.Two prepared sporangia; left sporangium shows a group of thick−walled cells near the stomium (dark de − pression); × 18. D. The spore of Osmundacidites type, preserved in situ, under SEM, × 800." figureDoi="http://doi.org/10.5281/zenodo.13173983" httpUri="https://zenodo.org/record/13173983/files/figure.png" pageId="2" pageNumber="471">Figs.3C</figureCitation>
,
<figureCitation id="137C63AECF68FFC48DA6F806FB6CF847" box="[1250,1265,1930,1954]" captionStart="A" captionStartId="4.[475,496,614,640]" captionText="A Fig.5. Todites lobulatus sp.nov. Intinskian Formation, Ufimian, Upper Permian.Holotype GIN 4846/102.Sporangium with open stomium and thick−walled cells located near its right side, photo− graph (A) and line drawing of the same sporangium portion (B). Scale bar 20 mm. B" pageId="2" pageNumber="471">5</figureCitation>
), which represents the opening mechanism of maturity.This structure is charac − teristic of recent osmundalean ferns, and fossil ones.
</paragraph>
<caption id="DF382FA3CF69FFC58922FA84FDDCFAB1" ID-DOI="http://doi.org/10.5281/zenodo.13173985" ID-Zenodo-Dep="13173985" httpUri="https://zenodo.org/record/13173985/files/figure.png" pageId="3" pageNumber="472" startId="3.[102,137,1288,1307]" targetBox="[102,1266,232,1364]" targetPageId="3" targetType="figure">
<paragraph id="8BF87F2BCF69FFC58922FA84FDDCFAB1" blockId="3.[102,608,1288,1364]" pageId="3" pageNumber="472">
Fig.4.
<taxonomicName id="4C4704A8CF69FFC589EDFA84FED9FAFE" authorityName="V.Naugolnykh" authorityYear="2002" box="[169,324,1288,1307]" class="Polypodiopsida" family="Osmundaceae" genus="Todites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Osmundales" pageId="3" pageNumber="472" phylum="Tracheophyta" rank="species" species="lobulatus" status="sp. nov.">
<emphasis id="B933A339CF69FFC589EDFA84FED9FAFE" box="[169,324,1288,1307]" italics="true" pageId="3" pageNumber="472">Todites lobulatus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2001E42CF69FFC58808FA84FE14FAFE" box="[332,393,1288,1307]" pageId="3" pageNumber="472" rank="species">sp.nov.</taxonomicNameLabel>
Intinskian Formation, Ufimian, Upper Permian.Holotype GIN 4846/102.Gen − eral morphology of a fertile pinna. Scale bar 0.5 cm.
</paragraph>
</caption>
<paragraph id="8BF87F2BCF69FFC589CFFA13FA8CFA32" blockId="3.[102,775,1439,2019]" lastBlockId="3.[812,1484,1439,1495]" pageId="3" pageNumber="472">
The uppermost fertile pinnule figured in
<figureCitation id="137C63AECF69FFC58B7BFA13FD10FA52" box="[575,653,1439,1463]" captionStart="Fig" captionStartId="2.[102,138,1630,1649]" captionTargetBox="[170,1417,226,1607]" captionTargetId="figure-155@2.[170,1417,226,1607]" captionTargetPageId="2" captionText="Fig.3. Todites lobulatus sp.nov.Intinskian Formation, Ufimian, Upper Permian.Holotype GIN 4846/102. A, B.General morphology of fertile pinna, bef − ore (A) and after (B) preparation; A, × 1; B, × 4. C.Two prepared sporangia; left sporangium shows a group of thick−walled cells near the stomium (dark de − pression); × 18. D. The spore of Osmundacidites type, preserved in situ, under SEM, × 800." figureDoi="http://doi.org/10.5281/zenodo.13173983" httpUri="https://zenodo.org/record/13173983/files/figure.png" pageId="3" pageNumber="472">Fig.3A</figureCitation>
and
<figureCitation id="137C63AECF69FFC58BF8FA13FD65FA52" box="[700,760,1439,1463]" captionStart="Fig" captionStartId="3.[102,137,1288,1307]" captionTargetBox="[102,1266,232,1364]" captionTargetId="figure-343@3.[316,1272,226,1361]" captionTargetPageId="3" captionText="Fig.4. Todites lobulatus sp.nov.Intinskian Formation, Ufimian, Upper Permian.Holotype GIN 4846/102.Gen − eral morphology of a fertile pinna. Scale bar 0.5 cm." figureDoi="http://doi.org/10.5281/zenodo.13173985" httpUri="https://zenodo.org/record/13173985/files/figure.png" pageId="3" pageNumber="472">Fig.4</figureCitation>
, right, was macerated.Numerous spores were extracted from the sporangia.During the maceration the sporangium walls were completely dissolved, but all spores were preserved
<emphasis id="B933A339CF69FFC58BB5F98EFF13F9DC" italics="true" pageId="3" pageNumber="472">in situ</emphasis>
as a round or spherical mass (
<figureCitation id="137C63AECF69FFC588A9F9ADFDF8F9DF" box="[493,613,1569,1594]" captionStart="Fig" captionStartId="5.[102,137,1856,1875]" captionTargetBox="[287,1295,227,1834]" captionTargetId="figure-9@5.[240,1348,227,1832]" captionTargetPageId="5" captionText="Fig.6. Todites lobulatus sp.nov.Intinskian Formation, Ufimian, Upper Permian.Macromorphology and in situ spores, based on holotype GIN 4846/102. A, B.Fertile pinnule reconstruction, upper (adaxial, A) and lower (abaxial, B) view, numerous sporangia attached to lower side of the pinnule surface (B). C.Individual sporangium with thick−walled cells on the top area. D.Granulate microstructure of the spore surface. E.General view on the isolated spore. F, G.Spore mass extracted from the sporangia, note different sizes (relatively small spores 1 and 2 and big spores 3 and 4 of F) and different outlines of the spores (round spores 2 and ovoid 1, 3 and 4 of F; round spore 1, rounded subtriangular spore 2 and ovoid spore 3 of G).Scale bars: A, B, 5 mm; C, 0.1 mm; E, F, G, 50 µm; D, 5 µm." figureDoi="http://doi.org/10.5281/zenodo.13173989" httpUri="https://zenodo.org/record/13173989/files/figure.png" pageId="3" pageNumber="472">Figs.6F, G</figureCitation>
,
<figureCitation id="137C63AECF69FFC58B37F9AEFD29F9DF" box="[627,692,1570,1594]" captionStart="Fig" captionStartId="6.[102,137,1751,1770]" captionTargetBox="[258,1327,226,1726]" captionTargetId="figure-70@6.[258,1330,226,1726]" captionTargetPageId="6" captionText="Fig.7. Todites lobulatus sp.nov.Intinskian Formation, Ufimian, Upper Permian. In situ spores extracted from the holotype GIN 4846/102. A.Spore mass repeating the initial round form of sporangium, × 50. B.Several spores of different size and form, × 200. C.Spore with clearly seen laesura (left part of the photograph), × 200. D. Spore mass of ovoid form from relatively elongated sporangium, × 50. E. Spore mass in detail, × 200. F. Detail of D, × 100." figureDoi="http://doi.org/10.5281/zenodo.13173991" httpUri="https://zenodo.org/record/13173991/files/figure.png" pageId="3" pageNumber="472">7A–F</figureCitation>
).This mass of spores has outlines corresponding to the original form of the sporangium.Separate spores are clearly seen at the periferal zone of the spore mass (
<figureCitation id="137C63AECF69FFC5888AF908FDA7F979" box="[462,570,1668,1692]" captionStart="Fig" captionStartId="6.[102,137,1751,1770]" captionTargetBox="[258,1327,226,1726]" captionTargetId="figure-70@6.[258,1330,226,1726]" captionTargetPageId="6" captionText="Fig.7. Todites lobulatus sp.nov.Intinskian Formation, Ufimian, Upper Permian. In situ spores extracted from the holotype GIN 4846/102. A.Spore mass repeating the initial round form of sporangium, × 50. B.Several spores of different size and form, × 200. C.Spore with clearly seen laesura (left part of the photograph), × 200. D. Spore mass of ovoid form from relatively elongated sporangium, × 50. E. Spore mass in detail, × 200. F. Detail of D, × 100." figureDoi="http://doi.org/10.5281/zenodo.13173991" httpUri="https://zenodo.org/record/13173991/files/figure.png" pageId="3" pageNumber="472">Fig.7B, C</figureCitation>
).The spores have a well developed tetrad scar (trilete mark or laesura), and the rays of the trilete mark almost reach the spore margin (
<figureCitation id="137C63AECF69FFC58B94F949FF34F91B" captionStart="Fig" captionStartId="5.[102,137,1856,1875]" captionTargetBox="[287,1295,227,1834]" captionTargetId="figure-9@5.[240,1348,227,1832]" captionTargetPageId="5" captionText="Fig.6. Todites lobulatus sp.nov.Intinskian Formation, Ufimian, Upper Permian.Macromorphology and in situ spores, based on holotype GIN 4846/102. A, B.Fertile pinnule reconstruction, upper (adaxial, A) and lower (abaxial, B) view, numerous sporangia attached to lower side of the pinnule surface (B). C.Individual sporangium with thick−walled cells on the top area. D.Granulate microstructure of the spore surface. E.General view on the isolated spore. F, G.Spore mass extracted from the sporangia, note different sizes (relatively small spores 1 and 2 and big spores 3 and 4 of F) and different outlines of the spores (round spores 2 and ovoid 1, 3 and 4 of F; round spore 1, rounded subtriangular spore 2 and ovoid spore 3 of G).Scale bars: A, B, 5 mm; C, 0.1 mm; E, F, G, 50 µm; D, 5 µm." figureDoi="http://doi.org/10.5281/zenodo.13173989" httpUri="https://zenodo.org/record/13173989/files/figure.png" pageId="3" pageNumber="472">Figs. 6E–G</figureCitation>
,
<figureCitation id="137C63AECF69FFC589F6F96AFF49F91B" box="[178,212,1766,1790]" captionStart="Fig" captionStartId="6.[102,137,1751,1770]" captionTargetBox="[258,1327,226,1726]" captionTargetId="figure-70@6.[258,1330,226,1726]" captionTargetPageId="6" captionText="Fig.7. Todites lobulatus sp.nov.Intinskian Formation, Ufimian, Upper Permian. In situ spores extracted from the holotype GIN 4846/102. A.Spore mass repeating the initial round form of sporangium, × 50. B.Several spores of different size and form, × 200. C.Spore with clearly seen laesura (left part of the photograph), × 200. D. Spore mass of ovoid form from relatively elongated sporangium, × 50. E. Spore mass in detail, × 200. F. Detail of D, × 100." figureDoi="http://doi.org/10.5281/zenodo.13173991" httpUri="https://zenodo.org/record/13173991/files/figure.png" pageId="3" pageNumber="472">7D</figureCitation>
).The spore surface is sculptured by very fine granu − late microstructure (
<figureCitation id="137C63AECF69FFC5887BF88AFE04F8FB" box="[319,409,1798,1822]" captionStart="Fig" captionStartId="5.[102,137,1856,1875]" captionTargetBox="[287,1295,227,1834]" captionTargetId="figure-9@5.[240,1348,227,1832]" captionTargetPageId="5" captionText="Fig.6. Todites lobulatus sp.nov.Intinskian Formation, Ufimian, Upper Permian.Macromorphology and in situ spores, based on holotype GIN 4846/102. A, B.Fertile pinnule reconstruction, upper (adaxial, A) and lower (abaxial, B) view, numerous sporangia attached to lower side of the pinnule surface (B). C.Individual sporangium with thick−walled cells on the top area. D.Granulate microstructure of the spore surface. E.General view on the isolated spore. F, G.Spore mass extracted from the sporangia, note different sizes (relatively small spores 1 and 2 and big spores 3 and 4 of F) and different outlines of the spores (round spores 2 and ovoid 1, 3 and 4 of F; round spore 1, rounded subtriangular spore 2 and ovoid spore 3 of G).Scale bars: A, B, 5 mm; C, 0.1 mm; E, F, G, 50 µm; D, 5 µm." figureDoi="http://doi.org/10.5281/zenodo.13173989" httpUri="https://zenodo.org/record/13173989/files/figure.png" pageId="3" pageNumber="472">Figs.6D</figureCitation>
,
<figureCitation id="137C63AECF69FFC588E7F88BFE7EF8FB" box="[419,483,1798,1822]" captionStart="Fig" captionStartId="6.[102,137,1751,1770]" captionTargetBox="[258,1327,226,1726]" captionTargetId="figure-70@6.[258,1330,226,1726]" captionTargetPageId="6" captionText="Fig.7. Todites lobulatus sp.nov.Intinskian Formation, Ufimian, Upper Permian. In situ spores extracted from the holotype GIN 4846/102. A.Spore mass repeating the initial round form of sporangium, × 50. B.Several spores of different size and form, × 200. C.Spore with clearly seen laesura (left part of the photograph), × 200. D. Spore mass of ovoid form from relatively elongated sporangium, × 50. E. Spore mass in detail, × 200. F. Detail of D, × 100." figureDoi="http://doi.org/10.5281/zenodo.13173991" httpUri="https://zenodo.org/record/13173991/files/figure.png" pageId="3" pageNumber="472">7B, D</figureCitation>
).Very similar, but slightly more developed and larger, spores were extracted from the matrix housing the
<taxonomicName id="4C4704A8CF69FFC58872F8C4FE6DF885" authorityName="V.Naugolnykh" authorityYear="2002" box="[310,496,1864,1888]" class="Polypodiopsida" family="Osmundaceae" genus="Todites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Osmundales" pageId="3" pageNumber="472" phylum="Tracheophyta" rank="species" species="lobulatus">
<emphasis id="B933A339CF69FFC58872F8C4FE6DF885" box="[310,496,1864,1888]" italics="true" pageId="3" pageNumber="472">Todites lobulatus</emphasis>
</taxonomicName>
fertile pinnae (
<figureCitation id="137C63AECF69FFC58BD7F8C4FD44F885" box="[659,729,1864,1888]" captionStart="Fig" captionStartId="7.[102,135,1915,1934]" captionTargetBox="[209,1377,227,1872]" captionTargetId="figure-8@7.[209,1377,227,1872]" captionTargetPageId="7" captionText="Fig.8.Spores of Osmundacidites type, extracted from the matrix containing the holotype of Todites lobulatus sp.nov.,GIN 4846/102. A, B, D, E.Different individual spores; A, B, D, × 1500; E, × 750. C, F. Granulate sculpture of spore surface, × 3000." figureDoi="http://doi.org/10.5281/zenodo.13173995" httpUri="https://zenodo.org/record/13173995/files/figure.png" pageId="3" pageNumber="472">Figs.8</figureCitation>
,
<figureCitation id="137C63AECF69FFC58BA7F8C4FD6EF885" box="[739,755,1864,1888]" captionStart="Fig" captionStartId="8.[102,134,1771,1790]" captionTargetBox="[238,1347,226,1747]" captionTargetId="figure-94@8.[238,1347,226,1747]" captionTargetPageId="8" captionText="Fig.9.Spores of Osmundacidites type, extracted from the matrix containing the holotype of Todites lobulatus sp.nov.,GIN 4846/102. A–C.Different indi − vidual spores, × 1500. D–F. Granulate sculpture of spore surface, × 3000." figureDoi="http://doi.org/10.5281/zenodo.13173997" httpUri="https://zenodo.org/record/13173997/files/figure.png" pageId="3" pageNumber="472">9</figureCitation>
). Most probably they belonged to the same species, but dispersed spores have a larger size because they are mature as opposed to those still in the sporangia.The dispersed spores of the same
<typeStatus id="54FCC189CF69FFC58993F847FE94F806" box="[215,265,1995,2019]" pageId="3" pageNumber="472">type</typeStatus>
, which can be attributed to
<taxonomicName id="4C4704A8CF69FFC58B12F846FC5EFA52" authority="Couper, 1953" authorityName="Couper" authorityYear="1953" box="[598,963,1439,2018]" class="Polypodiopsida" genus="Osmundacidites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Polypodiales" pageId="3" pageNumber="472" phylum="Tracheophyta" rank="genus">
<emphasis id="B933A339CF69FFC58B12F846FC9AF807" box="[598,775,1994,2018]" italics="true" pageId="3" pageNumber="472">Osmundacidites</emphasis>
<bibRefCitation id="EFD602DACF69FFC58A68FA13FC5EFA52" author="Couper, R. A." box="[812,963,1439,1463]" pageId="3" pageNumber="472" pagination="1 - 77" refId="ref3453" refString="Couper, R. A. 1953. Upper Mesozoic and Cenozoic spores and pollen grains from New Zealand. New Zealand Geological Survey Palaeontological Bulletin 22: 1 - 77." type="journal article" year="1953">Couper, 1953</bibRefCitation>
</taxonomicName>
, are known from the Upper Permian of other regions of the Urals and the Russian platform.
</paragraph>
</subSubSection>
<subSubSection id="C35D2CA0CF69FFC28A68FA67FD5DF916" lastPageId="4" lastPageNumber="473" pageId="3" pageNumber="472" type="discussion">
<paragraph id="8BF87F2BCF69FFC58A68FA67FCEAF807" blockId="3.[812,1485,1514,2018]" pageId="3" pageNumber="472">
<emphasis id="B933A339CF69FFC58A68FA67FBCEF9E7" box="[812,1107,1514,1538]" italics="true" pageId="3" pageNumber="472">Comparison and remarks</emphasis>
.—The new species differs from the type−species
<taxonomicName id="4C4704A8CF69FFC58ABCF986FC58F9A7" authority="(Brongniart, 1828) Seward, 1900" authorityName="Seward" authorityYear="1900" baseAuthorityName="Brongniart" baseAuthorityYear="1828" class="Polypodiopsida" family="Osmundaceae" genus="Todites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Osmundales" pageId="3" pageNumber="472" phylum="Tracheophyta" rank="species" species="williamsonii">
<emphasis id="B933A339CF69FFC58ABCF986FB7FF9C7" box="[1016,1250,1546,1570]" italics="true" pageId="3" pageNumber="472">Todites williamsonii</emphasis>
(
<bibRefCitation id="EFD602DACF69FFC58DBEF986FA58F9C7" author="Brongniart, A." box="[1274,1477,1546,1570]" pageId="3" pageNumber="472" refId="ref3427" refString="Brongniart, A. 1828. Prodrome d'une Histoire des vegetaux fossiles. 223 pp. G. Dufour et Ed. d'Ocagne, Paris." type="book" year="1828">Brongniart, 1828</bibRefCitation>
) Seward,
<quantity id="4CBFD2CECF69FFC58AC9F9A6FC58F9A7" box="[909,965,1578,1602]" metricMagnitude="1" metricUnit="m" metricValue="4.826" pageId="3" pageNumber="472" unit="in" value="1900.0">1900</quantity>
</taxonomicName>
in having deeply dissected pinnules and round pinnule apices; from
<taxonomicName id="4C4704A8CF69FFC58D5FF9C7FADDF987" authority="Barnard, 1965" authorityName="Barnard" authorityYear="1965" box="[1051,1344,1610,1634]" class="Polypodiopsida" family="Osmundaceae" genus="Todites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Osmundales" pageId="3" pageNumber="472" phylum="Tracheophyta" rank="species" species="crenatus">
<emphasis id="B933A339CF69FFC58D5FF9C7FB0AF987" box="[1051,1175,1611,1634]" italics="true" pageId="3" pageNumber="472">T. crenatus</emphasis>
<bibRefCitation id="EFD602DACF69FFC58DDBF9C6FADDF987" author="Barnard, P. D. W." box="[1183,1344,1610,1634]" pageId="3" pageNumber="472" pagination="1123 - 1168" refId="ref3388" refString="Barnard, P. D. W. 1965. Flora of the Shemshak Formation. Part 1. Liassic plants from Dorud. Revista Italiana di Paleontologia e Stratigrafia 71 (4): 1123 - 1168." type="journal article" year="1965">
Barnard,
<quantity id="4CBFD2CECF69FFC58C4CF9C6FADDF987" box="[1288,1344,1610,1634]" metricMagnitude="1" metricUnit="m" metricValue="4.9911" pageId="3" pageNumber="472" unit="in" value="1965.0">1965</quantity>
</bibRefCitation>
</taxonomicName>
in considerably shorter and wider pinnules; from
<taxonomicName id="4C4704A8CF69FFC58DA5F9E7FB14F947" authority="(Schmalhausen, 1879) Radczenko, 1955" authorityName="Radczenko" authorityYear="1955" baseAuthorityName="Schmalhausen" baseAuthorityYear="1879" class="Polypodiopsida" family="Osmundaceae" genus="Todites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Osmundales" pageId="3" pageNumber="472" phylum="Tracheophyta" rank="species" species="sibiricus">
<emphasis id="B933A339CF69FFC58DA5F9E7FAC3F967" box="[1249,1374,1642,1666]" italics="true" pageId="3" pageNumber="472">T. sibiricus</emphasis>
(Schmalhausen, 1879) Radczenko, 1955
</taxonomicName>
and
<taxonomicName id="4C4704A8CF69FFC58DFEF907FCF9F927" authority="Radczenko, 1955" authorityName="Radczenko" authorityYear="1955" class="Polypodiopsida" family="Osmundaceae" genus="Todites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Osmundales" pageId="3" pageNumber="472" phylum="Tracheophyta" rank="species" species="evenkensis">
<emphasis id="B933A339CF69FFC58DFEF907FAD5F947" box="[1210,1352,1674,1698]" italics="true" pageId="3" pageNumber="472">T. evenkensis</emphasis>
Radczenko,
<quantity id="4CBFD2CECF69FFC58A68F926FCF9F927" box="[812,868,1706,1730]" metricMagnitude="1" metricUnit="m" metricValue="4.9657" pageId="3" pageNumber="472" unit="in" value="1955.0">1955</quantity>
</taxonomicName>
in having well developed pinnule lobes, but shares with them the general structure of the fertile pinnules, e.g.,
<taxonomicName id="4C4704A8CF69FFC58CD3F946FCC9F8E7" form="and" pageId="3" pageNumber="472" rank="form">form and</taxonomicName>
position of sporangia; from
<taxonomicName id="4C4704A8CF69FFC58DC9F967FA9AF8E7" authorityName="Radczenko" authorityYear="1955" baseAuthorityName="Schmalhausen" baseAuthorityYear="1879" box="[1165,1287,1770,1794]" class="Polypodiopsida" family="Osmundaceae" genus="Todites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Osmundales" pageId="3" pageNumber="472" phylum="Tracheophyta" rank="species" species="sibiricus">
<emphasis id="B933A339CF69FFC58DC9F967FA9AF8E7" box="[1165,1287,1770,1794]" italics="true" pageId="3" pageNumber="472">T. sibiricus</emphasis>
</taxonomicName>
—in larger size of the pinnules (
<quantity id="4CBFD2CECF69FFC58A83F886FB8AF8C7" box="[967,1047,1802,1826]" metricMagnitude="-2" metricUnit="m" metricValue="1.5" pageId="3" pageNumber="472" unit="mm" value="15.0">15 mm</quantity>
instead
<quantity id="4CBFD2CECF69FFC58D33F886FB79F8C7" box="[1143,1252,1802,1826]" metricMagnitude="-3" metricUnit="m" metricValue="9.0" metricValueMax="10.0" metricValueMin="8.0" pageId="3" pageNumber="472" unit="mm" value="9.0" valueMax="10.0" valueMin="8.0">8–10 mm</quantity>
) and slightly curved pinnule (straight pinnules of
<taxonomicName id="4C4704A8CF69FFC58D2AF8A7FB77F8A7" authorityName="Radczenko" authorityYear="1955" baseAuthorityName="Schmalhausen" baseAuthorityYear="1879" box="[1134,1258,1834,1858]" class="Polypodiopsida" family="Osmundaceae" genus="Todites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Osmundales" pageId="3" pageNumber="472" phylum="Tracheophyta" rank="species" species="sibiricus">
<emphasis id="B933A339CF69FFC58D2AF8A7FB77F8A7" box="[1134,1258,1834,1858]" italics="true" pageId="3" pageNumber="472">T. sibiricus</emphasis>
</taxonomicName>
are disposed almost perpendicularly to the pinna rachis), from
<taxonomicName id="4C4704A8CF69FFC58C4CF8C7FA01F887" authorityName="Radczenko" authorityYear="1955" box="[1288,1436,1866,1890]" class="Polypodiopsida" family="Osmundaceae" genus="Todites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Osmundales" pageId="3" pageNumber="472" phylum="Tracheophyta" rank="species" species="evenkensis">
<emphasis id="B933A339CF69FFC58C4CF8C7FA01F887" box="[1288,1436,1866,1890]" italics="true" pageId="3" pageNumber="472">T. evenkensis</emphasis>
</taxonomicName>
—in round pinnule apex and wider angle of pinnule orientation on the last order pinna;
<taxonomicName id="4C4704A8CF69FFC58D5CF806FB45F847" authorityName="V.Naugolnykh" authorityYear="2002" box="[1048,1240,1930,1954]" class="Polypodiopsida" family="Osmundaceae" genus="Todites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Osmundales" pageId="3" pageNumber="472" phylum="Tracheophyta" rank="species" species="lobulatus">
<emphasis id="B933A339CF69FFC58D5CF806FB45F847" box="[1048,1240,1930,1954]" italics="true" pageId="3" pageNumber="472">Todites lobulatus</emphasis>
</taxonomicName>
has narrow wings on the rachis.In contrast,
<taxonomicName id="4C4704A8CF69FFC58D6EF827FB38F827" authorityName="Radczenko" authorityYear="1955" baseAuthorityName="Schmalhausen" baseAuthorityYear="1879" box="[1066,1189,1962,1986]" class="Polypodiopsida" family="Osmundaceae" genus="Todites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Osmundales" pageId="3" pageNumber="472" phylum="Tracheophyta" rank="species" species="sibiricus">
<emphasis id="B933A339CF69FFC58D6EF827FB38F827" box="[1066,1189,1962,1986]" italics="true" pageId="3" pageNumber="472">T. sibiricus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C4704A8CF69FFC58D98F827FAF2F827" authorityName="Radczenko" authorityYear="1955" box="[1244,1391,1962,1986]" class="Polypodiopsida" family="Osmundaceae" genus="Todites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Osmundales" pageId="3" pageNumber="472" phylum="Tracheophyta" rank="species" species="evenkensis">
<emphasis id="B933A339CF69FFC58D98F827FAF2F827" box="[1244,1391,1962,1986]" italics="true" pageId="3" pageNumber="472">T. evenkensis</emphasis>
</taxonomicName>
have no wings.
</paragraph>
<caption id="DF382FA3CF6EFFC28922FCCFFE98FBD3" ID-DOI="http://doi.org/10.5281/zenodo.13173987" ID-Zenodo-Dep="13173987" httpUri="https://zenodo.org/record/13173987/files/figure.png" pageId="4" pageNumber="473" startId="4.[102,138,835,854]" targetBox="[464,1485,226,1073]" targetPageId="4" targetType="figure">
<paragraph id="8BF87F2BCF6EFFC28922FCCFFE98FBD3" blockId="4.[102,404,835,1078]" pageId="4" pageNumber="473">
Fig.5.
<taxonomicName id="4C4704A8CF6EFFC289EDFCCFFED8FCB3" authorityName="V.Naugolnykh" authorityYear="2002" box="[169,325,835,854]" class="Polypodiopsida" family="Osmundaceae" genus="Todites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Osmundales" pageId="4" pageNumber="473" phylum="Tracheophyta" rank="species" species="lobulatus" status="sp. nov.">
<emphasis id="B933A339CF6EFFC289EDFCCFFED8FCB3" box="[169,325,835,854]" italics="true" pageId="4" pageNumber="473">Todites lobulatus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2001E42CF6EFFC28808FCCFFE10FCB3" box="[332,397,835,854]" pageId="4" pageNumber="473" rank="species">sp.nov.</taxonomicNameLabel>
Intinskian Formation, Ufimian, Upper Permian.Holotype GIN 4846/102.Sporangium with open stomium and thick−walled cells located near its right side, photograph (
<emphasis id="B933A339CF6EFFC289E2FC67FF2BFC1B" bold="true" box="[166,182,1003,1022]" pageId="4" pageNumber="473">A</emphasis>
) and line drawing of the same sporangium portion (
<emphasis id="B933A339CF6EFFC28833FB8BFE1BFBFF" bold="true" box="[375,390,1031,1050]" pageId="4" pageNumber="473">B</emphasis>
). Scale bar 20 mm.
</paragraph>
</caption>
<paragraph id="8BF87F2BCF6EFFC289CFFBF5FDFEFA54" blockId="4.[102,775,1145,1779]" pageId="4" pageNumber="473">
<bibRefCitation id="EFD602DACF6EFFC289CFFBF5FEABFB74" author="Fefilova, L. A." box="[139,310,1145,1169]" pageId="4" pageNumber="473" refId="ref3660" refString="Fefilova, L. A. 1973. Paporotnikovidnye permi severa Preduralskogo progiba. 192 pp. Nauka, Leningrad." type="book" year="1973">Fefilova (1973)</bibRefCitation>
described several sterile and fertile pinnae from the Upper Permian (Ufimian and Kazanian) deposits of Kosju, Big Synja and Perebor river basins (Pechora Cis−Urals).These pinnae were attributed to
<taxonomicName id="4C4704A8CF6EFFC28B56FB55FD1EFB14" box="[530,643,1241,1265]" class="Polypodiopsida" family="Osmundaceae" genus="Todites" kingdom="Plantae" order="Osmundales" pageId="4" pageNumber="473" phylum="Tracheophyta" rank="species" species="undetermined">
<emphasis id="B933A339CF6EFFC28B56FB55FDFDFB14" box="[530,608,1241,1265]" italics="true" pageId="4" pageNumber="473">Todites</emphasis>
sp.
</taxonomicName>
in open nomenclature.Some spores of
<taxonomicName id="4C4704A8CF6EFFC288D2FB75FF00FAD4" authority="Potonie and Kremp, 1954" authorityName="Potonie and Kremp" authorityYear="1954" genus="Lophotriletes" higherTaxonomySource="GBIF" kingdom="Plantae" pageId="4" pageNumber="473" rank="genus">
<emphasis id="B933A339CF6EFFC288D2FB75FDB5FAF4" box="[406,552,1273,1297]" italics="true" pageId="4" pageNumber="473">Lophotriletes</emphasis>
Potonie and Kremp, 1954
</taxonomicName>
were extracted from the sporangia.Later very similar fertile fronds of
<taxonomicName id="4C4704A8CF6EFFC28857FAB5FE1FFAB4" box="[275,386,1337,1361]" class="Polypodiopsida" family="Osmundaceae" genus="Todites" kingdom="Plantae" order="Osmundales" pageId="4" pageNumber="473" phylum="Tracheophyta" rank="species" species="undetermined">
<emphasis id="B933A339CF6EFFC28857FAB5FEFCFAB4" box="[275,353,1337,1361]" italics="true" pageId="4" pageNumber="473">Todites</emphasis>
sp.
</taxonomicName>
were reported from the Seidinskian and Talbeiskian Formations (presumably Kazanian in age, at least partly) of the Pechora coal district, in the Heyaha and Adzva rivers basin (Pukhonto, Fefilova 1983).
</paragraph>
<paragraph id="8BF87F2BCF6EFFC289CFFA36FF60F997" blockId="4.[102,775,1145,1779]" pageId="4" pageNumber="473">
Mogutcheva (1984) established the species
<taxonomicName id="4C4704A8CF6EFFC28B33FA36FF55FA17" class="Polypodiopsida" family="Osmundaceae" genus="Todites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Osmundales" pageId="4" pageNumber="473" phylum="Tracheophyta" rank="species" species="orulganensis">
<emphasis id="B933A339CF6EFFC28B33FA36FF55FA17" italics="true" pageId="4" pageNumber="473">Todites orulganensis</emphasis>
</taxonomicName>
from the Lower Triassic Surbeljakhskian Formation of North Verkhoyanie (Siberia).This species is similar to
<taxonomicName id="4C4704A8CF6EFFC28922F997FF74F9D7" authorityName="V.Naugolnykh" authorityYear="2002" box="[102,233,1562,1586]" class="Polypodiopsida" family="Osmundaceae" genus="Todites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Osmundales" pageId="4" pageNumber="473" phylum="Tracheophyta" rank="species" species="lobulatus">
<emphasis id="B933A339CF6EFFC28922F997FF74F9D7" box="[102,233,1562,1586]" italics="true" pageId="4" pageNumber="473">T. lobulatus</emphasis>
</taxonomicName>
described above, but differs by entire margined pinnules.There is no information on the
<emphasis id="B933A339CF6EFFC28B71F9B7FDE6F9B7" box="[565,635,1595,1618]" italics="true" pageId="4" pageNumber="473">in situ</emphasis>
spores of
<taxonomicName id="4C4704A8CF6EFFC28BB6F9B7FF6AF997" class="Polypodiopsida" family="Osmundaceae" genus="Todites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Osmundales" pageId="4" pageNumber="473" phylum="Tracheophyta" rank="species" species="orulganensis">
<emphasis id="B933A339CF6EFFC28BB6F9B7FF6AF997" italics="true" pageId="4" pageNumber="473">T. orulganensis</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BF87F2BCF6EFFC289C8F9F0FD5DF916" blockId="4.[102,775,1145,1779]" pageId="4" pageNumber="473">
In general, the new species
<taxonomicName id="4C4704A8CF6EFFC28887F9F0FDDAF976" authorityName="V.Naugolnykh" authorityYear="2002" box="[451,583,1659,1683]" class="Polypodiopsida" family="Osmundaceae" genus="Todites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Osmundales" pageId="4" pageNumber="473" phylum="Tracheophyta" rank="species" species="lobulatus">
<emphasis id="B933A339CF6EFFC28887F9F0FDDAF976" box="[451,583,1659,1683]" italics="true" pageId="4" pageNumber="473">T. lobulatus</emphasis>
</taxonomicName>
differs considerably from other species of
<taxonomicName id="4C4704A8CF6EFFC288E6F917FE6FF956" authorityName="Seward" authorityYear="1900" box="[418,498,1691,1715]" class="Polypodiopsida" family="Osmundaceae" genus="Todites" higherTaxonomySource="GBIF" kingdom="Plantae" order="Osmundales" pageId="4" pageNumber="473" phylum="Tracheophyta" rank="genus">
<emphasis id="B933A339CF6EFFC288E6F917FE6FF956" box="[418,498,1691,1715]" italics="true" pageId="4" pageNumber="473">Todites</emphasis>
</taxonomicName>
known from the Upper Permian and Lower Triassic deposits of Angaraland by wider pinnules and extremely well developed pinnule lobes.
</paragraph>
</subSubSection>
</treatment>
</document>
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