385 lines
53 KiB
XML
385 lines
53 KiB
XML
<document ID-DOI="10.1038/nature07447" ID-GBIF-Dataset="3692529d-0b66-43c4-9039-f1774e260d10" ID-Zenodo-Dep="2688152" checkinTime="1557418285737" checkinUser="plazi" docAuthor="Zhang, Fucheng, Zhou, Zhonghe, Xu, Xing, Wang, Xiaolin & Sullivan, Corwin" docDate="2008" docId="7670974E8624FFADFC6A5340BC82D952" docLanguage="en" docName="nature.455.07447.pdf" docOrigin="Nature 455" docStyle="DocumentStyle{}" docTitle="Epidexipteryx hui Zhang & Zhou & Xu & Wang & Sullivan 2008, gen. et sp. nov." docType="treatment" docVersion="9" lastPageNumber="1107" masterDocId="8A49EF368624FFAFFFB6500CBF18DC71" masterDocTitle="A bizarre Jurassic maniraptoran from China with elongate ribbon-like feathers" masterLastPageNumber="1108" masterPageNumber="1105" pageNumber="1105" updateTime="1645565462435" updateUser="ExternalLinkService">
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<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:titleInfo>
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<mods:title>A bizarre Jurassic maniraptoran from China with elongate ribbon-like feathers</mods:title>
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</mods:titleInfo>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Zhang, Fucheng</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Zhou, Zhonghe</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Xu, Xing</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Wang, Xiaolin</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Sullivan, Corwin</mods:namePart>
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</mods:name>
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<mods:typeOfResource>text</mods:typeOfResource>
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<mods:relatedItem type="host">
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<mods:titleInfo>
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<mods:title>Nature</mods:title>
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</mods:titleInfo>
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<mods:part>
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<mods:date>2008</mods:date>
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<mods:detail type="pubDate">
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<mods:number>2008-10-23</mods:number>
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</mods:detail>
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<mods:detail type="volume">
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<mods:number>455</mods:number>
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</mods:detail>
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||
<mods:extent unit="page">
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<mods:start>1105</mods:start>
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<mods:end>1108</mods:end>
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||
</mods:extent>
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||
</mods:part>
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</mods:relatedItem>
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<mods:classification>journal article</mods:classification>
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<mods:identifier type="DOI">10.1038/nature07447</mods:identifier>
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<mods:identifier type="GBIF-Dataset">3692529d-0b66-43c4-9039-f1774e260d10</mods:identifier>
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<mods:identifier type="Zenodo-Dep">2688152</mods:identifier>
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</mods:mods>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.6230241" ID-GBIF-Taxon="157099809" ID-Zenodo-Dep="6230241" LSID="urn:lsid:plazi:treatment:7670974E8624FFADFC6A5340BC82D952" httpUri="http://treatment.plazi.org/id/7670974E8624FFADFC6A5340BC82D952" lastPageId="2" lastPageNumber="1107" pageId="0" pageNumber="1105">
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<subSubSection box="[988,1326,844,866]" pageId="0" pageNumber="1105" type="nomenclature">
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<paragraph blockId="0.[947,1367,700,866]" box="[988,1326,844,866]" pageId="0" pageNumber="1105">
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<taxonomicName authority="Zhang & Zhou & Xu & Wang & Sullivan, 2008" authorityName="Zhang & Zhou & Xu & Wang & Sullivan" authorityYear="2008" box="[988,1170,844,866]" class="Reptilia" family="Scansoriopterygidae" genus="Epidexipteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1105" phylum="Chordata" rank="species" status="gen. et sp. nov.">
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<emphasis bold="true" box="[988,1170,844,866]" italics="true" pageId="0" pageNumber="1105">Epidexipteryx hui</emphasis>
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</taxonomicName>
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<taxonomicNameLabel box="[1176,1326,845,866]" pageId="0" pageNumber="1105" rank="species">gen. et sp. nov.</taxonomicNameLabel>
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</paragraph>
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</subSubSection>
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||
<subSubSection lastPageId="1" lastPageNumber="1106" pageId="0" pageNumber="1105" type="etymology">
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<paragraph blockId="0.[821,1493,894,945]" pageId="0" pageNumber="1105">
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<emphasis bold="true" box="[821,937,894,915]" pageId="0" pageNumber="1105">Etymology.</emphasis>
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<emphasis box="[945,1017,894,915]" italics="true" pageId="0" pageNumber="1105">Epidexi</emphasis>
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(Greek): display;
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<emphasis box="[1199,1258,895,916]" italics="true" pageId="0" pageNumber="1105">pteryx</emphasis>
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(Greek): wing, feather;
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<emphasis box="[821,853,922,943]" italics="true" pageId="0" pageNumber="1105">hui</emphasis>
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, in honour of the late young palaeontologist Yaoming Hu, who
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</paragraph>
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<caption httpUri="https://zenodo.org/record/2688156/files/figure.png" pageId="0" pageNumber="1105" startId="0.[112,170,1868,1887]" targetBox="[158,1447,991,1846]" targetPageId="0">
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<paragraph blockId="0.[112,781,1866,1938]" lastBlockId="0.[821,1493,1868,1938]" pageId="0" pageNumber="1105">
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<emphasis bold="true" box="[112,185,1868,1887]" pageId="0" pageNumber="1105">Figure 1</emphasis>
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<emphasis bold="true" box="[191,195,1866,1888]" pageId="0" pageNumber="1105">|</emphasis>
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<taxonomicName authority="Zhang & Zhou & Xu & Wang & Sullivan, 2008" authorityName="Zhang & Zhou & Xu & Wang & Sullivan" authorityYear="2008" box="[200,355,1868,1887]" class="Reptilia" family="Scansoriopterygidae" genus="Epidexipteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1105" phylum="Chordata" rank="species" status="gen. et sp. nov.">
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<emphasis bold="true" box="[200,355,1868,1887]" italics="true" pageId="0" pageNumber="1105">Epidexipteryx hui</emphasis>
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</taxonomicName>
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<taxonomicNameLabel box="[360,503,1868,1887]" pageId="0" pageNumber="1105" rank="species">gen. et sp. nov.</taxonomicNameLabel>
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, IVPP V15471, main slab and close-up photos. a, Main slab; b, c, skull in main slab (b) and counterslab (c); d, four elongate ribbon-like tail feathers; b
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<emphasis bold="true" box="[529,536,1918,1938]" italics="true" pageId="0" pageNumber="1105">9</emphasis>
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, c
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<emphasis bold="true" box="[557,564,1918,1938]" italics="true" pageId="0" pageNumber="1105">9</emphasis>
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, line drawings of b and
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<emphasis bold="true" pageId="0" pageNumber="1105">c, respectively. Abbreviations: l1, l2 and l7, 1st, 2nd and 7th left teeth of upper jaw; l1</emphasis>
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9,
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<emphasis bold="true" box="[950,969,1894,1913]" pageId="0" pageNumber="1105">r1</emphasis>
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9
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<emphasis bold="true" box="[980,1036,1894,1913]" pageId="0" pageNumber="1105">and r5</emphasis>
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9,
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<emphasis bold="true" pageId="0" pageNumber="1105">1st left, 1st right and 5th right teeth of lower jaw; l2 and r2, 2nd left and right teeth of upper jaw.</emphasis>
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</paragraph>
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</caption>
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<footnote pageId="0" pageNumber="1105">
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<paragraph blockId="0.[112,1486,1960,2002]" pageId="0" pageNumber="1105">
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<superScript attach="right" box="[112,117,1960,1971]" fontSize="5" pageId="0" pageNumber="1105">1</superScript>
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Laboratory of Evolutionary Systematics of Vertebrates̡ Institute of Vertebrate Paleontology and Paleoanthropology̡ Chinese Academy of Sciences̡ PO Box 643̡ Beijing 100044̡ China.
