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528 lines
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<document ID-DOI="10.1206/0003-0090-411.1.1" ID-GBIF-Dataset="55e2a047-2c2e-42a8-a8d9-918a125d1cf8" ID-ISSN="0003-0090" ID-Zenodo-Dep="4612829" checkinTime="1615994159644" checkinUser="felipe" docAuthor="Pei, Rui, Li, Quanguo, Meng, Qingjin, Norell, Mark A. & Gao, Ke-Qin" docDate="2017" docId="13087C7B1F5E970DFD48222600D0FC26" docLanguage="en" docName="BulAmeMusNatHis.2017.411.1-66.pdf.imf" docOrigin="Bulletin of the American Museum of Natural History 2017 (411)" docSource="http://www.bioone.org/doi/10.1206/0003-0090-411.1.1" docStyle="DocumentStyle:915933466F796C9C739DF4DB6B8DCFA6.7:BulAmeMusNatHis.2011-.journal_article.1cover.type1" docStyleId="915933466F796C9C739DF4DB6B8DCFA6" docStyleName="BulAmeMusNatHis.2011-.journal_article.1cover.type1" docStyleVersion="7" docTitle="Anchiornis HUXLEYI" docType="treatment" docVersion="6" lastPageNumber="57" masterDocId="EF3104031F689734FFAA2078015DFFBE" masterDocTitle="New Specimens Of Anchiornis Huxleyi (Theropoda: Paraves) From The Late Jurassic Of Northeastern China" masterLastPageNumber="66" masterPageNumber="1" pageNumber="54" updateTime="1643485001803" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0">
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<mods:titleInfo>
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<mods:title>New Specimens Of Anchiornis Huxleyi (Theropoda: Paraves) From The Late Jurassic Of Northeastern China</mods:title>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Pei, Rui</mods:namePart>
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<mods:affiliation>Department of Earth Sciences, the University of Hong Kong, Hong Kong; Division of Paleontology, American Museum of Natural History</mods:affiliation>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Li, Quanguo</mods:namePart>
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<mods:affiliation>State Key Laboratory of Biogeology and Environmental Geology, China University of Geosciences, Beijing</mods:affiliation>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Meng, Qingjin</mods:namePart>
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<mods:affiliation>Beijing Museum of Natural History, Beijing</mods:affiliation>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Norell, Mark A.</mods:namePart>
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<mods:affiliation>Division of Paleontology, American Museum of Natural History</mods:affiliation>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Gao, Ke-Qin</mods:namePart>
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<mods:affiliation>School of Earth and Space Sciences, Peking University, Beijing</mods:affiliation>
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<mods:date>2017</mods:date>
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<mods:number>2017-04-13</mods:number>
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<mods:number>2017</mods:number>
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<mods:number>411</mods:number>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.4630978" ID-GBIF-Taxon="180670700" ID-Zenodo-Dep="4630978" LSID="urn:lsid:plazi:treatment:13087C7B1F5E970DFD48222600D0FC26" httpUri="http://treatment.plazi.org/id/13087C7B1F5E970DFD48222600D0FC26" lastPageId="57" lastPageNumber="57" pageId="54" pageNumber="54">
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<subSubSection box="[738,1159,606,630]" pageId="54" pageNumber="54" type="nomenclature">
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<paragraph blockId="54.[738,1159,606,630]" box="[738,1159,606,630]" pageId="54" pageNumber="54">
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<heading box="[738,1159,606,630]" centered="true" fontSize="10" level="2" pageId="54" pageNumber="54" reason="4">
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AFFILIATION OF
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<taxonomicName authority="HUXLEYI" box="[922,1159,606,630]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="54" pageNumber="54" phylum="Chordata" rank="genus">
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<emphasis box="[922,1159,606,630]" italics="true" pageId="54" pageNumber="54">ANCHIORNIS HUXLEYI</emphasis>
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</taxonomicName>
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</heading>
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</paragraph>
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</subSubSection>
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<subSubSection lastPageId="57" lastPageNumber="57" pageId="54" pageNumber="54" type="description">
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<paragraph blockId="54.[684,1213,655,1738]" box="[684,1106,655,679]" pageId="54" pageNumber="54">Affiliation to Maniraptora and Paraves</paragraph>
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<paragraph blockId="54.[684,1213,655,1738]" pageId="54" pageNumber="54">
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The morphology of
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<taxonomicName authorityName=", PKUP V" authorityYear="1068" box="[934,1135,688,712]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="54" pageNumber="54" phylum="Chordata" rank="species">
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<emphasis box="[934,1135,688,712]" italics="true" pageId="54" pageNumber="54">Anchiornis huxleyi</emphasis>
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</taxonomicName>
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can be confidently assigned to the Maniraptora. Several synapomorphies of Maniraptora are present in
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<taxonomicName authorityName=", PKUP V" authorityYear="1068" box="[684,900,788,812]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="54" pageNumber="54" phylum="Chordata" rank="species">
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<emphasis box="[684,900,788,812]" italics="true" pageId="54" pageNumber="54">Anchiornis huxleyi</emphasis>
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</taxonomicName>
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: broad triangular process along the lateral edge of the quadrate that contacts the squamosal and the quadratojugal; absence of the prefrontal, fused parietals, teeth constricted between the root and crown, reduced prezygapophyses on the distal caudal vertebrae, presence of a semilunate carpal; manual phalanx III-3 significantly elongate, and fibular shaπ narrowing abruptly below the iliofibularis tubercle.
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</paragraph>
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<paragraph blockId="54.[684,1213,655,1738]" lastBlockId="55.[108,638,226,580]" lastPageId="55" lastPageNumber="55" pageId="54" pageNumber="54">
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In many previous studies a monophyletic Paraves is recovered as a clade that includes
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<taxonomicName box="[684,884,1151,1175]" class="Reptilia" family="Dromaeosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="54" pageNumber="54" phylum="Chordata" rank="family">Dromaeosauridae</taxonomicName>
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,
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<taxonomicName authorityName="Gilmore" authorityYear="1924" box="[903,1056,1151,1175]" class="Reptilia" family="Troodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="54" pageNumber="54" phylum="Chordata" rank="family">Troodontidae</taxonomicName>
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, and Avialae (
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;
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;
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;
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).
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<taxonomicName authorityName=", PKUP V" authorityYear="1068" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="54" pageNumber="54" phylum="Chordata" rank="species">
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<emphasis italics="true" pageId="54" pageNumber="54">Anchiornis huxleyi</emphasis>
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</taxonomicName>
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can be referred to Paraves based on the following derived features: shallow maxilla and premaxilla below the external naris, T-shaped lacrimal; dentary symphyseal region in line with the main part of the buccal margin, chevrons anteriorly bifurcate on the distal part of the tail, nearly symmetric furcula, acromion margin of the scapula with a laterally everted anterior edge, coracoid inflected medially from the scapula, forming an L-shaped scapulocoracoid, humerus longer than the scapula, ilium with the pubic peduncle much larger than the ischiadic peduncle, femur with a posterior trochanter, and the distal end of metatarsal III ginglymoid.
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</paragraph>
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<paragraph blockId="55.[108,638,226,580]" pageId="55" pageNumber="55">
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Within Paraves, the affiliation of
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<taxonomicName box="[499,611,358,382]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="55" pageNumber="55" phylum="Chordata" rank="genus">
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<emphasis box="[499,611,358,382]" italics="true" pageId="55" pageNumber="55">Anchiornis</emphasis>
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</taxonomicName>
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is controversial (
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,
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<bibRefCitation author="Xu, X. & H. You & F. Han" box="[417,469,391,415]" pageId="55" pageNumber="55" pagination="465 - 470" refId="ref33141" refString="Xu, X., H. You, and F. Han. 2011. An Archaeopteryx - like theropod from China and the origin of Avialae. Nature 475: 465 - 470." type="journal article" year="2011">2011</bibRefCitation>
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;
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<bibRefCitation author="Hu, D. & L. Hou & L. Zhang & X. Xu" box="[479,633,391,415]" pageId="55" pageNumber="55" pagination="640 - 643" refId="ref30594" refString="Hu, D., L. Hou, L. Zhang, and X. Xu. 2009. A pre- Archaeopteryx troodontid theropod from China with long feathers on the metatarsal. Nature 461: 640 - 643." type="journal article" year="2009">Hu et al., 2009</bibRefCitation>
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;
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<bibRefCitation author="Turner, A. H. & P. J. Makovicky & M. A. Norell" box="[108,306,424,448]" pageId="55" pageNumber="55" pagination="1 - 207" refId="ref32607" refString="Turner, A. H., P. J. Makovicky, and M. A. Norell. 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History 371: 1 - 207." type="journal article" year="2012">Turner et al., 2012</bibRefCitation>
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;
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<bibRefCitation author="Senter, P. & J. I. Kirkland & D. D. DeBlieux & S. Madsen & N. Toth" box="[319,510,424,448]" pageId="55" pageNumber="55" pagination="36790" refId="ref32350" refString="Senter, P., J. I. Kirkland, D. D. DeBlieux, S. Madsen, N. Toth. 2012. New dromaeosaurids (Dinosauria: Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid tail. PLoS ONE 7 (5): e 36790." type="journal article" year="2012">Senter et al., 2012</bibRefCitation>
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;
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<bibRefCitation author="Foth, C. & H. Tischlinger & O. W. M. Rauhut" pageId="55" pageNumber="55" pagination="79 - 82" refId="ref30225" refString="Foth, C., H. Tischlinger, and O. W. M. Rauhut. 2014. New specimen of Archaeopteryx provides insights into the evolution of pennaceous feathers. Nature 511: 79 - 82." type="journal article" year="2014">Foth et al., 2014</bibRefCitation>
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). This confusion is caused mainly by the conservative morphology of basal paravians. However, careful examination of new specimens gives us better clue of the affiliation of
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<taxonomicName box="[392,502,556,580]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="55" pageNumber="55" phylum="Chordata" rank="genus">
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<emphasis box="[392,502,556,580]" italics="true" pageId="55" pageNumber="55">Anchiornis</emphasis>
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</taxonomicName>
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.
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</paragraph>
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<paragraph blockId="55.[108,637,622,1738]" box="[108,463,622,646]" pageId="55" pageNumber="55">
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Comparison with
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<taxonomicName baseAuthorityName="Wellnhofer" baseAuthorityYear="2009" box="[309,463,622,646]" class="Aves" family="Archaeopterygidae" genus="Archaeopteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Archaeopterygiformes" pageId="55" pageNumber="55" phylum="Chordata" rank="genus">
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<emphasis box="[309,463,622,646]" italics="true" pageId="55" pageNumber="55">Archaeopteryx</emphasis>
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</taxonomicName>
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</paragraph>
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<paragraph blockId="55.[108,637,622,1738]" pageId="55" pageNumber="55">
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<taxonomicName authorityName=", PKUP V" authorityYear="1068" box="[140,349,655,679]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="55" pageNumber="55" phylum="Chordata" rank="species">
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<emphasis box="[140,349,655,679]" italics="true" pageId="55" pageNumber="55">Anchiornis huxleyi</emphasis>
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</taxonomicName>
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shares many features in common with
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<taxonomicName baseAuthorityName="Wellnhofer" baseAuthorityYear="2009" box="[267,419,688,712]" class="Aves" family="Archaeopterygidae" genus="Archaeopteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Archaeopterygiformes" pageId="55" pageNumber="55" phylum="Chordata" rank="genus">
|
|
<emphasis box="[267,419,688,712]" italics="true" pageId="55" pageNumber="55">Archaeopteryx</emphasis>
|
|
</taxonomicName>
|
|
that are not present in dromaeosaurids and troodontids.
|
|
<taxonomicName box="[520,636,722,746]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="55" pageNumber="55" phylum="Chordata" rank="genus">
|
|
<emphasis box="[520,636,722,746]" italics="true" pageId="55" pageNumber="55">Anchiornis</emphasis>
|
|
</taxonomicName>
|
|
resembles
|
|
<taxonomicName baseAuthorityName="Wellnhofer" baseAuthorityYear="2009" box="[220,372,755,779]" class="Aves" family="Archaeopterygidae" genus="Archaeopteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Archaeopterygiformes" pageId="55" pageNumber="55" phylum="Chordata" rank="genus">
|
|
<emphasis box="[220,372,755,779]" italics="true" pageId="55" pageNumber="55">Archaeopteryx</emphasis>
|
|
</taxonomicName>
|
|
in having: straight nasal process of the premaxilla, external naris posterior to all premaxillary teeth, fewer than 20 dentary teeth, absence of the coranoid eminence, absence of lateral exposure of the splenial, and absence of an external mandibular fenestra in lateral view. In the postcranial skeleton,
|
|
<taxonomicName box="[395,512,953,977]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="55" pageNumber="55" phylum="Chordata" rank="genus">
|
|
<emphasis box="[395,512,953,977]" italics="true" pageId="55" pageNumber="55">Anchiornis</emphasis>
|
|
</taxonomicName>
|
|
shares the following features with
|
|
<taxonomicName baseAuthorityName="Wellnhofer" baseAuthorityYear="2009" box="[382,538,986,1010]" class="Aves" family="Archaeopterygidae" genus="Archaeopteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Archaeopterygiformes" pageId="55" pageNumber="55" phylum="Chordata" rank="genus">
|
|
<emphasis box="[382,538,986,1010]" italics="true" pageId="55" pageNumber="55">Archaeopteryx</emphasis>
|
|
</taxonomicName>
|
|
: slightly bowed metacarpal III, pubic peduncle more ventrally expressed than the ischiadic peduncle, constricted base of the distally located obturator process, absence of a slit on the pubic apron, and proximal end of metatarsal III unconstricted. All of these features are found to be primitive for avialans, either as synapomorphies that are unique for avialans or synapomorphies of a more inclusive coelurosaurian clade that are secondarily lost in deinonychosaurians.
|
|
</paragraph>
|
|
<paragraph blockId="55.[108,637,622,1738]" lastBlockId="55.[684,1213,226,283]" pageId="55" pageNumber="55">
|
|
<taxonomicName box="[141,257,1350,1374]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="55" pageNumber="55" phylum="Chordata" rank="genus">
|
|
<emphasis box="[141,257,1350,1374]" italics="true" pageId="55" pageNumber="55">Anchiornis</emphasis>
|
|
</taxonomicName>
|
|
also shares the following features with
|
|
<taxonomicName baseAuthorityName="Wellnhofer" baseAuthorityYear="2009" box="[163,317,1383,1407]" class="Aves" family="Archaeopterygidae" genus="Archaeopteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Archaeopterygiformes" pageId="55" pageNumber="55" phylum="Chordata" rank="genus">
|
|
<emphasis box="[163,317,1383,1407]" italics="true" pageId="55" pageNumber="55">Archaeopteryx</emphasis>
|
|
</taxonomicName>
|
|
: enlarged orbit, presence of a shallow premaxilla, presence of a promaxillary fenestra, enlarged maxillary fenestra, lacrimal with slender and subequal anterior and posterior processes, dorsal vertebrae with lateral concavities but not pneumatic foramina, scapula significantly shorter and slimmer than humerus, manual phalanx III-3 significantly longer than III-1 and III-2 combined, supraacetabular crest, and ischium with a distally located obturator process. However, these features are also present in many basal troodontids and dromaeosaurids and thus are primitive for Paraves.
|
|
</paragraph>
|
|
<paragraph blockId="55.[684,1213,325,1738]" box="[685,1033,325,349]" pageId="55" pageNumber="55">
|
|
Comparison with
|
|
<taxonomicName authorityName="Gilmore" authorityYear="1924" box="[885,1033,325,349]" class="Reptilia" family="Troodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="55" pageNumber="55" phylum="Chordata" rank="family">Troodontidae</taxonomicName>
|
|
</paragraph>
|
|
<paragraph blockId="55.[684,1213,325,1738]" pageId="55" pageNumber="55">
|
|
Previous studies have assigned
|
|
<taxonomicName box="[1065,1183,358,382]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="55" pageNumber="55" phylum="Chordata" rank="genus">
|
|
<emphasis box="[1065,1183,358,382]" italics="true" pageId="55" pageNumber="55">Anchiornis</emphasis>
|
|
</taxonomicName>
|
|
as a basal troodontid (
|
|
<bibRefCitation author="Hu, D. & L. Hou & L. Zhang & X. Xu" box="[900,1062,391,415]" pageId="55" pageNumber="55" pagination="640 - 643" refId="ref30594" refString="Hu, D., L. Hou, L. Zhang, and X. Xu. 2009. A pre- Archaeopteryx troodontid theropod from China with long feathers on the metatarsal. Nature 461: 640 - 643." type="journal article" year="2009">Hu et al., 2009</bibRefCitation>
|
|
;
|
|
<bibRefCitation author="Turner, A. H. & P. J. Makovicky & M. A. Norell" pageId="55" pageNumber="55" pagination="1 - 207" refId="ref32607" refString="Turner, A. H., P. J. Makovicky, and M. A. Norell. 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History 371: 1 - 207." type="journal article" year="2012">Turner et al., 2012</bibRefCitation>
|
|
;
|
|
<bibRefCitation author="Pei, R. & Q. Li & Q. Meng & K. Gao & M. A. Norell" box="[756,938,424,448]" pageId="55" pageNumber="55" pagination="1 - 28" refId="ref32127" refString="Pei, R., Q. Li, Q. Meng, K. Gao, and M. A. Norell. 2014. A new specimen of Microraptor (Theropoda: Dromaeosauridae) from the Lower Cretaceous of western Liaoning, China. American Museum Novitates 3821: 1 - 28." type="journal article" year="2014">Pei et al., 2014</bibRefCitation>
|
|
).
|
|
<bibRefCitation author="Hu, D. & L. Hou & L. Zhang & X. Xu" box="[962,1152,424,448]" pageId="55" pageNumber="55" pagination="640 - 643" refId="ref30594" refString="Hu, D., L. Hou, L. Zhang, and X. Xu. 2009. A pre- Archaeopteryx troodontid theropod from China with long feathers on the metatarsal. Nature 461: 640 - 643." type="journal article" year="2009">Hu et al. (2009)</bibRefCitation>
|
|
suggested
|
|
<taxonomicName box="[763,881,457,481]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="55" pageNumber="55" phylum="Chordata" rank="genus">
|
|
<emphasis box="[763,881,457,481]" italics="true" pageId="55" pageNumber="55">Anchiornis</emphasis>
|
|
</taxonomicName>
|
|
and others troodontids have the following features in common: large maxillary fenestra, labial surface of dentary bearing a distinct groove, closely packed premaxillary and dentary teeth in the symphyseal region, and dorsal vertebrae and anteriormost caudal vertebrae bearing relatively long and slender transverse processes.
|
|
<bibRefCitation author="Hu, D. & L. Hou & L. Zhang & X. Xu" box="[868,1045,688,713]" pageId="55" pageNumber="55" pagination="640 - 643" refId="ref30594" refString="Hu, D., L. Hou, L. Zhang, and X. Xu. 2009. A pre- Archaeopteryx troodontid theropod from China with long feathers on the metatarsal. Nature 461: 640 - 643." type="journal article" year="2009">Hu et al. (2009)</bibRefCitation>
|
|
also suggested
|
|
<taxonomicName box="[686,807,722,746]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="55" pageNumber="55" phylum="Chordata" rank="genus">
|
|
<emphasis box="[686,807,722,746]" italics="true" pageId="55" pageNumber="55">Anchiornis</emphasis>
|
|
</taxonomicName>
|
|
shares with
|
|
<emphasis box="[968,1011,722,745]" italics="true" pageId="55" pageNumber="55">Mei</emphasis>
|
|
a large external naris extending posteriorly beyond and dorsal to the anterior border of antorbital fossa, a longitudinal groove along dorsomedial margin of the jugal, unserrated teeth, and a maxillary toothrow approaching the preorbital bar posteriorly. However, a large maxillary fenestra, a large external naris extending posteriorly beyond the anterior border of antorbital fossa, a longitudinal groove along dorsomedial margin of the jugal, unserrated teeth, and long transverse processes on dorsal vertebrae and anteriormost caudal vertebrae, are also present in
|
|
<taxonomicName baseAuthorityName="Wellnhofer" baseAuthorityYear="2009" box="[722,884,1151,1175]" class="Aves" family="Archaeopterygidae" genus="Archaeopteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Archaeopterygiformes" pageId="55" pageNumber="55" phylum="Chordata" rank="genus">
|
|
<emphasis box="[722,884,1151,1175]" italics="true" pageId="55" pageNumber="55">Archaeopteryx</emphasis>
|
|
</taxonomicName>
|
|
and basal dromaeosaurids (
|
|
<bibRefCitation author="Wellnhofer, P." box="[691,883,1184,1209]" pageId="55" pageNumber="55" pagination="169 - 216" refId="ref32701" refString="Wellnhofer, P. 1974. Das funπe Skelettexemplar von Archaeopteryx. Palaeontographica Abteilung A Palaeozoologie-Stratigraphie 147: 169 - 216." type="journal article" year="1974">Wellnhofer, 1974</bibRefCitation>
|
|
;
|
|
<bibRefCitation author="Mayr, G. & B. Pohl & D. S. Peters" box="[894,1080,1184,1209]" pageId="55" pageNumber="55" pagination="1483 - 1486" refId="ref31447" refString="Mayr, G., B. Pohl, and D. S. Peters. 2005. A well-preserved Archaeopteryx specimen with theropod features. Science 310: 1483 - 1486." type="journal article" year="2005">Mayr et al., 2005</bibRefCitation>
|
|
;
|
|
<bibRefCitation author="Makovicky, P. J. & S. Apesteguia & F. L. Agnolin" pageId="55" pageNumber="55" pagination="1007 - 1011" refId="ref31412" refString="Makovicky, P. J., S. Apesteguia, and F. L. Agnolin. 2005. The earliest dromaeosaurid theropod from South America. Nature 437: 1007 - 1011." type="journal article" year="2005">Makovicky et al., 2005</bibRefCitation>
|
|
;
|
|
<bibRefCitation author="Foth, C. & H. Tischlinger & O. W. M. Rauhut" box="[825,1014,1218,1242]" pageId="55" pageNumber="55" pagination="79 - 82" refId="ref30225" refString="Foth, C., H. Tischlinger, and O. W. M. Rauhut. 2014. New specimen of Archaeopteryx provides insights into the evolution of pennaceous feathers. Nature 511: 79 - 82." type="journal article" year="2014">Foth et al., 2014</bibRefCitation>
|
|
;
|
|
<bibRefCitation author="Pei, R. & Q. Li & Q. Meng & K. Gao & M. A. Norell" box="[1028,1201,1218,1242]" pageId="55" pageNumber="55" pagination="1 - 28" refId="ref32127" refString="Pei, R., Q. Li, Q. Meng, K. Gao, and M. A. Norell. 2014. A new specimen of Microraptor (Theropoda: Dromaeosauridae) from the Lower Cretaceous of western Liaoning, China. American Museum Novitates 3821: 1 - 28." type="journal article" year="2014">Pei et al., 2014</bibRefCitation>
|
|
). The primitive pennaraptoran
|
|
<taxonomicName box="[1012,1162,1251,1275]" class="Reptilia" family="Scansoriopterygidae" genus="Epidexipteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="55" pageNumber="55" phylum="Chordata" rank="genus">
|
|
<emphasis box="[1012,1162,1251,1275]" italics="true" pageId="55" pageNumber="55">Epidexipteryx</emphasis>
|
|
</taxonomicName>
|
|
also has closely packed premaxillary and dentary teeth in the symphyseal region (
|
|
<bibRefCitation author="Zhang, F. & Z. Zhou & X. Xu & X. Wang & C. Sullivan" pageId="55" pageNumber="55" pagination="1105 - 1108" refId="ref33293" refString="Zhang, F., Z. Zhou, X. Xu, X. Wang, and C. Sullivan. 2008. A bizarre Jurassic maniraptoran from China with elongate ribbon-like feathers. Nature 455: 1105 - 1108." type="journal article" year="2008">Zhang et al., 2008</bibRefCitation>
|
|
). The lateral groove on the dentary and maxillary toothrow approaching preorbital bar are observed in
|
|
<taxonomicName baseAuthorityName="Wellnhofer" baseAuthorityYear="2009" box="[870,1028,1416,1440]" class="Aves" family="Archaeopterygidae" genus="Archaeopteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Archaeopterygiformes" pageId="55" pageNumber="55" phylum="Chordata" rank="genus">
|
|
<emphasis box="[870,1028,1416,1440]" italics="true" pageId="55" pageNumber="55">Archaeopteryx</emphasis>
|
|
</taxonomicName>
|
|
and other nontroodontid basal deinonychosaurians such as
|
|
<taxonomicName authorityName="Makovicky, Apesteguia & Agnolin" authorityYear="2005" box="[684,820,1483,1506]" class="Reptilia" family="Dromaeosauridae" genus="Buitreraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="55" pageNumber="55" phylum="Chordata" rank="genus">
|
|
<emphasis box="[684,820,1483,1506]" italics="true" pageId="55" pageNumber="55">Buitreraptor</emphasis>
|
|
</taxonomicName>
|
|
(
|
|
<bibRefCitation author="Wellnhofer, P." box="[839,1040,1482,1506]" pageId="55" pageNumber="55" pagination="169 - 216" refId="ref32701" refString="Wellnhofer, P. 1974. Das funπe Skelettexemplar von Archaeopteryx. Palaeontographica Abteilung A Palaeozoologie-Stratigraphie 147: 169 - 216." type="journal article" year="1974">Wellnhofer, 1974</bibRefCitation>
|
|
;
|
|
<bibRefCitation author="Makovicky, P. J. & S. Apesteguia & F. L. Agnolin" pageId="55" pageNumber="55" pagination="1007 - 1011" refId="ref31412" refString="Makovicky, P. J., S. Apesteguia, and F. L. Agnolin. 2005. The earliest dromaeosaurid theropod from South America. Nature 437: 1007 - 1011." type="journal article" year="2005">Makovicky et al., 2005</bibRefCitation>
|
|
). Thus, these features are all primitive to a more inclusive coelurosaurian clade.
|
|
</paragraph>
|
|
<paragraph blockId="55.[684,1213,325,1738]" lastBlockId="56.[108,638,226,1506]" lastPageId="56" lastPageNumber="56" pageId="55" pageNumber="55">
|
|
In the phylogenetic analysis of
|
|
<bibRefCitation author="Turner, A. H. & P. J. Makovicky & M. A. Norell" pageId="55" pageNumber="55" pagination="1 - 207" refId="ref32607" refString="Turner, A. H., P. J. Makovicky, and M. A. Norell. 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History 371: 1 - 207." type="journal article" year="2012">Turner et al. (2012)</bibRefCitation>
|
|
, three characters were recovered to support
|
|
<taxonomicName box="[747,870,1647,1671]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="55" pageNumber="55" phylum="Chordata" rank="genus">
|
|
<emphasis box="[747,870,1647,1671]" italics="true" pageId="55" pageNumber="55">Anchiornis</emphasis>
|
|
</taxonomicName>
|
|
at the basalmost branch of
|
|
<taxonomicName authorityName="Gilmore" authorityYear="1924" box="[684,841,1680,1704]" class="Reptilia" family="Troodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="55" pageNumber="55" phylum="Chordata" rank="family">Troodontidae</taxonomicName>
|
|
, which include a dorsoventrally flattened internarial bar, the anteroventrally inclined quadrate and a subarctometatarsalian pes. In scoring based on a new
|
|
<taxonomicName baseAuthorityName="Wellnhofer" baseAuthorityYear="2009" box="[475,636,259,283]" class="Aves" family="Archaeopterygidae" genus="Archaeopteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Archaeopterygiformes" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[475,636,259,283]" italics="true" pageId="56" pageNumber="56">Archaeopteryx</emphasis>
|
|
</taxonomicName>
|
|
specimen (
|
|
<bibRefCitation author="Foth, C. & H. Tischlinger & O. W. M. Rauhut" box="[237,442,292,316]" pageId="56" pageNumber="56" pagination="79 - 82" refId="ref30225" refString="Foth, C., H. Tischlinger, and O. W. M. Rauhut. 2014. New specimen of Archaeopteryx provides insights into the evolution of pennaceous feathers. Nature 511: 79 - 82." type="journal article" year="2014">Foth et al., 2014</bibRefCitation>
|
|
), the internarial bar of
|
|
<taxonomicName baseAuthorityName="Wellnhofer" baseAuthorityYear="2009" box="[192,355,325,349]" class="Aves" family="Archaeopterygidae" genus="Archaeopteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Archaeopterygiformes" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[192,355,325,349]" italics="true" pageId="56" pageNumber="56">Archaeopteryx</emphasis>
|
|
</taxonomicName>
|
|
is scored as dorsoventrally flat as in troodontids like
|
|
<taxonomicName box="[495,629,358,381]" class="Reptilia" family="Troodontidae" genus="Sinovenator" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[495,629,358,381]" italics="true" pageId="56" pageNumber="56">Sinovenator</emphasis>
|
|
</taxonomicName>
|
|
,
|
|
<emphasis box="[108,266,392,415]" italics="true" pageId="56" pageNumber="56">
|
|
<taxonomicName box="[108,262,392,415]" class="Reptilia" family="Troodontidae" genus="Byronosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">Byronosaurus</taxonomicName>
|
|
,
|
|
</emphasis>
|
|
and
|
|
<taxonomicName box="[322,442,391,415]" class="Reptilia" family="Troodontidae" genus="Zanabazar" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[322,442,391,415]" italics="true" pageId="56" pageNumber="56">Zanabazar</emphasis>
|
|
</taxonomicName>
|
|
, and a dorsoventrally flat internarial bar is also observed in the dromaeosaurid
|
|
<taxonomicName authority="(Pei et al., 2014)" baseAuthorityName="Pei" baseAuthorityYear="2014" box="[291,632,457,481]" class="Reptilia" family="Dromaeosauridae" genus="Microraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[291,426,458,481]" italics="true" pageId="56" pageNumber="56">Microraptor</emphasis>
|
|
(
|
|
<bibRefCitation author="Pei, R. & Q. Li & Q. Meng & K. Gao & M. A. Norell" box="[443,623,457,481]" pageId="56" pageNumber="56" pagination="1 - 28" refId="ref32127" refString="Pei, R., Q. Li, Q. Meng, K. Gao, and M. A. Norell. 2014. A new specimen of Microraptor (Theropoda: Dromaeosauridae) from the Lower Cretaceous of western Liaoning, China. American Museum Novitates 3821: 1 - 28." type="journal article" year="2014">Pei et al., 2014</bibRefCitation>
|
|
)
|
|
</taxonomicName>
|
|
, so this feature is no longer restricted to
|
|
<taxonomicName authorityName="Gilmore" authorityYear="1924" box="[108,268,523,547]" class="Reptilia" family="Troodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="family">Troodontidae</taxonomicName>
|
|
. Both dorsoventrally flat (e.g., Eichstätt and Thermopolis specimens) and rodlike (e.g., Solnhofen specimen) internarial bars are actually observed in
|
|
<taxonomicName baseAuthorityName="Wellnhofer" baseAuthorityYear="2009" box="[466,629,622,646]" class="Aves" family="Archaeopterygidae" genus="Archaeopteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Archaeopterygiformes" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[466,629,622,646]" italics="true" pageId="56" pageNumber="56">Archaeopteryx</emphasis>
|
|
</taxonomicName>
|
|
. The anteroventrally inclined quadrate is also observed in
|
|
<taxonomicName baseAuthorityName="Wellnhofer" baseAuthorityYear="2009" box="[246,405,688,712]" class="Aves" family="Archaeopterygidae" genus="Archaeopteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Archaeopterygiformes" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[246,405,688,712]" italics="true" pageId="56" pageNumber="56">Archaeopteryx</emphasis>
|
|
</taxonomicName>
|
|
and more basal taxa such as
|
|
<taxonomicName authorityName="Ji & Ji" authorityYear="1996" box="[201,380,722,745]" class="Reptilia" family="Compsognathidae" genus="Sinosauropteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[201,380,722,745]" italics="true" pageId="56" pageNumber="56">Sinosauropteryx</emphasis>
|
|
</taxonomicName>
|
|
and
|
|
<taxonomicName authorityName="Marsh" authorityYear="1890" box="[439,603,722,746]" class="Reptilia" family="Ornithomimidae" genus="Ornithomimus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[439,603,722,746]" italics="true" pageId="56" pageNumber="56">Ornithomimus</emphasis>
|
|
</taxonomicName>
|
|
(
|
|
<bibRefCitation author="Ji, Q. & S. Ji" pageId="56" pageNumber="56" pagination="30 - 33" refId="ref30685" refString="Ji, Q., and S. Ji. 1996. On discovery of the earliest bird fossil in China and the origin of birds. Chinese Geology 10: 30 - 33." type="journal article" year="1996">Ji and Ji, 1996</bibRefCitation>
|
|
;
|
|
<bibRefCitation author="Wellnhofer, P." box="[259,456,755,779]" pageId="56" pageNumber="56" refId="ref32765" refString="Wellnhofer, P. 2009. Archaeopteryx: the icon of evolution. Munich: Verlag F. Pfeil." type="book" year="2009">Wellnhofer, 2009</bibRefCitation>
|
|
;
|
|
<bibRefCitation author="Rauhut, O. W. M." box="[468,623,755,779]" pageId="56" pageNumber="56" pagination="211 - 221" refId="ref32232" refString="Rauhut, O. W. M. 2014. New observations on the skull of Archaeopteryx. Palaontologische Zeitschriπ 88: 211 - 221." type="journal article" year="2014">Rauhut, 2014</bibRefCitation>
|
|
), and thus it is a synapomorphy of a larger clade. The arctometatarsalian pes of
|
|
<taxonomicName box="[481,604,821,845]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[481,604,821,845]" italics="true" pageId="56" pageNumber="56">Anchiornis</emphasis>
|
|
</taxonomicName>
|
|
is dubious, and new specimens revealed the pes of
|
|
<taxonomicName box="[147,272,887,911]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[147,272,887,911]" italics="true" pageId="56" pageNumber="56">Anchiornis</emphasis>
|
|
</taxonomicName>
|
|
resembles more the normal pedal configuration of
|
|
<emphasis box="[383,546,920,944]" italics="true" pageId="56" pageNumber="56">Archaeopteyrx</emphasis>
|
|
than it does the arctometatarsalian pes of
|
|
<taxonomicName box="[503,637,954,977]" class="Reptilia" family="Dromaeosauridae" genus="Microraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[503,637,954,977]" italics="true" pageId="56" pageNumber="56">Microraptor</emphasis>
|
|
</taxonomicName>
|
|
and
|
|
<taxonomicName box="[165,300,987,1010]" class="Reptilia" family="Troodontidae" genus="Sinovenator" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[165,300,987,1010]" italics="true" pageId="56" pageNumber="56">Sinovenator</emphasis>
|
|
</taxonomicName>
|
|
. As discussed above, these three characters no longer adequately support
|
|
<taxonomicName box="[110,231,1052,1076]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[110,231,1052,1076]" italics="true" pageId="56" pageNumber="56">Anchiornis</emphasis>
|
|
</taxonomicName>
|
|
as a member of
|
|
<taxonomicName authorityName="Gilmore" authorityYear="1924" box="[422,580,1052,1076]" class="Reptilia" family="Troodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="family">Troodontidae</taxonomicName>
|
|
.
|
|
</paragraph>
|
|
<paragraph blockId="56.[108,638,226,1506]" pageId="56" pageNumber="56">
|
|
As a Jurassic basal paravian,
|
|
<taxonomicName authorityName=", PKUP V" authorityYear="1068" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="species">
|
|
<emphasis italics="true" pageId="56" pageNumber="56">Anchiornis huxleyi</emphasis>
|
|
</taxonomicName>
|
|
differs from troodontids in several features. These features include: posteriorly tapering ventral ramus of the maxilla, elongation of the forelimb, middle and posterior maxillary teeth sparsely spaced, fewer than 20 dentary teeth, presence of a basisphenoid recess, wide and dorsoventrally low foramen magnum, transition of elongate caudal vertebrae occurring before the 10th caudal vertebra and symmetric metatarsus. These features are primitive for paravians and are found in basal dromaeosaurids and/or basal avialans, but absent in troodontids.
|
|
</paragraph>
|
|
<paragraph blockId="56.[108,636,1548,1738]" box="[108,506,1548,1572]" pageId="56" pageNumber="56">Comparison with Deinonychosauria</paragraph>
|
|
<paragraph blockId="56.[108,636,1548,1738]" lastBlockId="56.[684,1214,226,1738]" pageId="56" pageNumber="56">
|
|
Because
|
|
<taxonomicName box="[238,355,1581,1605]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[238,355,1581,1605]" italics="true" pageId="56" pageNumber="56">Anchiornis</emphasis>
|
|
</taxonomicName>
|
|
displays many primitive paravians features that are present in both troodontids and the basal avialan
|
|
<taxonomicName baseAuthorityName="Wellnhofer" baseAuthorityYear="2009" box="[476,630,1647,1671]" class="Aves" family="Archaeopterygidae" genus="Archaeopteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Archaeopterygiformes" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[476,630,1647,1671]" italics="true" pageId="56" pageNumber="56">Archaeopteryx</emphasis>
|
|
</taxonomicName>
|
|
, it is necessary to look at derived features of Deinonychosauria in order to distinguish
|
|
<taxonomicName box="[521,636,1714,1738]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[521,636,1714,1738]" italics="true" pageId="56" pageNumber="56">Anchiornis</emphasis>
|
|
</taxonomicName>
|
|
from
|
|
<taxonomicName authorityName="Gilmore" authorityYear="1924" box="[747,898,226,250]" class="Reptilia" family="Troodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="family">Troodontidae</taxonomicName>
|
|
.
|
|
<taxonomicName box="[911,1027,226,250]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[911,1027,226,250]" italics="true" pageId="56" pageNumber="56">Anchiornis</emphasis>
|
|
</taxonomicName>
|
|
does not have a lateral exposure of the splenial and also lacks a specialized raptorial digit II, which differentiates it from deinonychosaurians.
|
|
</paragraph>
|
|
<paragraph blockId="56.[684,1214,226,1738]" pageId="56" pageNumber="56">
|
|
Some features that are regarded as derived characters in deinonychosaurians, such as a subarctometatarsalian pes and a raptorial pedal digit II, were considered to be present in
|
|
<taxonomicName box="[1096,1212,457,481]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[1096,1212,457,481]" italics="true" pageId="56" pageNumber="56">Anchiornis</emphasis>
|
|
</taxonomicName>
|
|
and used as evidence for a troodontid affiliation of this taxon (
|
|
<bibRefCitation author="Hu, D. & L. Hou & L. Zhang & X. Xu" box="[841,1010,523,547]" pageId="56" pageNumber="56" pagination="640 - 643" refId="ref30594" refString="Hu, D., L. Hou, L. Zhang, and X. Xu. 2009. A pre- Archaeopteryx troodontid theropod from China with long feathers on the metatarsal. Nature 461: 640 - 643." type="journal article" year="2009">Hu et al., 2009</bibRefCitation>
|
|
). However, careful examination of
|
|
<taxonomicName box="[863,980,556,580]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[863,980,556,580]" italics="true" pageId="56" pageNumber="56">Anchiornis</emphasis>
|
|
</taxonomicName>
|
|
specimens indicates
|
|
<taxonomicName box="[684,804,589,613]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[684,804,589,613]" italics="true" pageId="56" pageNumber="56">Anchiornis</emphasis>
|
|
</taxonomicName>
|
|
resembles
|
|
<taxonomicName baseAuthorityName="Wellnhofer" baseAuthorityYear="2009" box="[943,1102,589,613]" class="Aves" family="Archaeopterygidae" genus="Archaeopteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Archaeopterygiformes" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[943,1102,589,613]" italics="true" pageId="56" pageNumber="56">Archaeopteryx</emphasis>
|
|
</taxonomicName>
|
|
in these pedal characters.
|
|
</paragraph>
|
|
<paragraph blockId="56.[684,1214,226,1738]" lastBlockId="57.[108,637,226,920]" lastPageId="57" lastPageNumber="57" pageId="56" pageNumber="56">
|
|
The pes of
|
|
<taxonomicName box="[842,959,655,679]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[842,959,655,679]" italics="true" pageId="56" pageNumber="56">Anchiornis</emphasis>
|
|
</taxonomicName>
|
|
is reported as subarctometatarsalian (
|
|
<bibRefCitation author="Zheng, X. & H. You & X. Xu & Z. Dong" box="[868,1028,688,713]" pageId="56" pageNumber="56" pagination="333 - 336" refId="ref33369" refString="Zheng, X., H. You, X. Xu, and Z. Dong. 2009. An Early Cretaceous heterodontosaurid dinosaur with filamentous integumentary structures. Nature 458: 333 - 336." type="journal article" year="2009">Xu et al., 2009</bibRefCitation>
|
|
;
|
|
<bibRefCitation author="Hu, D. & L. Hou & L. Zhang & X. Xu" box="[1038,1201,688,713]" pageId="56" pageNumber="56" pagination="640 - 643" refId="ref30594" refString="Hu, D., L. Hou, L. Zhang, and X. Xu. 2009. A pre- Archaeopteryx troodontid theropod from China with long feathers on the metatarsal. Nature 461: 640 - 643." type="journal article" year="2009">Hu et al., 2009</bibRefCitation>
|
|
). But close examination and comparison of new
|
|
<taxonomicName box="[684,801,755,779]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[684,801,755,779]" italics="true" pageId="56" pageNumber="56">Anchiornis</emphasis>
|
|
</taxonomicName>
|
|
specimens indicate that the proximal end of metatarsal III is not significantly constricted. Instead, the proximal end of metatarsal III has a subequal width as metatarsals II and IV in anterior view, which resembles
|
|
<taxonomicName baseAuthorityName="Wellnhofer" baseAuthorityYear="2009" box="[1057,1213,887,911]" class="Aves" family="Archaeopterygidae" genus="Archaeopteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Archaeopterygiformes" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[1057,1213,887,911]" italics="true" pageId="56" pageNumber="56">Archaeopteryx</emphasis>
|
|
</taxonomicName>
|
|
and most nontroodontid theropods, but different from the subarctometatarsus of basal troodontids (e.g.
|
|
<taxonomicName box="[879,1007,987,1010]" class="Reptilia" family="Troodontidae" genus="Sinovenator" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[879,1007,987,1010]" italics="true" pageId="56" pageNumber="56">Sinovenator</emphasis>
|
|
</taxonomicName>
|
|
and
|
|
<emphasis box="[1067,1109,987,1010]" italics="true" pageId="56" pageNumber="56">Mei</emphasis>
|
|
) and the
|
|
<typeStatus box="[684,731,1020,1044]" pageId="56" pageNumber="56">type</typeStatus>
|
|
specimen of
|
|
<taxonomicName box="[871,1052,1019,1043]" class="Reptilia" family="Dromaeosauridae" genus="Sinornithosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[871,1052,1019,1043]" italics="true" pageId="56" pageNumber="56">Sinornithosaurus</emphasis>
|
|
</taxonomicName>
|
|
and
|
|
<taxonomicName class="Reptilia" family="Dromaeosauridae" genus="Microraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis italics="true" pageId="56" pageNumber="56">Microraptor</emphasis>
|
|
</taxonomicName>
|
|
, where the proximal end of metatarsal III is significantly reduced in width in anterior view compared to metatarsal II and metatarsal IV (
|
|
<bibRefCitation author="Xu, X. & X. Wang & X. Wu" pageId="56" pageNumber="56" pagination="262 - 266" refId="ref32936" refString="Xu, X., X. Wang, and X. Wu. 1999. A dromaeosaurid dinosaur with a filamentous integument from the Jiufotang Formation of China. Nature 401: 262 - 266." type="journal article" year="1999">Xu et al., 1999</bibRefCitation>
|
|
, 2000, 2002; Xu, 2002;
|
|
<bibRefCitation author="Xu, X. & M. A. Norell" pageId="56" pageNumber="56" pagination="838 - 841" refId="ref32869" refString="Xu, X., and M. A. Norell. 2004. A new troodontid dinosaur from China with avian-like sleeping posture. Nature 431: 838 - 841." type="journal article" year="2004">Xu and Norell, 2004</bibRefCitation>
|
|
).
|
|
<taxonomicName box="[764,883,1184,1208]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[764,883,1184,1208]" italics="true" pageId="56" pageNumber="56">Anchiornis</emphasis>
|
|
</taxonomicName>
|
|
was also reported to have a deinonychosaurian-like pedal digit II (
|
|
<bibRefCitation author="Zheng, X. & H. You & X. Xu & Z. Dong" pageId="56" pageNumber="56" pagination="333 - 336" refId="ref33369" refString="Zheng, X., H. You, X. Xu, and Z. Dong. 2009. An Early Cretaceous heterodontosaurid dinosaur with filamentous integumentary structures. Nature 458: 333 - 336." type="journal article" year="2009">Xu et al., 2009</bibRefCitation>
|
|
;
|
|
<bibRefCitation author="Hu, D. & L. Hou & L. Zhang & X. Xu" box="[755,929,1251,1275]" pageId="56" pageNumber="56" pagination="640 - 643" refId="ref30594" refString="Hu, D., L. Hou, L. Zhang, and X. Xu. 2009. A pre- Archaeopteryx troodontid theropod from China with long feathers on the metatarsal. Nature 461: 640 - 643." type="journal article" year="2009">Hu et al., 2009</bibRefCitation>
|
|
). Careful examination of more specimens of
|
|
<taxonomicName box="[921,1041,1284,1308]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[921,1041,1284,1308]" italics="true" pageId="56" pageNumber="56">Anchiornis</emphasis>
|
|
</taxonomicName>
|
|
indicates that pedal digit II of these specimens lacks the typical specialized raptorial condition of deinonychosaurians. In
|
|
<taxonomicName box="[857,974,1383,1407]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[857,974,1383,1407]" italics="true" pageId="56" pageNumber="56">Anchiornis</emphasis>
|
|
</taxonomicName>
|
|
, pedal phalanx II-2 is not reduced in length and lacks a well-developed ventral heel or a constriction at midshaπ, which is different from deinonychosaurians (
|
|
<bibRefCitation author="Gauthier, J. A." pageId="56" pageNumber="56" pagination="1 - 55" refId="ref30373" refString="Gauthier, J. A. 1986. Saurischian monophyly and the origin of birds. In K. Padian (editor), The origin of birds and evolution of flight. 1 - 55. San Francisco: California Academy of Sciences Memoir." type="book chapter" year="1986">Gauthier, 1986</bibRefCitation>
|
|
;
|
|
<bibRefCitation author="Turner, A. H. & P. J. Makovicky & M. A. Norell" box="[752,960,1515,1539]" pageId="56" pageNumber="56" pagination="1 - 207" refId="ref32607" refString="Turner, A. H., P. J. Makovicky, and M. A. Norell. 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History 371: 1 - 207." type="journal article" year="2012">Turner et al., 2012</bibRefCitation>
|
|
). Ungual phalange II-3 of
|
|
<taxonomicName box="[716,834,1548,1572]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[716,834,1548,1572]" italics="true" pageId="56" pageNumber="56">Anchiornis</emphasis>
|
|
</taxonomicName>
|
|
is only slightly larger than other pedal ungual phalanges, not as distinctly enlarged as in deinonychosaurians. Although the pedal digit II of
|
|
<taxonomicName box="[914,1031,1647,1671]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="56" phylum="Chordata" rank="genus">
|
|
<emphasis box="[914,1031,1647,1671]" italics="true" pageId="56" pageNumber="56">Anchiornis</emphasis>
|
|
</taxonomicName>
|
|
is more derived than the configuration of basal maniraptoriforms (such as ornithomimosaurians, alva- rezsaurids, and oviraptorosaurians), it still exhibits the primitive features (as mentioned above) of paravians that is also observed in
|
|
<taxonomicName baseAuthorityName="Wellnhofer" baseAuthorityYear="2009" box="[108,267,326,350]" class="Aves" family="Archaeopterygidae" genus="Archaeopteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Archaeopterygiformes" pageId="57" pageNumber="57" phylum="Chordata" rank="genus">
|
|
<emphasis box="[108,267,326,350]" italics="true" pageId="57" pageNumber="57">Archaeopteryx</emphasis>
|
|
</taxonomicName>
|
|
(see
|
|
<bibRefCitation author="Turner, A. H. & P. J. Makovicky & M. A. Norell" box="[335,560,326,350]" pageId="57" pageNumber="57" pagination="1 - 207" refId="ref32607" refString="Turner, A. H., P. J. Makovicky, and M. A. Norell. 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History 371: 1 - 207." type="journal article" year="2012">Turner et al., 2012</bibRefCitation>
|
|
). As a contrast in deinonychosaurians, even the basal members of each lineage, such as
|
|
<taxonomicName box="[479,588,393,417]" class="Reptilia" family="Dromaeosauridae" genus="Mahakala" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="57" pageNumber="57" phylum="Chordata" rank="genus">
|
|
<emphasis box="[479,588,393,417]" italics="true" pageId="57" pageNumber="57">Mahakala</emphasis>
|
|
</taxonomicName>
|
|
and
|
|
<taxonomicName box="[109,237,427,450]" class="Reptilia" family="Troodontidae" genus="Sinovenator" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="57" pageNumber="57" phylum="Chordata" rank="genus">
|
|
<emphasis box="[109,237,427,450]" italics="true" pageId="57" pageNumber="57">Sinovenator</emphasis>
|
|
</taxonomicName>
|
|
have highly modified and hyperextensible pedal digit II that is different from
|
|
<taxonomicName box="[108,226,494,518]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="57" pageNumber="57" phylum="Chordata" rank="genus">
|
|
<emphasis box="[108,226,494,518]" italics="true" pageId="57" pageNumber="57">Anchiornis</emphasis>
|
|
</taxonomicName>
|
|
and
|
|
<taxonomicName baseAuthorityName="Wellnhofer" baseAuthorityYear="2009" box="[287,444,494,518]" class="Aves" family="Archaeopterygidae" genus="Archaeopteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Archaeopterygiformes" pageId="57" pageNumber="57" phylum="Chordata" rank="genus">
|
|
<emphasis box="[287,444,494,518]" italics="true" pageId="57" pageNumber="57">Archaeopteryx</emphasis>
|
|
</taxonomicName>
|
|
(
|
|
<bibRefCitation author="Xu, X. & M. A. Norell & X. Wang & P. J. Makovicky & X. Wu" box="[461,632,494,518]" pageId="57" pageNumber="57" pagination="780 - 784" refId="ref33003" refString="Xu, X., M. A. Norell, X. Wang, P. J. Makovicky, and X. Wu. 2002. A basal troodontid from the Early Cretaceous of China. Nature 415: 780 - 784." type="journal article" year="2002">Xu et al., 2002</bibRefCitation>
|
|
;
|
|
<bibRefCitation author="Turner, A. H. & D. Pol & J. A. Clarke & G. Erickson & M. A. Norell" box="[108,314,527,551]" pageId="57" pageNumber="57" pagination="1378 - 1381" refId="ref32515" refString="Turner, A. H., D. Pol., J. A. Clarke, G. Erickson, and M. A. Norell. 2007. A basal dromaeosaurid and size evolution preceding avian flight. Science 317: 1378 - 1381." type="journal article" year="2007">Turner et al., 2007</bibRefCitation>
|
|
,
|
|
<bibRefCitation author="Turner, A. H. & D. Pol & M. A. Norell" box="[325,380,527,551]" pageId="57" pageNumber="57" pagination="1 - 66" refId="ref32563" refString="Turner, A. H., D. Pol, and M. A. Norell. 2011. Anatomy of Mahakala omnogovae (Theropoda: Dromaeosauridae), Togrogiin Shiree, Mongolia. American Museum Novitates 3722: 1 - 66." type="journal article" year="2011">2011</bibRefCitation>
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,
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<bibRefCitation author="Turner, A. H. & P. J. Makovicky & M. A. Norell" box="[392,450,527,551]" pageId="57" pageNumber="57" pagination="1 - 207" refId="ref32607" refString="Turner, A. H., P. J. Makovicky, and M. A. Norell. 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History 371: 1 - 207." type="journal article" year="2012">2012</bibRefCitation>
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).
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</paragraph>
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<paragraph blockId="57.[108,637,226,920]" pageId="57" pageNumber="57">
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We consider
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<taxonomicName authorityName=", PKUP V" authorityYear="1068" box="[296,509,561,585]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="57" pageNumber="57" phylum="Chordata" rank="species">
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<emphasis box="[296,509,561,585]" italics="true" pageId="57" pageNumber="57">Anchiornis huxleyi</emphasis>
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</taxonomicName>
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as a basal avialan based on derived characters it shares with other avialans (
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<bibRefCitation author="Zheng, X. & H. You & X. Xu & Z. Dong" box="[331,491,628,652]" pageId="57" pageNumber="57" pagination="333 - 336" refId="ref33369" refString="Zheng, X., H. You, X. Xu, and Z. Dong. 2009. An Early Cretaceous heterodontosaurid dinosaur with filamentous integumentary structures. Nature 458: 333 - 336." type="journal article" year="2009">Xu et al., 2009</bibRefCitation>
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;
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<bibRefCitation author="Godefroit, P. & A. Cau & Hu D. - Y. & F. Escuillie & W. Wu & G. Dyke" pageId="57" pageNumber="57" pagination="1394" refId="ref30490" refString="Godefroit, P., A. Cau, Hu D. - Y., F. Escuillie, W. Wu, and G. Dyke. 2013 a. Reduced plumage and flight ability of a new paravian theropod from China. Nature Communications 4: 1394." type="journal article" year="2013">Godefroit et al., 2013a</bibRefCitation>
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;
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<bibRefCitation author="Foth, C. & H. Tischlinger & O. W. M. Rauhut" box="[230,418,661,685]" pageId="57" pageNumber="57" pagination="79 - 82" refId="ref30225" refString="Foth, C., H. Tischlinger, and O. W. M. Rauhut. 2014. New specimen of Archaeopteryx provides insights into the evolution of pennaceous feathers. Nature 511: 79 - 82." type="journal article" year="2014">Foth et al., 2014</bibRefCitation>
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), such as a straight nasal process of the premaxilla and the absence of an external mandibular fenestra in lateral view.
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<taxonomicName box="[172,289,762,786]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="57" pageNumber="57" phylum="Chordata" rank="genus">
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<emphasis box="[172,289,762,786]" italics="true" pageId="57" pageNumber="57">Anchiornis</emphasis>
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</taxonomicName>
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|
also lacks typical deinonychosaurian synapomorphies such as lateral exposure of the splenial, and a specialized raptorial pedal digit II, which differentiates
|
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<taxonomicName box="[517,636,862,886]" class="Reptilia" family="Troodontidae" genus="Anchiornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="57" pageNumber="57" phylum="Chordata" rank="genus">
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|
<emphasis box="[517,636,862,886]" italics="true" pageId="57" pageNumber="57">Anchiornis</emphasis>
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</taxonomicName>
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from deinonychosaurians.
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</paragraph>
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</subSubSection>
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</treatment>
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</document> |