530 lines
122 KiB
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530 lines
122 KiB
XML
<document id="96150B7A3DE033BA4B899C644ABE4091" ID-CLB-Dataset="53898" ID-DOI="10.1038/s41586-023-06359-z" ID-GBIF-Dataset="ed3047b8-f9a5-4287-96aa-06b104502866" ID-Zenodo-Dep="8263099" IM.bibliography_approvedBy="felipe" IM.illustrations_approvedBy="felipe" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="felipe" checkinTime="1692364486276" checkinUser="felipe" docAuthor="Müller, Rodrigo T., Ezcurra, Martín D., Garcia, Mauricio S., Agnolín, Federico L., Stocker, Michelle R., Novas, Fernando E., Soares, Marina B., Kellner, Alexander W. A. & Nesbitt, Sterling J." docDate="2023" docId="03C1A9535C12FF9DBAB0F9DFECBCFAD6" docLanguage="en" docName="Nature.620.589-594.pdf.imf" docOrigin="Nature 620 (7974)" docSource="http://dx.doi.org/10.1038/s41586-023-06359-z" docTitle="Venetoraptor gassenae Müller et al. 2023" docType="treatment" docVersion="3" lastPageNumber="593" masterDocId="FFF8D12B5C12FF99B979FFC0E858FFC5" masterDocTitle="New reptile shows dinosaurs and pterosaurs evolved among diverse precursors" masterLastPageNumber="594" masterPageNumber="589" pageNumber="589" updateTime="1692365273934" updateUser="ExternalLinkService">
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<mods:title id="5600B169E3349CD67D6479CD7472770D">New reptile shows dinosaurs and pterosaurs evolved among diverse precursors</mods:title>
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<mods:namePart id="44FEF80A86145EB3E3FA1330B6D72C67">Müller, Rodrigo T.</mods:namePart>
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<mods:namePart id="3C14D4AF74DCCF69B1B448187880D8BA">Ezcurra, Martín D.</mods:namePart>
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<mods:namePart id="C379DA7F88EDE76AC343D70AE04511EA">Garcia, Mauricio S.</mods:namePart>
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<mods:namePart id="EE6490CA1128EF10A50CB839C2238051">Agnolín, Federico L.</mods:namePart>
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<mods:namePart id="6473B12DD2684E898937DBCCED5C7BCF">Stocker, Michelle R.</mods:namePart>
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<mods:namePart id="68171EB4CCFD3ECD0AB3311E8F8E99E0">Novas, Fernando E.</mods:namePart>
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<mods:namePart id="5A0115FC4721DB923D4577042CCFACA8">Soares, Marina B.</mods:namePart>
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<treatment id="03C1A9535C12FF9DBAB0F9DFECBCFAD6" ID-DOI="http://doi.org/10.5281/zenodo.8263145" ID-Zenodo-Dep="8263145" LSID="urn:lsid:plazi:treatment:03C1A9535C12FF9DBAB0F9DFECBCFAD6" httpUri="http://treatment.plazi.org/id/03C1A9535C12FF9DBAB0F9DFECBCFAD6" lastPageId="4" lastPageNumber="593" pageId="0" pageNumber="589">
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<subSubSection id="C3724BCE5C12FF99BAB0F9DFED19F9F7" box="[969,1345,1566,1587]" pageId="0" pageNumber="589" type="nomenclature">
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<paragraph id="8BD718455C12FF99BAB0F9DFED19F9F7" blockId="0.[969,1345,1480,1587]" box="[969,1345,1566,1587]" pageId="0" pageNumber="589">
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<heading id="D09FAF295C12FF99BAB0F9DFED19F9F7" box="[969,1345,1566,1587]" level="2" pageId="0" pageNumber="589">
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<taxonomicName id="4C6863C65C12FF99BAB0F9DFED19F9F7" authority="Müller et al., 2023" authorityName="Müller et al." authorityYear="2023" box="[969,1345,1566,1587]" clade="Pterosauromorpha" family="Lagerpetidae" genus="Venetoraptor" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="0" pageNumber="589" phylum="Chordata" rank="species" species="gassenae" status="gen. et sp. nov.">Venetoraptor gassenae gen.et sp. nov.</taxonomicName>
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<subSubSection id="C3724BCE5C12FF99BA49F998EB98F8D2" pageId="0" pageNumber="589" type="etymology">
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Etymology. The genus name combines the word raptor,plunderer (Latin) in reference to its raptorial beak and grasping hands and the word Veneto in reference to ‘Vale Vêneto’, a touristic locality within the municipality of São João do Polêsine,
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<collectingRegion id="49ACD6A75C12FF99BC65F96DED8FF904" box="[1308,1495,1709,1730]" country="Brazil" name="Rio Grande do Sul" pageId="0" pageNumber="589">Rio Grande do Sul</collectingRegion>
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,
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<collectingCountry id="F37F58D55C12FF99BA49F90AEB28F91B" box="[816,880,1738,1758]" name="Brazil" pageId="0" pageNumber="589">Brazil</collectingCountry>
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. The specific epithet honours Mrs
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<taxonomicName id="4C6863C65C12FF99BDA3F90AEB26F93E" authority="MARIA BULEGON GASSEN" authorityName="MARIA BULEGON GASSEN" class="Globothalamea" family="Orbitolinidae" genus="Valserina" kingdom="Chromista" order="Loftusiida" pageId="0" pageNumber="589" phylum="Foraminifera" rank="genus">Valserina Maria Bulegon Gassen</taxonomicName>
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, one of the main people responsible for the foundation of
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/
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.
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<footnote id="E873044B5C12FF99B913F8F9EB66F807" pageId="0" pageNumber="589">
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Centro de Apoio à Pesquisa Paleontológica da Quarta Colônia, Universidade Federal de Santa Maria,São João do Polêsine,Brazil.
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<bibRefCitation id="EFF965B45C12FF99BD61F8F9EC45F887" author="Prentice, K. C. & Ruta, M. & Benton, M. J." box="[1048,1053,1849,1858]" journalOrPublisher="J. Syst. Palaeontol." pageId="0" pageNumber="589" pagination="337 - 353" part="9" refId="ref4319" refString="2. Prentice, K. C., Ruta, M. & Benton, M. J. Evolution of morphological disparity in pterosaurs. J. Syst. Palaeontol. 9, 337 - 353 (2011)." title="Evolution of morphological disparity in pterosaurs" type="journal article" year="2011">
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Programa de Pós-Graduação em Biodiversidade Animal, Universidade Federal de Santa Maria, Santa Maria,Brazil.
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<bibRefCitation id="EFF965B45C12FF99BB7EF891EA55F89F" author="Brusatte, S. L. & Butler, R. J. & Prieto-Marquez, A. & Norell, M. A." box="[519,525,1873,1882]" pageId="0" pageNumber="589" pagination="804" refId="ref4361" refString="3. Brusatte, S. L., Butler, R. J., Prieto-Marquez, A. & Norell, M. A. Dinosaur morphological diversity and the end-Cretaceous extinction. Nat. Commun. 3, 804 (2012)." type="journal article" year="2012">
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Sección Paleontología de Vertebrados CONICET−Museo Argentino de Ciencias Naturales ̋Bernardino Rivadavia̋,Buenos Aires, Argentina.
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<bibRefCitation id="EFF965B45C12FF99B9C0F8A9E8E7F8B7" author="Langer, M. C. & Ezcurra, M. D. & Bittencourt, J. S. & Novas, F. E." box="[185,191,1897,1906]" pageId="0" pageNumber="589" pagination="55 - 110" refId="ref4407" refString="4. Langer, M. C., Ezcurra, M. D., Bittencourt, J. S. & Novas, F. E. The origin and early evolution of dinosaurs. Biol. Rev. 85, 55 - 110 (2010)." type="journal article" year="2010">
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Laboratorio de Anatomía Comparada y Evolución de los Vertebrados CONICET−Museo Argentino de Ciencias Naturales ̋Bernardino Rivadavia̋,Buenos Aires,Argentina.
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<bibRefCitation id="EFF965B45C12FF99B910F841E837F84F" author="Andres, B." box="[105,111,1921,1930]" pageId="0" pageNumber="589" pagination="1356 - 1365" refId="ref4457" refString="5. Andres, B. The early evolutionary history and adaptive radiation of the Pterosauria. Acta Geol. Sin. 86, 1356 - 1365 (2012)." type="journal article" year="2012">
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Fundación de Historia Natural ‘Félix de Azara̍,Departamento de Ciencias Naturales y Antropología, Universidad Maimónides,Buenos Aires, Argentina.
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<bibRefCitation id="EFF965B45C12FF99BDCAF841ECE1F84F" author="Nesbitt, S. J." box="[1203,1209,1921,1930]" pageId="0" pageNumber="589" pagination="95 - 98" refId="ref4488" refString="6. Nesbitt, S. J. et al. Ecologically distinct dinosaurian sister group shows early diversification of Ornithodira. Nature 464, 95 - 98 (2010)." type="journal article" year="2010">
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Department of Geosciences,Virginia Tech, Blacksburg,VA,USA.
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<bibRefCitation id="EFF965B45C12FF99B848F859E96EF867" author="Cabreira, S. F." box="[305,310,1945,1954]" pageId="0" pageNumber="589" pagination="3090 - 3095" refId="ref4520" refString="7. Cabreira, S. F. et al. A unique Late Triassic dinosauromorph assemblage reveals dinosaur ancestral anatomy and diet. Curr. Biol. 26, 3090 - 3095 (2016)." type="journal article" year="2016">
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<superScript id="7C1DB50D5C12FF99B848F859E96EF867" attach="right" box="[305,310,1945,1954]" fontSize="4" pageId="0" pageNumber="589">7</superScript>
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Laboratório de Sistemática e Tafonomia de Vertebrados Fósseis,Setor de Paleovertebrados,Departamento de Geologia e Paleontologia,Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro,Brazil.✉e-mail: rodrigotmuller@hotmail.com
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</paragraph>
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</footnote>
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<caption id="DF1748CD5C13FF98B913FB55ED5FFA74" ID-DOI="http://doi.org/10.5281/zenodo.8263101" ID-Zenodo-Dep="8263101" httpUri="https://zenodo.org/record/8263101/files/figure.png" pageId="1" pageNumber="590" startId="1.[106,137,1173,1190]" targetBox="[90,1551,118,1158]" targetPageId="1" targetType="figure">
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<paragraph id="8BD718455C13FF98B913FB55EAB4FB33" blockId="1.[106,783,1173,1457]" pageId="1" pageNumber="590">Fig.1 | Skeletal anatomy of V.gassenae gen. et sp. nov. (CAPPA/UFSM 0356). a, Skull reconstruction,with preserved elements coloured orange(the presence of teeth is hypothetical;see Supplementary Information for further discussion).b,c, Left premaxilla.d, Anterior tip of right dentary (reversed).</paragraph>
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<paragraph id="8BD718455C13FF98B913FAC0EAB8FAA4" blockId="1.[106,783,1173,1457]" pageId="1" pageNumber="590">e, Left orbitotemporal region of the skull.f,g, Braincase.h, Axis.i, Cervical vertebra.j, Dorsal vertebra.k, Caudal vertebra.l, Skeletal reconstruction,with preserved elements coloured orange(the presence of teeth is hypothetical). m, Right forearm and manus.n, Proximal portion of right radius and ulna.</paragraph>
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<paragraph id="8BD718455C13FF98B913FAABEAA0FA74" blockId="1.[106,783,1173,1457]" pageId="1" pageNumber="590">o, Manual ungual phalanges.p, Right pubis.q,r,s, Right femur.Orientations, a,c− e,g,h,j− m,o− q, lateral;b,i,r, ventral;f, posterior;n, medial;s, anterior. Scale bars,1 cm(a,m,q), 2 mm (b–d,h–k), 5 mm (e–g,n–p,r,s)and 100 mm (l).</paragraph>
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<paragraph id="8BD718455C13FF98BA49FB55ED9AFAE9" blockId="1.[816,1499,1172,1457]" pageId="1" pageNumber="590">4t,fourth trochanter;alr,anterolateral ridge;ap,ambiens process;at,anterior trochanter;crs,crenulated surface;crtf,crista tibiofibularis;ef,extensor fossa;f,frontal;fm,foramen magnum;fo,foramen;ft,flexor tubercle; J,Jugal;lc,lateral condyle;lgr,lateral groove;ltf,laterotemporal fenestra; mc,medial condyle;mc I–V,metacarpal I–V;nfo,narial fossa;ns,neural spine; o,orbit;oc,occipital condyle;of,obturator foramen;olp,olecranon process;</paragraph>
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<paragraph id="8BD718455C13FF98BA49FAF5ED5FFA74" blockId="1.[816,1499,1172,1457]" pageId="1" pageNumber="590">p, parietal;pa, parapophysis;pbs,parabasisphenoid;po,postorbital; pof,postfrontal;poz,postzygapophysis;pp,paroccipital process;prf,prefrontal; pro,prootic;prz,prezygapophysis;q,quadrate;qJ,quadratoJugal;r,radius; so,supraoccipital;sq,squamosal;stf,supratemporal fenestra;tp,transverse process;ts,trochanteric shelf;u,ulna;vk,ventral keel.</paragraph>
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</caption>
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<subSubSection id="C3724BCE5C13FF98B913F9C2EA5EF8D2" pageId="1" pageNumber="590" type="materials_examined">
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<materialsCitation id="3B0012185C13FF98B913F9C2EA5EF8D2" collectionCode="UFSM" country="Brazil" latitude="-29.6595" location="Sao Joao do Polesine" longLatPrecision="1" longitude="-53.429836" pageId="1" pageNumber="590" specimenCount="1" stateProvince="Rio Grande do Sul" typeStatus="holotype">
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<typeStatus id="54D3A6E75C13FF98B913F9C2E895F9D0" box="[106,205,1538,1557]" pageId="1" pageNumber="590">Holotype</typeStatus>
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. CAPPA/
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<collectionCode id="ED7980805C13FF98B85CF9C1E938F9D0" box="[293,352,1537,1557]" collectionName="UFSM" pageId="1" pageNumber="590">UFSM</collectionCode>
|
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(Centro de Apoio à Pesquisa Paleontológica da Quarta Colônia da Universidade Federal de Santa Maria) 0356, a partial skeleton of a single individual,including cranial and postcranial remains (
|
||
<figureCitation id="135304C05C13FF98B9BAF997E8A0F9AE" box="[195,248,1623,1644]" captionStart="Fig" captionStartId="1.[106,137,1173,1190]" captionTargetBox="[90,1551,118,1158]" captionTargetId="figure-314@1.[125,1486,114,1166]" captionTargetPageId="1" captionText="Fig.1 | Skeletal anatomy ofV.gassenae gen. etsp. nov. (CAPPA/UFSM 0356). a, Skull reconstruction,withpreserved elements coloured orange(the presence of teethis hypothetical;see Supplementary Information forfurther discussion).b,c, Left premaxilla.d, Anteriortip of rightdentary (reversed). e, Left orbitotemporal regionof the skull.f,g, Braincase.h, Axis.i, Cervical vertebra.j, Dorsal vertebra.k, Caudal vertebra.l, Skeletal reconstruction,with preserved elementscoloured orange(the presence of teethis hypothetical). m, Rightforearm and manus.n, Proximalportionof right radius andulna. o, Manual ungualphalanges.p, Right pubis.q,r,s, Right femur.Orientations, a,c−e,g,h,j−m,o−q, lateral;b,i,r, ventral;f, posterior;n, medial;s, anterior. Scale bars,1 cm(a,m,q), 2 mm (b–d,h–k), 5 mm (e–g,n–p,r,s)and 100 mm (l). 4t,fourth trochanter;alr,anterolateral ridge;ap,ambiens process;at,anterior trochanter;crs,crenulated surface;crtf,crista tibiofibularis;ef,extensor fossa;f,frontal;fm,foramen magnum;fo,foramen;ft,flexor tubercle; J,Jugal;lc,lateral condyle;lgr,lateral groove;ltf,laterotemporal fenestra; mc,medial condyle;mcI–V,metacarpal I–V;nfo,narial fossa;ns,neural spine; o,orbit;oc,occipitalcondyle;of,obturator foramen;olp,olecranon process; p, parietal;pa, parapophysis;pbs,parabasisphenoid;po,postorbital; pof,postfrontal;poz,postzygapophysis;pp,paroccipitalprocess;prf,prefrontal; pro,prootic;prz,prezygapophysis;q,quadrate;qJ,quadratoJugal;r,radius; so,supraoccipital;sq,squamosal;stf,supratemporal fenestra;tp,transverse process;ts,trochantericshelf;u,ulna;vk,ventral keel." figureDoi="http://doi.org/10.5281/zenodo.8263101" httpUri="https://zenodo.org/record/8263101/files/figure.png" pageId="1" pageNumber="590">Fig. 1</figureCitation>
|
||
). Locality and horizon. Burial site (
|
||
<geoCoordinate id="EE5C7E825C13FF98B8B4F9B4EA3CF94D" box="[461,612,1652,1673]" degrees="29" direction="south" minutes="39" orientation="latitude" pageId="1" pageNumber="590" precision="1" seconds="34.2" value="-29.6595">29° 39′ 34.2″ S</geoCoordinate>
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,
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<geoCoordinate id="EE5C7E825C13FF98BB14F9B4EB5FF94D" box="[621,775,1652,1673]" degrees="53" direction="west" minutes="25" orientation="longitude" pageId="1" pageNumber="590" precision="1" seconds="47.4" value="-53.429836">53° 25′ 47.4″ W</geoCoordinate>
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),
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<location id="8EB74E9E5C13FF98B913F950E961F963" LSID="urn:lsid:plazi:treatment:03C1A9535C12FF9DBAB0F9DFECBCFAD6:8EB74E9E5C13FF98B913F950E961F963" box="[106,313,1680,1702]" country="Brazil" latitude="-29.6595" longLatPrecision="1" longitude="-53.429836" name="Sao Joao do Polesine" pageId="1" pageNumber="590" stateProvince="Rio Grande do Sul">São João do Polêsine</location>
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,
|
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<collectingRegion id="49ACD6A75C13FF98B846F950E9ABF961" box="[319,499,1680,1701]" country="Brazil" name="Rio Grande do Sul" pageId="1" pageNumber="590">Rio Grande do Sul</collectingRegion>
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,
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<collectingCountry id="F37F58D55C13FF98B880F951EA6BF960" box="[505,563,1681,1701]" name="Brazil" pageId="1" pageNumber="590">Brazil</collectingCountry>
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(Extended Data
|
||
<figureCitation id="135304C05C13FF98BBAAF950EB5EF960" box="[723,774,1680,1701]" captionStart="Fig" captionStartId="1.[106,137,1173,1190]" captionTargetBox="[90,1551,118,1158]" captionTargetId="figure-314@1.[125,1486,114,1166]" captionTargetPageId="1" captionText="Fig.1 | Skeletal anatomy ofV.gassenae gen. etsp. nov. (CAPPA/UFSM 0356). a, Skull reconstruction,withpreserved elements coloured orange(the presence of teethis hypothetical;see Supplementary Information forfurther discussion).b,c, Left premaxilla.d, Anteriortip of rightdentary (reversed). e, Left orbitotemporal regionof the skull.f,g, Braincase.h, Axis.i, Cervical vertebra.j, Dorsal vertebra.k, Caudal vertebra.l, Skeletal reconstruction,with preserved elementscoloured orange(the presence of teethis hypothetical). m, Rightforearm and manus.n, Proximalportionof right radius andulna. o, Manual ungualphalanges.p, Right pubis.q,r,s, Right femur.Orientations, a,c−e,g,h,j−m,o−q, lateral;b,i,r, ventral;f, posterior;n, medial;s, anterior. Scale bars,1 cm(a,m,q), 2 mm (b–d,h–k), 5 mm (e–g,n–p,r,s)and 100 mm (l). 4t,fourth trochanter;alr,anterolateral ridge;ap,ambiens process;at,anterior trochanter;crs,crenulated surface;crtf,crista tibiofibularis;ef,extensor fossa;f,frontal;fm,foramen magnum;fo,foramen;ft,flexor tubercle; J,Jugal;lc,lateral condyle;lgr,lateral groove;ltf,laterotemporal fenestra; mc,medial condyle;mcI–V,metacarpal I–V;nfo,narial fossa;ns,neural spine; o,orbit;oc,occipitalcondyle;of,obturator foramen;olp,olecranon process; p, parietal;pa, parapophysis;pbs,parabasisphenoid;po,postorbital; pof,postfrontal;poz,postzygapophysis;pp,paroccipitalprocess;prf,prefrontal; pro,prootic;prz,prezygapophysis;q,quadrate;qJ,quadratoJugal;r,radius; so,supraoccipital;sq,squamosal;stf,supratemporal fenestra;tp,transverse process;ts,trochantericshelf;u,ulna;vk,ventral keel." figureDoi="http://doi.org/10.5281/zenodo.8263101" httpUri="https://zenodo.org/record/8263101/files/figure.png" pageId="1" pageNumber="590">Fig. 1</figureCitation>
|
||
). The outcrop strata belong to the Santa Maria Formation of the Paraná Basin. The maximum depositional age of biostratigraphically corre- lated beds from a nearby site was recorded as Carnian (233.23 ± 0.73), Ischigualastian reptile age,
|
||
<geologicalTimeScale id="F94898EE5C13FF98B801F8C3E9ACF8D2" box="[376,500,1795,1815]" pageId="1" pageNumber="590">Late Triassic</geologicalTimeScale>
|
||
<bibRefCitation id="EFF965B45C13FF98B88AF8C0EA58F8C8" author="Langer, M. C. & Ramezani, J. & Da Rosa, A. A." box="[499,512,1792,1805]" pageId="1" pageNumber="590" pagination="133 - 140" refId="ref4779" refString="13. Langer, M. C., Ramezani, J. & Da Rosa, A. A. U-Pb age constraints on dinosaur rise from south Brazil. Gondwana Res. 57, 133 - 140 (2018)." type="journal article" unsafe="true" year="2018">
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<superScript id="7C1DB50D5C13FF98B88AF8C0EA58F8C8" attach="left" box="[499,512,1792,1805]" fontSize="5" pageId="1" pageNumber="590">13</superScript>
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</bibRefCitation>
|
||
.
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||
</materialsCitation>
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||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="C3724BCE5C13FF98B913F8E1ED09F94D" pageId="1" pageNumber="590" type="diagnosis">
|
||
<paragraph id="8BD718455C13FF98B913F8E1EB49F806" blockId="1.[105,787,1537,1987]" pageId="1" pageNumber="590">Diagnosis. Differs from other lagerpetids in the following combination of character states (*indicates local autapomorphies):prefrontal contacting postfrontal*;occlusal margin of anterior end of dentary crenulated*;sharply tapering odontoid process of the axis*;dorsal vertebrae with lateral fossa on centrum;metacarpal IVlongerthanmetacarpal III*; pubis with well-developed ambiens process;femur with anterolateral</paragraph>
|
||
<paragraph id="8BD718455C13FF98BA56F9C1ED09F94D" blockId="1.[815,1498,1537,1672]" pageId="1" pageNumber="590">ridge (≠dorsolateral trochanter) on proximal portion*; trochanteric shelf;ridge for attachment of caudofemoralis muscles (=fourth trochanter); deep extensor fossa on anterior surface of distal portion; concave surface on posterolateral surface of crista tibiofibularis; and fibula with transversely compressed proximal end.</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="C3724BCE5C13FF9ABA49F902EB95F9D0" lastPageId="3" lastPageNumber="592" pageId="1" pageNumber="590" type="description">
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||
<paragraph id="8BD718455C13FF98BA49F902EB9CF91F" blockId="1.[816,1497,1730,1987]" box="[816,964,1730,1754]" pageId="1" pageNumber="590">
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<heading id="D09FAF295C13FF98BA49F902EB9CF91F" bold="true" box="[816,964,1730,1754]" fontSize="10" level="5" pageId="1" pageNumber="590" reason="0">Description</heading>
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||
</paragraph>
|
||
<paragraph id="8BD718455C13FF9BBA49F927E952F84F" blockId="1.[816,1497,1730,1987]" lastBlockId="2.[105,787,1422,1988]" lastPageId="2" lastPageNumber="591" pageId="1" pageNumber="590">
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||
The premaxilla is edentulous (that is, there are no alveoli;
|
||
<figureCitation id="135304C05C13FF98BC00F926ED92F93F" box="[1401,1482,1766,1787]" captionStart="Fig" captionStartId="1.[106,137,1173,1190]" captionTargetBox="[90,1551,118,1158]" captionTargetId="figure-314@1.[125,1486,114,1166]" captionTargetPageId="1" captionText="Fig.1 | Skeletal anatomy ofV.gassenae gen. etsp. nov. (CAPPA/UFSM 0356). a, Skull reconstruction,withpreserved elements coloured orange(the presence of teethis hypothetical;see Supplementary Information forfurther discussion).b,c, Left premaxilla.d, Anteriortip of rightdentary (reversed). e, Left orbitotemporal regionof the skull.f,g, Braincase.h, Axis.i, Cervical vertebra.j, Dorsal vertebra.k, Caudal vertebra.l, Skeletal reconstruction,with preserved elementscoloured orange(the presence of teethis hypothetical). m, Rightforearm and manus.n, Proximalportionof right radius andulna. o, Manual ungualphalanges.p, Right pubis.q,r,s, Right femur.Orientations, a,c−e,g,h,j−m,o−q, lateral;b,i,r, ventral;f, posterior;n, medial;s, anterior. Scale bars,1 cm(a,m,q), 2 mm (b–d,h–k), 5 mm (e–g,n–p,r,s)and 100 mm (l). 4t,fourth trochanter;alr,anterolateral ridge;ap,ambiens process;at,anterior trochanter;crs,crenulated surface;crtf,crista tibiofibularis;ef,extensor fossa;f,frontal;fm,foramen magnum;fo,foramen;ft,flexor tubercle; J,Jugal;lc,lateral condyle;lgr,lateral groove;ltf,laterotemporal fenestra; mc,medial condyle;mcI–V,metacarpal I–V;nfo,narial fossa;ns,neural spine; o,orbit;oc,occipitalcondyle;of,obturator foramen;olp,olecranon process; p, parietal;pa, parapophysis;pbs,parabasisphenoid;po,postorbital; pof,postfrontal;poz,postzygapophysis;pp,paroccipitalprocess;prf,prefrontal; pro,prootic;prz,prezygapophysis;q,quadrate;qJ,quadratoJugal;r,radius; so,supraoccipital;sq,squamosal;stf,supratemporal fenestra;tp,transverse process;ts,trochantericshelf;u,ulna;vk,ventral keel." figureDoi="http://doi.org/10.5281/zenodo.8263101" httpUri="https://zenodo.org/record/8263101/files/figure.png" pageId="1" pageNumber="590">Fig. 1b,c</figureCitation>
|
||
), with a sharp occlusal margin and a ventrally directed anterior end, resulting in a raptorial beak (Extended Data
|
||
<figureCitation id="135304C05C13FF98BD8BF8E0ED72F8F1" box="[1266,1322,1824,1845]" captionStart="Fig" captionStartId="2.[106,137,1182,1199]" captionTargetBox="[49,1497,119,1168]" captionTargetId="figure-450@2.[122,1477,129,1158]" captionTargetPageId="2" captionText="Fig.2 | Results ofphylogenetic and biogeographical analyses. a, Time- calibrated reduced strict consensus treedepicting thephylogenetic position of V.gassenaegen.et sp.nov.and other pterosauromorphs,and their estimated biogeographic history.Absolute bootstrap frequencies (left) and Bremer support values (right) are indicated on each branch.b, Geographical distribution of lagerpetids during theMiddle to Late Triassic.Arrowsindicate dispersion events.c, Life reconstruction of V.gassenaegen.et sp.nov.by Caio Fantini.d, Biplot of accumulatedamount of latitudinal dispersionversus time for Pan-Aves during theTriassic.The pterosauromorph silhouette was adapted from ref.10 (CC BY 4.0;https://creativecommons.org/licenses/ by/4.0/). The silesaurid silhouette wasadapted from ref.43 (CC BY 4.0; https://creativecommons.org/licenses/by/4.0/). Maps wereadapted from The Paleobiology Database(CCBY 4.0; https://creativecommons.org/ licenses/by/4.0/)." figureDoi="http://doi.org/10.5281/zenodo.8263104" httpUri="https://zenodo.org/record/8263104/files/figure.png" pageId="1" pageNumber="590">Fig. 2</figureCitation>
|
||
) resembling that of some birds such as phorusrhacids and falconiforms.The lateral surface of the premaxilla is striated,indicating the potential support of a keratinous covering or rhamphotheca
|
||
<superScript id="7C1DB50D5C13FF98BD98F8B2ED58F8BA" attach="left" box="[1249,1280,1906,1919]" fontSize="5" pageId="1" pageNumber="590">
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<bibRefCitation id="EFF965B45C13FF98BD98F8B2ECB6F8BA" author="Morhardt, A. C." box="[1249,1262,1906,1919]" pageId="1" pageNumber="590" refId="ref5001" refString="18. Morhardt, A. C. Dinosaur Smiles: Do the Texture and Morphology of the Premaxilla, Maxilla, and Dentary Bones of Sauropsids Provide Osteological Correlates for Inferring Extra-oral Structures Reliably in Dinosaurs? Master ̍ s thesis, Western Illinois Univ. (2009)." type="book" year="2009">18</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFF965B45C13FF98BD8BF8B2ED58F8BA" author="Hieronymus, T. L. & Witmer, L. M. & Tanke, D. H. & Currie, P. J." box="[1266,1280,1906,1919]" pageId="1" pageNumber="590" pagination="1370 - 1396" refId="ref5052" refString="19. Hieronymus, T. L., Witmer, L. M., Tanke, D. H. & Currie, P. J. The facial integument of centrosaurine ceratopsids: morphological and histological correlates of novel skin structures. Anat. Rec. 292, 1370 - 1396 (2009)." type="journal article" year="2009">19</bibRefCitation>
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</superScript>
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and contrasting with the smooth surface in early pterosaurs
|
||
<superScript id="7C1DB50D5C13FF98BDC3F84FEC92F859" attach="left" box="[1210,1226,1935,1948]" fontSize="5" pageId="1" pageNumber="590">20</superScript>
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||
and dinosauromorphs
|
||
<superScript id="7C1DB50D5C13FF98BCCAF84FED8BF859" attach="left" box="[1459,1491,1935,1948]" fontSize="5" pageId="1" pageNumber="590">
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<bibRefCitation id="EFF965B45C13FF98BCCAF84FED98F859" author="Ezcurra, M. D. & Nesbitt, S. J. & Fiorelli, L. E. & Desojo, J. B." box="[1459,1472,1935,1948]" pageId="1" pageNumber="590" pagination="1393 - 1438" refId="ref5134" refString="21. Ezcurra, M. D., Nesbitt, S. J., Fiorelli, L. E. & Desojo, J. B. New specimen sheds light on the anatomy and taxonomy of the early Late Triassic dinosauriforms from the Chanares Formation, NW Argentina. Anat. Rec. 303, 1393 - 1438 (2020)." type="journal article" year="2020">21</bibRefCitation>
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||
,
|
||
<bibRefCitation id="EFF965B45C13FF98BCBDF84FED8BF859" author="Nesbitt, S. J. & Langer, M. C. & Ezcurra, M. D." box="[1476,1491,1935,1948]" pageId="1" pageNumber="590" pagination="813 - 873" refId="ref5199" refString="22. Nesbitt, S. J., Langer, M. C. & Ezcurra, M. D. The anatomy of Asilisaurus kongwe, a dinosauriform from the Lifua Member of the Manda Beds (~ Middle Triassic) of Africa. Anat. Rec. 303, 813 - 873 (2020)." type="journal article" year="2020">22</bibRefCitation>
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</superScript>
|
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. A narial fossa occupies most of the base of the internarial process,and four large foramina pierce the palatal surface of the bone.The external naris is not retracted,contrasting with the condition in Triassic pterosaurs
|
||
<superScript id="7C1DB50D5C10FF9BB9E6FA04E8F6FA14" attach="left" box="[159,174,1476,1489]" fontSize="5" pageId="2" pageNumber="591">20</superScript>
|
||
. The orbit is anteroposteriorly longer than high.The prefrontal is unusually elongated, contacting the parietal posteromedially and the postfrontal posterolaterally,thus excluding the frontal from the external orbital rim (
|
||
<figureCitation id="135304C05C10FF9BB84DF9DEE92BF9F7" box="[308,371,1566,1587]" captionStart="Fig" captionStartId="1.[106,137,1173,1190]" captionTargetBox="[90,1551,118,1158]" captionTargetId="figure-314@1.[125,1486,114,1166]" captionTargetPageId="1" captionText="Fig.1 | Skeletal anatomy ofV.gassenae gen. etsp. nov. (CAPPA/UFSM 0356). a, Skull reconstruction,withpreserved elements coloured orange(the presence of teethis hypothetical;see Supplementary Information forfurther discussion).b,c, Left premaxilla.d, Anteriortip of rightdentary (reversed). e, Left orbitotemporal regionof the skull.f,g, Braincase.h, Axis.i, Cervical vertebra.j, Dorsal vertebra.k, Caudal vertebra.l, Skeletal reconstruction,with preserved elementscoloured orange(the presence of teethis hypothetical). m, Rightforearm and manus.n, Proximalportionof right radius andulna. o, Manual ungualphalanges.p, Right pubis.q,r,s, Right femur.Orientations, a,c−e,g,h,j−m,o−q, lateral;b,i,r, ventral;f, posterior;n, medial;s, anterior. Scale bars,1 cm(a,m,q), 2 mm (b–d,h–k), 5 mm (e–g,n–p,r,s)and 100 mm (l). 4t,fourth trochanter;alr,anterolateral ridge;ap,ambiens process;at,anterior trochanter;crs,crenulated surface;crtf,crista tibiofibularis;ef,extensor fossa;f,frontal;fm,foramen magnum;fo,foramen;ft,flexor tubercle; J,Jugal;lc,lateral condyle;lgr,lateral groove;ltf,laterotemporal fenestra; mc,medial condyle;mcI–V,metacarpal I–V;nfo,narial fossa;ns,neural spine; o,orbit;oc,occipitalcondyle;of,obturator foramen;olp,olecranon process; p, parietal;pa, parapophysis;pbs,parabasisphenoid;po,postorbital; pof,postfrontal;poz,postzygapophysis;pp,paroccipitalprocess;prf,prefrontal; pro,prootic;prz,prezygapophysis;q,quadrate;qJ,quadratoJugal;r,radius; so,supraoccipital;sq,squamosal;stf,supratemporal fenestra;tp,transverse process;ts,trochantericshelf;u,ulna;vk,ventral keel." figureDoi="http://doi.org/10.5281/zenodo.8263101" httpUri="https://zenodo.org/record/8263101/files/figure.png" pageId="2" pageNumber="591">Fig.1e</figureCitation>
|
||
), as probably also occurs in the lagerpetid ‘
|
||
<taxonomicName id="4C6863C65C10FF9BB916F9FBE8AEF98B" authorityName="Irmis, Nesbitt, Padian, Smith, Turner, Woody & Downs" authorityYear="2007" box="[111,246,1595,1614]" class="Reptilia" family="Lagerpetidae" genus="Dromomeron" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="2" pageNumber="591" phylum="Chordata" rank="genus">Dromomeron</taxonomicName>
|
||
’ gregorii. The triradiate postfrontal is proportionally larger than in most archosauriforms,resembling other lagerpetids
|
||
<bibRefCitation id="EFF965B45C10FF9BBA7DF994EB54F9A4" author="Ezcurra, M. D." box="[772,780,1620,1633]" pageId="2" pageNumber="591" pagination="445 - 449" refId="ref4557" refString="8. Ezcurra, M. D. et al. Enigmatic dinosaur precursors bridge the gap to the origin of Pterosauria. Nature 588, 445 - 449 (2020)." type="journal article" unsafe="true" year="2020">
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<superScript id="7C1DB50D5C10FF9BBA7DF994EB54F9A4" attach="left" box="[772,780,1620,1633]" fontSize="5" pageId="2" pageNumber="591">8</superScript>
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</bibRefCitation>
|
||
. This bone forms the posterodorsal margin of the orbit and the anterior margin of the supratemporal fenestra (Extended Data
|
||
<figureCitation id="135304C05C10FF9BBBEDF950EA84F960" box="[660,732,1680,1701]" captionStart="Fig" captionStartId="3.[106,137,1284,1301]" captionTargetBox="[90,1523,117,1260]" captionTargetId="figure-368@3.[143,1400,138,715]" captionTargetPageId="3" captionText="Fig.3 |Morphological disparitybetween earlyornithodirans.a,f,Gnathovorax cabreirai.b,g, Scleromochlus taylori.c, Ixalerpeton polesinensis.d,i, V.gassenae. e,j, Seazzadactylusvenieri. h, D.romeri. a–e, Skull in left lateral view, reconstruction.f–j, Right forelimbinlateral view.k,l, V.gassenae gen.et sp. nov.plottedinthe ornithodiran morphospace based onthe wholeskeleton (k) and skull(l). m,n, Sumof variances of thewhole skeleton(m) andskull (n); dots aremeans,and 95% confidence intervalswere generated using thetwo tails of valuesrecovered from 9,999 bootstrap technical replicatesof a dataset comprising n = 11 (whole skeleton)and n = 12 (skull)species of Ornithodiran precursors,n = 18 (whole skeletonand skull) species of Dinosauriaand n = 10 (whole skeleton andskull)species of Pterosauria.pt,Pteroid." figureDoi="http://doi.org/10.5281/zenodo.8263106" httpUri="https://zenodo.org/record/8263106/files/figure.png" pageId="2" pageNumber="591">Fig. 3a</figureCitation>
|
||
). The braincase (
|
||
<figureCitation id="135304C05C10FF9BB9ADF96DE979F907" box="[212,289,1709,1730]" captionStart="Fig" captionStartId="1.[106,137,1173,1190]" captionTargetBox="[90,1551,118,1158]" captionTargetId="figure-314@1.[125,1486,114,1166]" captionTargetPageId="1" captionText="Fig.1 | Skeletal anatomy ofV.gassenae gen. etsp. nov. (CAPPA/UFSM 0356). a, Skull reconstruction,withpreserved elements coloured orange(the presence of teethis hypothetical;see Supplementary Information forfurther discussion).b,c, Left premaxilla.d, Anteriortip of rightdentary (reversed). e, Left orbitotemporal regionof the skull.f,g, Braincase.h, Axis.i, Cervical vertebra.j, Dorsal vertebra.k, Caudal vertebra.l, Skeletal reconstruction,with preserved elementscoloured orange(the presence of teethis hypothetical). m, Rightforearm and manus.n, Proximalportionof right radius andulna. o, Manual ungualphalanges.p, Right pubis.q,r,s, Right femur.Orientations, a,c−e,g,h,j−m,o−q, lateral;b,i,r, ventral;f, posterior;n, medial;s, anterior. Scale bars,1 cm(a,m,q), 2 mm (b–d,h–k), 5 mm (e–g,n–p,r,s)and 100 mm (l). 4t,fourth trochanter;alr,anterolateral ridge;ap,ambiens process;at,anterior trochanter;crs,crenulated surface;crtf,crista tibiofibularis;ef,extensor fossa;f,frontal;fm,foramen magnum;fo,foramen;ft,flexor tubercle; J,Jugal;lc,lateral condyle;lgr,lateral groove;ltf,laterotemporal fenestra; mc,medial condyle;mcI–V,metacarpal I–V;nfo,narial fossa;ns,neural spine; o,orbit;oc,occipitalcondyle;of,obturator foramen;olp,olecranon process; p, parietal;pa, parapophysis;pbs,parabasisphenoid;po,postorbital; pof,postfrontal;poz,postzygapophysis;pp,paroccipitalprocess;prf,prefrontal; pro,prootic;prz,prezygapophysis;q,quadrate;qJ,quadratoJugal;r,radius; so,supraoccipital;sq,squamosal;stf,supratemporal fenestra;tp,transverse process;ts,trochantericshelf;u,ulna;vk,ventral keel." figureDoi="http://doi.org/10.5281/zenodo.8263101" httpUri="https://zenodo.org/record/8263101/files/figure.png" pageId="2" pageNumber="591">Fig.1f,g</figureCitation>
|
||
) preserves the posterior margin of a deep anterior tympanic recess and the basal tubera are broadly connected to each other medially,as in other lagerpetids
|
||
<bibRefCitation id="EFF965B45C10FF9BB894F923E9ADF935" author="Ezcurra, M. D." box="[493,501,1763,1776]" pageId="2" pageNumber="591" pagination="445 - 449" refId="ref4557" refString="8. Ezcurra, M. D. et al. Enigmatic dinosaur precursors bridge the gap to the origin of Pterosauria. Nature 588, 445 - 449 (2020)." type="journal article" unsafe="true" year="2020">
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<superScript id="7C1DB50D5C10FF9BB894F923E9ADF935" attach="left" box="[493,501,1763,1776]" fontSize="5" pageId="2" pageNumber="591">8</superScript>
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||
</bibRefCitation>
|
||
. The anterior tip of the dentary is edentulous but the occlusal margin is crenulated (
|
||
<figureCitation id="135304C05C10FF9BBBE1F8C3EA82F8D2" box="[664,730,1795,1816]" captionStart="Fig" captionStartId="1.[106,137,1173,1190]" captionTargetBox="[90,1551,118,1158]" captionTargetId="figure-314@1.[125,1486,114,1166]" captionTargetPageId="1" captionText="Fig.1 | Skeletal anatomy ofV.gassenae gen. etsp. nov. (CAPPA/UFSM 0356). a, Skull reconstruction,withpreserved elements coloured orange(the presence of teethis hypothetical;see Supplementary Information forfurther discussion).b,c, Left premaxilla.d, Anteriortip of rightdentary (reversed). e, Left orbitotemporal regionof the skull.f,g, Braincase.h, Axis.i, Cervical vertebra.j, Dorsal vertebra.k, Caudal vertebra.l, Skeletal reconstruction,with preserved elementscoloured orange(the presence of teethis hypothetical). m, Rightforearm and manus.n, Proximalportionof right radius andulna. o, Manual ungualphalanges.p, Right pubis.q,r,s, Right femur.Orientations, a,c−e,g,h,j−m,o−q, lateral;b,i,r, ventral;f, posterior;n, medial;s, anterior. Scale bars,1 cm(a,m,q), 2 mm (b–d,h–k), 5 mm (e–g,n–p,r,s)and 100 mm (l). 4t,fourth trochanter;alr,anterolateral ridge;ap,ambiens process;at,anterior trochanter;crs,crenulated surface;crtf,crista tibiofibularis;ef,extensor fossa;f,frontal;fm,foramen magnum;fo,foramen;ft,flexor tubercle; J,Jugal;lc,lateral condyle;lgr,lateral groove;ltf,laterotemporal fenestra; mc,medial condyle;mcI–V,metacarpal I–V;nfo,narial fossa;ns,neural spine; o,orbit;oc,occipitalcondyle;of,obturator foramen;olp,olecranon process; p, parietal;pa, parapophysis;pbs,parabasisphenoid;po,postorbital; pof,postfrontal;poz,postzygapophysis;pp,paroccipitalprocess;prf,prefrontal; pro,prootic;prz,prezygapophysis;q,quadrate;qJ,quadratoJugal;r,radius; so,supraoccipital;sq,squamosal;stf,supratemporal fenestra;tp,transverse process;ts,trochantericshelf;u,ulna;vk,ventral keel." figureDoi="http://doi.org/10.5281/zenodo.8263101" httpUri="https://zenodo.org/record/8263101/files/figure.png" pageId="2" pageNumber="591">Fig. 1d</figureCitation>
|
||
). The lateral surface of the bone adJacent to the occlusal margin is striated and densely pitted ventrally,also indicating a probable keratinous covering.The dentition posterior to the beak, if present,is unknown in Venetoraptor.
|
||
</paragraph>
|
||
<caption id="DF1748CD5C10FF9BB913FB5EEB91FA95" ID-DOI="http://doi.org/10.5281/zenodo.8263104" ID-Zenodo-Dep="8263104" httpUri="https://zenodo.org/record/8263104/files/figure.png" pageId="2" pageNumber="591" startId="2.[106,137,1182,1199]" subCaptionStartIDs="2.[1240,1287,1288,1306]" subCaptionStarts="MAPS " targetBox="[49,1497,119,1168]" targetPageId="2" targetType="figure">
|
||
<paragraph id="8BD718455C10FF9BB913FB5EEDC6FB6A" blockId="2.[106,786,1182,1360]" lastBlockId="2.[816,1460,1181,1360]" pageId="2" pageNumber="591">Fig.2 | Results of phylogenetic and biogeographical analyses. a, Time- calibrated reduced strict consensus tree depicting the phylogenetic position of V.gassenae gen.et sp.nov.and other pterosauromorphs,and their estimated biogeographic history.Absolute bootstrap frequencies (left) and Bremer support values (right) are indicated on each branch.b, Geographical distribution of lagerpetids during the Middle to Late Triassic.Arrows indicate dispersion events.c, Life reconstruction of V.gassenae gen.et sp.nov.by Caio Fantini.d, Biplot of accumulated amount of latitudinal dispersion versus</paragraph>
|
||
<paragraph id="8BD718455C10FF9BBA49FB78EDF6FADF" blockId="2.[816,1460,1181,1360]" pageId="2" pageNumber="591">time for Pan-Aves during the Triassic.The pterosauromorph silhouette was adapted from ref.10 (CC BY 4.0;https://creativecommons.org/licenses/ by/4.0/). The silesaurid silhouette was adapted from ref.43 (CC BY 4.0; https://creativecommons.org/licenses/by/4.0/). Maps were adapted from</paragraph>
|
||
<paragraph id="8BD718455C10FF9BBA49FAE3EB91FA95" blockId="2.[816,1460,1181,1360]" pageId="2" pageNumber="591">The Paleobiology Database(CC BY 4.0; https://creativecommons.org/ licenses/by/4.0/).</paragraph>
|
||
</caption>
|
||
<paragraph id="8BD718455C10FF9BB9F8F852EC74F98B" blockId="2.[105,787,1422,1988]" lastBlockId="2.[815,1497,1422,1987]" pageId="2" pageNumber="591">
|
||
As in other lagerpetids the vertebrae are proportionately short along the vertebral column,resembling the condition in early pterosaurs
|
||
<emphasis id="B91CC4575C10FF9BBB84F86CEB55F87C" bold="true" box="[765,781,1964,1977]" pageId="2" pageNumber="591">
|
||
<superScript id="7C1DB50D5C10FF9BBB84F86CEB55F87C" attach="right" box="[765,781,1964,1977]" fontSize="5" pageId="2" pageNumber="591">20</superScript>
|
||
</emphasis>
|
||
. The axis has a well-developed ventral keel(
|
||
<figureCitation id="135304C05C10FF9BBDBBFA4EED58FA66" box="[1218,1280,1422,1443]" captionStart="Fig" captionStartId="1.[106,137,1173,1190]" captionTargetBox="[90,1551,118,1158]" captionTargetId="figure-314@1.[125,1486,114,1166]" captionTargetPageId="1" captionText="Fig.1 | Skeletal anatomy ofV.gassenae gen. etsp. nov. (CAPPA/UFSM 0356). a, Skull reconstruction,withpreserved elements coloured orange(the presence of teethis hypothetical;see Supplementary Information forfurther discussion).b,c, Left premaxilla.d, Anteriortip of rightdentary (reversed). e, Left orbitotemporal regionof the skull.f,g, Braincase.h, Axis.i, Cervical vertebra.j, Dorsal vertebra.k, Caudal vertebra.l, Skeletal reconstruction,with preserved elementscoloured orange(the presence of teethis hypothetical). m, Rightforearm and manus.n, Proximalportionof right radius andulna. o, Manual ungualphalanges.p, Right pubis.q,r,s, Right femur.Orientations, a,c−e,g,h,j−m,o−q, lateral;b,i,r, ventral;f, posterior;n, medial;s, anterior. Scale bars,1 cm(a,m,q), 2 mm (b–d,h–k), 5 mm (e–g,n–p,r,s)and 100 mm (l). 4t,fourth trochanter;alr,anterolateral ridge;ap,ambiens process;at,anterior trochanter;crs,crenulated surface;crtf,crista tibiofibularis;ef,extensor fossa;f,frontal;fm,foramen magnum;fo,foramen;ft,flexor tubercle; J,Jugal;lc,lateral condyle;lgr,lateral groove;ltf,laterotemporal fenestra; mc,medial condyle;mcI–V,metacarpal I–V;nfo,narial fossa;ns,neural spine; o,orbit;oc,occipitalcondyle;of,obturator foramen;olp,olecranon process; p, parietal;pa, parapophysis;pbs,parabasisphenoid;po,postorbital; pof,postfrontal;poz,postzygapophysis;pp,paroccipitalprocess;prf,prefrontal; pro,prootic;prz,prezygapophysis;q,quadrate;qJ,quadratoJugal;r,radius; so,supraoccipital;sq,squamosal;stf,supratemporal fenestra;tp,transverse process;ts,trochantericshelf;u,ulna;vk,ventral keel." figureDoi="http://doi.org/10.5281/zenodo.8263101" httpUri="https://zenodo.org/record/8263101/files/figure.png" pageId="2" pageNumber="591">Fig.1h</figureCitation>
|
||
).The atlantal centrum and intercentrum are fused to the axis, the former tapering strongly anteriorly. There is a moderately deep fossa on the lateral surface of the dorsal centra (
|
||
<figureCitation id="135304C05C10FF9BBA99FA24EC41FA3C" box="[992,1049,1508,1529]" captionStart="Fig" captionStartId="1.[106,137,1173,1190]" captionTargetBox="[90,1551,118,1158]" captionTargetId="figure-314@1.[125,1486,114,1166]" captionTargetPageId="1" captionText="Fig.1 | Skeletal anatomy ofV.gassenae gen. etsp. nov. (CAPPA/UFSM 0356). a, Skull reconstruction,withpreserved elements coloured orange(the presence of teethis hypothetical;see Supplementary Information forfurther discussion).b,c, Left premaxilla.d, Anteriortip of rightdentary (reversed). e, Left orbitotemporal regionof the skull.f,g, Braincase.h, Axis.i, Cervical vertebra.j, Dorsal vertebra.k, Caudal vertebra.l, Skeletal reconstruction,with preserved elementscoloured orange(the presence of teethis hypothetical). m, Rightforearm and manus.n, Proximalportionof right radius andulna. o, Manual ungualphalanges.p, Right pubis.q,r,s, Right femur.Orientations, a,c−e,g,h,j−m,o−q, lateral;b,i,r, ventral;f, posterior;n, medial;s, anterior. Scale bars,1 cm(a,m,q), 2 mm (b–d,h–k), 5 mm (e–g,n–p,r,s)and 100 mm (l). 4t,fourth trochanter;alr,anterolateral ridge;ap,ambiens process;at,anterior trochanter;crs,crenulated surface;crtf,crista tibiofibularis;ef,extensor fossa;f,frontal;fm,foramen magnum;fo,foramen;ft,flexor tubercle; J,Jugal;lc,lateral condyle;lgr,lateral groove;ltf,laterotemporal fenestra; mc,medial condyle;mcI–V,metacarpal I–V;nfo,narial fossa;ns,neural spine; o,orbit;oc,occipitalcondyle;of,obturator foramen;olp,olecranon process; p, parietal;pa, parapophysis;pbs,parabasisphenoid;po,postorbital; pof,postfrontal;poz,postzygapophysis;pp,paroccipitalprocess;prf,prefrontal; pro,prootic;prz,prezygapophysis;q,quadrate;qJ,quadratoJugal;r,radius; so,supraoccipital;sq,squamosal;stf,supratemporal fenestra;tp,transverse process;ts,trochantericshelf;u,ulna;vk,ventral keel." figureDoi="http://doi.org/10.5281/zenodo.8263101" httpUri="https://zenodo.org/record/8263101/files/figure.png" pageId="2" pageNumber="591">Fig.1J</figureCitation>
|
||
), as in the posterior dorsal vertebrae of
|
||
<taxonomicName id="4C6863C65C10FF9BBCE3FA26EB33F9D0" class="Reptilia" family="Lagerpetidae" genus="Lagerpeton" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="2" pageNumber="591" phylum="Chordata" rank="genus">Lagerpeton</taxonomicName>
|
||
. The preserved dorsal vertebrae have anterior and posterior centrodiapophyseal and postzygodiapophyseal laminae. There is no hyposphene-hypantrum.
|
||
</paragraph>
|
||
<paragraph id="8BD718455C10FF9BBA3EF997EB9FF806" blockId="2.[815,1497,1422,1987]" pageId="2" pageNumber="591">
|
||
The olecranon process of the ulna is robust, enlarged and covered by striations (
|
||
<figureCitation id="135304C05C10FF9BBAC3F9B4EBA5F94D" box="[954,1021,1652,1673]" captionStart="Fig" captionStartId="1.[106,137,1173,1190]" captionTargetBox="[90,1551,118,1158]" captionTargetId="figure-314@1.[125,1486,114,1166]" captionTargetPageId="1" captionText="Fig.1 | Skeletal anatomy ofV.gassenae gen. etsp. nov. (CAPPA/UFSM 0356). a, Skull reconstruction,withpreserved elements coloured orange(the presence of teethis hypothetical;see Supplementary Information forfurther discussion).b,c, Left premaxilla.d, Anteriortip of rightdentary (reversed). e, Left orbitotemporal regionof the skull.f,g, Braincase.h, Axis.i, Cervical vertebra.j, Dorsal vertebra.k, Caudal vertebra.l, Skeletal reconstruction,with preserved elementscoloured orange(the presence of teethis hypothetical). m, Rightforearm and manus.n, Proximalportionof right radius andulna. o, Manual ungualphalanges.p, Right pubis.q,r,s, Right femur.Orientations, a,c−e,g,h,j−m,o−q, lateral;b,i,r, ventral;f, posterior;n, medial;s, anterior. Scale bars,1 cm(a,m,q), 2 mm (b–d,h–k), 5 mm (e–g,n–p,r,s)and 100 mm (l). 4t,fourth trochanter;alr,anterolateral ridge;ap,ambiens process;at,anterior trochanter;crs,crenulated surface;crtf,crista tibiofibularis;ef,extensor fossa;f,frontal;fm,foramen magnum;fo,foramen;ft,flexor tubercle; J,Jugal;lc,lateral condyle;lgr,lateral groove;ltf,laterotemporal fenestra; mc,medial condyle;mcI–V,metacarpal I–V;nfo,narial fossa;ns,neural spine; o,orbit;oc,occipitalcondyle;of,obturator foramen;olp,olecranon process; p, parietal;pa, parapophysis;pbs,parabasisphenoid;po,postorbital; pof,postfrontal;poz,postzygapophysis;pp,paroccipitalprocess;prf,prefrontal; pro,prootic;prz,prezygapophysis;q,quadrate;qJ,quadratoJugal;r,radius; so,supraoccipital;sq,squamosal;stf,supratemporal fenestra;tp,transverse process;ts,trochantericshelf;u,ulna;vk,ventral keel." figureDoi="http://doi.org/10.5281/zenodo.8263101" httpUri="https://zenodo.org/record/8263101/files/figure.png" pageId="2" pageNumber="591">Fig. 1n</figureCitation>
|
||
). The distal articular surface of the ulna is con- vex whereas the distal end of the radius is concave.The metacarpals increase sequentially in length from digits I to IV (
|
||
<figureCitation id="135304C05C10FF9BBC63F96DED3AF904" box="[1306,1378,1709,1730]" captionStart="Fig" captionStartId="1.[106,137,1173,1190]" captionTargetBox="[90,1551,118,1158]" captionTargetId="figure-314@1.[125,1486,114,1166]" captionTargetPageId="1" captionText="Fig.1 | Skeletal anatomy ofV.gassenae gen. etsp. nov. (CAPPA/UFSM 0356). a, Skull reconstruction,withpreserved elements coloured orange(the presence of teethis hypothetical;see Supplementary Information forfurther discussion).b,c, Left premaxilla.d, Anteriortip of rightdentary (reversed). e, Left orbitotemporal regionof the skull.f,g, Braincase.h, Axis.i, Cervical vertebra.j, Dorsal vertebra.k, Caudal vertebra.l, Skeletal reconstruction,with preserved elementscoloured orange(the presence of teethis hypothetical). m, Rightforearm and manus.n, Proximalportionof right radius andulna. o, Manual ungualphalanges.p, Right pubis.q,r,s, Right femur.Orientations, a,c−e,g,h,j−m,o−q, lateral;b,i,r, ventral;f, posterior;n, medial;s, anterior. Scale bars,1 cm(a,m,q), 2 mm (b–d,h–k), 5 mm (e–g,n–p,r,s)and 100 mm (l). 4t,fourth trochanter;alr,anterolateral ridge;ap,ambiens process;at,anterior trochanter;crs,crenulated surface;crtf,crista tibiofibularis;ef,extensor fossa;f,frontal;fm,foramen magnum;fo,foramen;ft,flexor tubercle; J,Jugal;lc,lateral condyle;lgr,lateral groove;ltf,laterotemporal fenestra; mc,medial condyle;mcI–V,metacarpal I–V;nfo,narial fossa;ns,neural spine; o,orbit;oc,occipitalcondyle;of,obturator foramen;olp,olecranon process; p, parietal;pa, parapophysis;pbs,parabasisphenoid;po,postorbital; pof,postfrontal;poz,postzygapophysis;pp,paroccipitalprocess;prf,prefrontal; pro,prootic;prz,prezygapophysis;q,quadrate;qJ,quadratoJugal;r,radius; so,supraoccipital;sq,squamosal;stf,supratemporal fenestra;tp,transverse process;ts,trochantericshelf;u,ulna;vk,ventral keel." figureDoi="http://doi.org/10.5281/zenodo.8263101" httpUri="https://zenodo.org/record/8263101/files/figure.png" pageId="2" pageNumber="591">
|
||
Fig.
|
||
<quantity id="4C90B5A05C10FF9BBC3CF96DED3AF904" box="[1349,1378,1709,1729]" metricMagnitude="0" metricUnit="m" metricValue="1.0" pageId="2" pageNumber="591" unit="m" value="1.0">1m</quantity>
|
||
</figureCitation>
|
||
), as in most pterosaurs and Saltopus
|
||
<bibRefCitation id="EFF965B45C10FF9BBD50F906EC60F916" author="Benton, M. J. & Walker, A. D." box="[1065,1080,1734,1747]" pageId="2" pageNumber="591" pagination="285 - 299" refId="ref5258" refString="23. Benton, M. J. & Walker, A. D. Saltopus, a dinosauriform from the Upper Triassic of Scotland. Earth Environ. Sci. Trans. R. Soc. Edinb. 101, 285 - 299 (2011)." type="journal article" year="2011">
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<superScript id="7C1DB50D5C10FF9BBD50F906EC60F916" attach="left" box="[1065,1080,1734,1747]" fontSize="5" pageId="2" pageNumber="591">23</superScript>
|
||
</bibRefCitation>
|
||
. Conversely,metacarpal III is the longest in
|
||
<taxonomicName id="4C6863C65C10FF9BBA31F927EB96F93F" authorityName="Irmis, Nesbitt, Padian, Smith, Turner, Woody & Downs" authorityYear="2007" box="[840,974,1767,1786]" class="Reptilia" family="Lagerpetidae" genus="Dromomeron" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="2" pageNumber="591" phylum="Chordata" rank="genus">Dromomeron</taxonomicName>
|
||
romeri
|
||
<bibRefCitation id="EFF965B45C10FF9BBD6FF923EC46F935" author="Ezcurra, M. D." box="[1046,1054,1763,1776]" pageId="2" pageNumber="591" pagination="445 - 449" refId="ref4557" refString="8. Ezcurra, M. D. et al. Enigmatic dinosaur precursors bridge the gap to the origin of Pterosauria. Nature 588, 445 - 449 (2020)." type="journal article" unsafe="true" year="2020">
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<superScript id="7C1DB50D5C10FF9BBD6FF923EC46F935" attach="left" box="[1046,1054,1763,1776]" fontSize="5" pageId="2" pageNumber="591">8</superScript>
|
||
</bibRefCitation>
|
||
and Scleromochlus taylori
|
||
<bibRefCitation id="EFF965B45C10FF9BBC5FF923ED6BF935" author="Foffa, D." box="[1318,1331,1763,1776]" pageId="2" pageNumber="591" pagination="313 - 318" refId="ref4752" refString="12. Foffa, D. et al. Scleromochlus and the early evolution of Pterosauromorpha. Nature 610, 313 - 318 (2022)." type="journal article" year="2022">
|
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<superScript id="7C1DB50D5C10FF9BBC5FF923ED6BF935" attach="left" box="[1318,1331,1763,1776]" fontSize="5" pageId="2" pageNumber="591">12</superScript>
|
||
</bibRefCitation>
|
||
. Metacarpal
|
||
<collectionCode id="ED7980805C10FF9BBCCFF927ED9CF93E" box="[1462,1476,1767,1787]" country="Canada" lsid="urn:lsid:biocol.org:col:13946" name="Royal British Columbia Museum - Herbarium" pageId="2" pageNumber="591" type="Museum">V</collectionCode>
|
||
is reduced,slender,positioned ventral to the others and articulates with at least one phalanx (Extended Data
|
||
<figureCitation id="135304C05C10FF9BBDE1F8E0EC85F8F1" box="[1176,1245,1824,1845]" captionStart="Extended" captionStartId="12.[106,194,1551,1567]" captionTargetBox="[122,1480,132,1525]" captionTargetId="figure-3@12.[121,1481,131,1526]" captionText="Extended DataFig.4 |Additional boneelements of Venetoraptor gassenae (CAPPA/UFSM 0356). a, Right manus inlateral view.Proximalportionof the right fibula in(b)lateral,(c)anterior,and(d)proximal views.e, Rightmetatarsal IV in anteriorview.f, Right metatarsalIII inanterior view.g, Digit III of theright pes inmedial view.ef,extensorfossa;mc,metacarpal;ph,phalanx;uph,ungual phalanx.Scale bars:1 cm." figureDoi="http://doi.org/10.5281/zenodo.8263116" httpUri="https://zenodo.org/record/8263116/files/figure.png" pageId="2" pageNumber="591">Fig. 4a</figureCitation>
|
||
). There is a deep fossa on the anterior surface of the proximal portion of the metacarpals and a shallow extensor fossa on their distal ends.The ungual phalanges are proportionally long and trenchant (
|
||
<figureCitation id="135304C05C10FF9BBDF5F8B6EC95F84F" box="[1164,1229,1910,1931]" captionStart="Fig" captionStartId="1.[106,137,1173,1190]" captionTargetBox="[90,1551,118,1158]" captionTargetId="figure-314@1.[125,1486,114,1166]" captionTargetPageId="1" captionText="Fig.1 | Skeletal anatomy ofV.gassenae gen. etsp. nov. (CAPPA/UFSM 0356). a, Skull reconstruction,withpreserved elements coloured orange(the presence of teethis hypothetical;see Supplementary Information forfurther discussion).b,c, Left premaxilla.d, Anteriortip of rightdentary (reversed). e, Left orbitotemporal regionof the skull.f,g, Braincase.h, Axis.i, Cervical vertebra.j, Dorsal vertebra.k, Caudal vertebra.l, Skeletal reconstruction,with preserved elementscoloured orange(the presence of teethis hypothetical). m, Rightforearm and manus.n, Proximalportionof right radius andulna. o, Manual ungualphalanges.p, Right pubis.q,r,s, Right femur.Orientations, a,c−e,g,h,j−m,o−q, lateral;b,i,r, ventral;f, posterior;n, medial;s, anterior. Scale bars,1 cm(a,m,q), 2 mm (b–d,h–k), 5 mm (e–g,n–p,r,s)and 100 mm (l). 4t,fourth trochanter;alr,anterolateral ridge;ap,ambiens process;at,anterior trochanter;crs,crenulated surface;crtf,crista tibiofibularis;ef,extensor fossa;f,frontal;fm,foramen magnum;fo,foramen;ft,flexor tubercle; J,Jugal;lc,lateral condyle;lgr,lateral groove;ltf,laterotemporal fenestra; mc,medial condyle;mcI–V,metacarpal I–V;nfo,narial fossa;ns,neural spine; o,orbit;oc,occipitalcondyle;of,obturator foramen;olp,olecranon process; p, parietal;pa, parapophysis;pbs,parabasisphenoid;po,postorbital; pof,postfrontal;poz,postzygapophysis;pp,paroccipitalprocess;prf,prefrontal; pro,prootic;prz,prezygapophysis;q,quadrate;qJ,quadratoJugal;r,radius; so,supraoccipital;sq,squamosal;stf,supratemporal fenestra;tp,transverse process;ts,trochantericshelf;u,ulna;vk,ventral keel." figureDoi="http://doi.org/10.5281/zenodo.8263101" httpUri="https://zenodo.org/record/8263101/files/figure.png" pageId="2" pageNumber="591">Fig. 1o</figureCitation>
|
||
and Extended Data
|
||
<figureCitation id="135304C05C10FF9BBCF5F8B6ED97F84F" box="[1420,1487,1910,1931]" captionStart="Fig" captionStartId="3.[106,137,1284,1301]" captionTargetBox="[90,1523,117,1260]" captionTargetId="figure-368@3.[143,1400,138,715]" captionTargetPageId="3" captionText="Fig.3 |Morphological disparitybetween earlyornithodirans.a,f,Gnathovorax cabreirai.b,g, Scleromochlus taylori.c, Ixalerpeton polesinensis.d,i, V.gassenae. e,j, Seazzadactylusvenieri. h, D.romeri. a–e, Skull in left lateral view, reconstruction.f–j, Right forelimbinlateral view.k,l, V.gassenae gen.et sp. nov.plottedinthe ornithodiran morphospace based onthe wholeskeleton (k) and skull(l). m,n, Sumof variances of thewhole skeleton(m) andskull (n); dots aremeans,and 95% confidence intervalswere generated using thetwo tails of valuesrecovered from 9,999 bootstrap technical replicatesof a dataset comprising n = 11 (whole skeleton)and n = 12 (skull)species of Ornithodiran precursors,n = 18 (whole skeletonand skull) species of Dinosauriaand n = 10 (whole skeleton andskull)species of Pterosauria.pt,Pteroid." figureDoi="http://doi.org/10.5281/zenodo.8263106" httpUri="https://zenodo.org/record/8263106/files/figure.png" pageId="2" pageNumber="591">Fig.3c</figureCitation>
|
||
), bearing a large extensor tubercle that is perforated ventrally by a fossa as in D. romeri
|
||
<bibRefCitation id="EFF965B45C10FF9BBAC0F86CEB99F87C" author="Ezcurra, M. D." box="[953,961,1964,1977]" pageId="2" pageNumber="591" pagination="445 - 449" refId="ref4557" refString="8. Ezcurra, M. D. et al. Enigmatic dinosaur precursors bridge the gap to the origin of Pterosauria. Nature 588, 445 - 449 (2020)." type="journal article" year="2020">
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||
<superScript id="7C1DB50D5C10FF9BBAC0F86CEB99F87C" attach="right" box="[953,961,1964,1977]" fontSize="5" pageId="2" pageNumber="591">8</superScript>
|
||
</bibRefCitation>
|
||
.
|
||
</paragraph>
|
||
<caption id="DF1748CD5C11FF9AB913FAC4ED65FA45" ID-DOI="http://doi.org/10.5281/zenodo.8263106" ID-Zenodo-Dep="8263106" httpUri="https://zenodo.org/record/8263106/files/figure.png" pageId="3" pageNumber="592" startId="3.[106,137,1284,1301]" targetBox="[90,1523,117,1260]" targetPageId="3" targetType="figure">
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||
<paragraph id="8BD718455C11FF9AB913FAC4ED65FA45" blockId="3.[106,787,1283,1434]" lastBlockId="3.[817,1469,1283,1408]" pageId="3" pageNumber="592">
|
||
Fig.3 |Morphological disparity between early ornithodirans. a,f,Gnathovorax cabreirai.b,g, Scleromochlus taylori.c,
|
||
<taxonomicName id="4C6863C65C11FF9AB8CFFADEEA40FAEA" box="[438,536,1310,1327]" class="Reptilia" family="Lagerpetidae" genus="Ixalerpeton" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="592" phylum="Chordata" rank="genus">Ixalerpeton</taxonomicName>
|
||
polesinensis.d,i, V.gassenae. e,j, Seazzadactylus venieri. h, D. romeri. a–e, Skull in left lateral view, reconstruction.f–j, Right forelimb in lateral view.k,l, V.gassenae gen.et sp. nov.plotted in the ornithodiran morphospace based on the whole skeleton (k) and skull(l). m,n, Sum of variances of the whole skeleton(m) and skull (n); dots are means,and 95% confidence intervals were generated using the two tails of values recovered from 9,999 bootstrap technical replicates of a dataset comprising n = 11 (whole skeleton)and n = 12 (skull)species of Ornithodiran precursors,n = 18 (whole skeleton and skull) species of Dinosauria and n = 10 (whole skeleton and skull)species of Pterosauria.pt,Pteroid.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8BD718455C11FF9AB9F8FA25EB95F9D0" blockId="3.[105,788,1508,1988]" lastBlockId="3.[816,1496,1508,1557]" pageId="3" pageNumber="592">
|
||
The pubis is short and has a distinct dorsolaterally facing anti- trochanteric surface (
|
||
<figureCitation id="135304C05C11FF9AB83DF9C1E9DFF9D0" box="[324,391,1537,1558]" captionStart="Fig" captionStartId="1.[106,137,1173,1190]" captionTargetBox="[90,1551,118,1158]" captionTargetId="figure-314@1.[125,1486,114,1166]" captionTargetPageId="1" captionText="Fig.1 | Skeletal anatomy ofV.gassenae gen. etsp. nov. (CAPPA/UFSM 0356). a, Skull reconstruction,withpreserved elements coloured orange(the presence of teethis hypothetical;see Supplementary Information forfurther discussion).b,c, Left premaxilla.d, Anteriortip of rightdentary (reversed). e, Left orbitotemporal regionof the skull.f,g, Braincase.h, Axis.i, Cervical vertebra.j, Dorsal vertebra.k, Caudal vertebra.l, Skeletal reconstruction,with preserved elementscoloured orange(the presence of teethis hypothetical). m, Rightforearm and manus.n, Proximalportionof right radius andulna. o, Manual ungualphalanges.p, Right pubis.q,r,s, Right femur.Orientations, a,c−e,g,h,j−m,o−q, lateral;b,i,r, ventral;f, posterior;n, medial;s, anterior. Scale bars,1 cm(a,m,q), 2 mm (b–d,h–k), 5 mm (e–g,n–p,r,s)and 100 mm (l). 4t,fourth trochanter;alr,anterolateral ridge;ap,ambiens process;at,anterior trochanter;crs,crenulated surface;crtf,crista tibiofibularis;ef,extensor fossa;f,frontal;fm,foramen magnum;fo,foramen;ft,flexor tubercle; J,Jugal;lc,lateral condyle;lgr,lateral groove;ltf,laterotemporal fenestra; mc,medial condyle;mcI–V,metacarpal I–V;nfo,narial fossa;ns,neural spine; o,orbit;oc,occipitalcondyle;of,obturator foramen;olp,olecranon process; p, parietal;pa, parapophysis;pbs,parabasisphenoid;po,postorbital; pof,postfrontal;poz,postzygapophysis;pp,paroccipitalprocess;prf,prefrontal; pro,prootic;prz,prezygapophysis;q,quadrate;qJ,quadratoJugal;r,radius; so,supraoccipital;sq,squamosal;stf,supratemporal fenestra;tp,transverse process;ts,trochantericshelf;u,ulna;vk,ventral keel." figureDoi="http://doi.org/10.5281/zenodo.8263101" httpUri="https://zenodo.org/record/8263101/files/figure.png" pageId="3" pageNumber="592">Fig. 1p</figureCitation>
|
||
), as in
|
||
<taxonomicName id="4C6863C65C11FF9AB8B2F9C2EB4AF9D0" authority="AND EARLY PTEROSAURS" authorityName="AND EARLY PTEROSAURS" box="[459,786,1537,1557]" class="Reptilia" family="Lagerpetidae" genus="Lagerpeton" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="592" phylum="Chordata" rank="genus">Lagerpeton and early pterosaurs</taxonomicName>
|
||
(for example, Dimorphodon). The distal tip of the pubis lacks any expansion.The femur (
|
||
<figureCitation id="135304C05C11FF9AB830F9FAE9C7F98B" box="[329,415,1594,1615]" captionStart="Fig" captionStartId="1.[106,137,1173,1190]" captionTargetBox="[90,1551,118,1158]" captionTargetId="figure-314@1.[125,1486,114,1166]" captionTargetPageId="1" captionText="Fig.1 | Skeletal anatomy ofV.gassenae gen. etsp. nov. (CAPPA/UFSM 0356). a, Skull reconstruction,withpreserved elements coloured orange(the presence of teethis hypothetical;see Supplementary Information forfurther discussion).b,c, Left premaxilla.d, Anteriortip of rightdentary (reversed). e, Left orbitotemporal regionof the skull.f,g, Braincase.h, Axis.i, Cervical vertebra.j, Dorsal vertebra.k, Caudal vertebra.l, Skeletal reconstruction,with preserved elementscoloured orange(the presence of teethis hypothetical). m, Rightforearm and manus.n, Proximalportionof right radius andulna. o, Manual ungualphalanges.p, Right pubis.q,r,s, Right femur.Orientations, a,c−e,g,h,j−m,o−q, lateral;b,i,r, ventral;f, posterior;n, medial;s, anterior. Scale bars,1 cm(a,m,q), 2 mm (b–d,h–k), 5 mm (e–g,n–p,r,s)and 100 mm (l). 4t,fourth trochanter;alr,anterolateral ridge;ap,ambiens process;at,anterior trochanter;crs,crenulated surface;crtf,crista tibiofibularis;ef,extensor fossa;f,frontal;fm,foramen magnum;fo,foramen;ft,flexor tubercle; J,Jugal;lc,lateral condyle;lgr,lateral groove;ltf,laterotemporal fenestra; mc,medial condyle;mcI–V,metacarpal I–V;nfo,narial fossa;ns,neural spine; o,orbit;oc,occipitalcondyle;of,obturator foramen;olp,olecranon process; p, parietal;pa, parapophysis;pbs,parabasisphenoid;po,postorbital; pof,postfrontal;poz,postzygapophysis;pp,paroccipitalprocess;prf,prefrontal; pro,prootic;prz,prezygapophysis;q,quadrate;qJ,quadratoJugal;r,radius; so,supraoccipital;sq,squamosal;stf,supratemporal fenestra;tp,transverse process;ts,trochantericshelf;u,ulna;vk,ventral keel." figureDoi="http://doi.org/10.5281/zenodo.8263101" httpUri="https://zenodo.org/record/8263101/files/figure.png" pageId="3" pageNumber="592">Fig.1q–s</figureCitation>
|
||
) differs from that of other lagerpetids in the presence of an anterolateral vertical ridge on the proximal portion. This ridge is positioned more proximally than the dorsolateral trochanter of other dinosauromorphs
|
||
<bibRefCitation id="EFF965B45C11FF9AB8A7F94DE9B4F95F" author="Nesbitt, S. J." box="[478,492,1677,1690]" pageId="3" pageNumber="592" pagination="1 - 292" refId="ref5306" refString="24. Nesbitt, S. J. The early evolution of archosaurs: relationships and the origin of major clades. Bull. Am. Mus. Nat. Hist. 352, 1 - 292 (2011)." type="journal article" unsafe="true" year="2011">
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||
<superScript id="7C1DB50D5C11FF9AB8A7F94DE9B4F95F" attach="right" box="[478,492,1677,1690]" fontSize="5" pageId="3" pageNumber="592">24</superScript>
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||
</bibRefCitation>
|
||
, reaching the proximal articular surface.There is a rugose anterior trochanter associated with a densely striated trochanteric shelf, as occurs in ‘D.’ gregorii
|
||
<bibRefCitation id="EFF965B45C11FF9ABBA8F906EA87F916" author="Nesbitt, S. J." box="[721,735,1734,1747]" pageId="3" pageNumber="592" pagination="498 - 516" refId="ref5347" refString="25. Nesbitt, S. J. et al. Hindlimb osteology and distribution of basal dinosauromorphs from the Late Triassic of North America. J. Vertebr. Paleontol. 29, 498 - 516 (2009)." type="journal article" year="2009">
|
||
<superScript id="7C1DB50D5C11FF9ABBA8F906EA87F916" attach="left" box="[721,735,1734,1747]" fontSize="5" pageId="3" pageNumber="592">25</superScript>
|
||
</bibRefCitation>
|
||
. The fourth trochanter is crest-like and symmetrical, contrasting with its absence in Scleromochlus, D. romeri,
|
||
<taxonomicName id="4C6863C65C11FF9AB891F8C4EA2AF8D2" authorityName="Irmis, Nesbitt, Padian, Smith, Turner, Woody & Downs" authorityYear="2007" box="[488,626,1796,1815]" class="Reptilia" family="Lagerpetidae" genus="Dromomeron" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="592" phylum="Chordata" rank="genus">Dromomeron</taxonomicName>
|
||
gigas and early pterosaurs
|
||
<superScript id="7C1DB50D5C11FF9AB9A5F8DCE8A5F8EC" attach="left" box="[220,253,1820,1833]" fontSize="5" pageId="3" pageNumber="592">
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<bibRefCitation id="EFF965B45C11FF9AB9A5F8DCE8B2F8EC" author="Nesbitt, S. J." box="[220,234,1820,1833]" pageId="3" pageNumber="592" pagination="498 - 516" refId="ref5347" refString="25. Nesbitt, S. J. et al. Hindlimb osteology and distribution of basal dinosauromorphs from the Late Triassic of North America. J. Vertebr. Paleontol. 29, 498 - 516 (2009)." type="journal article" year="2009">25</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFF965B45C11FF9AB997F8DCE8A5F8EC" author="Padian, K." box="[238,253,1820,1833]" pageId="3" pageNumber="592" pagination="1 - 44" refId="ref5388" refString="26. Padian, K. Osteology and functional morphology of Dimorphodon macronyx (Buckland) (Pterosauria: Rhamphorhynchoidea) based on new material in the Yale Peabody Museum. Postilla 189, 1 - 44 (1983)." type="journal article" year="1983">26</bibRefCitation>
|
||
</superScript>
|
||
. The distal end of the femur has a concave posterolateral surface as in
|
||
<taxonomicName id="4C6863C65C11FF9AB84EF8FDE998F895" authorityName="Irmis, Nesbitt, Padian, Smith, Turner, Woody & Downs" authorityYear="2007" box="[311,448,1853,1872]" class="Reptilia" family="Lagerpetidae" genus="Dromomeron" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="592" phylum="Chordata" rank="genus">Dromomeron</taxonomicName>
|
||
<bibRefCitation id="EFF965B45C11FF9AB8B8F8F9E997F883" author="Nesbitt, S. J." box="[449,463,1849,1862]" pageId="3" pageNumber="592" pagination="498 - 516" refId="ref5347" refString="25. Nesbitt, S. J. et al. Hindlimb osteology and distribution of basal dinosauromorphs from the Late Triassic of North America. J. Vertebr. Paleontol. 29, 498 - 516 (2009)." type="journal article" year="2009">
|
||
<superScript id="7C1DB50D5C11FF9AB8B8F8F9E997F883" attach="right" box="[449,463,1849,1862]" fontSize="5" pageId="3" pageNumber="592">25</superScript>
|
||
</bibRefCitation>
|
||
. The fibular condyle and crista tibiofibularis form an obtuse angle between each other in distal view, contrasting with the acute angle present in
|
||
<taxonomicName id="4C6863C65C11FF9ABB6AF8B6EADEF84C" box="[531,646,1910,1929]" class="Reptilia" family="Lagerpetidae" genus="Ixalerpeton" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="592" phylum="Chordata" rank="genus">Ixalerpeton</taxonomicName>
|
||
, D. romeri and D. gigas
|
||
<superScript id="7C1DB50D5C11FF9AB9C0F84FE88FF859" attach="left" box="[185,215,1935,1948]" fontSize="5" pageId="3" pageNumber="592">
|
||
<bibRefCitation id="EFF965B45C11FF9AB9C0F84FE89FF859" author="Nesbitt, S. J." box="[185,199,1935,1948]" pageId="3" pageNumber="592" pagination="498 - 516" refId="ref5347" refString="25. Nesbitt, S. J. et al. Hindlimb osteology and distribution of basal dinosauromorphs from the Late Triassic of North America. J. Vertebr. Paleontol. 29, 498 - 516 (2009)." type="journal article" year="2009">25</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFF965B45C11FF9AB9B3F84FE88FF859" author="Martinez, R. N. & Apaldetti, C. & Correa, G. A. & Abelin, D." box="[202,215,1935,1948]" pageId="3" pageNumber="592" pagination="1 - 13" refId="ref5429" refString="27. Martinez, R. N., Apaldetti, C., Correa, G. A. & Abelin, D. A Norian lagerpetid dinosauromorph from the Quebrada del Barro Formation, northwestern Argentina. Ameghiniana 53, 1 - 13 (2016)." type="journal article" year="2016">27</bibRefCitation>
|
||
</superScript>
|
||
. The pedal unguals are not strongly recurved and lack a well-developed extensor tubercle. These unguals are slightly larger than those of the manus.By contrast,manual unguals are much larger in pterosaurs
|
||
<superScript id="7C1DB50D5C11FF9ABACEFA3EEB9FF9CE" attach="right" box="[951,967,1534,1547]" fontSize="5" pageId="3" pageNumber="592">20</superScript>
|
||
.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="C3724BCE5C11FF9DBA49F98FECBCFAD6" lastPageId="4" lastPageNumber="593" pageId="3" pageNumber="592" type="discussion">
|
||
<paragraph id="8BD718455C11FF9ABA49F98FEC02F9A2" blockId="3.[815,1497,1615,1987]" box="[816,1114,1615,1639]" pageId="3" pageNumber="592">
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||
<heading id="D09FAF295C11FF9ABA49F98FEC02F9A2" bold="true" box="[816,1114,1615,1639]" fontSize="10" level="5" pageId="3" pageNumber="592" reason="0">Analyses and discussion</heading>
|
||
</paragraph>
|
||
<paragraph id="8BD718455C11FF9DBA49F9B4E900FD83" blockId="3.[815,1497,1615,1987]" lastBlockId="4.[105,787,132,1988]" lastPageId="4" lastPageNumber="593" pageId="3" pageNumber="592">
|
||
The inclusion of
|
||
<collectionCode id="ED7980805C11FF9ABAABF9B5EBB8F94D" box="[978,992,1653,1672]" country="Canada" lsid="urn:lsid:biocol.org:col:13946" name="Royal British Columbia Museum - Herbarium" pageId="3" pageNumber="592" type="Museum">V</collectionCode>
|
||
.gassenae in the most comprehensive phylogenetic analyses of archosauromorph relationships,using both maximum parsimony and Bayesian inference as optimality criteria,nests the new species within
|
||
<taxonomicName id="4C6863C65C11FF9ABAC4F90AED54F91B" authority="AND LENDS SUPPORT TO" authorityName="AND LENDS SUPPORT TO" box="[957,1292,1738,1759]" family="Lagerpetidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="592" phylum="Chordata" rank="family">Lagerpetidae and lends support to</taxonomicName>
|
||
the sister group relationship between
|
||
<taxonomicName id="4C6863C65C11FF9ABAA6F926ED59F93F" authority="AND PTEROSAURIA" authorityName="AND PTEROSAURIA" box="[991,1281,1766,1787]" family="Lagerpetidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="592" phylum="Chordata" rank="family">Lagerpetidae and Pterosauria</taxonomicName>
|
||
<superScript id="7C1DB50D5C11FF9ABC78F923ED71F935" attach="left" box="[1281,1321,1763,1776]" fontSize="5" pageId="3" pageNumber="592">
|
||
<bibRefCitation id="EFF965B45C11FF9ABC78F923ED50F935" author="Ezcurra, M. D." box="[1281,1288,1763,1776]" pageId="3" pageNumber="592" pagination="445 - 449" refId="ref4557" refString="8. Ezcurra, M. D. et al. Enigmatic dinosaur precursors bridge the gap to the origin of Pterosauria. Nature 588, 445 - 449 (2020)." type="journal article" year="2020">8</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFF965B45C11FF9ABC75F923ED4FF935" author="Nesbitt, S. J." box="[1292,1303,1763,1776]" pageId="3" pageNumber="592" pagination="484 - 487" refId="ref4967" refString="17. Nesbitt, S. J. et al. The earliest bird-line archosaurs and the assembly of the dinosaur body plan. Nature 544, 484 - 487 (2017)." type="journal article" year="2017">17</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFF965B45C11FF9ABC63F923ED71F935" author="Baron, M. G." box="[1306,1321,1763,1776]" pageId="3" pageNumber="592" pagination="103777" refId="ref5478" refString="28. Baron, M. G. The origin of Pterosaurs. Earth Sci. Rev. 221, 103777 (2021)." type="journal article" year="2021">28</bibRefCitation>
|
||
</superScript>
|
||
. Under maximum parsimony, Venetoraptor is the sister taxon to the North American ‘D.’ gregorii in the reduced strict consensus tree (
|
||
<figureCitation id="135304C05C11FF9ABC79F8E0ED1DF8F1" box="[1280,1349,1824,1845]" captionStart="Fig" captionStartId="2.[106,137,1182,1199]" captionTargetBox="[49,1497,119,1168]" captionTargetId="figure-450@2.[122,1477,129,1158]" captionTargetPageId="2" captionText="Fig.2 | Results ofphylogenetic and biogeographical analyses. a, Time- calibrated reduced strict consensus treedepicting thephylogenetic position of V.gassenaegen.et sp.nov.and other pterosauromorphs,and their estimated biogeographic history.Absolute bootstrap frequencies (left) and Bremer support values (right) are indicated on each branch.b, Geographical distribution of lagerpetids during theMiddle to Late Triassic.Arrowsindicate dispersion events.c, Life reconstruction of V.gassenaegen.et sp.nov.by Caio Fantini.d, Biplot of accumulatedamount of latitudinal dispersionversus time for Pan-Aves during theTriassic.The pterosauromorph silhouette was adapted from ref.10 (CC BY 4.0;https://creativecommons.org/licenses/ by/4.0/). The silesaurid silhouette wasadapted from ref.43 (CC BY 4.0; https://creativecommons.org/licenses/by/4.0/). Maps wereadapted from The Paleobiology Database(CCBY 4.0; https://creativecommons.org/ licenses/by/4.0/)." figureDoi="http://doi.org/10.5281/zenodo.8263104" httpUri="https://zenodo.org/record/8263104/files/figure.png" pageId="3" pageNumber="592">Fig. 2a</figureCitation>
|
||
and Extended Data
|
||
<figureCitation id="135304C05C11FF9ABA1CF8FCEBC5F894" box="[869,925,1852,1873]" captionStart="Extended" captionStartId="13.[106,194,1769,1785]" captionTargetBox="[178,1424,131,1744]" captionTargetId="figure-7@13.[178,1424,131,1744]" captionText="Extended DataFig.5 | Reducedstrict consensus treedepicting the phylogeneticposition of Venetoraptor gassenae gen. et sp. nov.Absolute (left) andGC (grouppresent/contradicted)(right) bootstrap frequencies and Bremer support valuesare shown aboveeach branch.The silesaurid silhouette has beenadapted from Müller & Garcia43 (CC BY 4.0 (https://creativecommons. org/licenses/by/4.0/)." figureDoi="http://doi.org/10.5281/zenodo.8263118" httpUri="https://zenodo.org/record/8263118/files/figure.png" pageId="3" pageNumber="592">Fig. 5</figureCitation>
|
||
). The coeval Brazilian
|
||
<taxonomicName id="4C6863C65C11FF9ABDF8F8FDED54F895" authority="IS" authorityName="IS" box="[1153,1292,1852,1872]" class="Reptilia" family="Lagerpetidae" genus="Ixalerpeton" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="592" phylum="Chordata" rank="genus">Ixalerpeton is</taxonomicName>
|
||
the sister taxon to a clade comprising D. gigas and D. romeri, from the Norian-Rhaetian of
|
||
<collectingCountry id="F37F58D55C11FF9ABA49F8B6EBCAF84E" box="[816,914,1910,1931]" name="Argentina" pageId="3" pageNumber="592">Argentina</collectingCountry>
|
||
and North America,respectively.The Bayesian 50% maJority rule tree has a lower resolution within
|
||
<taxonomicName id="4C6863C65C11FF9ABDD9F852EC06F806" authority=", AND Venetoraptor IS FOUND IN A POLYTOMY WITH ALL" family="Lagerpetidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="592" phylum="Chordata" rank="family">Lagerpetidae,and Venetoraptor is found in a polytomy with all</taxonomicName>
|
||
<taxonomicName id="4C6863C65C11FF9ABD1BF870ED60F806" box="[1122,1336,1967,1987]" class="Reptilia" family="Lagerpetidae" genus="Dromomeron" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="592" phylum="Chordata" rank="species" species="undetermined">Dromomeron species</taxonomicName>
|
||
(Extended Data
|
||
<figureCitation id="135304C05C16FF9DB913FF44E8C7FF5C" box="[106,159,132,153]" captionStart="Extended" captionStartId="14.[106,194,1769,1785]" captionTargetBox="[236,1366,131,1744]" captionTargetId="figure-3@14.[236,1366,131,1744]" captionText="Extended DataFig.6 |Majority rule tree recoveredfrom the unconstrained Bayesian phylogeneticanalysis depicting the positionof Venetoraptor gassenae gen. et sp. nov. Numbers at nodesindicate posterior probabilities and dotted redvertical lines indicate theboundaries between thePermian and Triassic,Triassic andJurassic,and Cretaceusand Paleogene geologicalperiods. Life reconstructionof Venetoraptor gassenae gen.et sp.nov.by Caio Fantini." figureDoi="http://doi.org/10.5281/zenodo.8263120" httpUri="https://zenodo.org/record/8263120/files/figure.png" pageId="4" pageNumber="593">Fig.6</figureCitation>
|
||
).
|
||
<taxonomicName id="4C6863C65C16FF9DB9D7FF45EA65FF5C" authority="IS MORE SHALLOWLY NESTED THAN" box="[174,573,132,153]" class="Reptilia" family="Lagerpetidae" genus="Ixalerpeton" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="593" phylum="Chordata" rank="genus">Ixalerpeton is more shallowly nested than</taxonomicName>
|
||
the above-mentioned taxa and in a polytomy that includes the Argentinian
|
||
<taxonomicName id="4C6863C65C16FF9DBB09FF62EAB9FF70" box="[624,737,162,181]" class="Reptilia" family="Lagerpetidae" genus="Lagerpeton" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="593" phylum="Chordata" rank="genus">Lagerpeton</taxonomicName>
|
||
. The occurrence of two North American species deeply nested with South American forms supports a scenario of at least two episodes of dispersion during the Late Triassic (
|
||
<figureCitation id="135304C05C16FF9DB8F7FF37E98CFECE" box="[398,468,247,268]" captionStart="Fig" captionStartId="2.[106,137,1182,1199]" captionTargetBox="[49,1497,119,1168]" captionTargetId="figure-450@2.[122,1477,129,1158]" captionTargetPageId="2" captionText="Fig.2 | Results ofphylogenetic and biogeographical analyses. a, Time- calibrated reduced strict consensus treedepicting thephylogenetic position of V.gassenaegen.et sp.nov.and other pterosauromorphs,and their estimated biogeographic history.Absolute bootstrap frequencies (left) and Bremer support values (right) are indicated on each branch.b, Geographical distribution of lagerpetids during theMiddle to Late Triassic.Arrowsindicate dispersion events.c, Life reconstruction of V.gassenaegen.et sp.nov.by Caio Fantini.d, Biplot of accumulatedamount of latitudinal dispersionversus time for Pan-Aves during theTriassic.The pterosauromorph silhouette was adapted from ref.10 (CC BY 4.0;https://creativecommons.org/licenses/ by/4.0/). The silesaurid silhouette wasadapted from ref.43 (CC BY 4.0; https://creativecommons.org/licenses/by/4.0/). Maps wereadapted from The Paleobiology Database(CCBY 4.0; https://creativecommons.org/ licenses/by/4.0/)." figureDoi="http://doi.org/10.5281/zenodo.8263104" httpUri="https://zenodo.org/record/8263104/files/figure.png" pageId="4" pageNumber="593">Fig. 2b</figureCitation>
|
||
). Our biogeographical analyses reconstructed both dispersions with a south-to-north direction from a Gondwanan ancestral area (
|
||
<figureCitation id="135304C05C16FF9DB8EAFEF0E980FE80" box="[403,472,304,325]" captionStart="Fig" captionStartId="2.[106,137,1182,1199]" captionTargetBox="[49,1497,119,1168]" captionTargetId="figure-450@2.[122,1477,129,1158]" captionTargetPageId="2" captionText="Fig.2 | Results ofphylogenetic and biogeographical analyses. a, Time- calibrated reduced strict consensus treedepicting thephylogenetic position of V.gassenaegen.et sp.nov.and other pterosauromorphs,and their estimated biogeographic history.Absolute bootstrap frequencies (left) and Bremer support values (right) are indicated on each branch.b, Geographical distribution of lagerpetids during theMiddle to Late Triassic.Arrowsindicate dispersion events.c, Life reconstruction of V.gassenaegen.et sp.nov.by Caio Fantini.d, Biplot of accumulatedamount of latitudinal dispersionversus time for Pan-Aves during theTriassic.The pterosauromorph silhouette was adapted from ref.10 (CC BY 4.0;https://creativecommons.org/licenses/ by/4.0/). The silesaurid silhouette wasadapted from ref.43 (CC BY 4.0; https://creativecommons.org/licenses/by/4.0/). Maps wereadapted from The Paleobiology Database(CCBY 4.0; https://creativecommons.org/ licenses/by/4.0/)." figureDoi="http://doi.org/10.5281/zenodo.8263104" httpUri="https://zenodo.org/record/8263104/files/figure.png" pageId="4" pageNumber="593">Fig. 2a</figureCitation>
|
||
and Extended Data
|
||
<figureCitation id="135304C05C16FF9DBBDBFEF0EA82FE80" box="[674,730,304,325]" captionStart="Extended" captionStartId="15.[106,194,1769,1785]" captionTargetBox="[169,721,131,1744]" captionTargetId="figure-7@15.[169,723,131,1744]" captionText="Extended DataFig.7 | Ancestral geographicareas reconstructed by the Dispersal-Extinction-Cladogenesismodel in the eucrocopodan region of the tree. Abbreviations:AR,Argentina;BR,Brazil,Uruguay,Namibia;CH,China, Thailand,Kyrgyzstan;eNA,eastern USA,Eastern Canada,Morocco andAlgeria; EU,Europe,Russia andGreenland;INT,India,Tanzania,Zambia,Madagascar, Israel andSaudi Arabia;sAF,South Africa,Lesotho,Zimbabwe;wNA,western USA,British Columbia,Mexico and Venezuela.Life reconstruction of Venetoraptor gassenae gen.et sp.nov.by CaioFantini." figureDoi="http://doi.org/10.5281/zenodo.8263122" httpUri="https://zenodo.org/record/8263122/files/figure.png" pageId="4" pageNumber="593">Fig. 7</figureCitation>
|
||
). The apex of dispersed latitudinal degrees for lagerpetids occurred during post-Carnian times, soon after the end of the Carnian Pluvial Event (
|
||
<figureCitation id="135304C05C16FF9DB909FE46E8EBFE5E" box="[112,179,390,411]" captionStart="Fig" captionStartId="2.[106,137,1182,1199]" captionTargetBox="[49,1497,119,1168]" captionTargetId="figure-450@2.[122,1477,129,1158]" captionTargetPageId="2" captionText="Fig.2 | Results ofphylogenetic and biogeographical analyses. a, Time- calibrated reduced strict consensus treedepicting thephylogenetic position of V.gassenaegen.et sp.nov.and other pterosauromorphs,and their estimated biogeographic history.Absolute bootstrap frequencies (left) and Bremer support values (right) are indicated on each branch.b, Geographical distribution of lagerpetids during theMiddle to Late Triassic.Arrowsindicate dispersion events.c, Life reconstruction of V.gassenaegen.et sp.nov.by Caio Fantini.d, Biplot of accumulatedamount of latitudinal dispersionversus time for Pan-Aves during theTriassic.The pterosauromorph silhouette was adapted from ref.10 (CC BY 4.0;https://creativecommons.org/licenses/ by/4.0/). The silesaurid silhouette wasadapted from ref.43 (CC BY 4.0; https://creativecommons.org/licenses/by/4.0/). Maps wereadapted from The Paleobiology Database(CCBY 4.0; https://creativecommons.org/ licenses/by/4.0/)." figureDoi="http://doi.org/10.5281/zenodo.8263104" httpUri="https://zenodo.org/record/8263104/files/figure.png" pageId="4" pageNumber="593">Fig.2d</figureCitation>
|
||
). This dispersion model closely resembles that of dinosaurs
|
||
<bibRefCitation id="EFF965B45C16FF9DBB87FE43EB54FE55" author="Griffin, C. T." box="[766,780,387,400]" pageId="4" pageNumber="593" pagination="313 - 319" refId="ref5503" refString="29. Griffin, C. T. et al. Africa ̍ s oldest dinosaurs reveal early suppression of dinosaur distribution. Nature 609, 313 - 319 (2022)." type="journal article" unsafe="true" year="2022">
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<superScript id="7C1DB50D5C16FF9DBB87FE43EB54FE55" attach="left" box="[766,780,387,400]" fontSize="5" pageId="4" pageNumber="593">29</superScript>
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</bibRefCitation>
|
||
, suggesting that the same climatic barriers affected the radiation of both clades.By contrast,silesaurids show a sustained amount of latitudinal dispersion between the Middle-early Late Triassic,and in pterosaurs the peak is delayed until well into the early-middle Norian. Thus,Triassic ornithodirans had a much more complex biogeographic history than previously appreciated.
|
||
</paragraph>
|
||
<paragraph id="8BD718455C16FF9DB9F8FD8FE9DEFBA2" blockId="4.[105,787,132,1988]" pageId="4" pageNumber="593">
|
||
Whereas the phylogenetic relationships of lagerpetids have received considerable attention recently
|
||
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<bibRefCitation id="EFF965B45C16FF9DB8E2FDA8E9FAFDB0" author="Ezcurra, M. D." box="[411,418,616,629]" pageId="4" pageNumber="593" pagination="445 - 449" refId="ref4557" refString="8. Ezcurra, M. D. et al. Enigmatic dinosaur precursors bridge the gap to the origin of Pterosauria. Nature 588, 445 - 449 (2020)." type="journal article" year="2020">8</bibRefCitation>
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||
,
|
||
<bibRefCitation id="EFF965B45C16FF9DB8DFFDA8E9E9FDB0" author="Muller, R. T. & Langer, M. C. & Dias-Da-Silva, S." box="[422,433,616,629]" pageId="4" pageNumber="593" pagination="149 - 158" refId="ref4701" refString="11. Muller, R. T., Langer, M. C. & Dias-Da-Silva, S. Ingroup relationships of Lagerpetidae (Avemetatarsalia: Dinosauromorpha): a further phylogenetic investigation on the understanding of dinosaur relatives. Zootaxa 4392, 149 - 158 (2018)." type="journal article" year="2018">11</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFF965B45C16FF9DB8CDFDA8E999FDB0" author="Irmis, R. B." box="[436,449,616,629]" pageId="4" pageNumber="593" pagination="358 - 361" refId="ref4932" refString="16. Irmis, R. B. et al. A Late Triassic dinosauromorph assemblage from New Mexico and the rise of dinosaurs. Science 317, 358 - 361 (2007)." type="journal article" year="2007">16</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFF965B45C16FF9DB8BDFDA8E98AFDB0" author="Baron, M. G." box="[452,466,616,629]" pageId="4" pageNumber="593" pagination="103777" refId="ref5478" refString="28. Baron, M. G. The origin of Pterosaurs. Earth Sci. Rev. 221, 103777 (2021)." type="journal article" year="2021">28</bibRefCitation>
|
||
</superScript>
|
||
,their biology and ecological role remain obscure.The presence of Venetoraptor in the same fossiliferous site that yielded
|
||
<taxonomicName id="4C6863C65C16FF9DB868FD66E9A0FD7C" authority="REPRESENTS" authorityName="REPRESENTS" box="[273,504,677,697]" class="Reptilia" family="Lagerpetidae" genus="Ixalerpeton" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="593" phylum="Chordata" rank="genus">Ixalerpeton represents</taxonomicName>
|
||
the first robust evidence of sympatric lagerpetid species.This is particularly interesting because this site, located within the Hyperodapedon Assemblage Zone, also includes one of the oldest dinosaurs(that is,Buriolestes
|
||
<bibRefCitation id="EFF965B45C16FF9DBBF0FD38EAC8FCC0" author="Cabreira, S. F." box="[649,656,760,773]" pageId="4" pageNumber="593" pagination="3090 - 3095" refId="ref4520" refString="7. Cabreira, S. F. et al. A unique Late Triassic dinosauromorph assemblage reveals dinosaur ancestral anatomy and diet. Curr. Biol. 26, 3090 - 3095 (2016)." type="journal article" year="2016">
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<superScript id="7C1DB50D5C16FF9DBBF0FD38EAC8FCC0" attach="left" box="[649,656,760,773]" fontSize="5" pageId="4" pageNumber="593">7</superScript>
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</bibRefCitation>
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) and demon- strates the presence of an already diverse ornithodiran assemblage by the Carnian.Moreover,the new species indicates a previously unappre- ciated ecological diversity among early ornithodirans (=pterosaur and dinosaur precursors, hereafter ‘ornithodiran precursors’),with medium- to small-sized species (femoral length of Venetoraptor is
|
||
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versus
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<taxonomicName id="4C6863C65C16FF9DB98EFC68E9DAFC7E" authority="AT" authorityName="AT" box="[247,386,935,955]" class="Reptilia" family="Lagerpetidae" genus="Ixalerpeton" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="593" phylum="Chordata" rank="genus">Ixalerpeton at</taxonomicName>
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<quantity id="4C90B5A05C16FF9DB8FCFC67E994FC7E" box="[389,460,935,955]" metricMagnitude="-2" metricUnit="m" metricValue="6.9" pageId="4" pageNumber="593" unit="mm" value="69.0">69 mm</quantity>
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) and different rostral morphologies (
|
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<figureCitation id="135304C05C16FF9DB9E7FC04E8AEFC1D" box="[158,246,964,985]" captionStart="Fig" captionStartId="3.[106,137,1284,1301]" captionTargetBox="[90,1523,117,1260]" captionTargetId="figure-368@3.[143,1400,138,715]" captionTargetPageId="3" captionText="Fig.3 |Morphological disparitybetween earlyornithodirans.a,f,Gnathovorax cabreirai.b,g, Scleromochlus taylori.c, Ixalerpeton polesinensis.d,i, V.gassenae. e,j, Seazzadactylusvenieri. h, D.romeri. a–e, Skull in left lateral view, reconstruction.f–j, Right forelimbinlateral view.k,l, V.gassenae gen.et sp. nov.plottedinthe ornithodiran morphospace based onthe wholeskeleton (k) and skull(l). m,n, Sumof variances of thewhole skeleton(m) andskull (n); dots aremeans,and 95% confidence intervalswere generated using thetwo tails of valuesrecovered from 9,999 bootstrap technical replicatesof a dataset comprising n = 11 (whole skeleton)and n = 12 (skull)species of Ornithodiran precursors,n = 18 (whole skeletonand skull) species of Dinosauriaand n = 10 (whole skeleton andskull)species of Pterosauria.pt,Pteroid." figureDoi="http://doi.org/10.5281/zenodo.8263106" httpUri="https://zenodo.org/record/8263106/files/figure.png" pageId="4" pageNumber="593">Fig. 3c,d</figureCitation>
|
||
). Associated with the discovery of other new species (for example,I.polesinensis
|
||
<bibRefCitation id="EFF965B45C16FF9DB803FC1DE9D9FC2F" author="Cabreira, S. F." box="[378,385,989,1002]" pageId="4" pageNumber="593" pagination="3090 - 3095" refId="ref4520" refString="7. Cabreira, S. F. et al. A unique Late Triassic dinosauromorph assemblage reveals dinosaur ancestral anatomy and diet. Curr. Biol. 26, 3090 - 3095 (2016)." type="journal article" year="2016">
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</bibRefCitation>
|
||
;
|
||
<taxonomicName id="4C6863C65C16FF9DB8F3FC21EA19FC31" authority="Kely" authorityName="Kely" box="[394,577,993,1012]" family="Lagerpetidae" genus="Kongonaphon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="593" phylum="Chordata" rank="genus">Kongonaphon Kely</taxonomicName>
|
||
<bibRefCitation id="EFF965B45C16FF9DBB38FC1DEA11FC2F" author="Kammerer, C. F. & Nesbitt, S. J. & Flynn, J. J. & Ranivoharimanana, L. & Wyss, A. R." box="[577,585,989,1002]" pageId="4" pageNumber="593" pagination="17932 - 17936" refId="ref4590" refString="9. Kammerer, C. F., Nesbitt, S. J., Flynn, J. J., Ranivoharimanana, L. & Wyss, A. R. A tiny ornithodiran archosaur from the Triassic of Madagascar and the role of miniaturization in dinosaur and pterosaur ancestry. Proc. Natl Acad. Sci. USA 117, 17932 - 17936 (2020)." type="journal article" year="2020">
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<superScript id="7C1DB50D5C16FF9DBB38FC1DEA11FC2F" attach="left" box="[577,585,989,1002]" fontSize="5" pageId="4" pageNumber="593">9</superScript>
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</bibRefCitation>
|
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) and systematic and morphological re-evaluation of some taxa (for example,I.polesinensis
|
||
<bibRefCitation id="EFF965B45C16FF9DBA7DFC3AEB54FBC2" author="Ezcurra, M. D." box="[772,780,1018,1031]" pageId="4" pageNumber="593" pagination="445 - 449" refId="ref4557" refString="8. Ezcurra, M. D. et al. Enigmatic dinosaur precursors bridge the gap to the origin of Pterosauria. Nature 588, 445 - 449 (2020)." type="journal article" year="2020">
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<superScript id="7C1DB50D5C16FF9DBA7DFC3AEB54FBC2" attach="left" box="[772,780,1018,1031]" fontSize="5" pageId="4" pageNumber="593">8</superScript>
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</bibRefCitation>
|
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; Faxinalipterus minimus
|
||
<bibRefCitation id="EFF965B45C16FF9DB837FBD6E904FBE6" author="Kellner, A. W." box="[334,348,1046,1059]" pageId="4" pageNumber="593" pagination="13276" refId="ref4661" refString="10. Kellner, A. W. et al. Reassessment of Faxinalipterus minimus, a purported Triassic pterosaur from southern Brazil with the description of a new taxon. PeerJ 10, e 13276 (2022)." type="journal article" year="2022">
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<superScript id="7C1DB50D5C16FF9DB837FBD6E904FBE6" attach="left" box="[334,348,1046,1059]" fontSize="5" pageId="4" pageNumber="593">10</superScript>
|
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</bibRefCitation>
|
||
; Scleromochlus taylori
|
||
<bibRefCitation id="EFF965B45C16FF9DBB4CFBD6EA19FBE6" author="Foffa, D." box="[565,577,1046,1059]" pageId="4" pageNumber="593" pagination="313 - 318" refId="ref4752" refString="12. Foffa, D. et al. Scleromochlus and the early evolution of Pterosauromorpha. Nature 610, 313 - 318 (2022)." type="journal article" year="2022">
|
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<superScript id="7C1DB50D5C16FF9DBB4CFBD6EA19FBE6" attach="right" box="[565,577,1046,1059]" fontSize="5" pageId="4" pageNumber="593">12</superScript>
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</bibRefCitation>
|
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), Venetoraptor shows a hidden ecomorphological disparity within
|
||
<taxonomicName id="4C6863C65C16FF9DBB5BFBF6EAFFFB8E" box="[546,679,1078,1099]" family="Lagerpetidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="593" phylum="Chordata" rank="family">Lagerpetidae</taxonomicName>
|
||
(
|
||
<figureCitation id="135304C05C16FF9DBBC8FBF6EAB0FB8E" box="[689,744,1078,1099]" captionStart="Fig" captionStartId="3.[106,137,1284,1301]" captionTargetBox="[90,1523,117,1260]" captionTargetId="figure-368@3.[143,1400,138,715]" captionTargetPageId="3" captionText="Fig.3 |Morphological disparitybetween earlyornithodirans.a,f,Gnathovorax cabreirai.b,g, Scleromochlus taylori.c, Ixalerpeton polesinensis.d,i, V.gassenae. e,j, Seazzadactylusvenieri. h, D.romeri. a–e, Skull in left lateral view, reconstruction.f–j, Right forelimbinlateral view.k,l, V.gassenae gen.et sp. nov.plottedinthe ornithodiran morphospace based onthe wholeskeleton (k) and skull(l). m,n, Sumof variances of thewhole skeleton(m) andskull (n); dots aremeans,and 95% confidence intervalswere generated using thetwo tails of valuesrecovered from 9,999 bootstrap technical replicatesof a dataset comprising n = 11 (whole skeleton)and n = 12 (skull)species of Ornithodiran precursors,n = 18 (whole skeletonand skull) species of Dinosauriaand n = 10 (whole skeleton andskull)species of Pterosauria.pt,Pteroid." figureDoi="http://doi.org/10.5281/zenodo.8263106" httpUri="https://zenodo.org/record/8263106/files/figure.png" pageId="4" pageNumber="593">Fig. 3</figureCitation>
|
||
and Extended Data
|
||
<figureCitation id="135304C05C16FF9DB879FB93E967FBA2" box="[256,319,1107,1128]" captionStart="Extended" captionStartId="16.[106,194,1730,1746]" captionTargetBox="[121,1481,131,1706]" captionTargetId="figure-3@16.[121,1481,131,1706]" captionText="Extended DataFig.8 |Results of the morphological disparity analyses. Bivariate plots using:a, Whole skeleton;b,Skull;c, Rostrum;d, Forelimb; e, Anterior zeugopodiumand autopodium;f,Hindlimb.Sum of variances: g, Whole skeleton;h, Skull;i, Rostrum;j, Forelimb;k, Anteriorzeugopodium andautopodium;l,Hindlimb.In the Sum of Variancesthe dots are means and the95% confidence intervals weregenerated using thetwo tails of values recoveredfrom 9,999bootstrap technical replicates of adataset composed of n = 11 (Whole skeleton,Forelimb),n = 12(Skull,Rostrum),n = 7 (Anterior zeugopodium andautopodium) andn = 17 (Hindlimb)species of Ornithodiran precursors,n = 18 (Whole skeleton,Skull,Rostrum),n = 16 (Forelimb),n = 14 (Anterior zeugopodiumand autopodium) and n = 19(Hindlimb)species of Dinosauria,and n = 10 (Whole skeleton,Skull,Rostrum),n = 7 (Forelimb, Anterior zeugopodiumand autopodium) and n = 6 (Hindlimb) species of Pterosauria.Thepterosauromorph silhouette has beenadapted from Kellner et al.10 (CC BY 4.0 (https://creativecommons.org/licenses/by/4.0/)." figureDoi="http://doi.org/10.5281/zenodo.8263124" httpUri="https://zenodo.org/record/8263124/files/figure.png" pageId="4" pageNumber="593">Figs.8</figureCitation>
|
||
and
|
||
<figureCitation id="135304C05C16FF9DB815FB93E922FBA2" box="[364,378,1107,1127]" captionStart="Extended" captionStartId="18.[106,194,133,149]" captionTargetId="figure-7@17.[121,1481,131,1705]" captionTargetPageId="17" captionText="Extended Data Fig. 9 | Additional results of the morphological disparity analyses. Bivariate plots using:a, Wholeskeleton;b, Skull;c, Rostrum; d, Forelimb;e, Anterior zeugopodium and autopodium;f, Hindlimb.Sum of variances:g, Whole skeleton;h, Skull;i, Rostrum;j, Forelimb;k, Anterior zeugopodium and autopodium;l, Hindlimb.In the Sum of Variances the dots are meansand the 95% confidence intervals were generated using the two tails of values recovered from 9,999 bootstrap technicalreplicates of a dataset composed of n = 5 (Whole skeleton,Skull),n = 4 (Rostrum,Forelimb),n = 3 (Anterior zeugopodium andautopodium) and n = 9 (Hindlimb) speciesof Lagerpetidae,n = 5(Whole skeleton),n = 6 (Skull,Forelimb),n = 7 (Rostrum, Hindlimb)and n = 3 (Anterior zeugopodium and autopodium) species of Silesauridae,n = 18 (Whole skeleton,Skull,Rostrum),n = 16 (Forelimb),n = 14 (Anterior zeugopodium and autopodium) and n = 19 (Hindlimb) speciesof Dinosauria,and n = 10 (Whole skeleton,Skull,Rostrum),n = 7 (Forelimb, Anterior zeugopodium and autopodium) and n = 6 (Hindlimb)species of Pterosauria.The pterosauromorph silhouette has been adapted from Kellner et al.10 (CC BY 4.0(https://creativecommons.org/licenses/by/4.0/). The silesaurid silhouette has been adapted fromMüller & Garcia43 (CC BY 4.0 (https://creativecommons.org/licenses/by/4.0/)." figureDoi="http://doi.org/10.5281/zenodo.8263126" httpUri="https://zenodo.org/record/8263126/files/figure.png" pageId="4" pageNumber="593">9</figureCitation>
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).
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||
</paragraph>
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<paragraph id="8BD718455C16FF9DB9F8FBB0EA7BF8A8" blockId="4.[105,787,132,1988]" pageId="4" pageNumber="593">
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One of the most conspicuous features of Venetoraptor is the presence of a raptorial-like beak.Whereas the main forces that drove edentulism in birds—one of the most studied clades of beaked animals—remain controversial,its functional spectrum is far wider than feeding and foraging
|
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<bibRefCitation id="EFF965B45C16FF9DB9B9FB1FE888FB29" author="Bright, J. A. & Marugan-Lobon, J. & Cobb, S. N. & Rayfield, E. J." box="[192,208,1247,1260]" pageId="4" pageNumber="593" pagination="5352 - 5357" refId="ref5536" refString="30. Bright, J. A., Marugan-Lobon, J., Cobb, S. N. & Rayfield, E. J. The shapes of bird beaks are highly controlled by nondietary factors. Proc. Natl Acad. Sci. USA 113, 5352 - 5357 (2016)." type="journal article" year="2016">30</bibRefCitation>
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,
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<bibRefCitation id="EFF965B45C16FF9DB9ADFB1FE8B9FB29" author="Navalon, G. & Bright, J. A. & Lobon, J. & Rayfield, E. J." box="[212,225,1247,1260]" pageId="4" pageNumber="593" pagination="422 - 435" refId="ref5592" refString="31. Navalon, G., Bright, J. A., Marugan- Lobon, J. & Rayfield, E. J. The evolutionary relationship among beak shape, mechanical advantage, and feeding ecology in modern birds. Evolution 73, 422 - 435 (2019)." type="journal article" year="2019">31</bibRefCitation>
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</superScript>
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. This structure plays roles in sexual display, vocalization and thermoregulation,among others
|
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<bibRefCitation id="EFF965B45C16FF9DB8A2FB3CE9B0FACC" author="Navalon, G. & Bright, J. A. & Lobon, J. & Rayfield, E. J." box="[475,488,1276,1289]" pageId="4" pageNumber="593" pagination="422 - 435" refId="ref5592" refString="31. Navalon, G., Bright, J. A., Marugan- Lobon, J. & Rayfield, E. J. The evolutionary relationship among beak shape, mechanical advantage, and feeding ecology in modern birds. Evolution 73, 422 - 435 (2019)." type="journal article" unsafe="true" year="2019">
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</bibRefCitation>
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. The Triassic Period witnessed several episodes of edentulism and origination of beak-like structures within Archosauromorpha
|
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<bibRefCitation id="EFF965B45C16FF9DB804FAF5E9DDFA87" author="Ezcurra, M. D." box="[381,389,1333,1346]" pageId="4" pageNumber="593" pagination="445 - 449" refId="ref4557" refString="8. Ezcurra, M. D. et al. Enigmatic dinosaur precursors bridge the gap to the origin of Pterosauria. Nature 588, 445 - 449 (2020)." type="journal article" year="2020">8</bibRefCitation>
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,
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||
<bibRefCitation id="EFF965B45C16FF9DB8F0FAF5E9CFFA87" author="Sues, H. D." box="[393,407,1333,1346]" pageId="4" pageNumber="593" pagination="635 - 649" refId="ref5645" refString="32. Sues, H. D. An unusual new archosauromorph reptile from the Upper Triassic Wolfville Formation of Nova Scotia. Can. J. Earth Sci. 40, 635 - 649 (2003)." type="journal article" year="2003">32</bibRefCitation>
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–
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<bibRefCitation id="EFF965B45C16FF9DB8E6FAF5E9F5FA87" author="Desojo, J. B." box="[415,429,1333,1346]" pageId="4" pageNumber="593" pagination="203 - 239" refId="ref5771" refString="35. Desojo, J. B. et al. Aetosauria: a clade of armoured pseudosuchians from the Upper Triassic continental beds. Geol. Soc. Spec. Publ. 379, 203 - 239 (2013)." type="journal article" year="2013">35</bibRefCitation>
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</superScript>
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(Extended Data
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<figureCitation id="135304C05C16FF9DBB2FFAF8EAC3FA88" box="[598,667,1336,1357]" captionStart="Extended" captionStartId="19.[106,194,1437,1453]" captionTargetBox="[121,1481,132,1412]" captionTargetId="figure-7@19.[121,1481,131,1412]" captionText="Extended DataFig.10 |Evolutionary tree of archosauromorphs (above) and dinosaurs (below) depicting distinct episodes of edentulism. a, Langobadisaurus pandolfii. b, Trilophosaurus buettneri. c, Teyumbaita sulcognathus.d,Aetosauroides scagliai. e, EffigiaoKeeffeae. f, Venetoraptor gassenaegen.et sp.nov.g, Seazzadactylus venieri. h, AsilisaurusKongwe." figureDoi="http://doi.org/10.5281/zenodo.8263128" httpUri="https://zenodo.org/record/8263128/files/figure.png" pageId="4" pageNumber="593">Fig. 10</figureCitation>
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). Neverthe- less, the most numerous episodes of independent beak acquisitions occurred within the avian line of Archosauria.Lagerpetids and silesaurids developed an edentulous and sharp anterior tip of the dentary
|
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<bibRefCitation id="EFF965B45C16FF9DBB81FA4BEB58FA5D" author="Ezcurra, M. D." box="[760,768,1419,1432]" pageId="4" pageNumber="593" pagination="445 - 449" refId="ref4557" refString="8. Ezcurra, M. D. et al. Enigmatic dinosaur precursors bridge the gap to the origin of Pterosauria. Nature 588, 445 - 449 (2020)." type="journal article" year="2020">8</bibRefCitation>
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,
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<bibRefCitation id="EFF965B45C16FF9DBA7AFA4BEB4AFA5D" author="Dzik, J." box="[771,786,1419,1432]" pageId="4" pageNumber="593" pagination="556 - 574" refId="ref5813" refString="36. Dzik, J. A beaked herbivorous archosaur with dinosaur affinities from the early Late Triassic of Poland. J. Vertebr. Paleontol. 23, 556 - 574 (2003)." type="journal article" year="2003">36</bibRefCitation>
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whereas at least one silesaurid lost part of the premaxillary dentition
|
||
<bibRefCitation id="EFF965B45C16FF9DBB87FA68EB54FA70" author="Nesbitt, S. J. & Langer, M. C. & Ezcurra, M. D." box="[766,780,1448,1461]" pageId="4" pageNumber="593" pagination="813 - 873" refId="ref5199" refString="22. Nesbitt, S. J., Langer, M. C. & Ezcurra, M. D. The anatomy of Asilisaurus kongwe, a dinosauriform from the Lifua Member of the Manda Beds (~ Middle Triassic) of Africa. Anat. Rec. 303, 813 - 873 (2020)." type="journal article" unsafe="true" year="2020">
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<superScript id="7C1DB50D5C16FF9DBB87FA68EB54FA70" attach="left" box="[766,780,1448,1461]" fontSize="5" pageId="4" pageNumber="593">22</superScript>
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</bibRefCitation>
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||
. However, Venetoraptor differs from all other early ornithodirans in the presence of a dorsoventrally taller,ornamented beak with pos- sible support of a rhamphotheca.Furthermore,during the Mesozoic Era distinct lineages of dinosaurs evolved a wide range of beak-like structures
|
||
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<bibRefCitation id="EFF965B45C16FF9DB9ABF9F7E886F981" author="Lautenschlager, S. & Witmer, L. M. & Altangerel, P. & Rayfield, E. J." box="[210,222,1591,1604]" pageId="4" pageNumber="593" pagination="20657 - 20662" refId="ref5849" refString="37. Lautenschlager, S., Witmer, L. M., Altangerel, P. & Rayfield, E. J. Edentulism, beaks, and biomechanical innovations in the evolution of theropod dinosaurs. Proc. Natl Acad. Sci. USA 110, 20657 - 20662 (2013)." type="journal article" year="2013">37</bibRefCitation>
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||
,
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||
<bibRefCitation id="EFF965B45C16FF9DB99BF9F7E8A9F981" author="de Souza, G. A." box="[226,241,1591,1604]" pageId="4" pageNumber="593" pagination="22281" refId="ref5905" refString="38. de Souza, G. A. et al. The first edentulous ceratosaur from South America. Sci. Rep. 11, 22281 (2021)." type="journal article" year="2021">38</bibRefCitation>
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</superScript>
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, including the bird’s beak
|
||
<bibRefCitation id="EFF965B45C16FF9DB894F9F7E9A3F981" author="Brocklehurst, N. & Field, D. J." box="[493,507,1591,1604]" pageId="4" pageNumber="593" pagination="102243" refId="ref5936" refString="39. Brocklehurst, N. & Field, D. J. Macroevolutionary dynamics of dentition in Mesozoic birds reveal no long-term selection towards tooth loss. iScience 24, 102243 (2021)." type="journal article" unsafe="true" year="2021">
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<superScript id="7C1DB50D5C16FF9DB894F9F7E9A3F981" attach="right" box="[493,507,1591,1604]" fontSize="5" pageId="4" pageNumber="593">39</superScript>
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</bibRefCitation>
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. Pterosaurs evolved distinct kinds of beak-like structures,some forms of which were entirely edentulous during the subsequent Jurassic and Cretaceous Periods
|
||
<bibRefCitation id="EFF965B45C16FF9DBBABF9B0EABAF9B8" author="Wellnhofer, P. & Kellner, A. W. A." box="[722,738,1648,1661]" pageId="4" pageNumber="593" pagination="89 - 106" refId="ref5972" refString="40. Wellnhofer, P. & Kellner, A. W. A. The skull of Tapejara wellnhoferi Kellner (Reptilia: Pterosauria) from the Lower Cretaceous Santana Formation of the Araripe Basin, Northeastern Brazil. Mitt. Bayer. Staatsslg. Palaont. Hist. Geol. 31, 89 - 106 (1991)." type="journal article" unsafe="true" year="1991">
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<superScript id="7C1DB50D5C16FF9DBBABF9B0EABAF9B8" attach="left" box="[722,738,1648,1661]" fontSize="5" pageId="4" pageNumber="593">40</superScript>
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</bibRefCitation>
|
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. The discovery of Venetoraptor expands the morphological spectrum of beaks within Pterosauromorpha,highlights the independent occurrence of this feature in the two main pterosauromorph lineages (that is,
|
||
<taxonomicName id="4C6863C65C16FF9DB9FCF926E9E3F93F" authority="AND PTEROSAURIA" authorityName="AND PTEROSAURIA" box="[133,443,1766,1787]" family="Lagerpetidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="593" phylum="Chordata" rank="family">Lagerpetidae and Pterosauria</taxonomicName>
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) and shows the first evidence of a raptorial-like beak in this clade. In birds, similar raptorial-like beaks are associated with disparate functions,such as tearing flesh and hard fruit consumption
|
||
<bibRefCitation id="EFF965B45C16FF9DB85BF8F9E977F883" author="McClure, C. J." box="[290,303,1849,1862]" pageId="4" pageNumber="593" pagination="419 - 430" refId="ref6033" refString="41. McClure, C. J. et al. Commentary: defining raptors and birds of prey. J. Raptor Res. 53, 419 - 430 (2019)." type="journal article" unsafe="true" year="2019">
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<superScript id="7C1DB50D5C16FF9DB85BF8F9E977F883" attach="right" box="[290,303,1849,1862]" fontSize="5" pageId="4" pageNumber="593">41</superScript>
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</bibRefCitation>
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. The ecological role and evolutionary advantage of such a beak in Venetoraptor are uncertain.
|
||
</paragraph>
|
||
<paragraph id="8BD718455C16FF9DB9F8F8B6ED4BFD59" blockId="4.[105,787,132,1988]" lastBlockId="4.[815,1498,132,1300]" pageId="4" pageNumber="593">
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This new species increases the ecomorphological diversity of the early ornithodiran forelimb.The hands of
|
||
<emphasis id="B91CC4575C16FF9DB887F853EA27F863" bold="true" box="[510,639,1939,1958]" italics="true" pageId="4" pageNumber="593">Venetoraptor</emphasis>
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||
are unique and highly specialized,being proportionally large and bearing scythe-like claws (
|
||
<figureCitation id="135304C05C16FF9DBA0BFF44EBEAFF5C" box="[882,946,132,153]" captionStart="Fig" captionStartId="3.[106,137,1284,1301]" captionTargetBox="[90,1523,117,1260]" captionTargetId="figure-368@3.[143,1400,138,715]" captionTargetPageId="3" captionText="Fig.3 |Morphological disparitybetween earlyornithodirans.a,f,Gnathovorax cabreirai.b,g, Scleromochlus taylori.c, Ixalerpeton polesinensis.d,i, V.gassenae. e,j, Seazzadactylusvenieri. h, D.romeri. a–e, Skull in left lateral view, reconstruction.f–j, Right forelimbinlateral view.k,l, V.gassenae gen.et sp. nov.plottedinthe ornithodiran morphospace based onthe wholeskeleton (k) and skull(l). m,n, Sumof variances of thewhole skeleton(m) andskull (n); dots aremeans,and 95% confidence intervalswere generated using thetwo tails of valuesrecovered from 9,999 bootstrap technical replicatesof a dataset comprising n = 11 (whole skeleton)and n = 12 (skull)species of Ornithodiran precursors,n = 18 (whole skeletonand skull) species of Dinosauriaand n = 10 (whole skeleton andskull)species of Pterosauria.pt,Pteroid." figureDoi="http://doi.org/10.5281/zenodo.8263106" httpUri="https://zenodo.org/record/8263106/files/figure.png" pageId="4" pageNumber="593">Fig. 3i</figureCitation>
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). Curiously, whereas other lagerpetids have a proportionally shorter metacarpal IV
|
||
<superScript id="7C1DB50D5C16FF9DBD12FF5EECDCFF6E" attach="left" box="[1131,1156,158,171]" fontSize="5" pageId="4" pageNumber="593">
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<bibRefCitation id="EFF965B45C16FF9DBD12FF5EEC2BFF6E" author="Ezcurra, M. D." box="[1131,1139,158,171]" pageId="4" pageNumber="593" pagination="445 - 449" refId="ref4557" refString="8. Ezcurra, M. D. et al. Enigmatic dinosaur precursors bridge the gap to the origin of Pterosauria. Nature 588, 445 - 449 (2020)." type="journal article" year="2020">8</bibRefCitation>
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,
|
||
<bibRefCitation id="EFF965B45C16FF9DBD0EFF5EECDCFF6E" author="Foffa, D." box="[1143,1156,158,171]" pageId="4" pageNumber="593" pagination="313 - 318" refId="ref4752" refString="12. Foffa, D. et al. Scleromochlus and the early evolution of Pterosauromorpha. Nature 610, 313 - 318 (2022)." type="journal article" year="2022">12</bibRefCitation>
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this is the longest metacarpal in
|
||
<collectionCode id="ED7980805C16FF9DBA49FF7FEB66FF17" box="[816,830,191,210]" country="Canada" lsid="urn:lsid:biocol.org:col:13946" name="Royal British Columbia Museum - Herbarium" pageId="4" pageNumber="593" type="Museum">V</collectionCode>
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. gassenae. The non-obligatory quadrupedalism is interpreted by us as one of the main forces that drove the evolution of forelimb diversity within Ornithodira during the Late Triassic, which ranged from the non-volant manus with variable number and size of digits to the pterosaur wing.This is clear when the anterior zeugopodium and autopodium morphospace area occupied by Pan-Aves (=Avemetatarsalia) is compared with the considerably smaller area occupied by other archosauriforms (Extended Data
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||
<figureCitation id="135304C05C16FF9DBD17FE46ECF7FE5E" box="[1134,1199,390,411]" captionStart="Extended" captionStartId="15.[106,194,1769,1785]" captionTargetBox="[169,721,131,1744]" captionTargetId="figure-7@15.[169,723,131,1744]" captionText="Extended DataFig.7 | Ancestral geographicareas reconstructed by the Dispersal-Extinction-Cladogenesismodel in the eucrocopodan region of the tree. Abbreviations:AR,Argentina;BR,Brazil,Uruguay,Namibia;CH,China, Thailand,Kyrgyzstan;eNA,eastern USA,Eastern Canada,Morocco andAlgeria; EU,Europe,Russia andGreenland;INT,India,Tanzania,Zambia,Madagascar, Israel andSaudi Arabia;sAF,South Africa,Lesotho,Zimbabwe;wNA,western USA,British Columbia,Mexico and Venezuela.Life reconstruction of Venetoraptor gassenae gen.et sp.nov.by CaioFantini." figureDoi="http://doi.org/10.5281/zenodo.8263122" httpUri="https://zenodo.org/record/8263122/files/figure.png" pageId="4" pageNumber="593">Fig.7e</figureCitation>
|
||
). Whereas the hindlimb of ornithodirans has been exhaustively investigated,new well-preserved and peculiar forelimbs such as that of Venetoraptor uncover an intriguing field of exploration.Such an enlarged manus bearing sharp claws with deep extensor tubers provides clues to the behaviour of this reptile. Some authors have suggested that the forelimbs of lagerpetids achieved functions other than ground-dwelling locomotion
|
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<bibRefCitation id="EFF965B45C16FF9DBC5AFDEFED73FDF9" author="Ezcurra, M. D." box="[1315,1323,559,572]" pageId="4" pageNumber="593" pagination="445 - 449" refId="ref4557" refString="8. Ezcurra, M. D. et al. Enigmatic dinosaur precursors bridge the gap to the origin of Pterosauria. Nature 588, 445 - 449 (2020)." type="journal article" unsafe="true" year="2020">
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, such as climbing and manual processing of food resources.Thus,the recurved beak and grasping hands may have been employed to deal with putative prey and/or scansoriality (that is, the ability to climb).
|
||
</paragraph>
|
||
<paragraph id="8BD718455C16FF9DBA3EFD65ECBCFAD6" blockId="4.[815,1498,132,1300]" pageId="4" pageNumber="593">
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Quantitative morphological disparity analyses performed here— including the most comprehensive dataset of Triassic Pan-Aves ever assembled—found that the body plan of pterosaurs and dinosaurs evolved as part of a broader morphological diversification of ornithodirans during the Late Triassic.The morphological disparity of the whole skeleton and the skull alone of ornithodiran precursors is signifi- cantly greater than that of Triassic dinosaurs (
|
||
<figureCitation id="135304C05C16FF9DBD97FC91ED11FCA0" box="[1262,1353,849,870]" captionStart="Fig" captionStartId="3.[106,137,1284,1301]" captionTargetBox="[90,1523,117,1260]" captionTargetId="figure-368@3.[143,1400,138,715]" captionTargetPageId="3" captionText="Fig.3 |Morphological disparitybetween earlyornithodirans.a,f,Gnathovorax cabreirai.b,g, Scleromochlus taylori.c, Ixalerpeton polesinensis.d,i, V.gassenae. e,j, Seazzadactylusvenieri. h, D.romeri. a–e, Skull in left lateral view, reconstruction.f–j, Right forelimbinlateral view.k,l, V.gassenae gen.et sp. nov.plottedinthe ornithodiran morphospace based onthe wholeskeleton (k) and skull(l). m,n, Sumof variances of thewhole skeleton(m) andskull (n); dots aremeans,and 95% confidence intervalswere generated using thetwo tails of valuesrecovered from 9,999 bootstrap technical replicatesof a dataset comprising n = 11 (whole skeleton)and n = 12 (skull)species of Ornithodiran precursors,n = 18 (whole skeletonand skull) species of Dinosauriaand n = 10 (whole skeleton andskull)species of Pterosauria.pt,Pteroid." figureDoi="http://doi.org/10.5281/zenodo.8263106" httpUri="https://zenodo.org/record/8263106/files/figure.png" pageId="4" pageNumber="593">Fig.3k–n</figureCitation>
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), resembling a previous result based on the whole skeleton
|
||
<bibRefCitation id="EFF965B45C16FF9DBDA6FCAAECB6FCB2" author="Brusatte, S. L. & Benton, M. J. & Ruta, M. & Lloyd, G. T." box="[1247,1262,874,887]" pageId="4" pageNumber="593" pagination="1485 - 1488" refId="ref6067" refString="42. Brusatte, S. L., Benton, M. J., Ruta, M. & Lloyd, G. T. Superiority, competition, and opportunism in the evolutionary radiation of dinosaurs. Science 321, 1485 - 1488 (2008)." type="journal article" unsafe="true" year="2023">
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</bibRefCitation>
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. This is partially a result of divergent morphologies present between lagerpetids and silesaurids because, when they are analysed as independent groups, their morphological disparity is more similar to that of Triassic dinosaurs (Extended Data
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||
<figureCitation id="135304C05C16FF9DBAA9FC20EC50FC30" box="[976,1032,992,1013]" captionStart="Extended" captionStartId="18.[106,194,133,149]" captionTargetId="figure-7@17.[121,1481,131,1705]" captionTargetPageId="17" captionText="Extended Data Fig. 9 | Additional results of the morphological disparity analyses. Bivariate plots using:a, Wholeskeleton;b, Skull;c, Rostrum; d, Forelimb;e, Anterior zeugopodium and autopodium;f, Hindlimb.Sum of variances:g, Whole skeleton;h, Skull;i, Rostrum;j, Forelimb;k, Anterior zeugopodium and autopodium;l, Hindlimb.In the Sum of Variances the dots are meansand the 95% confidence intervals were generated using the two tails of values recovered from 9,999 bootstrap technicalreplicates of a dataset composed of n = 5 (Whole skeleton,Skull),n = 4 (Rostrum,Forelimb),n = 3 (Anterior zeugopodium andautopodium) and n = 9 (Hindlimb) speciesof Lagerpetidae,n = 5(Whole skeleton),n = 6 (Skull,Forelimb),n = 7 (Rostrum, Hindlimb)and n = 3 (Anterior zeugopodium and autopodium) species of Silesauridae,n = 18 (Whole skeleton,Skull,Rostrum),n = 16 (Forelimb),n = 14 (Anterior zeugopodium and autopodium) and n = 19 (Hindlimb) speciesof Dinosauria,and n = 10 (Whole skeleton,Skull,Rostrum),n = 7 (Forelimb, Anterior zeugopodium and autopodium) and n = 6 (Hindlimb)species of Pterosauria.The pterosauromorph silhouette has been adapted from Kellner et al.10 (CC BY 4.0(https://creativecommons.org/licenses/by/4.0/). The silesaurid silhouette has been adapted fromMüller & Garcia43 (CC BY 4.0 (https://creativecommons.org/licenses/by/4.0/)." figureDoi="http://doi.org/10.5281/zenodo.8263126" httpUri="https://zenodo.org/record/8263126/files/figure.png" pageId="4" pageNumber="593">Fig. 9</figureCitation>
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). This reflects the wide gamut of feeding appa- ratus that evolved early in the evolutionary history of ornithodirans, ranging from the plesiomorphic ziphodont dentition to specialized beak-like structures
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<superScript id="7C1DB50D5C16FF9DBA8DFBF3EC4AFB85" attach="left" box="[1012,1042,1075,1088]" fontSize="5" pageId="4" pageNumber="593">
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<bibRefCitation id="EFF965B45C16FF9DBA8DFBF3EBA1FB85" author="Cabreira, S. F." box="[1012,1017,1075,1088]" pageId="4" pageNumber="593" pagination="3090 - 3095" refId="ref4520" refString="7. Cabreira, S. F. et al. A unique Late Triassic dinosauromorph assemblage reveals dinosaur ancestral anatomy and diet. Curr. Biol. 26, 3090 - 3095 (2016)." type="journal article" year="2016">7</bibRefCitation>
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,
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<bibRefCitation id="EFF965B45C16FF9DBA85FBF3EC5BFB85" author="Ezcurra, M. D." box="[1020,1027,1075,1088]" pageId="4" pageNumber="593" pagination="445 - 449" refId="ref4557" refString="8. Ezcurra, M. D. et al. Enigmatic dinosaur precursors bridge the gap to the origin of Pterosauria. Nature 588, 445 - 449 (2020)." type="journal article" year="2020">8</bibRefCitation>
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,
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<bibRefCitation id="EFF965B45C16FF9DBD7EFBF3EC4AFB85" author="Nesbitt, S. J." box="[1031,1042,1075,1088]" pageId="4" pageNumber="593" pagination="484 - 487" refId="ref4967" refString="17. Nesbitt, S. J. et al. The earliest bird-line archosaurs and the assembly of the dinosaur body plan. Nature 544, 484 - 487 (2017)." type="journal article" year="2017">17</bibRefCitation>
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.The high anatomical variation among Triassic ornithodiran precursors showsthat the flourishing ofTriassic avian-line archosaurs started in early-diverging forms and not after the origin of the volant pterosaurs and cursorial dinosaurs.Venetoraptor and other ornithodiran precursors are part of a previously unexpectedly rich pool of ecomorphological diversity from which were selected two of the most successful vertebrate body plans that ruled the ground and skies during the remainder of the Mesozoic.
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</paragraph>
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</subSubSection>
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</treatment>
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