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</paragraph>
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</footnote>
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<paragraph blockId="1.[93,766,133,662]" pageId="1" pageNumber="1106">contributed significantly to the study of Mesozoic mammals from China. Generic name pronounced ‘ep-id-ecks-IP-ter-icks’.</paragraph>
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</subSubSection>
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<subSubSection pageId="1" pageNumber="1106" type="materials_examined">
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<materialsCitation ID-GBIF-Occurrence="2239544483" collectionCode="IVPP" country="China" county="Ningcheng County" location="Middle Jurassic 7 to Early Cretaceous. However" pageId="1" pageNumber="1106" specimenCode="IVPP V15471" specimenCount="1" stateProvince="Inner Mongolia" typeStatus="holotype">
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<paragraph blockId="1.[93,766,133,662]" pageId="1" pageNumber="1106">
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<emphasis bold="true" box="[93,191,190,211]" pageId="1" pageNumber="1106">
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<typeStatus box="[93,187,190,211]" pageId="1" pageNumber="1106">Holotype</typeStatus>
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.
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</emphasis>
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A feathered pigeon-sized skeleton, preserved on part and counterpart slabs, and housed at the Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, collection number
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<specimenCode box="[175,306,274,297]" pageId="1" pageNumber="1106">IVPP V15471</specimenCode>
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(
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<figureCitation box="[318,373,274,297]" captionStart="Figure 1" captionStartId="0.[112,170,1868,1887]" captionTargetBox="[158,1447,991,1846]" captionTargetId="figure@0.[158,1447,991,1846]" captionTargetPageId="0" captionText="Figure 1 | Epidexipteryx hui gen. et sp. nov., IVPP V15471, main slab and close-up photos. a, Main slab; b, c, skull in main slab (b) and counterslab (c); d, four elongate ribbon-like tail feathers; b9, c9, line drawings of b and c, respectively. Abbreviations: l1, l2 and l7, 1st, 2nd and 7th left teeth of upper jaw; l19, r19 and r59, 1st left, 1st right and 5th right teeth of lower jaw; l2 and r2, 2nd left and right teeth of upper jaw" httpUri="https://zenodo.org/record/2688156/files/figure.png" pageId="1" pageNumber="1106">Fig. 1</figureCitation>
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; see Supplementary Information).
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</paragraph>
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<paragraph blockId="1.[93,766,133,662]" pageId="1" pageNumber="1106">
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<emphasis bold="true" box="[93,337,303,325]" pageId="1" pageNumber="1106">Locality and horizon.</emphasis>
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Daohugou,
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<collectingCounty box="[486,689,303,326]" pageId="1" pageNumber="1106">Ningcheng County</collectingCounty>
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,
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<collectingRegion country="China" name="Nei Mongol" pageId="1" pageNumber="1106">Inner Mongolia</collectingRegion>
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, north
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<collectingCountry box="[263,326,331,354]" name="China" pageId="1" pageNumber="1106">China</collectingCountry>
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. The age of the Daohugou sediments is contentious, with possible dates ranging from
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<location LSID="urn:lsid:plazi:treatment:7670974E8624FFADFC6A5340BC82D952:FB0670838625FFAEFDB1516BBE2BDDE8" country="China" county="Ningcheng County" name="Middle Jurassic 7 to Early Cretaceous. However" pageId="1" pageNumber="1106" stateProvince="Inner Mongolia">
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Middle Jurassic
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<bibRefCitation author="Gao ̡ K. - Q. & Ren ̡ D." box="[673,681,357,369]" journalOrPublisher="Acta Geol. Sin." pageId="1" pageNumber="1106" pagination="42 - 45" part="80" refId="ref2098" refString="7. Gao ̡ K. - Q. & Ren ̡ D. Radiometric dating of ignimbrite from Inner Mongolia provides no indication of a post-Middle Jurassic age for the Daohugou Beds. Acta Geol. Sin. 80. 42 - 45 (2006)." title="Radiometric dating of ignimbrite from Inner Mongolia provides no indication of a post-Middle Jurassic age for the Daohugou Beds" type="journal article" year="2006">
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<superScript attach="left" box="[673,681,357,369]" fontSize="5" pageId="1" pageNumber="1106">7</superScript>
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</bibRefCitation>
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to Early Cretaceous. However
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</location>
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, published radioisotopic dating results span a narrower range from 152 to 168 Myr (Middle to Late Jurassic)
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<superScript attach="none" box="[713,743,412,425]" fontSize="5" pageId="1" pageNumber="1106">
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<bibRefCitation author="Chen ̡ W." box="[713,720,412,425]" journalOrPublisher="Geol. Bull. Chin." pageId="1" pageNumber="1106" pagination="1165 - 1169" part="23" refId="ref2146" refString="8. Chen ̡ W. et al. Isotope geochronology of the fossil-bearing beds in the Daohugou area ̡ Ningcheng ̡ Inner Mongolia. Geol. Bull. Chin. 23. 1165 - 1169 (2004)." title="Isotope geochronology of the fossil-bearing beds in the Daohugou area ̡ Ningcheng ̡ Inner Mongolia" type="journal article" year="2004">8</bibRefCitation>
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–
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<bibRefCitation author="Liu ̡ Y. - X. ̡ & Liu ̡ Y. - Q & Zhang ̡ H" box="[728,743,412,425]" journalOrPublisher="Acta Geol. Sin." pageId="1" pageNumber="1106" pagination="733 - 742" part="80" refId="ref2246" refString="10. Liu ̡ Y. - X. ̡ Liu ̡ Y. - Q. & Zhang ̡ H. LA-ICPMS zircon U-Pb dating in the Jurassic Daohugou Beds and correlative strata in Ningcheng of Inner Mongolia. Acta Geol. Sin. 80. 733 - 742 (2006)." title="LA-ICPMS zircon U-Pb dating in the Jurassic Daohugou Beds and correlative strata in Ningcheng of Inner Mongolia" type="journal article" year="2006">10</bibRefCitation>
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</superScript>
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.
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</paragraph>
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</materialsCitation>
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||
</subSubSection>
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||
<subSubSection lastPageId="2" lastPageNumber="1107" pageId="1" pageNumber="1106" type="diagnosis">
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<paragraph blockId="1.[93,766,133,662]" lastBlockId="1.[802,1475,133,663]" pageId="1" pageNumber="1106">
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<emphasis bold="true" box="[93,199,444,466]" pageId="1" pageNumber="1106">Diagnosis.</emphasis>
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Medium-sized avialan with four elongate ribbon-like tail feathers (ETFs), highly procumbent and significantly enlarged anterior teeth, and a distally tapering pygostyle-like structure formed by ten unfused caudals at the end of the tail (
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<figureCitation box="[456,514,528,551]" captionStart="Figure 1" captionStartId="0.[112,170,1868,1887]" captionTargetBox="[158,1447,991,1846]" captionTargetId="figure@0.[158,1447,991,1846]" captionTargetPageId="0" captionText="Figure 1 | Epidexipteryx hui gen. et sp. nov., IVPP V15471, main slab and close-up photos. a, Main slab; b, c, skull in main slab (b) and counterslab (c); d, four elongate ribbon-like tail feathers; b9, c9, line drawings of b and c, respectively. Abbreviations: l1, l2 and l7, 1st, 2nd and 7th left teeth of upper jaw; l19, r19 and r59, 1st left, 1st right and 5th right teeth of lower jaw; l2 and r2, 2nd left and right teeth of upper jaw" httpUri="https://zenodo.org/record/2688156/files/figure.png" pageId="1" pageNumber="1106">Figs 1</figureCitation>
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and
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<figureCitation box="[561,575,528,549]" captionStart="Figure 2" captionStartId="1.[93,151,1805,1824]" captionTargetId="figure@1.[271,1293,714,1784]" captionTargetPageId="1" captionText="Figure 2 | Line drawings and close-up photographs of Epidexipteryx hui gen. et sp. nov. a, Skeleton and feather outline, based on both main slab and counterslab, showing that each shafted feather is formed by central rachis and two unbranched vanes (a9). b, d, d9, Proximal regions of non-shafted feathers; barbs are parallel and closely united as an unbranched membranous structure (d, d9), vanes are either layered, indicated by white arrows (b), or arranged irregularly (c). The red arrow indicates the proximal end of the feather (d). e, e9, f, Distal regions of non-shafted feathers, in which barbs appear loosely parallel. Abbreviations: as1ca, astragalus and calcaneum; ce, cervical vertebrae; co, coracoid; cv, caudal vertebrae; dt314, distal tarsals 3 and 4; f, frontal; fe, femur; fi, fibula; ga, gastralia; hu, humerus; il, ilium; is, ischium; ma, mandible; mtI–IV, metatarsals I–IV; p, phalanges or metacarpals; pa, parietal; pm, premaxilla; pu, pubis; q, quadrate; r, ribs; ra, radius; rc, rachis; sc, scapula; st, sternum; ti, tibia; tv, thoracic vertebrae; u, ungual phalanges; ul, ulna; uv, unbranched vane" httpUri="https://zenodo.org/record/2688335/files/figure.png" pageId="1" pageNumber="1106">2</figureCitation>
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; see Supplementary Information). Differs significantly from
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<taxonomicName box="[505,662,556,577]" class="Reptilia" family="Scansoriopterygidae" genus="Epidendrosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="1107" phylum="Chordata" rank="genus">
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<emphasis box="[505,662,556,577]" italics="true" pageId="1" pageNumber="1106">Epidendrosaurus</emphasis>
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</taxonomicName>
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, the only other known scansoriopterygid, in caudal morphology (tail 70% of trunk length in
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<taxonomicName box="[258,386,612,633]" class="Reptilia" family="Scansoriopterygidae" genus="Epidexipteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="1106" phylum="Chordata" rank="genus">
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<emphasis box="[258,386,612,633]" italics="true" pageId="1" pageNumber="1106">Epidexipteryx</emphasis>
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</taxonomicName>
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, compared with more than 300% in
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<taxonomicName box="[93,250,640,661]" class="Reptilia" family="Scansoriopterygidae" genus="Epidendrosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="1107" phylum="Chordata" rank="genus">
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<emphasis box="[93,250,640,661]" italics="true" pageId="1" pageNumber="1106">Epidendrosaurus</emphasis>
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</taxonomicName>
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; 16 caudal vertebrae in
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<taxonomicName box="[479,607,640,661]" class="Reptilia" family="Scansoriopterygidae" genus="Epidexipteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="1106" phylum="Chordata" rank="genus">
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||
<emphasis box="[479,607,640,661]" italics="true" pageId="1" pageNumber="1106">Epidexipteryx</emphasis>
|
||
</taxonomicName>
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||
, compared with more than 40 in
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<taxonomicName box="[964,1121,133,154]" class="Reptilia" family="Scansoriopterygidae" genus="Epidendrosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="1107" phylum="Chordata" rank="genus">
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||
<emphasis box="[964,1121,133,154]" italics="true" pageId="1" pageNumber="1106">Epidendrosaurus</emphasis>
|
||
</taxonomicName>
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||
; caudal prezygapophyses reduced in
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<taxonomicName box="[802,930,161,182]" class="Reptilia" family="Scansoriopterygidae" genus="Epidexipteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="1106" phylum="Chordata" rank="genus">
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||
<emphasis box="[802,930,161,182]" italics="true" pageId="1" pageNumber="1106">Epidexipteryx</emphasis>
|
||
</taxonomicName>
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but significantly elongated in
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<taxonomicName box="[1225,1382,161,182]" class="Reptilia" family="Scansoriopterygidae" genus="Epidendrosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="1107" phylum="Chordata" rank="genus">
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||
<emphasis box="[1225,1382,161,182]" italics="true" pageId="1" pageNumber="1106">Epidendrosaurus</emphasis>
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||
</taxonomicName>
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||
).
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</paragraph>
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<paragraph blockId="1.[802,1475,133,663]" pageId="1" pageNumber="1106">
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<emphasis bold="true" box="[802,1110,190,212]" pageId="1" pageNumber="1106">Description and comparisons.</emphasis>
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<taxonomicName box="[1115,1246,190,211]" class="Reptilia" family="Scansoriopterygidae" genus="Epidexipteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="1106" phylum="Chordata" rank="genus">
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||
<emphasis box="[1115,1246,190,211]" italics="true" pageId="1" pageNumber="1106">Epidexipteryx</emphasis>
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</taxonomicName>
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is estimated to be 164 g in body mass (see Supplementary Information), smaller than most other basal avialans
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<superScript attach="left" box="[995,1021,244,257]" fontSize="5" pageId="1" pageNumber="1106">3,11</superScript>
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. The holotype probably represents a subadult individual, because the ends of some of the long bones seem imperfectly ossified.
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</paragraph>
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<paragraph blockId="1.[802,1475,133,663]" pageId="1" pageNumber="1106">
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As in oviraptorosaurs
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<bibRefCitation author="Xu ̡ X." box="[1047,1062,329,342]" journalOrPublisher="Nature" pageId="1" pageNumber="1106" pagination="291 - 293" part="419" refId="ref2332" refString="12. Xu ̡ X. et al. An unusual oviraptorosaurian dinosaur from China. Nature 419. 291 - 293 (2002)." title="An unusual oviraptorosaurian dinosaur from China" type="journal article" year="2002">
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||
<superScript attach="left" box="[1047,1062,329,342]" fontSize="5" pageId="1" pageNumber="1106">12</superScript>
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</bibRefCitation>
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||
, the skull is high in lateral view (height about 60% of length), the external naris is positioned high on the snout, and the parietal is proportionally long. The teeth of both the upper and lower jaws are highly procumbent, a feature previously known only in the ceratosaurian
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<emphasis box="[1127,1272,444,465]" italics="true" pageId="1" pageNumber="1106">Masiakasaurus</emphasis>
|
||
among theropods
|
||
<bibRefCitation author="Sampson ̡ S. D. ̡ & Carrano ̡ M. T. & Forster ̡ C. A." box="[1453,1468,441,454]" journalOrPublisher="Nature" pageId="1" pageNumber="1106" pagination="504 - 505" part="409" refId="ref2358" refString="13. Sampson ̡ S. D. ̡ Carrano ̡ M. T. & Forster ̡ C. A. A bizarre predatory dinosaur from Madagascar: implications for the evolution of Gondwanan theropods. Nature 409. 504 - 505 (2001)." title="bizarre predatory dinosaur from Madagascar: implications for the evolution of Gondwanan theropods" type="journal article" year="2001">
|
||
<superScript attach="left" box="[1453,1468,441,454]" fontSize="5" pageId="1" pageNumber="1106">13</superScript>
|
||
</bibRefCitation>
|
||
. Furthermore, the anterior teeth are much larger than the posterior ones, as in basal oviraptorosaurs
|
||
<bibRefCitation author="Xu ̡ X." box="[1133,1148,497,510]" journalOrPublisher="Nature" pageId="1" pageNumber="1106" pagination="291 - 293" part="419" refId="ref2332" refString="12. Xu ̡ X. et al. An unusual oviraptorosaurian dinosaur from China. Nature 419. 291 - 293 (2002)." title="An unusual oviraptorosaurian dinosaur from China" type="journal article" year="2002">
|
||
<superScript attach="left" box="[1133,1148,497,510]" fontSize="5" pageId="1" pageNumber="1106">12</superScript>
|
||
</bibRefCitation>
|
||
, basal therizinosaurs
|
||
<bibRefCitation author="Kirkland ̡ J. I." box="[1361,1376,497,510]" journalOrPublisher="Nature" pageId="1" pageNumber="1106" pagination="84 - 87" part="435" refId="ref2405" refString="14. Kirkland ̡ J. I. et al. A primitive therizinosauroid dinosaur from the Early Cretaceous of Utah. Nature 435. 84 - 87 (2005)." title="A primitive therizinosauroid dinosaur from the Early Cretaceous of Utah" type="journal article" year="2005">
|
||
<superScript attach="left" box="[1361,1376,497,510]" fontSize="5" pageId="1" pageNumber="1106">14</superScript>
|
||
</bibRefCitation>
|
||
and probably
|
||
<taxonomicName box="[868,1029,528,549]" class="Reptilia" family="Scansoriopterygidae" genus="Epidendrosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="1107" phylum="Chordata" rank="genus">
|
||
<emphasis box="[868,1029,528,549]" italics="true" pageId="1" pageNumber="1106">Epidendrosaurus</emphasis>
|
||
</taxonomicName>
|
||
<superScript attach="left" box="[1029,1048,525,538]" fontSize="5" pageId="1" pageNumber="1106">4,5</superScript>
|
||
. There are probably 9 cervicals and 14 thoracic vertebrae, and the synsacrum is composed of 7 vertebrae. The caudal series is much shorter than in non-avian theropods or in other basal avialans. The anterior six caudal vertebrae are proportionally short and wide. The posterior ten caudals bear no transverse
|
||
</paragraph>
|
||
<subSection pageId="1" pageNumber="1106" type="multiple">
|
||
<caption httpUri="https://zenodo.org/record/2688335/files/figure.png" pageId="1" pageNumber="1106" subCaptionStartIDs="1.[1069,1197,1805,1824]" subCaptionStarts="Abbr" targetBox="[272,1292,715,1781]" targetPageId="1">
|
||
<paragraph blockId="1.[93,765,1803,2002]" lastBlockId="1.[802,1475,1804,1976]" pageId="1" pageNumber="1106">
|
||
<figureCitation box="[93,169,1805,1824]" captionStart="Figure 2" captionStartId="1.[93,151,1805,1824]" captionTargetId="figure@1.[271,1293,714,1784]" captionTargetPageId="1" captionText="Figure 2 | Line drawings and close-up photographs of Epidexipteryx hui gen. et sp. nov. a, Skeleton and feather outline, based on both main slab and counterslab, showing that each shafted feather is formed by central rachis and two unbranched vanes (a9). b, d, d9, Proximal regions of non-shafted feathers; barbs are parallel and closely united as an unbranched membranous structure (d, d9), vanes are either layered, indicated by white arrows (b), or arranged irregularly (c). The red arrow indicates the proximal end of the feather (d). e, e9, f, Distal regions of non-shafted feathers, in which barbs appear loosely parallel. Abbreviations: as1ca, astragalus and calcaneum; ce, cervical vertebrae; co, coracoid; cv, caudal vertebrae; dt314, distal tarsals 3 and 4; f, frontal; fe, femur; fi, fibula; ga, gastralia; hu, humerus; il, ilium; is, ischium; ma, mandible; mtI–IV, metatarsals I–IV; p, phalanges or metacarpals; pa, parietal; pm, premaxilla; pu, pubis; q, quadrate; r, ribs; ra, radius; rc, rachis; sc, scapula; st, sternum; ti, tibia; tv, thoracic vertebrae; u, ungual phalanges; ul, ulna; uv, unbranched vane" httpUri="https://zenodo.org/record/2688335/files/figure.png" pageId="1" pageNumber="1106">
|
||
<emphasis bold="true" box="[93,169,1805,1824]" pageId="1" pageNumber="1106">Figure 2</emphasis>
|
||
</figureCitation>
|
||
<emphasis bold="true" box="[175,179,1803,1825]" pageId="1" pageNumber="1106">|</emphasis>
|
||
Line drawings and close-up photographs of
|
||
<taxonomicName authority="Zhang & Zhou & Xu & Wang & Sullivan, 2008" authorityName="Zhang & Zhou & Xu & Wang & Sullivan" authorityYear="2008" box="[585,740,1805,1824]" class="Reptilia" family="Scansoriopterygidae" genus="Epidexipteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="1106" phylum="Chordata" rank="species" status="gen. et sp. nov.">
|
||
<emphasis bold="true" box="[585,740,1805,1824]" italics="true" pageId="1" pageNumber="1106">Epidexipteryx hui</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel box="[93,229,1830,1849]" pageId="1" pageNumber="1106" rank="species">gen. et sp. nov.</taxonomicNameLabel>
|
||
a, Skeleton and feather outline, based on both main slab and counterslab, showing that each shafted feather is formed by central rachis and two unbranched vanes (a
|
||
<emphasis bold="true" box="[360,367,1881,1901]" italics="true" pageId="1" pageNumber="1106">9</emphasis>
|
||
). b, d, d
|
||
<emphasis bold="true" box="[442,449,1881,1901]" italics="true" pageId="1" pageNumber="1106">9</emphasis>
|
||
, Proximal regions of non-shafted feathers; barbs are parallel and closely united as an unbranched membranous structure (d, d
|
||
<emphasis bold="true" box="[347,354,1931,1951]" italics="true" pageId="1" pageNumber="1106">9</emphasis>
|
||
), vanes are either layered, indicated by white arrows (b), or arranged irregularly (c). The red arrow indicates the proximal end of the feather (d). e, e
|
||
<emphasis bold="true" box="[317,324,1982,2002]" italics="true" pageId="1" pageNumber="1106">9</emphasis>
|
||
, f, Distal regions of non-shafted feathers, in which
|
||
<emphasis bold="true" box="[802,1224,1805,1825]" pageId="1" pageNumber="1106">barbs appear loosely parallel. Abbreviations: as</emphasis>
|
||
1
|
||
<emphasis bold="true" pageId="1" pageNumber="1106">ca, astragalus and calcaneum; ce, cervical vertebrae; co, coracoid; cv, caudal vertebrae; dt3</emphasis>
|
||
1
|
||
<emphasis bold="true" pageId="1" pageNumber="1106">4, distal tarsals 3 and 4; f, frontal; fe, femur; fi, fibula; ga, gastralia; hu, humerus; il, ilium; is, ischium; ma, mandible; mtI–IV, metatarsals I–IV; p, phalanges or metacarpals; pa, parietal; pm, premaxilla; pu, pubis; q, quadrate; r, ribs; ra, radius; rc, rachis; sc, scapula; st, sternum; ti, tibia; tv, thoracic vertebrae; u, ungual phalanges; ul, ulna; uv, unbranched vane.</emphasis>
|
||
</paragraph>
|
||
</caption>
|
||
</subSection>
|
||
<paragraph blockId="2.[112,785,133,1181]" pageId="2" pageNumber="1107">
|
||
processes. They form a structure similar to the elongated, incipient pygostyle in some basal birds
|
||
<superScript attach="left" box="[414,448,159,172]" fontSize="5" pageId="2" pageNumber="1107">15,16</superScript>
|
||
but are not fused to each other (
|
||
<figureCitation box="[118,189,190,213]" captionStart="Figure 1" captionStartId="0.[112,170,1868,1887]" captionTargetBox="[158,1447,991,1846]" captionTargetId="figure@0.[158,1447,991,1846]" captionTargetPageId="0" captionText="Figure 1 | Epidexipteryx hui gen. et sp. nov., IVPP V15471, main slab and close-up photos. a, Main slab; b, c, skull in main slab (b) and counterslab (c); d, four elongate ribbon-like tail feathers; b9, c9, line drawings of b and c, respectively. Abbreviations: l1, l2 and l7, 1st, 2nd and 7th left teeth of upper jaw; l19, r19 and r59, 1st left, 1st right and 5th right teeth of lower jaw; l2 and r2, 2nd left and right teeth of upper jaw" httpUri="https://zenodo.org/record/2688156/files/figure.png" pageId="2" pageNumber="1107">Figs 1a</figureCitation>
|
||
and
|
||
<figureCitation box="[239,264,190,211]" captionStart="Figure 2" captionStartId="1.[93,151,1805,1824]" captionTargetId="figure@1.[271,1293,714,1784]" captionTargetPageId="1" captionText="Figure 2 | Line drawings and close-up photographs of Epidexipteryx hui gen. et sp. nov. a, Skeleton and feather outline, based on both main slab and counterslab, showing that each shafted feather is formed by central rachis and two unbranched vanes (a9). b, d, d9, Proximal regions of non-shafted feathers; barbs are parallel and closely united as an unbranched membranous structure (d, d9), vanes are either layered, indicated by white arrows (b), or arranged irregularly (c). The red arrow indicates the proximal end of the feather (d). e, e9, f, Distal regions of non-shafted feathers, in which barbs appear loosely parallel. Abbreviations: as1ca, astragalus and calcaneum; ce, cervical vertebrae; co, coracoid; cv, caudal vertebrae; dt314, distal tarsals 3 and 4; f, frontal; fe, femur; fi, fibula; ga, gastralia; hu, humerus; il, ilium; is, ischium; ma, mandible; mtI–IV, metatarsals I–IV; p, phalanges or metacarpals; pa, parietal; pm, premaxilla; pu, pubis; q, quadrate; r, ribs; ra, radius; rc, rachis; sc, scapula; st, sternum; ti, tibia; tv, thoracic vertebrae; u, ungual phalanges; ul, ulna; uv, unbranched vane" httpUri="https://zenodo.org/record/2688335/files/figure.png" pageId="2" pageNumber="1107">2a</figureCitation>
|
||
; see Supplementary Information).
|
||
</paragraph>
|
||
<paragraph blockId="2.[112,785,133,1181]" pageId="2" pageNumber="1107">
|
||
The partly preserved sternum is small and convex anteriorly, and seems to comprise two incompletely fused sternal plates, as in the primitive bird
|
||
<emphasis box="[260,353,276,297]" italics="true" pageId="2" pageNumber="1107">Jeholornis</emphasis>
|
||
<bibRefCitation author="Zhou ̡ Z. & Zhang ̡ F." box="[353,368,273,286]" journalOrPublisher="Nature" pageId="2" pageNumber="1107" pagination="405 - 409" part="418" refId="ref2535" refString="17. Zhou ̡ Z. & Zhang ̡ F. A long-tailed ̡ seed-eating bird from the Early Cretaceous of China. Nature 418. 405 - 409 (2002)." title="long-tailed ̡ seed-eating bird from the Early Cretaceous of China" type="journal article" year="2002">
|
||
<superScript attach="left" box="[353,368,273,286]" fontSize="5" pageId="2" pageNumber="1107">17</superScript>
|
||
</bibRefCitation>
|
||
. The scapula is significantly shorter than the humerus, as in some derived maniraptorans, and the coracoid is sub-rectangular. The pelvis has an unexpected combination of features among theropods, as in
|
||
<taxonomicName box="[459,618,361,382]" class="Reptilia" family="Scansoriopterygidae" genus="Epidendrosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="1107" phylum="Chordata" rank="genus">
|
||
<emphasis box="[459,618,361,382]" italics="true" pageId="2" pageNumber="1107">Epidendrosaurus</emphasis>
|
||
</taxonomicName>
|
||
<superScript attach="left" box="[618,637,359,372]" fontSize="5" pageId="2" pageNumber="1107">4,5</superScript>
|
||
. The ilium is bird-like in having a long preacetabular process with a strongly convex anterior margin, but differs from the ilia of most non-avian theropods in having a small pubic peduncle. Unusually among theropods, the straight pubis is shorter than the ischium and is significantly shorter than the femur. It is anteroventrally oriented and lacks a pubic boot. The posteriorly curved ischium is laterally compressed, gradually widens towards the distal end, and lacks an obturator process. The humerus is about same length as the femur, proportionally longer than in most other basal avialans. The ulna is posteriorly bowed and the manus is significantly elongated (see Supplementary Information) as in birds and other derived theropods
|
||
<superScript attach="left" box="[519,557,671,684]" fontSize="5" pageId="2" pageNumber="1107">
|
||
<bibRefCitation author="Zhang ̡ F. - C. ̡ & Zhou ̡ Z. - H. & Hou ̡ L. - H" box="[519,533,671,684]" editor="Chang ̡ M. - M. ̡" journalOrPublisher="Shanghai Sci. Technol. Publ" pageId="2" pageNumber="1107" pagination="129 - 149" refId="ref2568" refString="18. Zhang ̡ F. - C. ̡ Zhou ̡ Z. - H. & Hou ̡ L. - H. in The Jehol Biota (eds Chang ̡ M. - M. ̡ Chen ̡ P. - J. ̡ Wang ̡ Y. - Q. & Wang ̡ Y.) 129 - 149 (Shanghai Sci. Technol. Publ. ̡ 2003)." type="book chapter" volumeTitle="The Jehol Biota" year="2003">18</bibRefCitation>
|
||
–
|
||
<bibRefCitation author="Xu ̡ X." box="[541,557,671,684]" editor="Rong ̡ J. - Y." journalOrPublisher="Sci. Press ̡ Beijing" pageId="2" pageNumber="1107" pagination="627 - 642" refId="ref2762" refString="22. Xu ̡ X. in Originationsi Radiations and Biodiversity Changes (eds Rong ̡ J. - Y. et al.) 627 - 642; 927 - 930 (Sci. Press ̡ Beijing ̡ 2006)." type="book chapter" volumeTitle="Originationsi Radiations and Biodiversity Changes" year="2006">22</bibRefCitation>
|
||
</superScript>
|
||
. The curvature of the manual claws falls within the range known for the pedal claws of ground-based foraging birds (see Supplementary Information). The femur is about 160% the length of the metatarsus, and 80% the length of the tibia (
|
||
<figureCitation box="[231,301,787,810]" captionStart="Figure 1" captionStartId="0.[112,170,1868,1887]" captionTargetBox="[158,1447,991,1846]" captionTargetId="figure@0.[158,1447,991,1846]" captionTargetPageId="0" captionText="Figure 1 | Epidexipteryx hui gen. et sp. nov., IVPP V15471, main slab and close-up photos. a, Main slab; b, c, skull in main slab (b) and counterslab (c); d, four elongate ribbon-like tail feathers; b9, c9, line drawings of b and c, respectively. Abbreviations: l1, l2 and l7, 1st, 2nd and 7th left teeth of upper jaw; l19, r19 and r59, 1st left, 1st right and 5th right teeth of lower jaw; l2 and r2, 2nd left and right teeth of upper jaw" httpUri="https://zenodo.org/record/2688156/files/figure.png" pageId="2" pageNumber="1107">Figs 1a</figureCitation>
|
||
and
|
||
<figureCitation box="[351,375,788,809]" captionStart="Figure 2" captionStartId="1.[93,151,1805,1824]" captionTargetId="figure@1.[271,1293,714,1784]" captionTargetPageId="1" captionText="Figure 2 | Line drawings and close-up photographs of Epidexipteryx hui gen. et sp. nov. a, Skeleton and feather outline, based on both main slab and counterslab, showing that each shafted feather is formed by central rachis and two unbranched vanes (a9). b, d, d9, Proximal regions of non-shafted feathers; barbs are parallel and closely united as an unbranched membranous structure (d, d9), vanes are either layered, indicated by white arrows (b), or arranged irregularly (c). The red arrow indicates the proximal end of the feather (d). e, e9, f, Distal regions of non-shafted feathers, in which barbs appear loosely parallel. Abbreviations: as1ca, astragalus and calcaneum; ce, cervical vertebrae; co, coracoid; cv, caudal vertebrae; dt314, distal tarsals 3 and 4; f, frontal; fe, femur; fi, fibula; ga, gastralia; hu, humerus; il, ilium; is, ischium; ma, mandible; mtI–IV, metatarsals I–IV; p, phalanges or metacarpals; pa, parietal; pm, premaxilla; pu, pubis; q, quadrate; r, ribs; ra, radius; rc, rachis; sc, scapula; st, sternum; ti, tibia; tv, thoracic vertebrae; u, ungual phalanges; ul, ulna; uv, unbranched vane" httpUri="https://zenodo.org/record/2688335/files/figure.png" pageId="2" pageNumber="1107">2a</figureCitation>
|
||
; see Supplementary Information).
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="2" pageNumber="1107" type="discussion">
|
||
<paragraph blockId="2.[112,785,133,1181]" pageId="2" pageNumber="1107">
|
||
<emphasis bold="true" box="[112,377,816,838]" pageId="2" pageNumber="1107">Phylogeny and affinity.</emphasis>
|
||
Phylogenetic analysis indicates that
|
||
<taxonomicName box="[112,243,845,866]" class="Reptilia" family="Scansoriopterygidae" genus="Epidexipteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="1107" phylum="Chordata" rank="genus">
|
||
<emphasis box="[112,243,845,866]" italics="true" pageId="2" pageNumber="1107">Epidexipteryx</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName box="[338,499,845,866]" class="Reptilia" family="Scansoriopterygidae" genus="Epidendrosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="1107" phylum="Chordata" rank="genus">
|
||
<emphasis box="[338,499,845,866]" italics="true" pageId="2" pageNumber="1107">Epidendrosaurus</emphasis>
|
||
</taxonomicName>
|
||
form a monophyletic
|
||
<taxonomicName authorityName="Czerkas et Yuan" authorityYear="2002" box="[112,312,873,896]" class="Reptilia" family="Scansoriopterygidae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="1107" phylum="Chordata" rank="family">Scansoriopterygidae</taxonomicName>
|
||
(see Supplementary Information), representing a bizarre lineage at the base of the Avialae
|
||
<bibRefCitation author="Padian ̡ K." box="[514,522,899,912]" editor="Weishampel ̡ D. B" journalOrPublisher="Univ. of California Press ̡ Berkeley" pageId="2" pageNumber="1107" pagination="210 - 231" refId="ref2052" refString="6. Padian ̡ K. in The Dinosauria 2 nd edn (eds Weishampel ̡ D. B. ̡ Dodson ̡ P. ̡ Osmolska ̡ H.) 210 - 231 (Univ. of California Press ̡ Berkeley ̡ 2004)." title="The Dinosauria 2 nd edn" type="book chapter" year="2004">
|
||
<superScript attach="left" box="[514,522,899,912]" fontSize="5" pageId="2" pageNumber="1107">6</superScript>
|
||
</bibRefCitation>
|
||
(
|
||
<figureCitation box="[533,589,901,924]" captionStart="Figure 3" captionStartId="2.[112,170,1806,1825]" captionTargetBox="[214,679,1241,1784]" captionTargetId="graphics@2.[214,515,1246,1775]" captionTargetPageId="2" captionText="Figure 3 | Phylogenetic relationships of Epidexipteryx hui gen. et sp. nov. The cladogram is simplified from the strict consensus of nine most parsimonious trees (tree length 1,255; consistency index 0.35; retention index 0.75; see Supplementary Information). Scansoriopterygidae is defined as the least inclusive clade including Epidendrosaurus and Epidexipteryx, Avialae as the most inclusive clade including Vultur gryphus but not Deinonychus antirrhopus, and Aves as the least inclusive clade including Archaeopteryx and Vultur gryphus." httpUri="https://zenodo.org/record/2688162/files/figure.png" pageId="2" pageNumber="1107">Fig. 3</figureCitation>
|
||
; see Supplementary Information).
|
||
</paragraph>
|
||
<paragraph blockId="2.[112,785,133,1181]" lastBlockId="2.[821,1493,133,1315]" pageId="2" pageNumber="1107">
|
||
Although possessing many derived features seen in birds, including a humerus as long as the femur, a long preacetabular process of the ilium with a strongly convex margin, and many other features,
|
||
<taxonomicName box="[112,243,1044,1065]" class="Reptilia" family="Scansoriopterygidae" genus="Epidexipteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="1107" phylum="Chordata" rank="genus">
|
||
<emphasis box="[112,243,1044,1065]" italics="true" pageId="2" pageNumber="1107">Epidexipteryx</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName box="[294,455,1044,1065]" class="Reptilia" family="Scansoriopterygidae" genus="Epidendrosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="1107" phylum="Chordata" rank="genus">
|
||
<emphasis box="[294,455,1044,1065]" italics="true" pageId="2" pageNumber="1107">Epidendrosaurus</emphasis>
|
||
</taxonomicName>
|
||
also show some striking similarities to oviraptorosaurs
|
||
<bibRefCitation author="Xu ̡ X." box="[339,354,1070,1083]" journalOrPublisher="Nature" pageId="2" pageNumber="1107" pagination="291 - 293" part="419" refId="ref2332" refString="12. Xu ̡ X. et al. An unusual oviraptorosaurian dinosaur from China. Nature 419. 291 - 293 (2002)." title="An unusual oviraptorosaurian dinosaur from China" type="journal article" year="2002">
|
||
<superScript attach="left" box="[339,354,1070,1083]" fontSize="5" pageId="2" pageNumber="1107">12</superScript>
|
||
</bibRefCitation>
|
||
and to a lesser degree therizinosauroids
|
||
<superScript attach="left" box="[744,778,1070,1083]" fontSize="5" pageId="2" pageNumber="1107">14,21</superScript>
|
||
, including a short and high skull, an external naris positioned high on the snout, an anteriorly downturned and strongly dorsally convex mandible, a large external mandibular fenestra, and enlarged anterior teeth. Furthermore, some pelvic features, such as a proportionally very short pubis and a distally widening ischium, are not known in any other theropod. The bizarre appearance of scansoriopterygids indicates that morphological disparity among maniraptorans close to the origin of birds is higher than has previously been assumed, and underscores the importance of Jurassic theropods for understanding avian origins.
|
||
</paragraph>
|
||
<paragraph blockId="2.[821,1493,133,1315]" pageId="2" pageNumber="1107">
|
||
<emphasis bold="true" box="[821,948,332,353]" pageId="2" pageNumber="1107">Integument.</emphasis>
|
||
The integument bears feather-like structures of two types, ETFs and non-ETFs, which are roughly comparable to shafted and non-shafted feathers, respectively
|
||
<bibRefCitation author="Zhang ̡ F. - C. & Zhou ̡ Z. - H." box="[1193,1208,385,398]" editor="Rong ̡ J. - Y." journalOrPublisher="Sci. Press ̡ Beijing" pageId="2" pageNumber="1107" pagination="611 - 625" refId="ref2804" refString="23. Zhang ̡ F. - C. & Zhou ̡ Z. - H. in Originationsi Radiations and Biodiversity Changes (eds Rong ̡ J. - Y. e t al.) 611 - 625; 923 - 925 (Sci. Press ̡ Beijing ̡ 2006)." type="book chapter" volumeTitle="in Originationsi Radiations and Biodiversity Changes" year="2006">
|
||
<superScript attach="left" box="[1193,1208,385,398]" fontSize="5" pageId="2" pageNumber="1107">23</superScript>
|
||
</bibRefCitation>
|
||
. The distal part of each non- ETF is composed of filamentous parallel barbs (
|
||
<figureCitation box="[1289,1377,416,439]" captionStart="Figure 2" captionStartId="1.[93,151,1805,1824]" captionTargetId="figure@1.[271,1293,714,1784]" captionTargetPageId="1" captionText="Figure 2 | Line drawings and close-up photographs of Epidexipteryx hui gen. et sp. nov. a, Skeleton and feather outline, based on both main slab and counterslab, showing that each shafted feather is formed by central rachis and two unbranched vanes (a9). b, d, d9, Proximal regions of non-shafted feathers; barbs are parallel and closely united as an unbranched membranous structure (d, d9), vanes are either layered, indicated by white arrows (b), or arranged irregularly (c). The red arrow indicates the proximal end of the feather (d). e, e9, f, Distal regions of non-shafted feathers, in which barbs appear loosely parallel. Abbreviations: as1ca, astragalus and calcaneum; ce, cervical vertebrae; co, coracoid; cv, caudal vertebrae; dt314, distal tarsals 3 and 4; f, frontal; fe, femur; fi, fibula; ga, gastralia; hu, humerus; il, ilium; is, ischium; ma, mandible; mtI–IV, metatarsals I–IV; p, phalanges or metacarpals; pa, parietal; pm, premaxilla; pu, pubis; q, quadrate; r, ribs; ra, radius; rc, rachis; sc, scapula; st, sternum; ti, tibia; tv, thoracic vertebrae; u, ungual phalanges; ul, ulna; uv, unbranched vane" httpUri="https://zenodo.org/record/2688335/files/figure.png" pageId="2" pageNumber="1107">Fig. 2e, e</figureCitation>
|
||
9, f), similar to the condition seen in the non-shafted feathers of other feathered dinosaurs and primitive birds
|
||
<superScript attach="left" box="[1111,1169,470,483]" fontSize="5" pageId="2" pageNumber="1107">18,22–25</superScript>
|
||
. However, the free distal barbs of
|
||
<taxonomicName box="[821,952,501,522]" class="Reptilia" family="Scansoriopterygidae" genus="Epidexipteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="1107" phylum="Chordata" rank="genus">
|
||
<emphasis box="[821,952,501,522]" italics="true" pageId="2" pageNumber="1107">Epidexipteryx</emphasis>
|
||
</taxonomicName>
|
||
arise from the edge of a membranous structure (
|
||
<figureCitation captionStart="Figure 2" captionStartId="1.[93,151,1805,1824]" captionTargetId="figure@1.[271,1293,714,1784]" captionTargetPageId="1" captionText="Figure 2 | Line drawings and close-up photographs of Epidexipteryx hui gen. et sp. nov. a, Skeleton and feather outline, based on both main slab and counterslab, showing that each shafted feather is formed by central rachis and two unbranched vanes (a9). b, d, d9, Proximal regions of non-shafted feathers; barbs are parallel and closely united as an unbranched membranous structure (d, d9), vanes are either layered, indicated by white arrows (b), or arranged irregularly (c). The red arrow indicates the proximal end of the feather (d). e, e9, f, Distal regions of non-shafted feathers, in which barbs appear loosely parallel. Abbreviations: as1ca, astragalus and calcaneum; ce, cervical vertebrae; co, coracoid; cv, caudal vertebrae; dt314, distal tarsals 3 and 4; f, frontal; fe, femur; fi, fibula; ga, gastralia; hu, humerus; il, ilium; is, ischium; ma, mandible; mtI–IV, metatarsals I–IV; p, phalanges or metacarpals; pa, parietal; pm, premaxilla; pu, pubis; q, quadrate; r, ribs; ra, radius; rc, rachis; sc, scapula; st, sternum; ti, tibia; tv, thoracic vertebrae; u, ungual phalanges; ul, ulna; uv, unbranched vane" httpUri="https://zenodo.org/record/2688335/files/figure.png" pageId="2" pageNumber="1107">Fig. 2b, c, d, d</figureCitation>
|
||
9), an arrangement that has never previously been reported.
|
||
</paragraph>
|
||
<paragraph blockId="2.[821,1493,133,1315]" pageId="2" pageNumber="1107">
|
||
The four ETFs are tightly attached to the last ten caudal segments (
|
||
<figureCitation box="[828,898,586,609]" captionStart="Figure 1" captionStartId="0.[112,170,1868,1887]" captionTargetBox="[158,1447,991,1846]" captionTargetId="figure@0.[158,1447,991,1846]" captionTargetPageId="0" captionText="Figure 1 | Epidexipteryx hui gen. et sp. nov., IVPP V15471, main slab and close-up photos. a, Main slab; b, c, skull in main slab (b) and counterslab (c); d, four elongate ribbon-like tail feathers; b9, c9, line drawings of b and c, respectively. Abbreviations: l1, l2 and l7, 1st, 2nd and 7th left teeth of upper jaw; l19, r19 and r59, 1st left, 1st right and 5th right teeth of lower jaw; l2 and r2, 2nd left and right teeth of upper jaw" httpUri="https://zenodo.org/record/2688156/files/figure.png" pageId="2" pageNumber="1107">Figs 1a</figureCitation>
|
||
and
|
||
<figureCitation box="[948,975,587,609]" captionStart="Figure 2" captionStartId="1.[93,151,1805,1824]" captionTargetId="figure@1.[271,1293,714,1784]" captionTargetPageId="1" captionText="Figure 2 | Line drawings and close-up photographs of Epidexipteryx hui gen. et sp. nov. a, Skeleton and feather outline, based on both main slab and counterslab, showing that each shafted feather is formed by central rachis and two unbranched vanes (a9). b, d, d9, Proximal regions of non-shafted feathers; barbs are parallel and closely united as an unbranched membranous structure (d, d9), vanes are either layered, indicated by white arrows (b), or arranged irregularly (c). The red arrow indicates the proximal end of the feather (d). e, e9, f, Distal regions of non-shafted feathers, in which barbs appear loosely parallel. Abbreviations: as1ca, astragalus and calcaneum; ce, cervical vertebrae; co, coracoid; cv, caudal vertebrae; dt314, distal tarsals 3 and 4; f, frontal; fe, femur; fi, fibula; ga, gastralia; hu, humerus; il, ilium; is, ischium; ma, mandible; mtI–IV, metatarsals I–IV; p, phalanges or metacarpals; pa, parietal; pm, premaxilla; pu, pubis; q, quadrate; r, ribs; ra, radius; rc, rachis; sc, scapula; st, sternum; ti, tibia; tv, thoracic vertebrae; u, ungual phalanges; ul, ulna; uv, unbranched vane" httpUri="https://zenodo.org/record/2688335/files/figure.png" pageId="2" pageNumber="1107">2a</figureCitation>
|
||
). These feathers are incomplete distally, but the preserved part of each ETF is identical to the corresponding structure in some primitive birds
|
||
<superScript attach="left" box="[1031,1088,640,653]" fontSize="5" pageId="2" pageNumber="1107">16,26–28</superScript>
|
||
, for example in having a similar central rachis and unbranched vanes (
|
||
<figureCitation box="[1114,1202,671,694]" captionStart="Figure 1" captionStartId="0.[112,170,1868,1887]" captionTargetBox="[158,1447,991,1846]" captionTargetId="figure@0.[158,1447,991,1846]" captionTargetPageId="0" captionText="Figure 1 | Epidexipteryx hui gen. et sp. nov., IVPP V15471, main slab and close-up photos. a, Main slab; b, c, skull in main slab (b) and counterslab (c); d, four elongate ribbon-like tail feathers; b9, c9, line drawings of b and c, respectively. Abbreviations: l1, l2 and l7, 1st, 2nd and 7th left teeth of upper jaw; l19, r19 and r59, 1st left, 1st right and 5th right teeth of lower jaw; l2 and r2, 2nd left and right teeth of upper jaw" httpUri="https://zenodo.org/record/2688156/files/figure.png" pageId="2" pageNumber="1107">Fig. 1a, d</figureCitation>
|
||
and
|
||
<figureCitation box="[1248,1291,671,693]" captionStart="Figure 2" captionStartId="1.[93,151,1805,1824]" captionTargetId="figure@1.[271,1293,714,1784]" captionTargetPageId="1" captionText="Figure 2 | Line drawings and close-up photographs of Epidexipteryx hui gen. et sp. nov. a, Skeleton and feather outline, based on both main slab and counterslab, showing that each shafted feather is formed by central rachis and two unbranched vanes (a9). b, d, d9, Proximal regions of non-shafted feathers; barbs are parallel and closely united as an unbranched membranous structure (d, d9), vanes are either layered, indicated by white arrows (b), or arranged irregularly (c). The red arrow indicates the proximal end of the feather (d). e, e9, f, Distal regions of non-shafted feathers, in which barbs appear loosely parallel. Abbreviations: as1ca, astragalus and calcaneum; ce, cervical vertebrae; co, coracoid; cv, caudal vertebrae; dt314, distal tarsals 3 and 4; f, frontal; fe, femur; fi, fibula; ga, gastralia; hu, humerus; il, ilium; is, ischium; ma, mandible; mtI–IV, metatarsals I–IV; p, phalanges or metacarpals; pa, parietal; pm, premaxilla; pu, pubis; q, quadrate; r, ribs; ra, radius; rc, rachis; sc, scapula; st, sternum; ti, tibia; tv, thoracic vertebrae; u, ungual phalanges; ul, ulna; uv, unbranched vane" httpUri="https://zenodo.org/record/2688335/files/figure.png" pageId="2" pageNumber="1107">2a, a</figureCitation>
|
||
9; see Supplementary Information). Elongate tail feathers are a normal component of the ornamental plumage in extant birds. In contrast to other feather types, ornamental plumage is used to send visual signals that are essential to a wide range of avian behaviour patterns, particularly relating to courtship
|
||
<bibRefCitation author="Gill ̡ F. B." box="[1027,1042,810,823]" journalOrPublisher="Freeman ̡" pageId="2" pageNumber="1107" refId="ref3061" refString="29. Gill ̡ F. B. Ornithology 2 nd edn (Freeman ̡ 1995)." title="Ornithology 2 nd edn" type="book" year="1995">
|
||
<superScript attach="left" box="[1027,1042,810,823]" fontSize="5" pageId="2" pageNumber="1107">29</superScript>
|
||
</bibRefCitation>
|
||
. For example, experiments have shown that, in some species, males with long tail plumage attract more mates than their short-tailed counterparts
|
||
<superScript attach="right" box="[1118,1133,866,879]" fontSize="5" pageId="2" pageNumber="1107">30</superScript>
|
||
. It is highly probable that the ETFs of
|
||
<taxonomicName box="[821,952,897,918]" class="Reptilia" family="Scansoriopterygidae" genus="Epidexipteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="1107" phylum="Chordata" rank="genus">
|
||
<emphasis box="[821,952,897,918]" italics="true" pageId="2" pageNumber="1107">Epidexipteryx</emphasis>
|
||
</taxonomicName>
|
||
similarly had display as their primary function, rather than serving other purposes such as flight or insulation
|
||
<bibRefCitation author="Gill ̡ F. B." box="[1389,1404,923,936]" journalOrPublisher="Freeman ̡" pageId="2" pageNumber="1107" refId="ref3061" refString="29. Gill ̡ F. B. Ornithology 2 nd edn (Freeman ̡ 1995)." title="Ornithology 2 nd edn" type="book" year="1995">
|
||
<superScript attach="left" box="[1389,1404,923,936]" fontSize="5" pageId="2" pageNumber="1107">29</superScript>
|
||
</bibRefCitation>
|
||
. Indeed, pennaceous feathers suitable for flight are not present in
|
||
<taxonomicName box="[821,952,982,1003]" class="Reptilia" family="Scansoriopterygidae" genus="Epidexipteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="1107" phylum="Chordata" rank="genus">
|
||
<emphasis box="[821,952,982,1003]" italics="true" pageId="2" pageNumber="1107">Epidexipteryx</emphasis>
|
||
</taxonomicName>
|
||
, even though the bones and integument are well preserved. Because pennaceous feathers are commonly encountered in other feathered maniraptorans
|
||
<superScript attach="left" box="[1144,1208,1036,1049]" fontSize="5" pageId="2" pageNumber="1107">2,21,22,25</superScript>
|
||
, their absence constitutes another highly unusual feature of
|
||
<taxonomicName box="[1166,1297,1067,1088]" class="Reptilia" family="Scansoriopterygidae" genus="Epidexipteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="1107" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1166,1297,1067,1088]" italics="true" pageId="2" pageNumber="1107">Epidexipteryx</emphasis>
|
||
</taxonomicName>
|
||
, as well as strongly implying that this taxon was non-volant.
|
||
<taxonomicName box="[1236,1367,1095,1116]" class="Reptilia" family="Scansoriopterygidae" genus="Epidexipteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="1107" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1236,1367,1095,1116]" italics="true" pageId="2" pageNumber="1107">Epidexipteryx</emphasis>
|
||
</taxonomicName>
|
||
is the oldest and most phylogenetically basal theropod known to possess display feathers, indicating that basal avialans experimented with integumentary ornament as early as the Middle to Late Jurassic. Unless
|
||
<taxonomicName box="[821,952,1208,1229]" class="Reptilia" family="Scansoriopterygidae" genus="Epidexipteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="1107" phylum="Chordata" rank="genus">
|
||
<emphasis box="[821,952,1208,1229]" italics="true" pageId="2" pageNumber="1107">Epidexipteryx</emphasis>
|
||
</taxonomicName>
|
||
is interpreted as secondarily flightless, the absence of pennaceous limb feathers in this taxon suggests that display feathers appeared before airfoil feathers and flight ability in basal avialan evolution.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |