414 lines
51 KiB
XML
414 lines
51 KiB
XML
<document ID-DOI="http://doi.org/10.5281/zenodo.3716970" ID-GBIF-Dataset="973fc06c-9ee3-425d-bb8a-b3368fb81ba2" ID-GBIF-Taxon="162645937" ID-Zenodo-Dep="3716970" LSID="urn:lsid:zoobank.org:act:8FE7BF29-AE0B-49B6-9EB2-0C0B3D13BAA3" checkinTime="1584628455610" checkinUser="plazi" docAuthor="Field, Daniel J., Benito, Juan, Chen, Albert, Jagt, John W. M. & Ksepka, Daniel T." docDate="2020" docId="039CDC2BFC5CFFEE36D2AD368BDDDAA0" docLanguage="en" docName="nature.s41586-020-2096-0.pdf" docOrigin="Nature 579" docStyle="DocumentStyle{}" docTitle="Asteriornis maastrichtensis Field & Benito & Chen & Jagt & Ksepka 2020, gen.et sp.nov." docType="treatment" docVersion="5" lastPageId="2" lastPageNumber="399" masterDocId="FFA5A453FC5CFFEC356DAA328E07DF59" masterDocTitle="Late Cretaceous neornithine from Europe illuminates the origins of crown birds" masterLastPageNumber="414" masterPageNumber="397" pageId="0" pageNumber="397" updateTime="1643440217241" 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>Late Cretaceous neornithine from Europe illuminates the origins of crown birds</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>Field, Daniel J.</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>Benito, Juan</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>Chen, Albert</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>Jagt, John W. M.</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>Ksepka, Daniel T.</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>2020</mods:date>
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<mods:detail type="pubDate">
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<mods:number>2020-03-18</mods:number>
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</mods:detail>
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<mods:detail type="volume">
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<mods:number>579</mods:number>
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</mods:detail>
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<mods:extent unit="page">
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<mods:start>397</mods:start>
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<mods:end>414</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/s41586-020-2096-0</mods:identifier>
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<mods:identifier type="GBIF-Dataset">973fc06c-9ee3-425d-bb8a-b3368fb81ba2</mods:identifier>
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<mods:identifier type="Zenodo-Dep">3716350</mods:identifier>
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</mods:mods>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.3716970" ID-GBIF-Taxon="162645937" ID-Zenodo-Dep="3716970" LSID="urn:lsid:zoobank.org:act:8FE7BF29-AE0B-49B6-9EB2-0C0B3D13BAA3" httpUri="http://treatment.plazi.org/id/039CDC2BFC5CFFEE36D2AD368BDDDAA0" lastPageId="2" lastPageNumber="399" pageId="0" pageNumber="397">
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<subSubSection box="[959,1378,1795,1816]" pageId="0" pageNumber="397" type="nomenclature">
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<paragraph blockId="0.[816,1497,1336,1902]" box="[959,1378,1795,1816]" pageId="0" pageNumber="397">
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<taxonomicName authority="Field & Benito & Chen & Jagt & Ksepka, 2020" authorityName="Field & Benito & Chen & Jagt & Ksepka" authorityYear="2020" box="[959,1228,1795,1816]" genus="Asteriornis" pageId="0" pageNumber="397" rank="species" species="maastrichtensis" status="gen.et sp.nov.">
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<emphasis box="[959,1228,1795,1816]" italics="true" pageId="0" pageNumber="397">Asteriornis maastrichtensis</emphasis>
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</taxonomicName>
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<emphasis bold="true" box="[1231,1378,1796,1816]" pageId="0" pageNumber="397">
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<taxonomicNameLabel box="[1231,1378,1796,1816]" pageId="0" pageNumber="397" rank="species">gen.et sp.nov.</taxonomicNameLabel>
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</emphasis>
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</paragraph>
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</subSubSection>
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<subSubSection lastPageId="1" lastPageNumber="398" pageId="0" pageNumber="397" type="discussion">
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<paragraph blockId="0.[816,1497,1336,1902]" lastBlockId="1.[106,788,1190,1988]" lastPageId="1" lastPageNumber="398" pageId="0" pageNumber="397">
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<emphasis bold="true" pageId="0" pageNumber="397">
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Remarks.We use Avialae to refer to theropods crownward of
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<taxonomicName class="Reptilia" family="Dromaeosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="397" phylum="Chordata" rank="family">Dromaeosauridae</taxonomicName>
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and
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<taxonomicName authorityName="Gilmore" authorityYear="1924" box="[945,1085,1881,1902]" class="Reptilia" family="Troodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="397" phylum="Chordata" rank="family">Troodontidae</taxonomicName>
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.
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<taxonomicName authorityName="Gadow" authorityYear="1892" box="[1090,1201,1881,1902]" class="Aves" higherTaxonomySource="GBIF" kingdom="Metazoa" pageId="0" pageNumber="397" phylum="Chordata" rank="subClass" subClass="Neornithes">Neornithes</taxonomicName>
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is equivalent to the bird crown
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</emphasis>
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<heading bold="true" box="[106,787,1190,1215]" centered="true" fontSize="8" level="3" pageId="1" pageNumber="398" reason="0">
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<emphasis bold="true" box="[106,427,1193,1214]" pageId="1" pageNumber="398">
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group (
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<taxonomicName authority="sensu Gauthier, 1986" authorityName="sensu Gauthier" authorityYear="1986" box="[174,427,1193,1214]" class="Aves" higherTaxonomySource="GBIF" kingdom="Metazoa" pageId="1" pageNumber="398" phylum="Chordata" rank="class">Aves sensu Gauthier,1986</taxonomicName>
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</emphasis>
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<bibRefCitation author="Gauthier, J. A. & de Queiroz, K." box="[428,442,1190,1203]" editor="Gauthier, J. & Gall, L. F." journalOrPublisher="Peabody Museum of Natural History, Yale University" pageId="1" pageNumber="398" pagination="7 - 41" refId="ref4673" refString="23. Gauthier, J. A. & de Queiroz, K. in New Perspectives on the Origin and Early Evolution of Birds: Proceedings of the International Symposium in Honor of John H. Ostrom (eds Gauthier, J. & Gall, L. F.) 7 - 41 (Peabody Museum of Natural History, Yale University, 2001)." type="book chapter" volumeTitle="New Perspectives on the Origin and Early Evolution of Birds: Proceedings of the International Symposium in Honor of John H. Ostrom" year="2001">
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<emphasis bold="true" box="[428,442,1190,1203]" pageId="1" pageNumber="398">
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<superScript attach="right" box="[428,442,1190,1203]" fontSize="5" pageId="1" pageNumber="398">23</superScript>
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</emphasis>
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</bibRefCitation>
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<emphasis bold="true" pageId="1" pageNumber="398">).Pangalloanserae defines the most inclusive clade including</emphasis>
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</heading>
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<taxonomicName authority="anser" box="[356,470,1223,1243]" class="Aves" family="Anatidae" genus="Anser" kingdom="Animalia" order="Anseriformes" pageId="1" pageNumber="398" phylum="Chordata" rank="species" species="anser">
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<emphasis box="[356,470,1223,1243]" italics="true" pageId="1" pageNumber="398">
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<taxonomicName box="[356,412,1223,1243]" class="Aves" family="Anatidae" genus="Anser" higherTaxonomySource="CoL" kingdom="Animalia" order="Anseriformes" pageId="1" pageNumber="398" phylum="Chordata" rank="genus">Anser</taxonomicName>
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anser
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</emphasis>
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</taxonomicName>
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<emphasis bold="true" box="[474,512,1222,1243]" pageId="1" pageNumber="398">and</emphasis>
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<taxonomicName authority="gallus" box="[515,642,1221,1243]" class="Aves" family="Phasianidae" genus="Gallus" kingdom="Animalia" order="Galliformes" pageId="1" pageNumber="398" phylum="Chordata" rank="species" species="gallus">
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<emphasis box="[515,642,1221,1243]" italics="true" pageId="1" pageNumber="398">Gallus gallus</emphasis>
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</taxonomicName>
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<emphasis bold="true" box="[646,719,1222,1243]" pageId="1" pageNumber="398">but not</emphasis>
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<taxonomicName baseAuthorityName="Linnaeus" baseAuthorityYear="1758" class="Aves" family="Passeridae" genus="Passer" kingdom="Animalia" order="Passeriformes" pageId="1" pageNumber="398" phylum="Chordata" rank="species" species="domesticus">
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<emphasis italics="true" pageId="1" pageNumber="398">Passer domesticus</emphasis>
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</taxonomicName>
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<emphasis bold="true" pageId="1" pageNumber="398">(that is,the galloanseran total group).Further phylogenetic definitions are presented in the Supplementary Information.</emphasis>
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</paragraph>
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</subSubSection>
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<footnote pageId="0" pageNumber="397">
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<paragraph blockId="0.[106,1465,1945,1987]" pageId="0" pageNumber="397">
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<superScript attach="right" box="[106,110,1945,1954]" fontSize="4" pageId="0" pageNumber="397">1</superScript>
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Department of Earth Sciences,University of Cambridge,Cambridge, UK.
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<bibRefCitation author="Jarvis, E. D." box="[639,644,1945,1954]" journalOrPublisher="Science" pageId="0" pageNumber="397" pagination="1320 - 1331" part="346" refId="ref3816" refString="2. Jarvis, E. D. et al. Whole-genome analyses resolve early branches in the tree of life of modern birds. Science 346, 1320 - 1331 (2014)." title="Whole-genome analyses resolve early branches in the tree of life of modern birds" type="journal article" year="2014">
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<superScript attach="right" box="[639,644,1945,1954]" fontSize="4" pageId="0" pageNumber="397">2</superScript>
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</bibRefCitation>
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Department of Biology & Biochemistry,Milner Centre for Evolution,University of Bath,Bath,UK.
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<bibRefCitation author="Claramunt, S. & Cracraft, J." box="[1338,1344,1945,1954]" journalOrPublisher="Sci. Adv." pageId="0" pageNumber="397" pagination="1501005" part="1" refId="ref3851" refString="3. Claramunt, S. & Cracraft, J. A new time tree reveals Earth history's imprint on the evolution of modern birds. Sci. Adv. 1, e 1501005 (2015)." title="A new time tree reveals Earth history's imprint on the evolution of modern birds" type="journal article" year="2015">
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<superScript attach="right" box="[1338,1344,1945,1954]" fontSize="4" pageId="0" pageNumber="397">3</superScript>
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</bibRefCitation>
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Natuurhistorisch Museum Maastricht,Maastricht,The Netherlands.
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<bibRefCitation author="Field, D. J." box="[471,477,1969,1978]" journalOrPublisher="Curr. Biol." pageId="0" pageNumber="397" pagination="1825 - 1831" part="28" refId="ref3889" refString="4. Field, D. J. et al. Early evolution of modern birds structured by global forest collapse at the end-Cretaceous mass extinction. Curr. Biol. 28, 1825 - 1831 (2018)." title="Early evolution of modern birds structured by global forest collapse at the end-Cretaceous mass extinction" type="journal article" year="2018">
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<superScript attach="right" box="[471,477,1969,1978]" fontSize="4" pageId="0" pageNumber="397">4</superScript>
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</bibRefCitation>
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Bruce Museum, Greenwich,CT,USA.
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<emphasis bold="true" box="[746,757,1967,1982]" pageId="0" pageNumber="397">✉</emphasis>
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e-mail:djf70@cam.ac.uk
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</paragraph>
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</footnote>
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<caption ID-DOI="http://doi.org/10.5281/zenodo.3716352" ID-Zenodo-Dep="3716352" httpUri="https://zenodo.org/record/3716352/files/figure.png" pageId="1" pageNumber="398" startId="1.[106,137,960,978]" targetBox="[144,1459,131,935]" targetPageId="1">
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<paragraph blockId="1.[106,774,960,1112]" pageId="1" pageNumber="398">
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Fig.1 | Digitally segmented skull of
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<taxonomicName authorityName="Field & Benito & Chen & Jagt & Ksepka" authorityYear="2020" box="[416,666,960,978]" genus="Asteriornis" pageId="1" pageNumber="398" rank="species" species="maastrichtensis">
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<emphasis bold="true" box="[416,666,960,978]" italics="true" pageId="1" pageNumber="398">Asteriornis maastrichtensis</emphasis>
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</taxonomicName>
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.
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<emphasis bold="true" box="[106,704,986,1005]" pageId="1" pageNumber="398">a–c,Left lateral view (a), right lateral view (b) and dorsal view (c) of the</emphasis>
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<taxonomicName authorityName="Field & Benito & Chen & Jagt & Ksepka" authorityYear="2020" box="[106,265,1013,1032]" genus="Asteriornis" pageId="1" pageNumber="398" rank="species" species="maastrichtensis">
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<emphasis box="[106,265,1013,1032]" italics="true" pageId="1" pageNumber="398">A.maastrichtensis</emphasis>
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</taxonomicName>
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<emphasis bold="true" pageId="1" pageNumber="398">skull.d,Comparison of retroarticular processes in lateral view of</emphasis>
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<emphasis box="[170,328,1040,1059]" italics="true" pageId="1" pageNumber="398">Anatalavis oxfordi</emphasis>
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<emphasis bold="true" box="[330,461,1040,1059]" pageId="1" pageNumber="398">(right side) and</emphasis>
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<taxonomicName box="[464,558,1041,1058]" genus="Asteriornis" pageId="1" pageNumber="398" rank="genus">
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<emphasis box="[464,558,1041,1058]" italics="true" pageId="1" pageNumber="398">Asteriornis</emphasis>
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</taxonomicName>
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<emphasis bold="true" pageId="1" pageNumber="398">
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(left side,reflected). Selected synapomorphies are denoted with numbers (red,
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<taxonomicName authorityName="Gadow" authorityYear="1892" box="[610,712,1066,1085]" class="Aves" higherTaxonomySource="GBIF" kingdom="Metazoa" pageId="1" pageNumber="398" phylum="Chordata" rank="subClass" subClass="Neornithes">Neornithes</taxonomicName>
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;green, Galloanserae
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</emphasis>
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<emphasis bold="true" box="[221,251,1091,1101]" pageId="1" pageNumber="398">
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<superScript attach="right" box="[221,251,1091,1101]" fontSize="4" pageId="1" pageNumber="398">
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<bibRefCitation author="Cracraft, J." box="[221,232,1091,1101]" editor="Benton, M. J." journalOrPublisher="Oxford Univ. Press" pageId="1" pageNumber="398" pagination="339 - 361" refId="ref5416" refString="37. Cracraft, J. in The Phylogeny and Classification of the Tetrapods Vol. 1 (ed. Benton, M. J.) 339 - 361 (Oxford Univ. Press, 1988)." type="book chapter" volumeTitle="The Phylogeny and Classification of the Tetrapods" year="1988">37</bibRefCitation>
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–
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<bibRefCitation author="Cracraft, J. & Clarke, J." box="[238,251,1091,1101]" editor="Gauthier, J. & Gall, L. F." journalOrPublisher="Peabody Museum of Natural History, Yale University" pageId="1" pageNumber="398" pagination="143 - 156" refId="ref5498" refString="39. Cracraft, J. & Clarke, J. The basal clades of modern birds. In New Perspectives on the Origin and Early Evolution of Birds: Proceedings of the International Symposium in Honor of John H. Ostrom (eds Gauthier, J. & Gall, L. F.) 143 - 156 (Peabody Museum of Natural History, Yale University, 2001)." title="The basal clades of modern birds" type="book chapter" volumeTitle="New Perspectives on the Origin and Early Evolution of Birds: Proceedings of the International Symposium in Honor of John H. Ostrom" year="2001">39</bibRefCitation>
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</superScript>
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</emphasis>
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<emphasis bold="true" box="[251,728,1093,1112]" pageId="1" pageNumber="398">):1,bony mandibular symphysis;2,toothless beak;3,no</emphasis>
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</paragraph>
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<paragraph blockId="1.[816,1491,960,1084]" pageId="1" pageNumber="398">
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<emphasis bold="true" pageId="1" pageNumber="398">coronoid bone;4,bicondylar mandibular process of quadrate;5,retroarticular process long,curving,strongly compressed mediolaterally;6,dorsally oriented internal articular process of mandible;7,maxillary process of premaxilla dorsoventrally deep and lateromedially compressed.Scale bars,</emphasis>
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<emphasis bold="true" box="[816,869,1067,1084]" pageId="1" pageNumber="398">5 mm.</emphasis>
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</paragraph>
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</caption>
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<subSubSection pageId="1" pageNumber="398" type="etymology">
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<paragraph blockId="1.[106,788,1190,1988]" pageId="1" pageNumber="398">
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<emphasis bold="true" box="[106,228,1309,1329]" pageId="1" pageNumber="398">Etymology.</emphasis>
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<taxonomicName box="[232,341,1309,1329]" genus="Asteriornis" pageId="1" pageNumber="398" rank="genus">
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<emphasis box="[232,341,1309,1329]" italics="true" pageId="1" pageNumber="398">Asteriornis</emphasis>
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</taxonomicName>
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,
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<emphasis bold="true" box="[351,710,1307,1329]" pageId="1" pageNumber="398">from the name of the Titan goddess</emphasis>
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<taxonomicName authority="Felder, 1874" authorityName="Felder" authorityYear="1874" box="[715,786,1309,1329]" class="Insecta" family="Limacodidae" genus="Asteria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="1" pageNumber="398" phylum="Arthropoda" rank="genus">
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<emphasis box="[715,786,1309,1329]" italics="true" pageId="1" pageNumber="398">Asteria</emphasis>
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</taxonomicName>
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<emphasis bold="true" box="[106,241,1337,1358]" pageId="1" pageNumber="398">and the Greek</emphasis>
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<emphasis box="[244,293,1338,1358]" italics="true" pageId="1" pageNumber="398">ornis</emphasis>
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||
<emphasis bold="true" pageId="1" pageNumber="398">
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||
for bird.In Greek mythology
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<taxonomicName authority="Felder, 1874" authorityName="Felder" authorityYear="1874" box="[576,646,1338,1358]" class="Insecta" family="Limacodidae" genus="Asteria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="1" pageNumber="398" phylum="Arthropoda" rank="genus">Asteria</taxonomicName>
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is the goddess of falling stars and transforms herself into a quail—attributes that are reflected by both the impending Cretaceous–Palaeogene(K–Pg) asteroid impact and the galloanseran affinities of
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||
</emphasis>
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||
<taxonomicName box="[548,655,1424,1444]" genus="Asteriornis" pageId="1" pageNumber="398" rank="genus">
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<emphasis box="[548,655,1424,1444]" italics="true" pageId="1" pageNumber="398">Asteriornis</emphasis>
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</taxonomicName>
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||
.
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||
<emphasis bold="true" pageId="1" pageNumber="398">The specific epithet</emphasis>
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||
<taxonomicName authorityName="Field & Benito & Chen & Jagt & Ksepka" authorityYear="2020" box="[183,341,1451,1472]" genus="Asteriornis" pageId="1" pageNumber="398" rank="species" species="maastrichtensis">
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||
<emphasis box="[183,341,1451,1472]" italics="true" pageId="1" pageNumber="398">maastrichtensis</emphasis>
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</taxonomicName>
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<emphasis bold="true" pageId="1" pageNumber="398">reflects the provenance of the holotype—the Maastricht Formation (the type locality of the Late Cretaceous Maastrichtian stage).</emphasis>
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</paragraph>
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</subSubSection>
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<subSubSection pageId="1" pageNumber="398" type="materials_examined">
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<materialsCitation ID-GBIF-Occurrence="2578927073" collectionCode="NHMM" country="Belgium" location="CBR-Romontbos Quarry, Eben-Emael, Liege" pageId="1" pageNumber="398" specimenCode="2013 008" specimenCount="1" typeStatus="holotype">
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||
<paragraph blockId="1.[106,788,1190,1988]" pageId="1" pageNumber="398">
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||
<emphasis bold="true" pageId="1" pageNumber="398">
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||
<typeStatus box="[106,207,1538,1558]" pageId="1" pageNumber="398">Holotype</typeStatus>
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. Natuurhistorisch Museum Maastricht (
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<collectionCode box="[600,677,1537,1558]" collectionName="Germany, Mainz, Naturhistorisches Museum Mainz" pageId="1" pageNumber="398">NHMM</collectionCode>
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)
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||
<specimenCode box="[687,783,1538,1558]" pageId="1" pageNumber="398">2013 008</specimenCode>
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, a nearly complete, articulated skull including mandibles and associated postcranial remains preserved in four blocks (
|
||
<figureCitation box="[624,677,1594,1616]" captionStart="Fig" captionStartId="1.[106,137,960,978]" captionTargetBox="[144,1459,131,935]" captionTargetId="figure@1.[519,896,488,847]" captionTargetPageId="1" captionText="Fig.1 | Digitallysegmented skull ofAsteriornis maastrichtensis. a–c,Left lateral view (a), rightlateral view (b) anddorsal view (c) of the A.maastrichtensisskull.d,Comparison of retroarticular processes inlateral view of Anatalavis oxfordi(rightside) and Asteriornis(left side,reflected). Selected synapomorphies aredenoted withnumbers (red,Neornithes;green, Galloanserae37–39):1,bony mandibular symphysis;2,toothless beak;3,no coronoid bone;4,bicondylar mandibularprocess of quadrate;5,retroarticular process long,curving,stronglycompressed mediolaterally;6,dorsally oriented internal articular process of mandible;7,maxillary process of premaxilla dorsoventrallydeep and lateromediallycompressed.Scale bars, 5 mm." figureDoi="http://doi.org/10.5281/zenodo.3716352" httpUri="https://zenodo.org/record/3716352/files/figure.png" pageId="1" pageNumber="398">Fig. 1</figureCitation>
|
||
, Extended Data
|
||
<figureCitation box="[157,244,1624,1644]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStart-3="Extended" captionStart-4="Extended" captionStart-5="Extended" captionStartId-0="1.[106,137,960,978]" captionStartId-1="2.[106,137,1074,1092]" captionStartId-2="2.[816,847,1044,1062]" captionStartId-3="11.[106,193,1769,1787]" captionStartId-4="12.[106,193,1769,1787]" captionStartId-5="13.[106,193,1753,1771]" captionTargetBox-0="[144,1459,131,935]" captionTargetBox-1="[111,781,130,1049]" captionTargetBox-2="[821,1469,136,1017]" captionTargetBox-3="[203,1399,132,1743]" captionTargetBox-4="[233,1370,131,1744]" captionTargetBox-5="[122,1481,131,1728]" captionTargetId-0="figure@1.[519,896,488,847]" captionTargetId-1="figure@2.[109,494,497,865]" captionTargetId-2="graphics@2.[834,1470,245,1013]" captionTargetId-3="figure@11.[203,1399,131,1744]" captionTargetId-4="figure@12.[233,1370,131,1744]" captionTargetId-5="figure@13.[121,1481,131,1729]" captionTargetPageId-0="1" captionTargetPageId-1="2" captionTargetPageId-2="2" captionTargetPageId-3="11" captionTargetPageId-4="12" captionText-0="Fig.1 | Digitallysegmented skull ofAsteriornis maastrichtensis. a–c,Left lateral view (a), rightlateral view (b) anddorsal view (c) of the A.maastrichtensisskull.d,Comparison of retroarticular processes inlateral view of Anatalavis oxfordi(rightside) and Asteriornis(left side,reflected). Selected synapomorphies aredenoted withnumbers (red,Neornithes;green, Galloanserae37–39):1,bony mandibular symphysis;2,toothless beak;3,no coronoid bone;4,bicondylar mandibularprocess of quadrate;5,retroarticular process long,curving,stronglycompressed mediolaterally;6,dorsally oriented internal articular process of mandible;7,maxillary process of premaxilla dorsoventrallydeep and lateromediallycompressed.Scale bars, 5 mm." captionText-1="Fig.2| Comparativequadrate and skull morphology ofselected total-group Galloanserae.a–d, Left quadrates(andreflected rightquadrate of Presbyornis) inmedial view (left) andcaudal view (middle).Skulls arein right lateral view,except Alectura which is inreflected left lateral view.The skull of PresbyornisUSNM 299846 is shown.Scale bars,5mm (quadrates);1cm (skulls). See Extended DataFig.4 for additional high-resolution images of these taxa andother extant and fossil Galloanserae." captionText-2="Fig.3 | Relationships ofA. maastrichtensisand stratigraphic provenanceof holotype.a, Cladogram showing the phylogeneticposition of Asteriornis and Vegavis inferred under parsimony(solid coloured lines) and tip-dated Bayesian analyses (dashedlines).A position forVegavis allied withAnseriformes,as previously found12,28,34, isalso shown (curved branch).Numbers withinclades denote extant species richness.The analysis is based ona newlymodified dataset;see Extended Data Fig.9 andSupplementary Information for full phylogenetic results.Extinct taxa aredenoted withdaggers.b, Simplified stratigraphy of the Maastricht Formation,exposed inthe vicinityof the holotype localityand surrounding areas24. c, Locationof theCBR-Romontbos Quarry near Eben-Emael,Belgium." captionText-3="Extended DataFig.5 | Detailed comparisonsofgalloanseran retroarticular morphology.Retroarticularregions of left mandiblesin lateral (left),medial (middle) and dorsal (right)views.Both the left andrightmandibles of Asteriornis areshown in dorsal view,as theretroarticular process is only preserved on theleft mandible andthe medial processis only preserved onthe right mandible.Imagesof Anatalavis are mirrored.Scalebars,1 cm." captionText-4="Extended DataFig.6 | Postcranial morphology of A.maastrichtensis(NHMM2013008). Theleft distal femur of Presbyornispervetus(UW 27596) is shownfor comparison." captionText-5="Extended DataFig.7| Internalcomposition ofNHMM 2013008 blocks. a, Block containing the left femur,left tibiotarsusand the mainportionof the skull,viewed from theside withthe femur exposed.b, Same blockas ina, viewed from theside containing thetibiotarsus.c, Block containing theright femur andtarsometatarsus.d, Block containing therightdistal radius,several unidentified bonefragments and a portionof thecranial roofnear the frontoparietal suture.e,Block containing the righttibiotarsus.Numerous fragments of fossil echinoderms andmolluscs are visiblewithin theblocks. Scale bars,1 cm." figureDoi-0="http://doi.org/10.5281/zenodo.3716352" figureDoi-1="http://doi.org/10.5281/zenodo.3716358" figureDoi-2="http://doi.org/10.5281/zenodo.3716354" figureDoi-3="http://doi.org/10.5281/zenodo.3716369" figureDoi-4="http://doi.org/10.5281/zenodo.3716371" figureDoi-5="http://doi.org/10.5281/zenodo.3716373" httpUri-0="https://zenodo.org/record/3716352/files/figure.png" httpUri-1="https://zenodo.org/record/3716358/files/figure.png" httpUri-2="https://zenodo.org/record/3716354/files/figure.png" httpUri-3="https://zenodo.org/record/3716369/files/figure.png" httpUri-4="https://zenodo.org/record/3716371/files/figure.png" httpUri-5="https://zenodo.org/record/3716373/files/figure.png" pageId="1" pageNumber="398">Figs.1–7</figureCitation>
|
||
;see Supplementary Information for videos,character information,measurements,additional description and discussion). Preserved elements include the premaxillae,maxillae,nasals,frontals, laterosphenoid,basisphenoid,mesethmoid, left quadrate,left jugal, right palatine and lower jaws.Associated postcranial elements include incomplete femora,tibiotarsi,tarsometatarsus and radius.
|
||
</emphasis>
|
||
</paragraph>
|
||
<paragraph blockId="1.[106,788,1190,1988]" pageId="1" pageNumber="398">
|
||
<emphasis bold="true" pageId="1" pageNumber="398">
|
||
Locality and age.
|
||
<location LSID="urn:lsid:plazi:treatment:039CDC2BFC5CFFEE36D2AD368BDDDAA0:8EEA3BE6FC5DFFED3479AD318CB1D840" box="[276,694,1795,1817]" country="Belgium" name="CBR-Romontbos Quarry, Eben-Emael, Liege" pageId="1" pageNumber="398">CBR-Romontbos Quarry,Eben-Emael,Liège</location>
|
||
,
|
||
<collectingCountry box="[700,784,1795,1817]" name="Belgium" pageId="1" pageNumber="398">Belgium</collectingCountry>
|
||
.
|
||
<location LSID="urn:lsid:plazi:treatment:039CDC2BFC5CFFEE36D2AD368BDDDAA0:8EEA3BE6FC5DFFED3507AD2D8FD3D808" country="Belgium" name="Valkenburg Member (66.8 - 66.7 million years old 24), Maastricht Formation, Late Maastrichtian, Cretaceous" pageId="1" pageNumber="398">
|
||
Valkenburg Member(66.8–66.7million years old
|
||
<bibRefCitation author="Keutgen, N." box="[576,590,1820,1833]" journalOrPublisher="Neth. J. Geosci." pageId="1" pageNumber="398" pagination="229 - 260" part="97" refId="ref4741" refString="24. Keutgen, N. A bioclast-based astronomical timescale for the Maastrichtian in the type area (southeast Netherlands, northeast Belgium) and stratigraphic implications: the legacy of PJ Felder. Neth. J. Geosci. 97, 229 - 260 (2018)." title="A bioclast-based astronomical timescale for the Maastrichtian in the type area (southeast Netherlands, northeast Belgium) and stratigraphic implications: the legacy of PJ Felder" type="journal article" year="2018">
|
||
<emphasis bold="true" box="[576,590,1820,1833]" pageId="1" pageNumber="398">
|
||
<superScript attach="left" box="[576,590,1820,1833]" fontSize="5" pageId="1" pageNumber="398">24</superScript>
|
||
</emphasis>
|
||
</bibRefCitation>
|
||
<emphasis bold="true" pageId="1" pageNumber="398">), Maastricht Formation,Late Maastrichtian,Cretaceous.Additional details regarding the locality and stratigraphic setting are presented in the Supplementary Information.</emphasis>
|
||
</location>
|
||
</emphasis>
|
||
</paragraph>
|
||
</materialsCitation>
|
||
</subSubSection>
|
||
<subSubSection pageId="1" pageNumber="398" type="diagnosis">
|
||
<paragraph blockId="1.[106,788,1190,1988]" lastBlockId="1.[816,1498,1193,1272]" pageId="1" pageNumber="398">
|
||
<emphasis bold="true" box="[106,216,1940,1960]" pageId="1" pageNumber="398">Diagnosis.</emphasis>
|
||
<taxonomicName box="[219,326,1940,1960]" genus="Asteriornis" pageId="1" pageNumber="398" rank="genus">
|
||
<emphasis box="[219,326,1940,1960]" italics="true" pageId="1" pageNumber="398">Asteriornis</emphasis>
|
||
</taxonomicName>
|
||
<emphasis bold="true" pageId="1" pageNumber="398">is unique among known taxa in exhibiting caudally pointed nasals that overlie the frontals and meet at the midline</emphasis>
|
||
<emphasis bold="true" pageId="1" pageNumber="398">of the skull,and a slightly rounded, unhooked tip of the premaxilla. Additional character combinations from phylogenetic analyses that dif- ferentiate</emphasis>
|
||
<taxonomicName box="[915,1020,1252,1272]" genus="Asteriornis" pageId="1" pageNumber="398" rank="genus">
|
||
<emphasis box="[915,1020,1252,1272]" italics="true" pageId="1" pageNumber="398">Asteriornis</emphasis>
|
||
</taxonomicName>
|
||
<emphasis bold="true" box="[1023,1498,1250,1272]" pageId="1" pageNumber="398">are presented in the Supplementary Information.</emphasis>
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection lastPageId="2" lastPageNumber="399" pageId="1" pageNumber="398" type="description">
|
||
<paragraph blockId="1.[816,1498,1329,1988]" box="[816,963,1329,1354]" pageId="1" pageNumber="398">
|
||
<heading bold="true" box="[816,963,1329,1354]" fontSize="10" level="1" pageId="1" pageNumber="398" reason="0">
|
||
<emphasis bold="true" box="[816,963,1329,1354]" pageId="1" pageNumber="398">Description</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="1.[816,1498,1329,1988]" pageId="1" pageNumber="398">
|
||
<taxonomicName authorityName="Field & Benito & Chen & Jagt & Ksepka" authorityYear="2020" box="[816,1090,1365,1386]" genus="Asteriornis" pageId="1" pageNumber="398" rank="species" species="maastrichtensis">
|
||
<emphasis box="[816,1090,1365,1386]" italics="true" pageId="1" pageNumber="398">
|
||
<taxonomicName box="[816,925,1366,1386]" genus="Asteriornis" pageId="1" pageNumber="398" rank="genus">Asteriornis</taxonomicName>
|
||
maastrichtensis
|
||
</emphasis>
|
||
</taxonomicName>
|
||
<emphasis bold="true" pageId="1" pageNumber="398">is a small pan-galloanseran, with an estimated mean body mass from hindlimb scaling regressions of 394 g (around the 21st percentile among extant Galloanserae</emphasis>
|
||
<bibRefCitation author="Field, D. J. & Lynner, C. & Brown, C. & Darroch, S. A. F." box="[1374,1388,1419,1432]" journalOrPublisher="PLoS One" pageId="1" pageNumber="398" pagination="82000" part="8" refId="ref4790" refString="25. Field, D. J., Lynner, C., Brown, C. & Darroch, S. A. F. Skeletal correlates for body mass estimation in modern and fossil flying birds. PLoS One 8, e 82000 (2013)." title="Skeletal correlates for body mass estimation in modern and fossil flying birds" type="journal article" year="2013">
|
||
<emphasis bold="true" box="[1374,1388,1419,1432]" pageId="1" pageNumber="398">
|
||
<superScript attach="left" box="[1374,1388,1419,1432]" fontSize="5" pageId="1" pageNumber="398">25</superScript>
|
||
</emphasis>
|
||
</bibRefCitation>
|
||
<emphasis bold="true" pageId="1" pageNumber="398">
|
||
; Extended Data
|
||
<figureCitation box="[865,919,1451,1472]" captionStart="Extended" captionStartId="14.[106,193,906,924]" captionTargetBox="[126,1479,136,878]" captionTargetId="figure@14.[121,1481,131,881]" captionTargetPageId="14" captionText="Extended DataFig.8 | Relativebody sizeofA.maastrichtensis. Estimateof themean body sizeof the Asteriornisholotype25 comparedwith extant Galloanserae82, rankedon the xaxis from smallest tolargest.Themean body- size estimatefor Asteriornis (394g)is closest tothat of male Anas crecca(392 g; 7.8thpercentile amongAnseriformes) andfemale Perdixperdix (393 g;33rd percentile among Galliformes)." figureDoi="http://doi.org/10.5281/zenodo.3716375" httpUri="https://zenodo.org/record/3716375/files/figure.png" pageId="1" pageNumber="398">Fig.8</figureCitation>
|
||
).Complete measurements of NHMM 2013 008 are provided in the Supplementary Information.
|
||
</emphasis>
|
||
</paragraph>
|
||
<paragraph blockId="1.[816,1498,1329,1988]" pageId="1" pageNumber="398">
|
||
<emphasis bold="true" pageId="1" pageNumber="398">
|
||
Most major cranial components are in their original anatomical positions.The general appearance of the premaxillary beak resembles that of extant
|
||
<taxonomicName box="[911,1034,1566,1587]" class="Aves" higherTaxonomySource="GBIF" kingdom="Animalia" order="Galliformes" pageId="1" pageNumber="398" phylum="Chordata" rank="order">Galliformes</taxonomicName>
|
||
,particularly in its gently down-curved tip and delicate construction,with no ossified joints among the rostral components
|
||
</emphasis>
|
||
<bibRefCitation author="Olson, S. L. & Feduccia, A." box="[900,915,1620,1633]" journalOrPublisher="Smithson. Contrib. Zool." pageId="1" pageNumber="398" pagination="1 - 24" part="323" refId="ref4840" refString="26. Olson, S. L. & Feduccia, A. Presbyornis and the origin of the Anseriformes (Aves: Charadriomorphae). Smithson. Contrib. Zool. 323, 1 - 24 (1980)." title="Presbyornis and the origin of the Anseriformes (Aves: Charadriomorphae)" type="journal article" year="1980">
|
||
<emphasis bold="true" box="[900,915,1620,1633]" pageId="1" pageNumber="398">
|
||
<superScript attach="right" box="[900,915,1620,1633]" fontSize="5" pageId="1" pageNumber="398">26</superScript>
|
||
</emphasis>
|
||
</bibRefCitation>
|
||
<emphasis bold="true" pageId="1" pageNumber="398">. The contralateral frontal processes of the premaxillae are unfused along their length,and the premaxillae and nasals are unfused at both their tomial and narial contacts. The beak tip is unhooked, which distinguishes</emphasis>
|
||
<taxonomicName box="[1013,1118,1710,1730]" genus="Asteriornis" pageId="1" pageNumber="398" rank="genus">
|
||
<emphasis box="[1013,1118,1710,1730]" italics="true" pageId="1" pageNumber="398">Asteriornis</emphasis>
|
||
</taxonomicName>
|
||
<emphasis bold="true" pageId="1" pageNumber="398">
|
||
from most Galloanserae except certain
|
||
<taxonomicName box="[816,906,1738,1759]" class="Aves" family="Anatidae" kingdom="Animalia" order="Anseriformes" pageId="1" pageNumber="398" phylum="Chordata" rank="family">Anatidae</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="Wetmore" authorityYear="1926" box="[952,1129,1738,1759]" class="Aves" family="Presbyornithidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anseriformes" pageId="1" pageNumber="398" phylum="Chordata" rank="family">Presbyornithidae</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
</paragraph>
|
||
<paragraph blockId="1.[816,1498,1329,1988]" pageId="1" pageNumber="398">
|
||
<emphasis bold="true" pageId="1" pageNumber="398">
|
||
The skull lacks a distinct nasofrontal hinge.As such,the architecture of this region more closely resembles that of extant
|
||
<taxonomicName box="[1269,1381,1795,1816]" class="Aves" higherTaxonomySource="GBIF" kingdom="Animalia" order="Galliformes" pageId="1" pageNumber="398" phylum="Chordata" rank="order">Galliformes</taxonomicName>
|
||
than
|
||
<taxonomicName class="Aves" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anseriformes" pageId="1" pageNumber="398" phylum="Chordata" rank="order">Anseriformes</taxonomicName>
|
||
.At the midpoint of the orbits the frontals are constricted,yielding an hourglass-shaped cranial roof with wider rostral and caudal extremities. Indorsolateralview,therightpostorbitalprocesssweepsstronglyventrally beforedeflectingrostrallytodefinepartoftheventralmarginoftheorbit.
|
||
</emphasis>
|
||
</paragraph>
|
||
<paragraph blockId="1.[816,1498,1329,1988]" pageId="1" pageNumber="398">
|
||
<emphasis bold="true" pageId="1" pageNumber="398">The left quadrate is well-preserved in three dimensions and is generally similar to the quadrate of the fossil pan-anseriform</emphasis>
|
||
<taxonomicName authorityName="Wetmore" authorityYear="1926" box="[1367,1482,1967,1988]" class="Aves" family="Presbyornithidae" genus="Presbyornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Eoenantiornithiformes" pageId="1" pageNumber="398" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1367,1482,1967,1988]" italics="true" pageId="1" pageNumber="398">Presbyornis</emphasis>
|
||
</taxonomicName>
|
||
<bibRefCitation author="Elzanowski, A. & Stidham, T. A." box="[1483,1497,1964,1977]" journalOrPublisher="J. Morphol." pageId="1" pageNumber="398" pagination="305 - 323" part="271" refId="ref4881" refString="27. Elzanowski, A. & Stidham, T. A. Morphology of the quadrate in the Eocene anseriform Presbyornis and extant galloanserine birds. J. Morphol. 271, 305 - 323 (2010)." title="Morphology of the quadrate in the Eocene anseriform Presbyornis and extant galloanserine birds" type="journal article" year="2010">
|
||
<emphasis bold="true" box="[1483,1497,1964,1977]" pageId="1" pageNumber="398">
|
||
<superScript attach="right" box="[1483,1497,1964,1977]" fontSize="5" pageId="1" pageNumber="398">27</superScript>
|
||
</emphasis>
|
||
</bibRefCitation>
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.3716354" ID-Zenodo-Dep="3716354" httpUri="https://zenodo.org/record/3716354/files/figure.png" pageId="2" pageNumber="399" startId="2.[816,847,1044,1062]" targetBox="[821,1469,136,1017]" targetPageId="2">
|
||
<paragraph blockId="2.[816,1493,1044,1329]" pageId="2" pageNumber="399">
|
||
<emphasis bold="true" pageId="2" pageNumber="399">
|
||
Fig.3 | Relationships of
|
||
<taxonomicName authorityName="Field & Benito & Chen & Jagt & Ksepka" authorityYear="2020" box="[1024,1193,1044,1062]" genus="Asteriornis" pageId="2" pageNumber="399" rank="species" species="maastrichtensis">
|
||
<emphasis bold="true" box="[1024,1193,1044,1062]" italics="true" pageId="2" pageNumber="399">A. maastrichtensis</emphasis>
|
||
</taxonomicName>
|
||
and stratigraphic provenance of holotype.a, Cladogram showing the phylogenetic position of
|
||
</emphasis>
|
||
<taxonomicName box="[1351,1445,1071,1088]" genus="Asteriornis" pageId="2" pageNumber="399" rank="genus">
|
||
<emphasis box="[1351,1445,1071,1088]" italics="true" pageId="2" pageNumber="399">Asteriornis</emphasis>
|
||
</taxonomicName>
|
||
<emphasis bold="true" box="[1448,1481,1070,1089]" pageId="2" pageNumber="399">and</emphasis>
|
||
<emphasis box="[816,881,1098,1115]" italics="true" pageId="2" pageNumber="399">Vegavis</emphasis>
|
||
<emphasis bold="true" pageId="2" pageNumber="399">inferred under parsimony(solid coloured lines) and tip-dated Bayesian analyses (dashed lines).A position for</emphasis>
|
||
<emphasis box="[1141,1206,1124,1141]" italics="true" pageId="2" pageNumber="399">Vegavis</emphasis>
|
||
<emphasis bold="true" pageId="2" pageNumber="399">
|
||
allied with
|
||
<taxonomicName box="[1302,1425,1123,1142]" class="Aves" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anseriformes" pageId="2" pageNumber="399" phylum="Chordata" rank="order">Anseriformes</taxonomicName>
|
||
,as previously found
|
||
</emphasis>
|
||
<emphasis bold="true" box="[964,1010,1148,1159]" pageId="2" pageNumber="399">
|
||
<superScript attach="left" box="[964,1010,1148,1159]" fontSize="4" pageId="2" pageNumber="399">
|
||
<bibRefCitation author="Clarke, J. A. & Tambussi, C. P. & Noriega, J. I. & Erickson, G. M. & Ketcham, R. A." box="[964,976,1148,1158]" journalOrPublisher="Nature" pageId="2" pageNumber="399" pagination="305 - 308" part="433" refId="ref4200" refString="12. Clarke, J. A., Tambussi, C. P., Noriega, J. I., Erickson, G. M. & Ketcham, R. A. Definitive fossil evidence for the extant avian radiation in the Cretaceous. Nature 433, 305 - 308 (2005)." title="Definitive fossil evidence for the extant avian radiation in the Cretaceous" type="journal article" year="2005">12</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Worthy, T. H. & Degrange, F. J. & Handley, W. D. & Lee, M. S. Y." box="[980,993,1148,1158]" journalOrPublisher="R. Soc. Open Sci." pageId="2" pageNumber="399" pagination="170975" part="4" refId="ref4921" refString="28. Worthy, T. H., Degrange, F. J., Handley, W. D. & Lee, M. S. Y. The evolution of giant flightless birds and novel phylogenetic relationships for extinct fowl (Aves, Galloanseres). R. Soc. Open Sci. 4, 170975 (2017)." title="The evolution of giant flightless birds and novel phylogenetic relationships for extinct fowl (Aves, Galloanseres)" type="journal article" year="2017">28</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Clarke, J. A." box="[997,1010,1148,1158]" journalOrPublisher="Nature" pageId="2" pageNumber="399" pagination="502 - 505" part="538" refId="ref5272" refString="34. Clarke, J. A. et al. Fossil evidence of the avian vocal organ from the Mesozoic. Nature 538, 502 - 505 (2016)." title="Fossil evidence of the avian vocal organ from the Mesozoic" type="journal article" year="2016">34</bibRefCitation>
|
||
</superScript>
|
||
</emphasis>
|
||
<emphasis bold="true" pageId="2" pageNumber="399">, is also shown (curved branch).Numbers within clades denote extant species richness.The analysis is based on a newly modified dataset;see Extended Data Fig.9 and Supplementary Information for full phylogenetic results.Extinct taxa are denoted with daggers.b, Simplified stratigraphy of the Maastricht Formation,exposed in the vicinity of the holotype locality and surrounding areas</emphasis>
|
||
<bibRefCitation author="Keutgen, N." box="[1162,1175,1282,1292]" journalOrPublisher="Neth. J. Geosci." pageId="2" pageNumber="399" pagination="229 - 260" part="97" refId="ref4741" refString="24. Keutgen, N. A bioclast-based astronomical timescale for the Maastrichtian in the type area (southeast Netherlands, northeast Belgium) and stratigraphic implications: the legacy of PJ Felder. Neth. J. Geosci. 97, 229 - 260 (2018)." title="A bioclast-based astronomical timescale for the Maastrichtian in the type area (southeast Netherlands, northeast Belgium) and stratigraphic implications: the legacy of PJ Felder" type="journal article" year="2018">
|
||
<emphasis bold="true" box="[1162,1175,1282,1292]" pageId="2" pageNumber="399">
|
||
<superScript attach="left" box="[1162,1175,1282,1292]" fontSize="4" pageId="2" pageNumber="399">24</superScript>
|
||
</emphasis>
|
||
</bibRefCitation>
|
||
. c, Location of the CBR-Romontbos Quarry near Eben-Emael,Belgium.
|
||
</paragraph>
|
||
</caption>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.3716358" ID-Zenodo-Dep="3716358" httpUri="https://zenodo.org/record/3716358/files/figure.png" pageId="2" pageNumber="399" startId="2.[106,137,1074,1092]" subCaptionStartIDs="2.[270,300,1207,1224]" subCaptionStarts="Fig.4 f" targetBox="[111,781,130,1049]" targetPageId="2">
|
||
<paragraph blockId="2.[106,782,1073,1252]" pageId="2" pageNumber="399">
|
||
<emphasis bold="true" pageId="2" pageNumber="399">Fig.2| Comparative quadrate and skull morphology of selected total-group Galloanserae.a–d, Left quadrates(and reflected right quadrate of</emphasis>
|
||
<taxonomicName authorityName="Wetmore" authorityYear="1926" box="[106,207,1126,1145]" class="Aves" family="Presbyornithidae" genus="Presbyornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Eoenantiornithiformes" pageId="2" pageNumber="399" phylum="Chordata" rank="genus">
|
||
<emphasis box="[106,207,1126,1145]" italics="true" pageId="2" pageNumber="399">Presbyornis</emphasis>
|
||
</taxonomicName>
|
||
<emphasis bold="true" pageId="2" pageNumber="399">) in medial view (left) and caudal view (middle).Skulls are in right lateral view,except</emphasis>
|
||
<taxonomicName box="[274,347,1153,1172]" class="Aves" family="Megapodiidae" genus="Alectura" kingdom="Animalia" order="Craciformes" pageId="2" pageNumber="399" phylum="Chordata" rank="genus">
|
||
<emphasis box="[274,347,1153,1172]" italics="true" pageId="2" pageNumber="399">Alectura</emphasis>
|
||
</taxonomicName>
|
||
<emphasis bold="true" box="[350,763,1153,1172]" pageId="2" pageNumber="399">which is in reflected left lateral view.The skull of</emphasis>
|
||
<taxonomicName authorityName="Wetmore" authorityYear="1926" box="[106,207,1179,1198]" class="Aves" family="Presbyornithidae" genus="Presbyornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Eoenantiornithiformes" pageId="2" pageNumber="399" phylum="Chordata" rank="genus">
|
||
<emphasis box="[106,207,1179,1198]" italics="true" pageId="2" pageNumber="399">Presbyornis</emphasis>
|
||
</taxonomicName>
|
||
<emphasis bold="true" pageId="2" pageNumber="399">USNM 299846 is shown.Scale bars,5mm (quadrates);1cm (skulls). See Extended Data Fig.4 for additional high-resolution images of these taxa and other extant and fossil Galloanserae.</emphasis>
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="2.[106,787,1336,1989]" pageId="2" pageNumber="399">
|
||
<emphasis bold="true" pageId="2" pageNumber="399">
|
||
(
|
||
<figureCitation box="[112,168,1336,1358]" captionStart="Fig" captionStartId="2.[106,137,1074,1092]" captionTargetBox="[111,781,130,1049]" captionTargetId="figure@2.[109,494,497,865]" captionTargetPageId="2" captionText="Fig.2| Comparativequadrate and skull morphology ofselected total-group Galloanserae.a–d, Left quadrates(andreflected rightquadrate of Presbyornis) inmedial view (left) andcaudal view (middle).Skulls arein right lateral view,except Alectura which is inreflected left lateral view.The skull of PresbyornisUSNM 299846 is shown.Scale bars,5mm (quadrates);1cm (skulls). See Extended DataFig.4 for additional high-resolution images of these taxa andother extant and fossil Galloanserae." figureDoi="http://doi.org/10.5281/zenodo.3716358" httpUri="https://zenodo.org/record/3716358/files/figure.png" pageId="2" pageNumber="399">Fig. 2</figureCitation>
|
||
, Extended Data
|
||
<figureCitation box="[333,391,1337,1358]" captionStart="Fig" captionStartId="2.[106,137,1074,1092]" captionTargetBox="[111,781,130,1049]" captionTargetId="figure@2.[109,494,497,865]" captionTargetPageId="2" captionText="Fig.2| Comparativequadrate and skull morphology ofselected total-group Galloanserae.a–d, Left quadrates(andreflected rightquadrate of Presbyornis) inmedial view (left) andcaudal view (middle).Skulls arein right lateral view,except Alectura which is inreflected left lateral view.The skull of PresbyornisUSNM 299846 is shown.Scale bars,5mm (quadrates);1cm (skulls). See Extended DataFig.4 for additional high-resolution images of these taxa andother extant and fossil Galloanserae." figureDoi="http://doi.org/10.5281/zenodo.3716358" httpUri="https://zenodo.org/record/3716358/files/figure.png" pageId="2" pageNumber="399">Fig. 4</figureCitation>
|
||
). The prootic and squamosal heads are divided by a well-developed incisure,as in almost all Neognathae;this contrasts with the condition in crownward stem birds such as
|
||
</emphasis>
|
||
<emphasis italics="true" pageId="2" pageNumber="399">Ichthyornis</emphasis>
|
||
,
|
||
<emphasis bold="true" pageId="2" pageNumber="399">in which the division between these condyles is poorly marked,and Palaeognathae,in which the condyles merge into a single head.Two pneumatic foramina pierce the quadrate:the foramen pneumaticum rostromediale and the foramen pneumaticum basiorbitale.A tuberculum subcapitulare is moderately developed on the lateral face of the otic process.This characteristic has been considered a derived feature of Galloanserae</emphasis>
|
||
<bibRefCitation author="Elzanowski, A. & Stidham, T. A." box="[259,273,1591,1604]" journalOrPublisher="J. Morphol." pageId="2" pageNumber="399" pagination="305 - 323" part="271" refId="ref4881" refString="27. Elzanowski, A. & Stidham, T. A. Morphology of the quadrate in the Eocene anseriform Presbyornis and extant galloanserine birds. J. Morphol. 271, 305 - 323 (2010)." title="Morphology of the quadrate in the Eocene anseriform Presbyornis and extant galloanserine birds" type="journal article" year="2010">
|
||
<emphasis bold="true" box="[259,273,1591,1604]" pageId="2" pageNumber="399">
|
||
<superScript attach="left" box="[259,273,1591,1604]" fontSize="5" pageId="2" pageNumber="399">27</superScript>
|
||
</emphasis>
|
||
</bibRefCitation>
|
||
<emphasis bold="true" pageId="2" pageNumber="399">; however,it is present in many Neoaves,and given its absence in Palaeognathae and</emphasis>
|
||
<emphasis box="[410,522,1623,1644]" italics="true" pageId="2" pageNumber="399">Ichthyornis</emphasis>
|
||
<emphasis bold="true" pageId="2" pageNumber="399">it might instead represent a synapomorphy of Neognathae.The cotyla quadratojugalis of the quadrate is fairly deep,with a complete,un-notched rim.The pterygoid articulation is more widely separated dorsally from the mandibular condyle than in most extant Galloanserae,and is very similar to that of</emphasis>
|
||
<taxonomicName authorityName="Wetmore" authorityYear="1926" box="[130,245,1767,1788]" class="Aves" family="Presbyornithidae" genus="Presbyornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Eoenantiornithiformes" pageId="2" pageNumber="399" phylum="Chordata" rank="genus">
|
||
<emphasis box="[130,245,1767,1788]" italics="true" pageId="2" pageNumber="399">Presbyornis</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<paragraph blockId="2.[106,787,1336,1989]" pageId="2" pageNumber="399">
|
||
<emphasis bold="true" box="[129,587,1795,1816]" pageId="2" pageNumber="399">As in all known Galloanserae,the mandible of</emphasis>
|
||
<taxonomicName box="[590,699,1796,1816]" genus="Asteriornis" pageId="2" pageNumber="399" rank="genus">
|
||
<emphasis box="[590,699,1796,1816]" italics="true" pageId="2" pageNumber="399">Asteriornis</emphasis>
|
||
</taxonomicName>
|
||
<emphasis bold="true" pageId="2" pageNumber="399">exhibits two cotylae for articulation with the bicondylar quadrate and a bladelike retroarticular process.This process is strongly hooked,and in its shape and proportions bears a strong resemblance to the condition in the pan-anseriform</emphasis>
|
||
<emphasis box="[303,486,1910,1931]" italics="true" pageId="2" pageNumber="399">Anatalavis oxfordi</emphasis>
|
||
<emphasis bold="true" pageId="2" pageNumber="399">
|
||
(
|
||
<figureCitation box="[495,561,1909,1931]" captionStart="Fig" captionStartId="1.[106,137,960,978]" captionTargetBox="[144,1459,131,935]" captionTargetId="figure@1.[519,896,488,847]" captionTargetPageId="1" captionText="Fig.1 | Digitallysegmented skull ofAsteriornis maastrichtensis. a–c,Left lateral view (a), rightlateral view (b) anddorsal view (c) of the A.maastrichtensisskull.d,Comparison of retroarticular processes inlateral view of Anatalavis oxfordi(rightside) and Asteriornis(left side,reflected). Selected synapomorphies aredenoted withnumbers (red,Neornithes;green, Galloanserae37–39):1,bony mandibular symphysis;2,toothless beak;3,no coronoid bone;4,bicondylar mandibularprocess of quadrate;5,retroarticular process long,curving,stronglycompressed mediolaterally;6,dorsally oriented internal articular process of mandible;7,maxillary process of premaxilla dorsoventrallydeep and lateromediallycompressed.Scale bars, 5 mm." figureDoi="http://doi.org/10.5281/zenodo.3716352" httpUri="https://zenodo.org/record/3716352/files/figure.png" pageId="2" pageNumber="399">Fig. 1d</figureCitation>
|
||
, Extended Data
|
||
<figureCitation box="[720,774,1910,1931]" captionStart="Extended" captionStartId="11.[106,193,1769,1787]" captionTargetBox="[203,1399,132,1743]" captionTargetId="figure@11.[203,1399,131,1744]" captionTargetPageId="11" captionText="Extended DataFig.5 | Detailed comparisonsofgalloanseran retroarticular morphology.Retroarticularregions of left mandiblesin lateral (left),medial (middle) and dorsal (right)views.Both the left andrightmandibles of Asteriornis areshown in dorsal view,as theretroarticular process is only preserved on theleft mandible andthe medial processis only preserved onthe right mandible.Imagesof Anatalavis are mirrored.Scalebars,1 cm." figureDoi="http://doi.org/10.5281/zenodo.3716369" httpUri="https://zenodo.org/record/3716369/files/figure.png" pageId="2" pageNumber="399">Fig. 5</figureCitation>
|
||
). An elongate,slightly dorsally oriented processus medialis is preserved on the right jaw,as in all Galloanserae.
|
||
</emphasis>
|
||
</paragraph>
|
||
<paragraph blockId="2.[816,1498,1422,1530]" pageId="2" pageNumber="399">
|
||
<emphasis bold="true" pageId="2" pageNumber="399">The left distal femur is well-preserved,exhibiting a medial condyle with a bluntly angular profile between the articular surface of the condyle and its cranial margin. A detailed anatomical description of the skull and postcranium is provided in the Supplementary Information.</emphasis>
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |