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<document ID-DOI="10.1111/joa.13588" ID-GBIF-Dataset="37542e68-bc2a-4824-ad1f-2fc1ae5253d1" ID-PMC="PMC8930807" ID-PubMed="34854094" ID-Zenodo-Dep="5808218" approvalRequired="1" approvalRequired_for_document="1" checkinTime="1640731186193" checkinUser="donat" docAuthor="Li, Zhiheng, Wang, Min, Stidham, Thomas A., Zhou, Zhonghe &amp; Clarke, Julia" docDate="2021" docId="03F18797C462FF9BFCC5F1F73507EF2C" docLanguage="en" docName="joa-13588.pdf" docOrigin="Journal of Anatomy 239 (6)" docSource="http://dx.doi.org/10.1111/joa.13588" docTitle="Brevirostruavis macrohyoideus Li &amp; Wang &amp; Stidham &amp; Zhou &amp; Clarke 2021, gen. et sp. nov." docType="treatment" docVersion="6" lastPageNumber="11" masterDocId="FFC8FFEFC463FF91FF90F25C343BEA73" masterDocTitle="Novel evolution of a hyperelongated tongue in a Cretaceous enantiornithine from China and the evolution of the hyolingual apparatus and feeding in birds" masterLastPageNumber="12" masterPageNumber="1" pageNumber="2" updateTime="1668134224990" updateUser="ExternalLinkService">
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<mods:titleInfo>
<mods:title>Novel evolution of a hyperelongated tongue in a Cretaceous enantiornithine from China and the evolution of the hyolingual apparatus and feeding in birds</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Li, Zhiheng</mods:namePart>
<mods:nameIdentifier type="ORCID">0000-0002-6968-6982</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Wang, Min</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Stidham, Thomas A.</mods:namePart>
<mods:nameIdentifier type="ORCID">0000-0003-4766-0041</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Zhou, Zhonghe</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Clarke, Julia</mods:namePart>
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<mods:title>Journal of Anatomy</mods:title>
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<mods:part>
<mods:date>2021</mods:date>
<mods:detail type="pubDate">
<mods:number>2021-12-31</mods:number>
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<mods:detail type="volume">
<mods:number>239</mods:number>
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<mods:number>6</mods:number>
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<mods:start>1</mods:start>
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<mods:url>http://dx.doi.org/10.1111/joa.13588</mods:url>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">10.1111/joa.13588</mods:identifier>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.5808228" ID-GBIF-Taxon="191793453" ID-Zenodo-Dep="5808228" LSID="urn:lsid:plazi:treatment:03F18797C462FF9BFCC5F1F73507EF2C" httpUri="http://treatment.plazi.org/id/03F18797C462FF9BFCC5F1F73507EF2C" lastPageId="10" lastPageNumber="11" pageId="1" pageNumber="2">
<subSubSection box="[853,1281,939,959]" pageId="1" pageNumber="2" type="nomenclature">
<paragraph blockId="1.[853,1281,870,959]" box="[853,1281,939,959]" pageId="1" pageNumber="2">
<taxonomicName authority="Li &amp; Wang &amp; Stidham &amp; Zhou &amp; Clarke, 2021" authorityName="Li &amp; Wang &amp; Stidham &amp; Zhou &amp; Clarke" authorityYear="2021" box="[853,1132,939,959]" class="Aves" genus="Brevirostruavis" kingdom="Animalia" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="macrohyoideus" status="gen. et sp. nov.">
<emphasis box="[853,1132,939,959]" italics="true" pageId="1" pageNumber="2">Brevirostruavis macrohyoideus</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[1137,1281,939,959]" pageId="1" pageNumber="2" rank="species">gen. et sp. nov.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection pageId="1" pageNumber="2" type="materials_examined">
<paragraph blockId="1.[818,856,1039,1064]" box="[818,1026,1039,1064]" lastBlockId="1.[913,1026,1039,1064]" pageId="1" pageNumber="2">
<emphasis bold="true" box="[818,856,1039,1064]" pageId="1" pageNumber="2">3.1</emphasis>
|
<heading bold="true" box="[913,1026,1039,1064]" fontSize="10" level="2" pageId="1" pageNumber="2" reason="0">
<emphasis bold="true" box="[913,1026,1039,1064]" pageId="1" pageNumber="2">Holotype</emphasis>
</heading>
</paragraph>
<paragraph blockId="1.[818,1467,1112,1271]" pageId="1" pageNumber="2">
<materialsCitation ID-GBIF-Occurrence="3422473301" collectionCode="IVPP" pageId="1" pageNumber="2" specimenCode="V13266" specimenCount="1" typeStatus="holotype">
<collectionCode box="[818,865,1112,1132]" collectionName="IVPP" pageId="1" pageNumber="2">IVPP</collectionCode>
(Institute of Vertebrate Paleontology and Paleoanthropology, Beijing, China)
<specimenCode box="[964,1040,1147,1167]" pageId="1" pageNumber="2">V13266</specimenCode>
is a nearly complete skeleton preserved on a single slab (missing part of the humerus and the pelvic elements), with associated traces of preserved integument around the body (
<figureCitation box="[823,937,1251,1271]" captionStart-0="FIGURE 1" captionStart-1="FIGURE 2" captionStart-2="FIGURE 3" captionStart-3="FIGURE 4" captionStartId-0="2.[122,214,967,988]" captionStartId-1="3.[121,213,1287,1308]" captionStartId-2="4.[122,214,1163,1184]" captionStartId-3="5.[122,214,625,646]" captionTargetBox-0="[122,1466,57,926]" captionTargetBox-1="[351,1235,123,1246]" captionTargetBox-2="[122,1466,57,1122]" captionTargetBox-3="[122,1465,57,584]" captionTargetId-0="figure-635@2.[129,1459,123,926]" captionTargetId-1="figure-435@3.[351,1235,123,1246]" captionTargetId-2="figure-510@4.[129,1459,123,1122]" captionTargetId-3="figure-839@5.[128,1459,123,585]" captionTargetPageId-0="2" captionTargetPageId-1="3" captionTargetPageId-2="4" captionTargetPageId-3="5" captionText-0="FIGURE 1 Photograph and line drawing of the body of the holotype specimen of Brevirostruavis macrohyoideus (IVPP V13266). Anatomical abbreviations: c, coracoid; cav, caudal vertebrate; cv, cervical; fe, femur; fi, fibula; fu, furcula; h, humerus; il, ilium; isch, ischium; int, integument; mc I, alular metacarpal; mc II, major metacarpal; pub, pubis; py, pygostyle; r, radius; ra, radiale; sk, skull; st, sternum; tbt, tibiotarsus; tv, thoracic vertebrate; tmt, tarsometatarsus; u, ulna; and ul, ulnare" captionText-1="FIGURE 2 Photograph and line drawing of the skull of the holotype specimen of Brevirostruavis macrohyoideus (IVPP V13266). Anatomical abbreviations: an, angular; ar, articular; at, atlas; ax, axis; bp, basisphenoid process; bs, basisphenoid; cb, ceratobranchial; cr, cervical rib; cv, cervical; de, dentary; dr, descending ramus; ep?, ectopterygoid?; fr, frontal; la, lacrimal; ju, jugal; ma, maxillae; na, nasal; op, occipital process; pa, parietal; pm, premaxilla; pt?, pterygoid?; qu, quadrate; sa, surangular; and to, tooth" captionText-2="FIGURE 3 Photograph and line drawing of the sternum and pelvic girdle of the holotype of Brevirostruavis macrohyoideus IVPP V13266. Anatomical abbreviations: cav, caudal vertebrate; cp, cranial lateral process; fe, femur; il, ilium; is, ischium; lt, lateral trabecula; mp, middle process; mt, medial trabecula; para, parapophysis; pb, pubic boot; pu, pubis; py, pygostyle; ra, radius; st, sternum; sy, synsacrum; tr, thoracic rib; and xp, xiphoid process" captionText-3="FIGURE 4 Photograph of cervical vertebrae (a), tarsometatarsus (b), and sternum (c) of the holotype of Brevirostruavis macrohyoideus (IVPP V13266) with features described in the text. Anatomical abbreviations: cr, cervical rib; cp, cranial lateral process; hc, hypocleidium; lp, lateral process; prf, facet of prezygapophysis; pof, facet of postzygapophysis; mt I, metatarsal I; mt II, metatarsal II; mp, medial process; od, odontoid process; pc, pedal claw; and xp, xiphoid process" figureDoi-0="http://doi.org/10.5281/zenodo.5808220" figureDoi-1="http://doi.org/10.5281/zenodo.5808224" figureDoi-2="http://doi.org/10.5281/zenodo.5808226" figureDoi-3="http://doi.org/10.5281/zenodo.5808230" httpUri-0="https://zenodo.org/record/5808220/files/figure.png" httpUri-1="https://zenodo.org/record/5808224/files/figure.png" httpUri-2="https://zenodo.org/record/5808226/files/figure.png" httpUri-3="https://zenodo.org/record/5808230/files/figure.png" pageId="1" pageNumber="2">Figures 14</figureCitation>
).
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="1" pageNumber="2" type="etymology">
<paragraph blockId="1.[818,856,1351,1376]" box="[818,1044,1351,1376]" lastBlockId="1.[915,1044,1351,1376]" pageId="1" pageNumber="2">
<emphasis bold="true" box="[818,856,1351,1376]" pageId="1" pageNumber="2">3.2</emphasis>
|
<heading bold="true" box="[915,1044,1351,1376]" fontSize="10" level="2" pageId="1" pageNumber="2" reason="0">
<emphasis bold="true" box="[915,1044,1351,1376]" pageId="1" pageNumber="2">Etymology</emphasis>
</heading>
</paragraph>
<paragraph blockId="1.[818,1465,1424,1479]" pageId="1" pageNumber="2">The genus name refers to its short rostrum (and bird), and the specific epithet refers to the particularly long hyoid apparatus.</paragraph>
</subSubSection>
<subSubSection pageId="1" pageNumber="2" type="materials_examined">
<paragraph blockId="1.[818,856,1559,1584]" box="[818,1157,1559,1584]" lastBlockId="1.[914,1157,1559,1584]" pageId="1" pageNumber="2">
<emphasis bold="true" box="[818,856,1559,1584]" pageId="1" pageNumber="2">3.3</emphasis>
|
<heading bold="true" box="[914,1157,1559,1584]" fontSize="10" level="2" pageId="1" pageNumber="2" reason="0">
<emphasis bold="true" box="[914,1157,1559,1584]" pageId="1" pageNumber="2">Locality and horizon</emphasis>
</heading>
</paragraph>
<paragraph blockId="1.[818,1467,1632,1722]" pageId="1" pageNumber="2">
<materialsCitation ID-GBIF-Occurrence="3422473302" country="China" county="Jianchang County" location="Xiaotaizi Village" municipality="Jiufotang Formation" pageId="1" pageNumber="2" specimenCount="1" stateProvince="Liaoning Province" typeStatus="holotype">
<location LSID="urn:lsid:plazi:treatment:03F18797C462FF9BFCC5F1F73507EF2C:8E87605AC462FF90FCA2F43C37ECEC07" box="[818,983,1632,1652]" country="China" county="Jianchang County" municipality="Jiufotang Formation" name="Xiaotaizi Village" pageId="1" pageNumber="2" stateProvince="Liaoning Province">Xiaotaizi Village</location>
,
<collectingCounty box="[1002,1185,1632,1652]" pageId="1" pageNumber="2">Jianchang County</collectingCounty>
,
<collectingRegion box="[1203,1386,1632,1652]" country="China" name="Liaoning" pageId="1" pageNumber="2">Liaoning Province</collectingRegion>
,
<collectingCountry box="[1405,1463,1632,1652]" name="China" pageId="1" pageNumber="2">China</collectingCountry>
;
<collectingMunicipality box="[818,1015,1667,1687]" pageId="1" pageNumber="2">Jiufotang Formation</collectingMunicipality>
,
<geologicalTimeScale box="[1022,1466,1667,1687]" pageId="1" pageNumber="2">Lower Cretaceous. Age approximately 120 Ma</geologicalTimeScale>
(
<bibRefCitation author="He, H. Y. &amp; Wang, X. L. &amp; Zhou, Z. H. &amp; Wang, F. &amp; Boven, A. &amp; Shi, G. H." box="[824,968,1702,1722]" journalOrPublisher="Geophysical Research Letters" pageId="1" pageNumber="2" pagination="1 - 4" part="31" refId="ref7180" refString="He, H. Y., Wang, X. L., Zhou, Z. H., Wang, F., Boven, A., Shi, G. H. et al. (2004) Timing of the Jiufotang Formation (Jehol Group) in Liaoning, northeastern China, and its implications. Geophysical Research Letters, 31, 1 - 4." title="Timing of the Jiufotang Formation (Jehol Group) in Liaoning, northeastern China, and its implications" type="journal article" year="2004">He et al., 2004</bibRefCitation>
).
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection lastPageId="2" lastPageNumber="3" pageId="1" pageNumber="2" type="diagnosis">
<paragraph blockId="1.[818,856,1802,1827]" box="[818,1030,1802,1827]" lastBlockId="1.[914,1030,1802,1827]" pageId="1" pageNumber="2">
<emphasis bold="true" box="[818,856,1802,1827]" pageId="1" pageNumber="2">3.4</emphasis>
|
<heading bold="true" box="[914,1030,1802,1827]" fontSize="10" level="2" pageId="1" pageNumber="2" reason="0">
<emphasis bold="true" box="[914,1030,1802,1827]" pageId="1" pageNumber="2">Diagnosis</emphasis>
</heading>
</paragraph>
<paragraph blockId="1.[818,1466,1875,1964]" lastBlockId="2.[122,770,1147,1548]" lastPageId="2" lastPageNumber="3" pageId="1" pageNumber="2">
A medium-sized enantiornithine that is distinguished from all known enantiornithines based on the unique combination of the following features: a short and pointed skull rostrum lined with small peg-shaped teeth; a pair of extremely long ceratobranchials, only slightly shorter than the skull length (see
<tableCitation box="[531,600,1182,1202]" captionStart="TABLE 1" captionStartId="6.[122,198,124,144]" captionTargetBox="[138,1438,175,1548]" captionTargetId="graphics-254@6.[122,1465,243,1554]" captionText="TABLE 1 Measurements of Brevirostruavis macrohyoideus (IVPP V 13266) in millimeters" httpUri="http://table.plazi.org/id/DF276609C465FF97FFEAF22037EAEAE3" pageId="2" pageNumber="3" tableUuid="DF276609C465FF97FFEAF22037EAEAE3">Table 1</tableCitation>
); a sternum with well-extended craniolateral processes; lateral trabeculae of the sternum with expanded triangular processes at the caudal ends;elongate prezygapophyses of the cranial cervical vertebrae; postzygapophyseal facet of the axis is tear-drop shaped; third cervical vertebra with sub-rounded articular facet of the postzygapophyses; ischium bearing a pronounced proximodorsal process; distal tibiotarsus with a knob on its cranial surface; length ratio between the fibula and tibiotarsus approximately 0.7; tarsometatarsus about half of the length of the tibiotarsus; medial rim of metatarsal trochlea III larger than the lateral rim; and pedal digit-I more robust than other pedal digits.
</paragraph>
</subSubSection>
<caption ID-DOI="http://doi.org/10.5281/zenodo.5808220" ID-Zenodo-Dep="5808220" httpUri="https://zenodo.org/record/5808220/files/figure.png" pageId="2" pageNumber="3" startId="2.[122,214,967,988]" subCaptionStartIDs="2.[235,368,997,1017]" subCaptionStarts="abbr" targetBox="[122,1466,57,926]" targetPageId="2">
<paragraph blockId="2.[122,1445,967,1076]" pageId="2" pageNumber="3">
<emphasis bold="true" box="[122,237,967,988]" pageId="2" pageNumber="3">FIGURE 1</emphasis>
Photograph and line drawing of the body of the holotype specimen of
<taxonomicName authorityName="Li &amp; Wang &amp; Stidham &amp; Zhou &amp; Clarke" authorityYear="2021" box="[927,1206,968,988]" genus="Brevirostruavis" pageId="2" pageNumber="3" rank="species" species="macrohyoideus">
<emphasis box="[927,1206,968,988]" italics="true" pageId="2" pageNumber="3">Brevirostruavis macrohyoideus</emphasis>
</taxonomicName>
(IVPP V13266). Anatomical abbreviations: c, coracoid; cav, caudal vertebrate; cv, cervical; fe, femur; fi, fibula; fu, furcula; h, humerus; il, ilium; isch, ischium; int, integument; mc I, alular metacarpal; mc II, major metacarpal; pub, pubis; py, pygostyle; r, radius; ra, radiale; sk, skull; st, sternum; tbt, tibiotarsus; tv, thoracic vertebrate; tmt, tarsometatarsus; u, ulna; and ul, ulnare
</paragraph>
</caption>
<subSubSection lastPageId="5" lastPageNumber="6" pageId="2" pageNumber="3" type="description">
<paragraph blockId="2.[122,137,1629,1654]" box="[122,717,1628,1654]" lastBlockId="2.[201,717,1628,1654]" pageId="2" pageNumber="3">
<emphasis bold="true" box="[122,137,1629,1654]" pageId="2" pageNumber="3">4</emphasis>
|
<emphasis bold="true" box="[201,717,1628,1654]" pageId="2" pageNumber="3">DESCRIPTION AND COMPARISONS</emphasis>
</paragraph>
<paragraph blockId="2.[122,769,1702,1964]" lastBlockId="2.[818,1467,1147,1964]" pageId="2" pageNumber="3">
Except for part of humerus and tarsometatarsus, nearly all of the skeletal elements are well-preserved, but a halo of feather impressions are poorly preserved surrounding the body. The whole skeleton is exposed ventrally except for the skull. The skull of IVPP V13266 is preserved mainly in lateroventral view, and it has been heavily compressed (Figure 2). The short premaxillae are fused only rostrally (Figure 2: pm). The nasal processes of the premaxillae are long and slim. The wide pterygoid has a wedge-shaped rostral margin (Figure 2). The jugal bar is dorsoventrally expanded, and it is deflected dorsally and narrows toward its caudal end, as in other enantiornithines like
<taxonomicName authority="(Li et al., 2014)" baseAuthorityName="Li" baseAuthorityYear="2014" box="[996,1241,1216,1236]" class="Aves" family="Bohaiornithidae" genus="Bohaiornis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis box="[996,1093,1216,1236]" italics="true" pageId="2" pageNumber="3">Bohaiornis</emphasis>
(
<bibRefCitation author="Li, Z. &amp; Zhou, Z. &amp; Wang, M. &amp; Clarke, J. A." box="[1104,1236,1216,1236]" journalOrPublisher="Journal of Paleontology" pageId="2" pageNumber="3" pagination="99 - 108" part="88" refId="ref7568" refString="Li, Z., Zhou, Z., Wang, M. &amp; Clarke, J. A. (2014) A new specimen of largebodied basal enantiornithine Bohaiornis from the Early Cretaceous of China and the inference of feeding ecology in Mesozoic birds. Journal of Paleontology, 88, 99 - 108." title="A new specimen of largebodied basal enantiornithine Bohaiornis from the Early Cretaceous of China and the inference of feeding ecology in Mesozoic birds" type="journal article" year="2014">Li et al., 2014</bibRefCitation>
)
</taxonomicName>
. The descending ramus of the right lacrimal is significantly longer (Figure 2: dr) and more robust than the dorsal ramus with its thin dorsal ridge. A tiny depression appears present in the center of the lacrimal. The nasal is morphologically similar to that of
<emphasis box="[1144,1269,1355,1375]" italics="true" pageId="2" pageNumber="3">Eoenantiornis</emphasis>
, with a pointed premaxillary process. A rounded square-shaped parietal is preserved caudal to the basicranium (Figure 2: pa). The left dentary is exposed in medial view, and has seven visible teeth. The rostral tip of the dentary seems to bend dorsally. There are four small teeth on each side of the premaxillae (Figure 2: to), and these teeth have a rounded crown and constricted base. Only three teeth are present in the right maxilla, and they are restricted to its rostral portion.
</paragraph>
<paragraph blockId="2.[818,1467,1147,1964]" lastBlockId="3.[122,770,1459,1964]" lastPageId="3" lastPageNumber="4" pageId="2" pageNumber="3">
The frontals are fused to one another, and only part of the left parietal is visible because of the significantly crushed preservation. The quadrate body is very slender as in most other enantiornithines, but the otic head is distinctly inflated relative to the mediolateral diameter of the quadrate body. The mandibular process is greatly enlarged medially and laterally relative to the quadrate body, and the lateral mandibular condyle is wider than the medial. Though broken and largely covered, there appears to be at least a partial lateral crest on the quadrate as in
<taxonomicName authority="(Stidham &amp; O'Connor, 2021)" baseAuthorityName="Stidham &amp; O'Connor" baseAuthorityYear="2021" box="[1079,1463,1910,1930]" class="Aves" family="Longipterygidae" genus="Longipteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Longipterygiformes" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis box="[1079,1186,1910,1930]" italics="true" pageId="2" pageNumber="3">Longipteryx</emphasis>
(
<bibRefCitation author="Stidham, T. A. &amp; O'Connor, J. K." box="[1197,1457,1910,1930]" journalOrPublisher="Journal of Anatomy" pageId="2" pageNumber="3" pagination="1066 - 1074" part="239" refId="ref8012" refString="Stidham, T. A. &amp; O'Connor, J. K. (2021) The evolutionary and functional implications of the unusual quadrate of Longipteryx chaoyangensis (Avialae: Enantiornithes) from the Cretaceous Jehol Biota of China. Journal of Anatomy, 239, 1066 - 1074." title="The evolutionary and functional implications of the unusual quadrate of Longipteryx chaoyangensis (Avialae: Enantiornithes) from the Cretaceous Jehol Biota of China" type="journal article" year="2021">Stidham &amp; O'Connor, 2021</bibRefCitation>
)
</taxonomicName>
. Adjacent to the mandibles, a pair of dorsally curved hyoid apparatus bones (
<figureCitation box="[190,272,1459,1479]" captionStart="FIGURE 2" captionStartId="3.[121,213,1287,1308]" captionTargetBox="[351,1235,123,1246]" captionTargetId="figure-435@3.[351,1235,123,1246]" captionTargetPageId="3" captionText="FIGURE 2 Photograph and line drawing of the skull of the holotype specimen of Brevirostruavis macrohyoideus (IVPP V13266). Anatomical abbreviations: an, angular; ar, articular; at, atlas; ax, axis; bp, basisphenoid process; bs, basisphenoid; cb, ceratobranchial; cr, cervical rib; cv, cervical; de, dentary; dr, descending ramus; ep?, ectopterygoid?; fr, frontal; la, lacrimal; ju, jugal; ma, maxillae; na, nasal; op, occipital process; pa, parietal; pm, premaxilla; pt?, pterygoid?; qu, quadrate; sa, surangular; and to, tooth" figureDoi="http://doi.org/10.5281/zenodo.5808224" httpUri="https://zenodo.org/record/5808224/files/figure.png" pageId="3" pageNumber="4">Figure 2</figureCitation>
: cb) are markedly elongate with spatula-shaped caudal ends. The rostral ends of these ceratobranchials have concave surfaces, and are separated from one another. While the caudal portion of the ceratobranchials curve significantly dorsally, their more rostral portion appears to have been possibly concave ventrally. The distorted mediolateral width of the foramen magnum is twice that of its dorsoventral height.
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.5808224" ID-Zenodo-Dep="5808224" httpUri="https://zenodo.org/record/5808224/files/figure.png" pageId="3" pageNumber="4" startId="3.[121,213,1287,1308]" subCaptionStartIDs="3.[122,254,1316,1336]" subCaptionStarts="abbr" targetBox="[351,1235,123,1246]" targetPageId="3">
<paragraph blockId="3.[121,1456,1287,1395]" pageId="3" pageNumber="4">
<emphasis bold="true" box="[121,237,1287,1308]" pageId="3" pageNumber="4">FIGURE 2</emphasis>
Photograph and line drawing of the skull of the holotype specimen of
<taxonomicName authorityName="Li &amp; Wang &amp; Stidham &amp; Zhou &amp; Clarke" authorityYear="2021" box="[917,1194,1287,1307]" genus="Brevirostruavis" pageId="3" pageNumber="4" rank="species" species="macrohyoideus">
<emphasis box="[917,1194,1287,1307]" italics="true" pageId="3" pageNumber="4">Brevirostruavis macrohyoideus</emphasis>
</taxonomicName>
(IVPP V13266). Anatomical abbreviations: an, angular; ar, articular; at, atlas; ax, axis; bp, basisphenoid process; bs, basisphenoid; cb, ceratobranchial; cr, cervical rib; cv, cervical; de, dentary; dr, descending ramus; ep?, ectopterygoid?; fr, frontal; la, lacrimal; ju, jugal; ma, maxillae; na, nasal; op, occipital process; pa, parietal; pm, premaxilla; pt?, pterygoid?; qu, quadrate; sa, surangular; and to, tooth
</paragraph>
</caption>
<paragraph blockId="3.[122,770,1459,1964]" lastBlockId="3.[818,1467,1459,1964]" pageId="3" pageNumber="4">
Ten cervical vertebrae are preserved, including the atlas and axis preserved in articulation in ventral view (
<figureCitation box="[517,607,1736,1756]" captionStart="FIGURE 1" captionStartId="2.[122,214,967,988]" captionTargetBox="[122,1466,57,926]" captionTargetId="figure-635@2.[129,1459,123,926]" captionText="FIGURE 1 Photograph and line drawing of the body of the holotype specimen of Brevirostruavis macrohyoideus (IVPP V13266). Anatomical abbreviations: c, coracoid; cav, caudal vertebrate; cv, cervical; fe, femur; fi, fibula; fu, furcula; h, humerus; il, ilium; isch, ischium; int, integument; mc I, alular metacarpal; mc II, major metacarpal; pub, pubis; py, pygostyle; r, radius; ra, radiale; sk, skull; st, sternum; tbt, tibiotarsus; tv, thoracic vertebrate; tmt, tarsometatarsus; u, ulna; and ul, ulnare" figureDoi="http://doi.org/10.5281/zenodo.5808220" httpUri="https://zenodo.org/record/5808220/files/figure.png" pageId="3" pageNumber="4">Figures 1</figureCitation>
and 2). Only the dorsal portion of the atlas is preserved with a slender pointed lateral process. The axis is markedly wider than its length, and the odontoid process overlies the ventral corpus of the atlas (Figure 2). A low ventral ridge is present on the axis. The articular facet of the axis postzygapophyses are crescent shaped, differing from the rounded shape in the more caudal cervicals (Figures 2 and
<figureCitation box="[611,625,1944,1964]" captionStart="FIGURE 3" captionStartId="4.[122,214,1163,1184]" captionTargetBox="[122,1466,57,1122]" captionTargetId="figure-510@4.[129,1459,123,1122]" captionText="FIGURE 3 Photograph and line drawing of the sternum and pelvic girdle of the holotype of Brevirostruavis macrohyoideus IVPP V13266. Anatomical abbreviations: cav, caudal vertebrate; cp, cranial lateral process; fe, femur; il, ilium; is, ischium; lt, lateral trabecula; mp, middle process; mt, medial trabecula; para, parapophysis; pb, pubic boot; pu, pubis; py, pygostyle; ra, radius; st, sternum; sy, synsacrum; tr, thoracic rib; and xp, xiphoid process" figureDoi="http://doi.org/10.5281/zenodo.5808226" httpUri="https://zenodo.org/record/5808226/files/figure.png" pageId="3" pageNumber="4">3</figureCitation>
). The variation in the shape of the articular facets between the second and more caudal postzygapophyses might be indicative of the evolution of an “S-shaped” curved neck evident among living birds (
<bibRefCitation author="Kambic, R. E. &amp; Biewener, A. A. &amp; Pierce, S. E." journalOrPublisher="Frontiers in Zoology" pageId="3" pageNumber="4" pagination="37" part="14" refId="ref7369" refString="Kambic, R. E., Biewener, A. A. &amp; Pierce, S. E. (2017) Experimental determination of three-dimensional cervical joint mobility in the avian neck. Frontiers in Zoology, 14, 37." title="Experimental determination of three-dimensional cervical joint mobility in the avian neck" type="journal article" year="2017">Kambic et al., 2017</bibRefCitation>
).
</paragraph>
<paragraph blockId="3.[818,1467,1459,1964]" lastBlockId="4.[122,771,1320,1964]" lastPageId="4" lastPageNumber="5" pageId="3" pageNumber="4">
The prezygapophyses of the 3rd and 4th cervicals project cranially beyond the centra by a distance that is approximately half of the craniocaudal length of the respective centrum, a feature unknown in other enantiornithines. Moving caudally along the cervicals, the prezygapophyses become shorter and are subequal with the length of the postzygapophyses. The 4th to the 7th cervicals are longer than both the cranial and caudal ones in the neck (
<figureCitation box="[1258,1336,1806,1826]" captionStart="FIGURE 1" captionStartId="2.[122,214,967,988]" captionTargetBox="[122,1466,57,926]" captionTargetId="figure-635@2.[129,1459,123,926]" captionText="FIGURE 1 Photograph and line drawing of the body of the holotype specimen of Brevirostruavis macrohyoideus (IVPP V13266). Anatomical abbreviations: c, coracoid; cav, caudal vertebrate; cv, cervical; fe, femur; fi, fibula; fu, furcula; h, humerus; il, ilium; isch, ischium; int, integument; mc I, alular metacarpal; mc II, major metacarpal; pub, pubis; py, pygostyle; r, radius; ra, radiale; sk, skull; st, sternum; tbt, tibiotarsus; tv, thoracic vertebrate; tmt, tarsometatarsus; u, ulna; and ul, ulnare" figureDoi="http://doi.org/10.5281/zenodo.5808220" httpUri="https://zenodo.org/record/5808220/files/figure.png" pageId="3" pageNumber="4">Figure 1</figureCitation>
). The ventral surfaces of the cervicals are keeled. Carotid processes are present on the 6th cervical, but the distribution of the processes among the other cervicals is obscured. Two short cervical ribs are displaced from the cervical axis (Figure 2). At least four paired long curved ribs are associated with the lateral edge of the sternum. These sternal ribs are quite robust, bearing a medium depth on the shaft. A few gastralia are present.
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.5808226" ID-Zenodo-Dep="5808226" httpUri="https://zenodo.org/record/5808226/files/figure.png" pageId="4" pageNumber="5" startId="4.[122,214,1163,1184]" subCaptionStartIDs="4.[235,368,1193,1213]" subCaptionStarts="abbr" targetBox="[122,1466,57,1122]" targetPageId="4">
<paragraph blockId="4.[122,1449,1163,1271]" pageId="4" pageNumber="5">
<emphasis bold="true" box="[122,237,1163,1184]" pageId="4" pageNumber="5">FIGURE 3</emphasis>
Photograph and line drawing of the sternum and pelvic girdle of the holotype of
<taxonomicName authority="IVPP V" authorityName="IVPP V" box="[1023,1375,1163,1183]" genus="Brevirostruavis" pageId="4" pageNumber="5" rank="species" species="macrohyoideus">
<emphasis box="[1023,1302,1163,1183]" italics="true" pageId="4" pageNumber="5">Brevirostruavis macrohyoideus</emphasis>
IVPP V
</taxonomicName>
13266. Anatomical abbreviations: cav, caudal vertebrate; cp, cranial lateral process; fe, femur; il, ilium; is, ischium; lt, lateral trabecula; mp, middle process; mt, medial trabecula; para, parapophysis; pb, pubic boot; pu, pubis; py, pygostyle; ra, radius; st, sternum; sy, synsacrum; tr, thoracic rib; and xp, xiphoid process
</paragraph>
</caption>
<paragraph blockId="4.[122,771,1320,1964]" pageId="4" pageNumber="5">
Four caudal thoracic vertebrae are preserved cranial to the synsacrum (
<figureCitation box="[237,320,1459,1479]" captionStart="FIGURE 3" captionStartId="4.[122,214,1163,1184]" captionTargetBox="[122,1466,57,1122]" captionTargetId="figure-510@4.[129,1459,123,1122]" captionText="FIGURE 3 Photograph and line drawing of the sternum and pelvic girdle of the holotype of Brevirostruavis macrohyoideus IVPP V13266. Anatomical abbreviations: cav, caudal vertebrate; cp, cranial lateral process; fe, femur; il, ilium; is, ischium; lt, lateral trabecula; mp, middle process; mt, medial trabecula; para, parapophysis; pb, pubic boot; pu, pubis; py, pygostyle; ra, radius; st, sternum; sy, synsacrum; tr, thoracic rib; and xp, xiphoid process" figureDoi="http://doi.org/10.5281/zenodo.5808226" httpUri="https://zenodo.org/record/5808226/files/figure.png" pageId="4" pageNumber="5">Figure 3</figureCitation>
). The parapophysis is centrally located on the lateral face of the centra, similar to the typical condition among Enantiornithes (
<figureCitation box="[272,353,1528,1548]" captionStart="FIGURE 3" captionStartId="4.[122,214,1163,1184]" captionTargetBox="[122,1466,57,1122]" captionTargetId="figure-510@4.[129,1459,123,1122]" captionTargetPageId="4" captionText="FIGURE 3 Photograph and line drawing of the sternum and pelvic girdle of the holotype of Brevirostruavis macrohyoideus IVPP V13266. Anatomical abbreviations: cav, caudal vertebrate; cp, cranial lateral process; fe, femur; il, ilium; is, ischium; lt, lateral trabecula; mp, middle process; mt, medial trabecula; para, parapophysis; pb, pubic boot; pu, pubis; py, pygostyle; ra, radius; st, sternum; sy, synsacrum; tr, thoracic rib; and xp, xiphoid process" figureDoi="http://doi.org/10.5281/zenodo.5808226" httpUri="https://zenodo.org/record/5808226/files/figure.png" pageId="4" pageNumber="5">Figure 3</figureCitation>
: para). The synsacrum is composed of about seven to eight vertebrae. Approximately five free caudal vertebrae are piled up together with markedly long transverse and ventral processes (
<figureCitation box="[196,280,1632,1652]" captionStart="FIGURE 3" captionStartId="4.[122,214,1163,1184]" captionTargetBox="[122,1466,57,1122]" captionTargetId="figure-510@4.[129,1459,123,1122]" captionText="FIGURE 3 Photograph and line drawing of the sternum and pelvic girdle of the holotype of Brevirostruavis macrohyoideus IVPP V13266. Anatomical abbreviations: cav, caudal vertebrate; cp, cranial lateral process; fe, femur; il, ilium; is, ischium; lt, lateral trabecula; mp, middle process; mt, medial trabecula; para, parapophysis; pb, pubic boot; pu, pubis; py, pygostyle; ra, radius; st, sternum; sy, synsacrum; tr, thoracic rib; and xp, xiphoid process" figureDoi="http://doi.org/10.5281/zenodo.5808226" httpUri="https://zenodo.org/record/5808226/files/figure.png" pageId="4" pageNumber="5">Figure 3</figureCitation>
: cav). The long pygostyle has a pair of dorsal processes and a pointed caudal end.
</paragraph>
<paragraph blockId="4.[122,771,1320,1964]" lastBlockId="4.[818,1467,1320,1964]" pageId="4" pageNumber="5">
The mediolateral width of the sternal plate is slightly longer than its craniocaudal length. The sternal keel is low, and diverges into two low ridges cranially, a feature only observed in enantiornithines (e.g., Martin 2011). Craniolateral processes project from the lateral edge of the sternum (
<figureCitation box="[268,349,1840,1860]" captionStart="FIGURE 3" captionStartId="4.[122,214,1163,1184]" captionTargetBox="[122,1466,57,1122]" captionTargetId="figure-510@4.[129,1459,123,1122]" captionText="FIGURE 3 Photograph and line drawing of the sternum and pelvic girdle of the holotype of Brevirostruavis macrohyoideus IVPP V13266. Anatomical abbreviations: cav, caudal vertebrate; cp, cranial lateral process; fe, femur; il, ilium; is, ischium; lt, lateral trabecula; mp, middle process; mt, medial trabecula; para, parapophysis; pb, pubic boot; pu, pubis; py, pygostyle; ra, radius; st, sternum; sy, synsacrum; tr, thoracic rib; and xp, xiphoid process" figureDoi="http://doi.org/10.5281/zenodo.5808226" httpUri="https://zenodo.org/record/5808226/files/figure.png" pageId="4" pageNumber="5">Figure 3</figureCitation>
: lp) with a gentle slope transitioning between this process and the carinal sulcus. The craniolateral processes are rarely developed in Early Cretaceous enantiornithines with a few exceptions, including
<emphasis box="[329,433,1944,1964]" italics="true" pageId="4" pageNumber="5">Pterygornis</emphasis>
and
<emphasis box="[480,571,1944,1964]" italics="true" pageId="4" pageNumber="5">Concornis</emphasis>
(
<bibRefCitation author="Wang, M. &amp; Hu, H. &amp; Li, Z." box="[582,767,1944,1964]" journalOrPublisher="Journal of Systematic Palaeontology" pageId="4" pageNumber="5" pagination="481 - 497" part="14" refId="ref8063" refString="Wang, M., Hu, H. &amp; Li, Z. (2016 a) A new small enantiornithine bird from the Jehol Biota, with implications for early evolution of avian skull morphology. Journal of Systematic Palaeontology, 14, 481 - 497." title="A new small enantiornithine bird from the Jehol Biota, with implications for early evolution of avian skull morphology" type="journal article" year="2016">Wang et al., 2016a</bibRefCitation>
,
<bibRefCitation author="Wang, M. &amp; Li, Z. &amp; Zhou, Z." box="[818,881,1320,1340]" journalOrPublisher="Proceedings of the National Academy of Sciences" pageId="4" pageNumber="5" pagination="11470 - 11475" part="114" refId="ref8112" refString="Wang, M., Li, Z. &amp; Zhou, Z. (2017 a) Insight into the growth pattern and bone fusion of basal birds from an Early Cretaceous enantiornithine bird. Proceedings of the National Academy of Sciences, 114, 11470 - 11475." title="Insight into the growth pattern and bone fusion of basal birds from an Early Cretaceous enantiornithine bird" type="journal article" year="2017">2017a</bibRefCitation>
;
<bibRefCitation author="Zheng, X. &amp; Wang, X. &amp; Jingmai, K. O. C. &amp; Zhou, Z." box="[891,1074,1320,1340]" journalOrPublisher="Nature Communications" pageId="4" pageNumber="5" pagination="1116" part="3" refId="ref8725" refString="Zheng, X., Wang, X., Jingmai, K. O. C. &amp; Zhou, Z. (2012) Insight into the early evolution of the avian sternum from juvenile enantiornithines. Nature Communications, 3, 1116." title="Insight into the early evolution of the avian sternum from juvenile enantiornithines" type="journal article" year="2012">Zheng et al., 2012</bibRefCitation>
), and they have been proposed to form from ossification centers separate from the main sternal body.
</paragraph>
<paragraph blockId="4.[818,1467,1320,1964]" pageId="4" pageNumber="5">
Lateral and intermediate trabeculae are present on the caudal margin of the sternum (
<figureCitation box="[1043,1125,1424,1444]" captionStart="FIGURE 3" captionStartId="4.[122,214,1163,1184]" captionTargetBox="[122,1466,57,1122]" captionTargetId="figure-510@4.[129,1459,123,1122]" captionText="FIGURE 3 Photograph and line drawing of the sternum and pelvic girdle of the holotype of Brevirostruavis macrohyoideus IVPP V13266. Anatomical abbreviations: cav, caudal vertebrate; cp, cranial lateral process; fe, femur; il, ilium; is, ischium; lt, lateral trabecula; mp, middle process; mt, medial trabecula; para, parapophysis; pb, pubic boot; pu, pubis; py, pygostyle; ra, radius; st, sternum; sy, synsacrum; tr, thoracic rib; and xp, xiphoid process" figureDoi="http://doi.org/10.5281/zenodo.5808226" httpUri="https://zenodo.org/record/5808226/files/figure.png" pageId="4" pageNumber="5">Figure 3</figureCitation>
). The lateral trabecula bears a large distal flare, and the intermediate one deflects toward the midline (
<figureCitation box="[823,907,1494,1514]" captionStart="FIGURE 3" captionStartId="4.[122,214,1163,1184]" captionTargetBox="[122,1466,57,1122]" captionTargetId="figure-510@4.[129,1459,123,1122]" captionText="FIGURE 3 Photograph and line drawing of the sternum and pelvic girdle of the holotype of Brevirostruavis macrohyoideus IVPP V13266. Anatomical abbreviations: cav, caudal vertebrate; cp, cranial lateral process; fe, femur; il, ilium; is, ischium; lt, lateral trabecula; mp, middle process; mt, medial trabecula; para, parapophysis; pb, pubic boot; pu, pubis; py, pygostyle; ra, radius; st, sternum; sy, synsacrum; tr, thoracic rib; and xp, xiphoid process" figureDoi="http://doi.org/10.5281/zenodo.5808226" httpUri="https://zenodo.org/record/5808226/files/figure.png" pageId="4" pageNumber="5">Figure 3</figureCitation>
: lt). The sternal keel extends as far caudally as the lateral trabecula, similar to the condition in
<emphasis box="[1193,1301,1528,1548]" italics="true" pageId="4" pageNumber="5">Protopteryx</emphasis>
and
<emphasis italics="true" pageId="4" pageNumber="5">Cathayornis yandica</emphasis>
(
<bibRefCitation author="Wang, M. &amp; Liu, D." box="[898,1070,1563,1583]" journalOrPublisher="Journal of Systematic Palaeontology" pageId="4" pageNumber="5" pagination="29 - 47" part="14" refId="ref8162" refString="Wang, M. &amp; Liu, D. (2016) Taxonomical reappraisal of Cathayornithidae (Aves: Enantiornithes). Journal of Systematic Palaeontology, 14, 29 - 47." title="Taxonomical reappraisal of Cathayornithidae (Aves: Enantiornithes)" type="journal article" year="2016">Wang &amp; Liu, 2016</bibRefCitation>
). The slim furcular rami form a sharp angle of approximately 45° (
<figureCitation box="[1026,1104,1598,1618]" captionStart="FIGURE 1" captionStartId="2.[122,214,967,988]" captionTargetBox="[122,1466,57,926]" captionTargetId="figure-635@2.[129,1459,123,926]" captionText="FIGURE 1 Photograph and line drawing of the body of the holotype specimen of Brevirostruavis macrohyoideus (IVPP V13266). Anatomical abbreviations: c, coracoid; cav, caudal vertebrate; cv, cervical; fe, femur; fi, fibula; fu, furcula; h, humerus; il, ilium; isch, ischium; int, integument; mc I, alular metacarpal; mc II, major metacarpal; pub, pubis; py, pygostyle; r, radius; ra, radiale; sk, skull; st, sternum; tbt, tibiotarsus; tv, thoracic vertebrate; tmt, tarsometatarsus; u, ulna; and ul, ulnare" figureDoi="http://doi.org/10.5281/zenodo.5808220" httpUri="https://zenodo.org/record/5808220/files/figure.png" pageId="4" pageNumber="5">Figure 1</figureCitation>
: fu). The hypocleidium is less than half of the length of the ramus. The strut-like coracoids have a rounded, slightly raised acrocoracoid, and lack a procoracoid process. The lateral margin of coracoids appears to be straight or slightly concave (
<figureCitation box="[823,900,1736,1756]" captionStart="FIGURE 1" captionStartId="2.[122,214,967,988]" captionTargetBox="[122,1466,57,926]" captionTargetId="figure-635@2.[129,1459,123,926]" captionText="FIGURE 1 Photograph and line drawing of the body of the holotype specimen of Brevirostruavis macrohyoideus (IVPP V13266). Anatomical abbreviations: c, coracoid; cav, caudal vertebrate; cv, cervical; fe, femur; fi, fibula; fu, furcula; h, humerus; il, ilium; isch, ischium; int, integument; mc I, alular metacarpal; mc II, major metacarpal; pub, pubis; py, pygostyle; r, radius; ra, radiale; sk, skull; st, sternum; tbt, tibiotarsus; tv, thoracic vertebrate; tmt, tarsometatarsus; u, ulna; and ul, ulnare" figureDoi="http://doi.org/10.5281/zenodo.5808220" httpUri="https://zenodo.org/record/5808220/files/figure.png" pageId="4" pageNumber="5">Figure 1</figureCitation>
). Only the proximal end of the scapula is exposed in medial view without any identifiable details (
<figureCitation box="[1175,1252,1771,1791]" captionStart="FIGURE 1" captionStartId="2.[122,214,967,988]" captionTargetBox="[122,1466,57,926]" captionTargetId="figure-635@2.[129,1459,123,926]" captionText="FIGURE 1 Photograph and line drawing of the body of the holotype specimen of Brevirostruavis macrohyoideus (IVPP V13266). Anatomical abbreviations: c, coracoid; cav, caudal vertebrate; cv, cervical; fe, femur; fi, fibula; fu, furcula; h, humerus; il, ilium; isch, ischium; int, integument; mc I, alular metacarpal; mc II, major metacarpal; pub, pubis; py, pygostyle; r, radius; ra, radiale; sk, skull; st, sternum; tbt, tibiotarsus; tv, thoracic vertebrate; tmt, tarsometatarsus; u, ulna; and ul, ulnare" figureDoi="http://doi.org/10.5281/zenodo.5808220" httpUri="https://zenodo.org/record/5808220/files/figure.png" pageId="4" pageNumber="5">Figure 1</figureCitation>
).
</paragraph>
<paragraph blockId="4.[818,1467,1320,1964]" lastBlockId="5.[122,771,800,1964]" lastPageId="5" lastPageNumber="6" pageId="4" pageNumber="5">
The humerus preserved in ventral view, and has a narrow deltopectoral crest. The humeral head appears to be strip-like with a shallow transverse groove. The dorsal condyle is rounded, and more clearly defined than the ventral one. A small dorsal supracondyle is present. The ulna has a narrow m. brachialis scar proximally, and the radial depression distally. The dorsal and the ventral rami of the ulnare are slightly differentiated, with the ventral one slightly larger. The semilunate carpal is fused with the proximal ends of the major and minor metacarpals. The carpometacarpi are preserved in ventral view and the infratrochlear fossa is visible. The alular metacarpal is short and partially fused with the major metacarpal. The proximal phalanx of the alular digit is short, less than half of the length of the major metacarpal. The minor metacarpal is slightly bowed and extends distally beyond that of the major metacarpal. The major and minor metacarpals have only a slim intermetacarpal space separating them. There are two small ungual claws associated with the first two manual digits. The forelimb is marginally longer than the hindlimb with a length ratio (humerus+ulna+carpometacarpus /femur+tibiotarsus+tarsometatarsus = ratio) of 1.05, similar to
<taxonomicName box="[673,765,1251,1271]" class="Aves" family="Longipterygidae" genus="Rapaxavis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Longipterygiformes" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[673,765,1251,1271]" italics="true" pageId="5" pageNumber="6">Rapaxavis</emphasis>
</taxonomicName>
,
<taxonomicName box="[122,212,1286,1306]" class="Aves" family="Bohaiornithidae" genus="Zhouornis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[122,212,1286,1306]" italics="true" pageId="5" pageNumber="6">Zhouornis</emphasis>
</taxonomicName>
, and
<emphasis box="[269,378,1286,1306]" italics="true" pageId="5" pageNumber="6">Cathayornis</emphasis>
(
<tableCitation box="[391,463,1286,1306]" captionStart="TABLE 2" captionStartId="7.[122,198,124,144]" captionTargetBox="[138,1449,175,949]" captionTargetId="graphics-546@7.[122,1465,243,956]" captionText="TABLE 2 Limb measurements of the Brevirostruavis macrohyoideus with comparison of other avialans (Adopted from Zhang et al., 2013)" httpUri="http://table.plazi.org/id/DF276609C464FF96FFEAF22031A7EAE3" pageId="5" pageNumber="6" tableUuid="DF276609C464FF96FFEAF22031A7EAE3">Table 2</tableCitation>
for measurements), and differing from other enantiornithines (
<tableCitation box="[434,514,1320,1340]" captionStart="TABLE 1" captionStartId="6.[122,198,124,144]" captionTargetBox="[138,1438,175,1548]" captionTargetId="graphics-254@6.[122,1465,243,1554]" captionText="TABLE 1 Measurements of Brevirostruavis macrohyoideus (IVPP V 13266) in millimeters" httpUri="http://table.plazi.org/id/DF276609C465FF97FFEAF22037EAEAE3" pageId="5" pageNumber="6" tableUuid="DF276609C465FF97FFEAF22037EAEAE3">Tables 1</tableCitation>
and
<tableCitation box="[562,576,1320,1340]" captionStart="TABLE 2" captionStartId="7.[122,198,124,144]" captionTargetBox="[138,1449,175,949]" captionTargetId="graphics-546@7.[122,1465,243,956]" captionText="TABLE 2 Limb measurements of the Brevirostruavis macrohyoideus with comparison of other avialans (Adopted from Zhang et al., 2013)" httpUri="http://table.plazi.org/id/DF276609C464FF96FFEAF22031A7EAE3" pageId="5" pageNumber="6" tableUuid="DF276609C464FF96FFEAF22031A7EAE3">2</tableCitation>
). The ilium, ischium, and pubis are not fused with one another around the acetabulum, and they are preserved in lateral view. The pre-acetabular portion of the ilium is longer and dorsoventrally wider than the caudal portion. There is a small crest on the cranial portion of the ilium. The proximodorsal process of the ischium is well projected (
<figureCitation box="[576,657,1494,1514]" captionStart="FIGURE 3" captionStartId="4.[122,214,1163,1184]" captionTargetBox="[122,1466,57,1122]" captionTargetId="figure-510@4.[129,1459,123,1122]" captionText="FIGURE 3 Photograph and line drawing of the sternum and pelvic girdle of the holotype of Brevirostruavis macrohyoideus IVPP V13266. Anatomical abbreviations: cav, caudal vertebrate; cp, cranial lateral process; fe, femur; il, ilium; is, ischium; lt, lateral trabecula; mp, middle process; mt, medial trabecula; para, parapophysis; pb, pubic boot; pu, pubis; py, pygostyle; ra, radius; st, sternum; sy, synsacrum; tr, thoracic rib; and xp, xiphoid process" figureDoi="http://doi.org/10.5281/zenodo.5808226" httpUri="https://zenodo.org/record/5808226/files/figure.png" pageId="5" pageNumber="6">Figure 3</figureCitation>
), approaching, but not contacting, the ventral margin of the ilium. The pubis is rounded in cross-section proximally, lateromedially flattened distally, and flared at its distal end, forming a pubic boot (
<figureCitation box="[585,666,1598,1618]" captionStart="FIGURE 3" captionStartId="4.[122,214,1163,1184]" captionTargetBox="[122,1466,57,1122]" captionTargetId="figure-510@4.[129,1459,123,1122]" captionText="FIGURE 3 Photograph and line drawing of the sternum and pelvic girdle of the holotype of Brevirostruavis macrohyoideus IVPP V13266. Anatomical abbreviations: cav, caudal vertebrate; cp, cranial lateral process; fe, femur; il, ilium; is, ischium; lt, lateral trabecula; mp, middle process; mt, medial trabecula; para, parapophysis; pb, pubic boot; pu, pubis; py, pygostyle; ra, radius; st, sternum; sy, synsacrum; tr, thoracic rib; and xp, xiphoid process" figureDoi="http://doi.org/10.5281/zenodo.5808226" httpUri="https://zenodo.org/record/5808226/files/figure.png" pageId="5" pageNumber="6">Figure 3</figureCitation>
: pb).
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.5808230" ID-Zenodo-Dep="5808230" httpUri="https://zenodo.org/record/5808230/files/figure.png" pageId="5" pageNumber="6" startId="5.[122,214,625,646]" subCaptionStartIDs="5.[721,854,655,675]" subCaptionStarts="abbr" targetBox="[122,1465,57,584]" targetPageId="5">
<paragraph blockId="5.[122,1451,625,734]" pageId="5" pageNumber="6">
<emphasis bold="true" box="[122,237,625,646]" pageId="5" pageNumber="6">FIGURE 4</emphasis>
Photograph of cervical vertebrae (a), tarsometatarsus (b), and sternum (c) of the holotype of
<taxonomicName box="[1136,1415,626,646]" genus="Brevirostruavis" pageId="5" pageNumber="6" rank="species" species="macrohyoideus">
<emphasis box="[1136,1415,626,646]" italics="true" pageId="5" pageNumber="6">Brevirostruavis macrohyoideus</emphasis>
</taxonomicName>
(IVPP V13266) with features described in the text. Anatomical abbreviations: cr, cervical rib; cp, cranial lateral process; hc, hypocleidium; lp, lateral process; prf, facet of prezygapophysis; pof, facet of postzygapophysis; mt I, metatarsal I; mt II, metatarsal II; mp, medial process; od, odontoid process; pc, pedal claw; and xp, xiphoid process
</paragraph>
</caption>
<paragraph blockId="5.[122,771,800,1964]" lastBlockId="5.[818,1467,800,1340]" pageId="5" pageNumber="6">
The right femur preserved in medial view is slightly bowed with the pit for the lig. capitis present on the medial surface of the femoral head. The periosteal surface of the distal femur appears to be rough, and the condyles are not well developed. The femur is 70% of the tibiotarsus length. The right and left tibiotarsi are preserved in cranial and caudal view, respectively (
<figureCitation box="[514,592,1806,1826]" captionStart="FIGURE 1" captionStartId="2.[122,214,967,988]" captionTargetBox="[122,1466,57,926]" captionTargetId="figure-635@2.[129,1459,123,926]" captionText="FIGURE 1 Photograph and line drawing of the body of the holotype specimen of Brevirostruavis macrohyoideus (IVPP V13266). Anatomical abbreviations: c, coracoid; cav, caudal vertebrate; cv, cervical; fe, femur; fi, fibula; fu, furcula; h, humerus; il, ilium; isch, ischium; int, integument; mc I, alular metacarpal; mc II, major metacarpal; pub, pubis; py, pygostyle; r, radius; ra, radiale; sk, skull; st, sternum; tbt, tibiotarsus; tv, thoracic vertebrate; tmt, tarsometatarsus; u, ulna; and ul, ulnare" figureDoi="http://doi.org/10.5281/zenodo.5808220" httpUri="https://zenodo.org/record/5808220/files/figure.png" pageId="5" pageNumber="6">Figure 1</figureCitation>
). The medial condyle of the tibiotarsus is wider than the lateral one. The distal end of the tibiotarsus shaft is not significantly wider than the mid-shaft. The fibula is proportionally longer than many other enantiornithines. Although the distal end is missing, the preserved length of the left fibula is more than 70% of that of the tibiotarsus. The tubercle for the insertion of the m. iliofibularis is caudolaterally directed. The popliteal tubercle is present on the proximal face of the tibiotarsus. The lateral cnemial crest and the fibular crest are present on the tibiotarsus. The distal tarsals are completely fused with the proximal metatarsals, forming a true tarsometatarsus in dorsal view. The tarsometatarsus is extremely short and measures only about half of the length of the tibiotarsus (
<tableCitation box="[1066,1136,1043,1063]" captionStart="TABLE 1" captionStartId="6.[122,198,124,144]" captionTargetBox="[138,1438,175,1548]" captionTargetId="graphics-254@6.[122,1465,243,1554]" captionText="TABLE 1 Measurements of Brevirostruavis macrohyoideus (IVPP V 13266) in millimeters" httpUri="http://table.plazi.org/id/DF276609C465FF97FFEAF22037EAEAE3" pageId="5" pageNumber="6" tableUuid="DF276609C465FF97FFEAF22037EAEAE3">Table 1</tableCitation>
). Metatarsals II-IV are fused with each other proximally, but not distally. Metatarsal II is slightly longer than metatarsal IV. Metatarsal trochlea III has a medial trochlear rim that extends further distally than the lateral rim (
<figureCitation box="[1276,1357,1147,1167]" captionStart="FIGURE 4" captionStartId="5.[122,214,625,646]" captionTargetBox="[122,1465,57,584]" captionTargetId="figure-839@5.[128,1459,123,585]" captionTargetPageId="5" captionText="FIGURE 4 Photograph of cervical vertebrae (a), tarsometatarsus (b), and sternum (c) of the holotype of Brevirostruavis macrohyoideus (IVPP V13266) with features described in the text. Anatomical abbreviations: cr, cervical rib; cp, cranial lateral process; hc, hypocleidium; lp, lateral process; prf, facet of prezygapophysis; pof, facet of postzygapophysis; mt I, metatarsal I; mt II, metatarsal II; mp, medial process; od, odontoid process; pc, pedal claw; and xp, xiphoid process" figureDoi="http://doi.org/10.5281/zenodo.5808230" httpUri="https://zenodo.org/record/5808230/files/figure.png" pageId="5" pageNumber="6">Figure 4</figureCitation>
: mp and lp). Metatarsal IV is slightly narrower than metatarsals II and III, a typical condition within Enantiornithes. The ungual phalanx of the middle pedal digit is the longest. The proximal two phalanges of pedal digit IV are much shorter than the distal ones. The size of the ungual claws is larger than most the proximal phalanges.
</paragraph>
</subSubSection>
<subSubSection lastPageId="10" lastPageNumber="11" pageId="5" pageNumber="6" type="discussion">
<paragraph blockId="5.[818,833,1421,1446]" box="[818,1083,1420,1446]" lastBlockId="5.[897,1083,1420,1446]" pageId="5" pageNumber="6">
<emphasis bold="true" box="[818,833,1421,1446]" pageId="5" pageNumber="6">5</emphasis>
|
<heading allCaps="true" bold="true" box="[897,1083,1420,1446]" fontSize="10" level="1" pageId="5" pageNumber="6" reason="0">
<emphasis bold="true" box="[897,1083,1420,1446]" pageId="5" pageNumber="6">DISCUSSION</emphasis>
</heading>
</paragraph>
<paragraph blockId="5.[818,1466,1494,1964]" pageId="5" pageNumber="6">
IVPP V13266 can be referred to the Enantiornithes on basis of the presence of the following synapomorphies corroborated by our phylogenetic analysis: a “Y”-shaped furcula with a long hypocleidium, and minor metacarpal extending distal to the major metacarpal. The phylogenetic analysis produced 230 most parsimonious trees (MPTs) with a length of 1087 (Consistency index = 0.338, Retention index = 0.661). The strict consensus is poorly resolved, and most enantiornithines including
<taxonomicName genus="Brevirostruavis" pageId="5" pageNumber="6" rank="species" species="macrohyoideus">
<emphasis italics="true" pageId="5" pageNumber="6">Brevirostruavis macrohyoideus</emphasis>
</taxonomicName>
form a large polytomy with a few derived clades resolved (
<figureCitation box="[895,996,1806,1826]" captionStart="FIGURE 5" captionStartId="8.[123,215,1343,1364]" captionTargetBox="[122,1466,57,1301]" captionTargetId="figure-360@8.[129,1459,123,1302]" captionText="FIGURE 5 Phylogenetic position of Brevirostruavis macrohyoideus (IVPP V 13266) in Avialae. (a) Strict consensus tree from current phylogenetic analysis; (b) reduced consensus tree with ten most unstable taxa removed. The bootstrap and absolute Bremer support values are denoted in normal and bold Italic fonts" figureDoi="http://doi.org/10.5281/zenodo.5808232" httpUri="https://zenodo.org/record/5808232/files/figure.png" pageId="5" pageNumber="6">Figure 5A</figureCitation>
). The poorly resolved strict consensus tree results largely from unstable taxa that occupy different positions in the MPTs. In order to extract a consensus of phylogenetic information, we performed a reduced consensus analysis using TNT (
<bibRefCitation author="Pol, D. &amp; Escapa, I. H." box="[876,1078,1944,1964]" journalOrPublisher="Cladistics" pageId="5" pageNumber="6" pagination="515 - 527" part="25" refId="ref7932" refString="Pol, D. &amp; Escapa, I. H. (2009) Unstable taxa in cladistic analysis: identification and the assessment of relevant characters. Cladistics, 25, 515 - 527." title="Unstable taxa in cladistic analysis: identification and the assessment of relevant characters" type="journal article" year="2009">Pol &amp; Escapa, 2009</bibRefCitation>
). The ten most unstable taxa (
<taxonomicName authorityName="Chiappe, Suzuki, Dyke, Watabe, Tsogtbaatar &amp; Barsbold" authorityYear="2007" box="[1390,1460,1944,1964]" class="Aves" genus="Elsornis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[1390,1460,1944,1964]" italics="true" pageId="5" pageNumber="6">Elsornis</emphasis>
</taxonomicName>
,
</paragraph>
<caption ID-Table-UUID="DF276609C465FF97FFEAF22037EAEAE3" box="[122,977,124,144]" httpUri="http://table.plazi.org/id/DF276609C465FF97FFEAF22037EAEAE3" pageId="6" pageNumber="7" startId="6.[122,198,124,144]" targetBox="[138,1438,175,1548]" targetIsTable="true" targetPageId="6">
<paragraph blockId="6.[122,222,124,144]" box="[122,222,124,144]" pageId="6" pageNumber="7">
<emphasis bold="true" box="[122,222,124,144]" pageId="6" pageNumber="7">TABLE 1</emphasis>
</paragraph>
<paragraph blockId="6.[241,977,124,144]" box="[241,977,124,144]" pageId="6" pageNumber="7">
Measurements of
<taxonomicName box="[414,693,124,144]" genus="Brevirostruavis" pageId="6" pageNumber="7" rank="species" species="macrohyoideus">
<emphasis box="[414,693,124,144]" italics="true" pageId="6" pageNumber="7">Brevirostruavis macrohyoideus</emphasis>
</taxonomicName>
(IVPP V 13266) in millimeters
</paragraph>
</caption>
<paragraph pageId="6" pageNumber="7">
<table box="[138,1438,175,1548]" gridcols="5" gridrows="25" pageId="6" pageNumber="7">
<tr box="[138,1438,175,195]" gridrow="0" pageId="6" pageNumber="7">
<th box="[138,1438,175,195]" colspan="5" colspanRight="4" gridcol="0" gridrow="0" pageId="6" pageNumber="7">
<emphasis bold="true" box="[138,223,175,195]" pageId="6" pageNumber="7">Elements</emphasis>
</th>
</tr>
<tr box="[138,1438,218,238]" gridrow="1" pageId="6" pageNumber="7" rowspan-2="1" rowspan-3="1" rowspan-4="1">
<th box="[138,310,218,238]" gridcol="0" gridrow="1" pageId="6" pageNumber="7">Skull length</th>
<td box="[504,561,218,238]" gridcol="1" gridrow="1" pageId="6" pageNumber="7">40.0</td>
</tr>
<tr box="[138,1438,255,275]" gridrow="2" pageId="6" pageNumber="7">
<th box="[138,310,255,275]" gridcol="0" gridrow="2" pageId="6" pageNumber="7">Ceratobranchials</th>
<td box="[504,561,255,275]" gridcol="1" gridrow="2" pageId="6" pageNumber="7">Left</td>
<td box="[770,917,255,275]" gridcol="2" gridrow="2" pageId="6" pageNumber="7">25.0</td>
<td box="[1036,1166,255,275]" gridcol="3" gridrow="2" pageId="6" pageNumber="7">Right</td>
<td box="[1290,1438,255,275]" gridcol="4" gridrow="2" pageId="6" pageNumber="7">28.4</td>
</tr>
<tr box="[138,1438,293,313]" gridrow="3" pageId="6" pageNumber="7" rowspan-3="1" rowspan-4="1">
<th box="[138,310,293,313]" gridcol="0" gridrow="3" pageId="6" pageNumber="7">Humerus length</th>
<td box="[504,561,293,313]" gridcol="1" gridrow="3" pageId="6" pageNumber="7">Left</td>
<td box="[770,917,293,313]" gridcol="2" gridrow="3" pageId="6" pageNumber="7">39.2</td>
</tr>
<tr box="[138,1438,330,350]" gridrow="4" pageId="6" pageNumber="7" rowspan-3="1" rowspan-4="1">
<th box="[138,310,330,350]" gridcol="0" gridrow="4" pageId="6" pageNumber="7">Ulna length</th>
<td box="[504,561,330,350]" gridcol="1" gridrow="4" pageId="6" pageNumber="7">Left</td>
<td box="[770,917,330,350]" gridcol="2" gridrow="4" pageId="6" pageNumber="7">38.6</td>
</tr>
<tr box="[138,1438,368,388]" gridrow="5" pageId="6" pageNumber="7">
<th box="[138,310,368,388]" gridcol="0" gridrow="5" pageId="6" pageNumber="7">Alular metacarpal</th>
<td box="[504,561,368,388]" gridcol="1" gridrow="5" pageId="6" pageNumber="7">Left</td>
<td box="[770,917,368,388]" gridcol="2" gridrow="5" pageId="6" pageNumber="7">2.6</td>
<td box="[1036,1166,368,388]" gridcol="3" gridrow="5" pageId="6" pageNumber="7">Right</td>
<td box="[1290,1438,368,388]" gridcol="4" gridrow="5" pageId="6" pageNumber="7">2.9</td>
</tr>
<tr box="[138,1438,405,425]" gridrow="6" pageId="6" pageNumber="7">
<th box="[138,310,405,425]" gridcol="0" gridrow="6" pageId="6" pageNumber="7">Metacarpal II</th>
<td box="[504,561,405,425]" gridcol="1" gridrow="6" pageId="6" pageNumber="7">Left</td>
<td box="[770,917,405,425]" gridcol="2" gridrow="6" pageId="6" pageNumber="7">15.0</td>
<td box="[1036,1166,405,425]" gridcol="3" gridrow="6" pageId="6" pageNumber="7">Right</td>
<td box="[1290,1438,405,425]" gridcol="4" gridrow="6" pageId="6" pageNumber="7">14.9</td>
</tr>
<tr box="[138,1438,442,462]" gridrow="7" pageId="6" pageNumber="7">
<th box="[138,310,442,462]" gridcol="0" gridrow="7" pageId="6" pageNumber="7">Metacarpal III</th>
<td box="[504,561,442,462]" gridcol="1" gridrow="7" pageId="6" pageNumber="7">Left</td>
<td box="[770,917,442,462]" gridcol="2" gridrow="7" pageId="6" pageNumber="7">17.7</td>
<td box="[1036,1166,442,462]" gridcol="3" gridrow="7" pageId="6" pageNumber="7">Right</td>
<td box="[1290,1438,442,462]" gridcol="4" gridrow="7" pageId="6" pageNumber="7">15.8</td>
</tr>
<tr box="[138,1438,480,500]" gridrow="8" pageId="6" pageNumber="7">
<th box="[138,310,480,500]" gridcol="0" gridrow="8" pageId="6" pageNumber="7">Phalanx I-1</th>
<td box="[504,561,480,500]" gridcol="1" gridrow="8" pageId="6" pageNumber="7">Left</td>
<td box="[770,917,480,500]" gridcol="2" gridrow="8" pageId="6" pageNumber="7">6.9</td>
<td box="[1036,1166,480,500]" gridcol="3" gridrow="8" pageId="6" pageNumber="7">Right</td>
<td box="[1290,1438,480,500]" gridcol="4" gridrow="8" pageId="6" pageNumber="7">6.6</td>
</tr>
<tr box="[138,1438,517,537]" gridrow="9" pageId="6" pageNumber="7">
<th box="[138,310,517,537]" gridcol="0" gridrow="9" pageId="6" pageNumber="7">Phalanx II-1</th>
<td box="[504,561,517,537]" gridcol="1" gridrow="9" pageId="6" pageNumber="7">Left</td>
<td box="[770,917,517,537]" gridcol="2" gridrow="9" pageId="6" pageNumber="7">9.6</td>
<td box="[1036,1166,517,537]" gridcol="3" gridrow="9" pageId="6" pageNumber="7">Right</td>
<td box="[1290,1438,517,537]" gridcol="4" gridrow="9" pageId="6" pageNumber="7">8.8</td>
</tr>
<tr box="[138,1438,555,575]" gridrow="10" pageId="6" pageNumber="7">
<th box="[138,310,555,575]" gridcol="0" gridrow="10" pageId="6" pageNumber="7">Phalanx II-2</th>
<td box="[504,561,555,575]" gridcol="1" gridrow="10" pageId="6" pageNumber="7">Left</td>
<td box="[770,917,555,575]" gridcol="2" gridrow="10" pageId="6" pageNumber="7">6.3</td>
<td box="[1036,1166,555,575]" gridcol="3" gridrow="10" pageId="6" pageNumber="7">Right</td>
<td box="[1290,1438,555,575]" gridcol="4" gridrow="10" pageId="6" pageNumber="7">5.5</td>
</tr>
<tr box="[138,1438,592,612]" gridrow="11" pageId="6" pageNumber="7">
<th box="[138,310,592,612]" gridcol="0" gridrow="11" pageId="6" pageNumber="7">Phalanx III-1</th>
<td box="[504,561,592,612]" gridcol="1" gridrow="11" pageId="6" pageNumber="7">Left</td>
<td box="[770,917,592,612]" gridcol="2" gridrow="11" pageId="6" pageNumber="7">4.6</td>
<td box="[1036,1166,592,612]" gridcol="3" gridrow="11" pageId="6" pageNumber="7">Right</td>
<td box="[1290,1438,592,612]" gridcol="4" gridrow="11" pageId="6" pageNumber="7">4.6</td>
</tr>
<tr box="[138,1438,630,650]" gridrow="12" pageId="6" pageNumber="7">
<th box="[138,310,630,650]" gridcol="0" gridrow="12" pageId="6" pageNumber="7">Manual Ungual</th>
<td box="[504,561,630,650]" gridcol="1" gridrow="12" pageId="6" pageNumber="7">D-2</td>
<td box="[770,917,630,650]" gridcol="2" gridrow="12" pageId="6" pageNumber="7">3.3</td>
<td box="[1036,1166,630,650]" gridcol="3" gridrow="12" pageId="6" pageNumber="7">D-1</td>
<td box="[1290,1438,630,650]" gridcol="4" gridrow="12" pageId="6" pageNumber="7">3.5</td>
</tr>
<tr box="[138,1438,667,687]" gridrow="13" pageId="6" pageNumber="7">
<th box="[138,310,667,687]" gridcol="0" gridrow="13" pageId="6" pageNumber="7">Sternum</th>
<td box="[504,561,667,687]" gridcol="1" gridrow="13" pageId="6" pageNumber="7">Width</td>
<td box="[770,917,667,687]" gridcol="2" gridrow="13" pageId="6" pageNumber="7">17.5</td>
<td box="[1036,1166,667,687]" gridcol="3" gridrow="13" pageId="6" pageNumber="7">Midline length</td>
<td box="[1290,1438,667,687]" gridcol="4" gridrow="13" pageId="6" pageNumber="7">25.3</td>
</tr>
<tr box="[138,1438,705,725]" gridrow="14" pageId="6" pageNumber="7" rowspan-3="1" rowspan-4="1">
<th box="[138,310,705,725]" gridcol="0" gridrow="14" pageId="6" pageNumber="7">Coracoid</th>
<td box="[504,561,705,725]" gridcol="1" gridrow="14" pageId="6" pageNumber="7">Left</td>
<td box="[770,917,705,725]" gridcol="2" gridrow="14" pageId="6" pageNumber="7">22.3 (estimated)</td>
</tr>
<tr box="[138,1438,742,762]" gridrow="15" pageId="6" pageNumber="7" rowspan-3="1" rowspan-4="1">
<th box="[138,310,742,762]" gridcol="0" gridrow="15" pageId="6" pageNumber="7">Femur length</th>
<td box="[504,561,742,762]" gridcol="1" gridrow="15" pageId="6" pageNumber="7">Left</td>
<td box="[770,917,742,762]" gridcol="2" gridrow="15" pageId="6" pageNumber="7">28.3</td>
</tr>
<tr box="[138,1438,779,799]" gridrow="16" pageId="6" pageNumber="7" rowspan-3="1" rowspan-4="1">
<th box="[138,310,779,799]" gridcol="0" gridrow="16" pageId="6" pageNumber="7">Tibiotarsus length</th>
<td box="[504,561,779,799]" gridcol="1" gridrow="16" pageId="6" pageNumber="7">Left</td>
<td box="[770,917,779,799]" gridcol="2" gridrow="16" pageId="6" pageNumber="7">40.0</td>
</tr>
<tr box="[138,1438,817,837]" gridrow="17" pageId="6" pageNumber="7" rowspan-1="1" rowspan-2="1">
<th box="[138,310,817,837]" gridcol="0" gridrow="17" pageId="6" pageNumber="7">Fibula length</th>
<td box="[1036,1166,817,837]" gridcol="3" gridrow="17" pageId="6" pageNumber="7">Right</td>
<td box="[1290,1438,817,837]" gridcol="4" gridrow="17" pageId="6" pageNumber="7">29.3 (preserved)</td>
</tr>
<tr box="[138,1438,854,874]" gridrow="18" pageId="6" pageNumber="7" rowspan-3="1" rowspan-4="1">
<th box="[138,310,854,874]" gridcol="0" gridrow="18" pageId="6" pageNumber="7">Metatarsal I</th>
<td box="[504,561,854,874]" gridcol="1" gridrow="18" pageId="6" pageNumber="7">Left</td>
<td box="[770,917,854,874]" gridcol="2" gridrow="18" pageId="6" pageNumber="7">5.1</td>
</tr>
<tr box="[138,1438,892,912]" gridrow="19" pageId="6" pageNumber="7" rowspan-1="1" rowspan-3="1" rowspan-4="1">
<th box="[138,310,892,912]" gridcol="0" gridrow="19" pageId="6" pageNumber="7">Metatarsal II</th>
<td box="[770,917,892,912]" gridcol="2" gridrow="19" pageId="6" pageNumber="7">18.8</td>
</tr>
<tr box="[138,1438,929,949]" gridrow="20" pageId="6" pageNumber="7" rowspan-1="1" rowspan-3="1" rowspan-4="1">
<th box="[138,310,929,949]" gridcol="0" gridrow="20" pageId="6" pageNumber="7">Metatarsal III</th>
<td box="[770,917,929,949]" gridcol="2" gridrow="20" pageId="6" pageNumber="7">20.4</td>
</tr>
<tr box="[138,1438,967,987]" gridrow="21" pageId="6" pageNumber="7" rowspan-1="1" rowspan-3="1" rowspan-4="1">
<th box="[138,310,967,987]" gridcol="0" gridrow="21" pageId="6" pageNumber="7">Metatarsal IV</th>
<td box="[770,917,967,987]" gridcol="2" gridrow="21" pageId="6" pageNumber="7">18.7</td>
</tr>
<tr box="[138,1438,1004,1024]" gridrow="22" pageId="6" pageNumber="7" rowspan-1="1" rowspan-3="1" rowspan-4="1">
<th box="[138,310,1004,1024]" gridcol="0" gridrow="22" pageId="6" pageNumber="7">Pygostyle length</th>
<td box="[770,917,1004,1024]" gridcol="2" gridrow="22" pageId="6" pageNumber="7">19.8</td>
</tr>
<tr box="[138,1438,1042,1398]" gridrow="23" pageId="6" pageNumber="7" rowspan-3="1" rowspan-4="1">
<th box="[138,310,1042,1398]" gridcol="0" gridrow="23" pageId="6" pageNumber="7">Pedal phalanx (left)</th>
<td box="[504,561,1042,1398]" gridcol="1" gridrow="23" pageId="6" pageNumber="7">I-1 II-1 II-2 III-1 III-2 III-3 IV-1 IV-2 IV-3 IV-4</td>
<td box="[770,917,1042,1398]" gridcol="2" gridrow="23" pageId="6" pageNumber="7">5.3 5.0 6.2 6.8 5.3 5.9 2.6 3.1 3.4 4.3</td>
</tr>
<tr box="[138,1438,1416,1548]" gridrow="24" pageId="6" pageNumber="7" rowspan-3="1" rowspan-4="1">
<th box="[138,310,1416,1548]" gridcol="0" gridrow="24" pageId="6" pageNumber="7">Pedal Ungual (left)</th>
<td box="[504,561,1416,1548]" gridcol="1" gridrow="24" pageId="6" pageNumber="7">D-1 D-2 D-3 D-4</td>
<td box="[770,917,1416,1548]" gridcol="2" gridrow="24" pageId="6" pageNumber="7">8.3 7.7 8.4 6.8</td>
</tr>
</table>
</paragraph>
<paragraph blockId="6.[122,770,1632,1964]" lastBlockId="7.[122,770,1112,1964]" lastPageId="7" lastPageNumber="8" pageId="6" pageNumber="7">
<taxonomicName authority=", Iberomesornis" authorityName="Iberomesornis" box="[122,371,1632,1652]" class="Aves" family="Gobipterygidae" genus="Gobipteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Gobipterygiformes" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis box="[122,224,1632,1652]" italics="true" pageId="6" pageNumber="7">Gobipteryx</emphasis>
,
<emphasis box="[242,371,1632,1652]" italics="true" pageId="6" pageNumber="7">Iberomesornis</emphasis>
</taxonomicName>
,
<taxonomicName box="[390,519,1632,1652]" class="Aves" family="Longipterygidae" genus="Longirostravis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Longipterygiformes" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis box="[390,519,1632,1652]" italics="true" pageId="6" pageNumber="7">Longirostravis</emphasis>
</taxonomicName>
,
<taxonomicName box="[536,629,1632,1652]" class="Aves" family="Longipterygidae" genus="Rapaxavis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Longipterygiformes" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis box="[536,629,1632,1652]" italics="true" pageId="6" pageNumber="7">Rapaxavis</emphasis>
</taxonomicName>
,
<taxonomicName authority=", Vescornis" authorityName="Vescornis" class="Aves" family="Longipterygidae" genus="Shanweiniao" higherTaxonomySource="GBIF" kingdom="Animalia" order="Longipterygiformes" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis box="[648,765,1632,1652]" italics="true" pageId="6" pageNumber="7">Shanweiniao</emphasis>
,
<emphasis box="[122,209,1667,1687]" italics="true" pageId="6" pageNumber="7">Vescornis</emphasis>
</taxonomicName>
,
<taxonomicName authorityName="Ji, Atterholt, O'Connor, Lamanna, Harris, Li, You &amp; Dodson" authorityYear="2011" box="[223,295,1667,1687]" class="Aves" genus="Qiliania" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis box="[223,295,1667,1687]" italics="true" pageId="6" pageNumber="7">Qiliania</emphasis>
</taxonomicName>
,
<taxonomicName box="[309,430,1667,1687]" class="Aves" genus="Fortunguavis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis box="[309,430,1667,1687]" italics="true" pageId="6" pageNumber="7">Fortunguavis</emphasis>
</taxonomicName>
, and
<emphasis box="[489,612,1667,1687]" italics="true" pageId="6" pageNumber="7">Songlingornis</emphasis>
) were removed from the reduced consensus analysis, and the reduced consensus tree is relatively better resolved (
<figureCitation box="[458,557,1736,1756]" captionStart="FIGURE 5" captionStartId="8.[123,215,1343,1364]" captionTargetBox="[122,1466,57,1301]" captionTargetId="figure-360@8.[129,1459,123,1302]" captionText="FIGURE 5 Phylogenetic position of Brevirostruavis macrohyoideus (IVPP V 13266) in Avialae. (a) Strict consensus tree from current phylogenetic analysis; (b) reduced consensus tree with ten most unstable taxa removed. The bootstrap and absolute Bremer support values are denoted in normal and bold Italic fonts" figureDoi="http://doi.org/10.5281/zenodo.5808232" httpUri="https://zenodo.org/record/5808232/files/figure.png" pageId="6" pageNumber="7">Figure 5B</figureCitation>
). However, the interrelationships among the Enantiornithes are still obscured by a few polytomies, and
<taxonomicName box="[286,425,1806,1826]" genus="Brevirostruavis" pageId="6" pageNumber="7" rank="genus">
<emphasis box="[286,425,1806,1826]" italics="true" pageId="6" pageNumber="7">Brevirostruavis</emphasis>
</taxonomicName>
falls into one of them (
<figureCitation box="[661,760,1806,1826]" captionStart="FIGURE 5" captionStartId="8.[123,215,1343,1364]" captionTargetBox="[122,1466,57,1301]" captionTargetId="figure-360@8.[129,1459,123,1302]" captionText="FIGURE 5 Phylogenetic position of Brevirostruavis macrohyoideus (IVPP V 13266) in Avialae. (a) Strict consensus tree from current phylogenetic analysis; (b) reduced consensus tree with ten most unstable taxa removed. The bootstrap and absolute Bremer support values are denoted in normal and bold Italic fonts" figureDoi="http://doi.org/10.5281/zenodo.5808232" httpUri="https://zenodo.org/record/5808232/files/figure.png" pageId="6" pageNumber="7">Figure 5B</figureCitation>
). A majority rule consensus tree was produced, and its topology is consistent with most recent studies in the composition of major groupings (
<bibRefCitation author="O'Connor, J. K. &amp; Wang, X. &amp; Chiappe, L. M. &amp; Gao, C. &amp; Meng, Q. &amp; Cheng, X." box="[228,439,1910,1930]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="6" pageNumber="7" pagination="188 - 204" part="29" refId="ref7756" refString="O'Connor, J. K., Wang, X., Chiappe, L. M., Gao, C., Meng, Q., Cheng, X. et al. (2009) Phylogenetic support for a specialized clade of Cretaceous enantiornithine birds with information from a new species. Journal of Vertebrate Paleontology, 29, 188 - 204." title="Phylogenetic support for a specialized clade of Cretaceous enantiornithine birds with information from a new species" type="journal article" year="2009">OConnor et al., 2009</bibRefCitation>
;
<bibRefCitation author="Wang, M. &amp; Li, Z. &amp; Zhou, Z." box="[447,628,1910,1930]" journalOrPublisher="Proceedings of the National Academy of Sciences" pageId="6" pageNumber="7" pagination="11470 - 11475" part="114" refId="ref8112" refString="Wang, M., Li, Z. &amp; Zhou, Z. (2017 a) Insight into the growth pattern and bone fusion of basal birds from an Early Cretaceous enantiornithine bird. Proceedings of the National Academy of Sciences, 114, 11470 - 11475." title="Insight into the growth pattern and bone fusion of basal birds from an Early Cretaceous enantiornithine bird" type="journal article" year="2017">Wang et al., 2017a</bibRefCitation>
,
<bibRefCitation author="Wang, M. &amp; O'Connor, J. K. &amp; Pan, Y. &amp; Zhou, Z." box="[636,701,1910,1930]" journalOrPublisher="Nature Communications" pageId="6" pageNumber="7" pagination="14141" part="8" refId="ref8195" refString="Wang, M., O'Connor, J. K., Pan, Y. &amp; Zhou, Z. (2017 b) A bizarre Early Cretaceous enantiornithine bird with unique crural feathers and an ornithuromorph plough-shaped pygostyle. Nature Communications, 8, 14141." title="A bizarre Early Cretaceous enantiornithine bird with unique crural feathers and an ornithuromorph plough-shaped pygostyle" type="journal article" year="2017">2017b</bibRefCitation>
). In the majority rule consensus tree,
<emphasis box="[409,501,1944,1964]" italics="true" pageId="6" pageNumber="7">Eoalulavis</emphasis>
,
<emphasis box="[513,600,1944,1964]" italics="true" pageId="6" pageNumber="7">Vescornis</emphasis>
,
<emphasis box="[611,722,1944,1964]" italics="true" pageId="6" pageNumber="7">Cathayornis</emphasis>
, and
<taxonomicName box="[818,1102,1632,1652]" genus="Brevirostruavis" pageId="6" pageNumber="7" rank="species" species="macrohyoideus">
<emphasis box="[818,1102,1632,1652]" italics="true" pageId="6" pageNumber="7">Brevirostruavis macrohyoideus</emphasis>
</taxonomicName>
are recovered as the successive outgroups to the
<taxonomicName authority="Concornis" authorityName="Chiappe &amp; Calvo" authorityYear="1994" box="[956,1084,1667,1687]" class="Aves" family="Avisauridae" genus="Neuquenornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Enantiornithiformes" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis box="[956,1084,1667,1687]" italics="true" pageId="6" pageNumber="7">Neuquenornis</emphasis>
</taxonomicName>
+ Concornis clade. In comparison with other enantiornithines (e.g.,
<emphasis box="[1098,1209,1702,1722]" italics="true" pageId="6" pageNumber="7">Cathayornis</emphasis>
), the new specimen has a proportionally shorter rostrum, reminiscent of, but more rounded than that of
<emphasis box="[937,1063,1771,1791]" italics="true" pageId="6" pageNumber="7">Eoenantiornis</emphasis>
(
<bibRefCitation author="Zhou, Z. &amp; Zhang, F." box="[1073,1275,1771,1791]" journalOrPublisher="Canadian Journal of Earth Sciences" pageId="6" pageNumber="7" pagination="731 - 747" part="40" refId="ref8879" refString="Zhou, Z. &amp; Zhang, F. (2003) Anatomy of the primitive bird Sapeornis chaoyangensis from the Early Cretaceous of Liaoning, China. Canadian Journal of Earth Sciences, 40, 731 - 747." title="Anatomy of the primitive bird Sapeornis chaoyangensis from the Early Cretaceous of Liaoning, China" type="journal article" year="2003">Zhou &amp; Zhang, 2003</bibRefCitation>
). In regards to postcranial features,
<taxonomicName box="[993,1282,1806,1826]" genus="Brevirostruavis" pageId="6" pageNumber="7" rank="species" species="macrohyoideus">
<emphasis box="[993,1282,1806,1826]" italics="true" pageId="6" pageNumber="7">Brevirostruavis macrohyoideus</emphasis>
</taxonomicName>
is different from
<emphasis box="[818,944,1840,1860]" italics="true" pageId="6" pageNumber="7">Eoenantiornis</emphasis>
in sternal morphology, including the extension of midline projection and the large craniolateral process. This process extends further dorsally in the new specimen than in most other taxa (
<bibRefCitation author="Zheng, X. &amp; Wang, X. &amp; Jingmai, K. O. C. &amp; Zhou, Z." box="[875,1065,1944,1964]" journalOrPublisher="Nature Communications" pageId="6" pageNumber="7" pagination="1116" part="3" refId="ref8725" refString="Zheng, X., Wang, X., Jingmai, K. O. C. &amp; Zhou, Z. (2012) Insight into the early evolution of the avian sternum from juvenile enantiornithines. Nature Communications, 3, 1116." title="Insight into the early evolution of the avian sternum from juvenile enantiornithines" type="journal article" year="2012">Zheng et al., 2012</bibRefCitation>
). The relative width proportions of the metatarsals (slightly narrower metatarsal IV in
<taxonomicName box="[567,705,1112,1132]" genus="Brevirostruavis" pageId="7" pageNumber="8" rank="genus">
<emphasis box="[567,705,1112,1132]" italics="true" pageId="7" pageNumber="8">Brevirostruavis</emphasis>
</taxonomicName>
) differ from
<emphasis box="[174,273,1147,1167]" italics="true" pageId="7" pageNumber="8">Cathyornis</emphasis>
and
<emphasis box="[321,412,1147,1167]" italics="true" pageId="7" pageNumber="8">Boluochia</emphasis>
, in which the metatarsal IV is significantly narrower than the other metatarsals. IVPP V13266 is distinct in its smaller and less robust teeth, in comparison to
<taxonomicName box="[630,728,1216,1236]" class="Aves" family="Bohaiornithidae" genus="Bohaiornis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
<emphasis box="[630,728,1216,1236]" italics="true" pageId="7" pageNumber="8">Bohaiornis</emphasis>
</taxonomicName>
and other closely related taxa in
<taxonomicName box="[396,554,1251,1271]" class="Aves" family="Bohaiornithidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="8" phylum="Chordata" rank="family">Bohaiornithidae</taxonomicName>
.
</paragraph>
<caption ID-Table-UUID="DF276609C464FF96FFEAF22031A7EAE3" box="[122,1436,124,144]" httpUri="http://table.plazi.org/id/DF276609C464FF96FFEAF22031A7EAE3" pageId="7" pageNumber="8" startId="7.[122,198,124,144]" targetBox="[138,1449,175,949]" targetIsTable="true" targetPageId="7">
<paragraph blockId="7.[122,222,124,144]" box="[122,222,124,144]" pageId="7" pageNumber="8">
<emphasis bold="true" box="[122,222,124,144]" pageId="7" pageNumber="8">TABLE 2</emphasis>
</paragraph>
<paragraph blockId="7.[241,1436,124,144]" box="[241,1436,124,144]" pageId="7" pageNumber="8">
Limb measurements of the
<taxonomicName box="[501,780,124,144]" genus="Brevirostruavis" pageId="7" pageNumber="8" rank="species" species="macrohyoideus">
<emphasis box="[501,780,124,144]" italics="true" pageId="7" pageNumber="8">Brevirostruavis macrohyoideus</emphasis>
</taxonomicName>
with comparison of other avialans (Adopted from
<bibRefCitation author="Zhang, Z. &amp; Chiappe, L. M. &amp; Han, G. &amp; Chinsamy, A." box="[1259,1431,124,144]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="7" pageNumber="8" pagination="1176 - 1189" part="33" refId="ref8545" refString="Zhang, Z., Chiappe, L. M., Han, G. &amp; Chinsamy, A. (2013) A large bird from the Early Cretaceous of China: new information on the skull of enantiornithines. Journal of Vertebrate Paleontology, 33, 1176 - 1189." title="A large bird from the Early Cretaceous of China: new information on the skull of enantiornithines" type="journal article" year="2013">Zhang et al., 2013</bibRefCitation>
)
</paragraph>
</caption>
<paragraph pageId="7" pageNumber="8">
<table box="[138,1449,175,949]" gridcols="8" gridrows="21" pageId="7" pageNumber="8">
<tr box="[138,1449,175,195]" gridrow="0" pageId="7" pageNumber="8">
<th box="[138,537,175,195]" gridcol="0" gridrow="0" pageId="7" pageNumber="8">
<emphasis bold="true" box="[138,179,175,195]" italics="true" pageId="7" pageNumber="8">Taxa</emphasis>
</th>
<th box="[580,622,175,195]" gridcol="1" gridrow="0" pageId="7" pageNumber="8">
<emphasis bold="true" box="[580,595,175,195]" pageId="7" pageNumber="8">H</emphasis>
</th>
<th box="[712,753,175,195]" gridcol="2" gridrow="0" pageId="7" pageNumber="8">
<emphasis bold="true" box="[712,727,175,195]" pageId="7" pageNumber="8">U</emphasis>
</th>
<th box="[850,892,175,195]" gridcol="3" gridrow="0" pageId="7" pageNumber="8">
<emphasis bold="true" box="[850,890,175,195]" pageId="7" pageNumber="8">Cmc</emphasis>
</th>
<th box="[954,995,175,195]" gridcol="4" gridrow="0" pageId="7" pageNumber="8">
<emphasis bold="true" box="[954,976,175,195]" pageId="7" pageNumber="8">Fe</emphasis>
</th>
<th box="[1091,1133,175,195]" gridcol="5" gridrow="0" pageId="7" pageNumber="8">
<emphasis bold="true" box="[1091,1122,175,195]" pageId="7" pageNumber="8">Tbt</emphasis>
</th>
<th box="[1184,1225,175,195]" gridcol="6" gridrow="0" pageId="7" pageNumber="8">
<emphasis bold="true" box="[1184,1219,175,195]" pageId="7" pageNumber="8">Tmt</emphasis>
</th>
<th box="[1288,1449,175,195]" gridcol="7" gridrow="0" pageId="7" pageNumber="8">
<emphasis bold="true" box="[1288,1449,175,195]" italics="true" pageId="7" pageNumber="8">Forelimb/hindlimb</emphasis>
</th>
</tr>
<tr box="[138,1449,218,238]" gridrow="1" pageId="7" pageNumber="8">
<th box="[138,537,218,238]" gridcol="0" gridrow="1" pageId="7" pageNumber="8">
<taxonomicName box="[138,312,218,238]" class="Aves" family="Songlingornithidae" genus="Aberratiodontus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Chaoyangiformes" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="wui">
<emphasis box="[138,312,218,238]" italics="true" pageId="7" pageNumber="8">Aberratiodontus wui</emphasis>
</taxonomicName>
(LHV0001)
</th>
<td box="[580,622,218,238]" gridcol="1" gridrow="1" pageId="7" pageNumber="8"></td>
<td box="[712,753,218,238]" gridcol="2" gridrow="1" pageId="7" pageNumber="8"></td>
<td box="[850,892,218,238]" gridcol="3" gridrow="1" pageId="7" pageNumber="8"></td>
<td box="[954,995,218,238]" gridcol="4" gridrow="1" pageId="7" pageNumber="8">55.0</td>
<td box="[1091,1133,218,238]" gridcol="5" gridrow="1" pageId="7" pageNumber="8">66.7</td>
<td box="[1184,1225,218,238]" gridcol="6" gridrow="1" pageId="7" pageNumber="8">33.0</td>
<td box="[1288,1449,218,238]" gridcol="7" gridrow="1" pageId="7" pageNumber="8">NA</td>
</tr>
<tr box="[138,1449,255,275]" gridrow="2" pageId="7" pageNumber="8">
<th box="[138,537,255,275]" gridcol="0" gridrow="2" pageId="7" pageNumber="8">
<taxonomicName box="[138,282,255,275]" class="Aves" family="Longipterygidae" genus="Boluochia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="zhengi">
<emphasis box="[138,282,255,275]" italics="true" pageId="7" pageNumber="8">Boluochia zhengi</emphasis>
</taxonomicName>
(IVPP 9770)
</th>
<td box="[580,622,255,275]" gridcol="1" gridrow="2" pageId="7" pageNumber="8"></td>
<td box="[712,753,255,275]" gridcol="2" gridrow="2" pageId="7" pageNumber="8"></td>
<td box="[850,892,255,275]" gridcol="3" gridrow="2" pageId="7" pageNumber="8"></td>
<td box="[954,995,255,275]" gridcol="4" gridrow="2" pageId="7" pageNumber="8"></td>
<td box="[1091,1133,255,275]" gridcol="5" gridrow="2" pageId="7" pageNumber="8">37.0</td>
<td box="[1184,1225,255,275]" gridcol="6" gridrow="2" pageId="7" pageNumber="8">17.5</td>
<td box="[1288,1449,255,275]" gridcol="7" gridrow="2" pageId="7" pageNumber="8">NA</td>
</tr>
<tr box="[138,1449,293,313]" gridrow="3" pageId="7" pageNumber="8">
<th box="[138,537,293,313]" gridcol="0" gridrow="3" pageId="7" pageNumber="8">
<taxonomicName box="[138,400,293,313]" genus="Brevirostruavis" pageId="7" pageNumber="8" rank="species" species="macrohyoideus">
<emphasis box="[138,400,293,313]" italics="true" pageId="7" pageNumber="8">Brevirostruavis macrohyoideus</emphasis>
</taxonomicName>
(IVPP V13266)
</th>
<td box="[580,622,293,313]" gridcol="1" gridrow="3" pageId="7" pageNumber="8">39.2</td>
<td box="[712,753,293,313]" gridcol="2" gridrow="3" pageId="7" pageNumber="8">38.6</td>
<td box="[850,892,293,313]" gridcol="3" gridrow="3" pageId="7" pageNumber="8">15.0</td>
<td box="[954,995,293,313]" gridcol="4" gridrow="3" pageId="7" pageNumber="8">28.3</td>
<td box="[1091,1133,293,313]" gridcol="5" gridrow="3" pageId="7" pageNumber="8">40.0</td>
<td box="[1184,1225,293,313]" gridcol="6" gridrow="3" pageId="7" pageNumber="8">20.4</td>
<td box="[1288,1449,293,313]" gridcol="7" gridrow="3" pageId="7" pageNumber="8">1.05</td>
</tr>
<tr box="[138,1449,330,350]" gridrow="4" pageId="7" pageNumber="8">
<th box="[138,537,330,350]" gridcol="0" gridrow="4" pageId="7" pageNumber="8">
<taxonomicName box="[138,340,330,350]" pageId="7" pageNumber="8">
<emphasis box="[138,340,330,350]" italics="true" pageId="7" pageNumber="8">Cathayornis chabuensis</emphasis>
</taxonomicName>
(BMNHC-Ph000110)
</th>
<td box="[580,622,330,350]" gridcol="1" gridrow="4" pageId="7" pageNumber="8">31.5</td>
<td box="[712,753,330,350]" gridcol="2" gridrow="4" pageId="7" pageNumber="8">32.1</td>
<td box="[850,892,330,350]" gridcol="3" gridrow="4" pageId="7" pageNumber="8">14.0</td>
<td box="[954,995,330,350]" gridcol="4" gridrow="4" pageId="7" pageNumber="8">30.7</td>
<td box="[1091,1133,330,350]" gridcol="5" gridrow="4" pageId="7" pageNumber="8">35.9</td>
<td box="[1184,1225,330,350]" gridcol="6" gridrow="4" pageId="7" pageNumber="8">17.2</td>
<td box="[1288,1449,330,350]" gridcol="7" gridrow="4" pageId="7" pageNumber="8">0.93</td>
</tr>
<tr box="[138,1449,368,388]" gridrow="5" pageId="7" pageNumber="8">
<th box="[138,537,368,388]" gridcol="0" gridrow="5" pageId="7" pageNumber="8">
<taxonomicName box="[138,313,368,388]" class="Aves" genus="Cathayornis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="yandica">
<emphasis box="[138,313,368,388]" italics="true" pageId="7" pageNumber="8">Cathayornis yandica</emphasis>
</taxonomicName>
(IVPP V9769)
</th>
<td box="[580,622,368,388]" gridcol="1" gridrow="5" pageId="7" pageNumber="8">33.0</td>
<td box="[712,753,368,388]" gridcol="2" gridrow="5" pageId="7" pageNumber="8">34.0</td>
<td box="[850,892,368,388]" gridcol="3" gridrow="5" pageId="7" pageNumber="8">18.0</td>
<td box="[954,995,368,388]" gridcol="4" gridrow="5" pageId="7" pageNumber="8">28.0</td>
<td box="[1091,1133,368,388]" gridcol="5" gridrow="5" pageId="7" pageNumber="8">34.0</td>
<td box="[1184,1225,368,388]" gridcol="6" gridrow="5" pageId="7" pageNumber="8">21.0</td>
<td box="[1288,1449,368,388]" gridcol="7" gridrow="5" pageId="7" pageNumber="8">1.02</td>
</tr>
<tr box="[138,1449,405,425]" gridrow="6" pageId="7" pageNumber="8">
<th box="[138,537,405,425]" gridcol="0" gridrow="6" pageId="7" pageNumber="8">
<taxonomicName box="[138,305,405,425]" class="Aves" genus="Dalingheornis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="liweii">
<emphasis box="[138,305,405,425]" italics="true" pageId="7" pageNumber="8">Dalingheornis liweii</emphasis>
</taxonomicName>
(CNU VB2005001)
</th>
<td box="[580,622,405,425]" gridcol="1" gridrow="6" pageId="7" pageNumber="8">14.0</td>
<td box="[712,753,405,425]" gridcol="2" gridrow="6" pageId="7" pageNumber="8">14.0</td>
<td box="[850,892,405,425]" gridcol="3" gridrow="6" pageId="7" pageNumber="8">6.0</td>
<td box="[954,995,405,425]" gridcol="4" gridrow="6" pageId="7" pageNumber="8">11.0</td>
<td box="[1091,1133,405,425]" gridcol="5" gridrow="6" pageId="7" pageNumber="8">16.0</td>
<td box="[1184,1225,405,425]" gridcol="6" gridrow="6" pageId="7" pageNumber="8">9.0</td>
<td box="[1288,1449,405,425]" gridcol="7" gridrow="6" pageId="7" pageNumber="8">0.94</td>
</tr>
<tr box="[138,1449,442,462]" gridrow="7" pageId="7" pageNumber="8">
<th box="[138,537,442,462]" gridcol="0" gridrow="7" pageId="7" pageNumber="8">
<taxonomicName box="[138,397,442,462]" class="Aves" family="Eoenantiornithidae" genus="Dapingfangornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Eoenantiornithiformes" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="sentisorhinus">
<emphasis box="[138,397,442,462]" italics="true" pageId="7" pageNumber="8">Dapingfangornis sentisorhinus</emphasis>
</taxonomicName>
(LPM00039)
</th>
<td box="[580,622,442,462]" gridcol="1" gridrow="7" pageId="7" pageNumber="8">22.0</td>
<td box="[712,753,442,462]" gridcol="2" gridrow="7" pageId="7" pageNumber="8">27.0</td>
<td box="[850,892,442,462]" gridcol="3" gridrow="7" pageId="7" pageNumber="8">11.0</td>
<td box="[954,995,442,462]" gridcol="4" gridrow="7" pageId="7" pageNumber="8">23.0</td>
<td box="[1091,1133,442,462]" gridcol="5" gridrow="7" pageId="7" pageNumber="8">29.0</td>
<td box="[1184,1225,442,462]" gridcol="6" gridrow="7" pageId="7" pageNumber="8">16.0</td>
<td box="[1288,1449,442,462]" gridcol="7" gridrow="7" pageId="7" pageNumber="8">0.88</td>
</tr>
<tr box="[138,1449,480,500]" gridrow="8" pageId="7" pageNumber="8">
<th box="[138,537,480,500]" gridcol="0" gridrow="8" pageId="7" pageNumber="8">
<taxonomicName authorityName="Zhou" authorityYear="2002" box="[138,325,480,500]" class="Aves" family="Cathayornithidae" genus="Eocathayornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cathayornithiformes" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="walkeri">
<emphasis box="[138,325,480,500]" italics="true" pageId="7" pageNumber="8">Eocathayornis walkeri</emphasis>
</taxonomicName>
(IVPP 10916)
</th>
<td box="[580,622,480,500]" gridcol="1" gridrow="8" pageId="7" pageNumber="8">23.5</td>
<td box="[712,753,480,500]" gridcol="2" gridrow="8" pageId="7" pageNumber="8">26.0</td>
<td box="[850,892,480,500]" gridcol="3" gridrow="8" pageId="7" pageNumber="8">14.0</td>
<td box="[954,995,480,500]" gridcol="4" gridrow="8" pageId="7" pageNumber="8"></td>
<td box="[1091,1133,480,500]" gridcol="5" gridrow="8" pageId="7" pageNumber="8"></td>
<td box="[1184,1225,480,500]" gridcol="6" gridrow="8" pageId="7" pageNumber="8"></td>
<td box="[1288,1449,480,500]" gridcol="7" gridrow="8" pageId="7" pageNumber="8">NA</td>
</tr>
<tr box="[138,1449,517,537]" gridrow="9" pageId="7" pageNumber="8">
<th box="[138,537,517,537]" gridcol="0" gridrow="9" pageId="7" pageNumber="8">
<taxonomicName box="[138,319,517,537]" class="Aves" genus="Eoenantiornis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="buhleri">
<emphasis box="[138,319,517,537]" italics="true" pageId="7" pageNumber="8">Eoenantiornis buhleri</emphasis>
</taxonomicName>
(IVPP V11537)
</th>
<td box="[580,622,517,537]" gridcol="1" gridrow="9" pageId="7" pageNumber="8">29.5</td>
<td box="[712,753,517,537]" gridcol="2" gridrow="9" pageId="7" pageNumber="8">31.0</td>
<td box="[850,892,517,537]" gridcol="3" gridrow="9" pageId="7" pageNumber="8">12.0</td>
<td box="[954,995,517,537]" gridcol="4" gridrow="9" pageId="7" pageNumber="8">26.5</td>
<td box="[1091,1133,517,537]" gridcol="5" gridrow="9" pageId="7" pageNumber="8">31.0</td>
<td box="[1184,1225,517,537]" gridcol="6" gridrow="9" pageId="7" pageNumber="8">22.3</td>
<td box="[1288,1449,517,537]" gridcol="7" gridrow="9" pageId="7" pageNumber="8">0.91</td>
</tr>
<tr box="[138,1449,555,575]" gridrow="10" pageId="7" pageNumber="8">
<th box="[138,537,555,575]" gridcol="0" gridrow="10" pageId="7" pageNumber="8">
<taxonomicName baseAuthorityName="Stidham &amp; O'Connor" baseAuthorityYear="2021" box="[138,367,555,575]" class="Aves" family="Longipterygidae" genus="Longipteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Longipterygiformes" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="chaoyangensis">
<emphasis box="[138,367,555,575]" italics="true" pageId="7" pageNumber="8">Longipteryx chaoyangensis</emphasis>
</taxonomicName>
(IVPP V12325)
</th>
<td box="[580,622,555,575]" gridcol="1" gridrow="10" pageId="7" pageNumber="8">45.0</td>
<td box="[712,753,555,575]" gridcol="2" gridrow="10" pageId="7" pageNumber="8">47.0</td>
<td box="[850,892,555,575]" gridcol="3" gridrow="10" pageId="7" pageNumber="8">19.0</td>
<td box="[954,995,555,575]" gridcol="4" gridrow="10" pageId="7" pageNumber="8">31.0</td>
<td box="[1091,1133,555,575]" gridcol="5" gridrow="10" pageId="7" pageNumber="8">32.0</td>
<td box="[1184,1225,555,575]" gridcol="6" gridrow="10" pageId="7" pageNumber="8">21.0</td>
<td box="[1288,1449,555,575]" gridcol="7" gridrow="10" pageId="7" pageNumber="8">1.32</td>
</tr>
<tr box="[138,1449,592,612]" gridrow="11" pageId="7" pageNumber="8">
<th box="[138,537,592,612]" gridcol="0" gridrow="11" pageId="7" pageNumber="8">
<taxonomicName box="[138,299,592,612]" class="Aves" family="Longipterygidae" genus="Longirostravis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Longipterygiformes" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="hani">
<emphasis box="[138,299,592,612]" italics="true" pageId="7" pageNumber="8">Longirostravis hani</emphasis>
</taxonomicName>
(IVPP11309)
</th>
<td box="[580,622,592,612]" gridcol="1" gridrow="11" pageId="7" pageNumber="8">24.4</td>
<td box="[712,753,592,612]" gridcol="2" gridrow="11" pageId="7" pageNumber="8">25.5</td>
<td box="[850,892,592,612]" gridcol="3" gridrow="11" pageId="7" pageNumber="8"></td>
<td box="[954,995,592,612]" gridcol="4" gridrow="11" pageId="7" pageNumber="8">20.0</td>
<td box="[1091,1133,592,612]" gridcol="5" gridrow="11" pageId="7" pageNumber="8">25.5</td>
<td box="[1184,1225,592,612]" gridcol="6" gridrow="11" pageId="7" pageNumber="8">14.0</td>
<td box="[1288,1449,592,612]" gridcol="7" gridrow="11" pageId="7" pageNumber="8">NA</td>
</tr>
<tr box="[138,1449,630,650]" gridrow="12" pageId="7" pageNumber="8">
<th box="[138,537,630,650]" gridcol="0" gridrow="12" pageId="7" pageNumber="8">
<taxonomicName box="[138,342,630,650]" class="Aves" genus="Paraprotopteryx" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="gracilis">
<emphasis box="[138,342,630,650]" italics="true" pageId="7" pageNumber="8">Paraprotopteryx gracilis</emphasis>
</taxonomicName>
(STM V001)
</th>
<td box="[580,622,630,650]" gridcol="1" gridrow="12" pageId="7" pageNumber="8">22.6</td>
<td box="[712,753,630,650]" gridcol="2" gridrow="12" pageId="7" pageNumber="8">23.5</td>
<td box="[850,892,630,650]" gridcol="3" gridrow="12" pageId="7" pageNumber="8">11.5</td>
<td box="[954,995,630,650]" gridcol="4" gridrow="12" pageId="7" pageNumber="8">22.2</td>
<td box="[1091,1133,630,650]" gridcol="5" gridrow="12" pageId="7" pageNumber="8">26.3</td>
<td box="[1184,1225,630,650]" gridcol="6" gridrow="12" pageId="7" pageNumber="8">15.7</td>
<td box="[1288,1449,630,650]" gridcol="7" gridrow="12" pageId="7" pageNumber="8">0.90</td>
</tr>
<tr box="[138,1449,667,687]" gridrow="13" pageId="7" pageNumber="8">
<th box="[138,537,667,687]" gridcol="0" gridrow="13" pageId="7" pageNumber="8">
<taxonomicName box="[138,263,667,687]" class="Aves" family="Pengornithidae" genus="Pengornis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="houi">
<emphasis box="[138,263,667,687]" italics="true" pageId="7" pageNumber="8">Pengornis houi</emphasis>
</taxonomicName>
(IVPP V15336)
</th>
<td box="[580,622,667,687]" gridcol="1" gridrow="13" pageId="7" pageNumber="8">64.3</td>
<td box="[712,753,667,687]" gridcol="2" gridrow="13" pageId="7" pageNumber="8">70.7</td>
<td box="[850,892,667,687]" gridcol="3" gridrow="13" pageId="7" pageNumber="8">34.3</td>
<td box="[954,995,667,687]" gridcol="4" gridrow="13" pageId="7" pageNumber="8">48.0</td>
<td box="[1091,1133,667,687]" gridcol="5" gridrow="13" pageId="7" pageNumber="8">50.4</td>
<td box="[1184,1225,667,687]" gridcol="6" gridrow="13" pageId="7" pageNumber="8">26.5</td>
<td box="[1288,1449,667,687]" gridcol="7" gridrow="13" pageId="7" pageNumber="8">1.36</td>
</tr>
<tr box="[138,1449,705,725]" gridrow="14" pageId="7" pageNumber="8">
<th box="[138,537,705,725]" gridcol="0" gridrow="14" pageId="7" pageNumber="8">
<taxonomicName box="[138,360,705,725]" class="Aves" genus="Protopteryx" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="fengningensis">
<emphasis box="[138,360,705,725]" italics="true" pageId="7" pageNumber="8">Protopteryx fengningensis</emphasis>
</taxonomicName>
(IVPP11665)
</th>
<td box="[580,622,705,725]" gridcol="1" gridrow="14" pageId="7" pageNumber="8">27.5</td>
<td box="[712,753,705,725]" gridcol="2" gridrow="14" pageId="7" pageNumber="8">27.9</td>
<td box="[850,892,705,725]" gridcol="3" gridrow="14" pageId="7" pageNumber="8">14.7</td>
<td box="[954,995,705,725]" gridcol="4" gridrow="14" pageId="7" pageNumber="8">22.6</td>
<td box="[1091,1133,705,725]" gridcol="5" gridrow="14" pageId="7" pageNumber="8">29.1</td>
<td box="[1184,1225,705,725]" gridcol="6" gridrow="14" pageId="7" pageNumber="8">16.5</td>
<td box="[1288,1449,705,725]" gridcol="7" gridrow="14" pageId="7" pageNumber="8">1.03</td>
</tr>
<tr box="[138,1449,742,762]" gridrow="15" pageId="7" pageNumber="8">
<th box="[138,537,742,762]" gridcol="0" gridrow="15" pageId="7" pageNumber="8">
<taxonomicName box="[138,266,742,762]" class="Aves" family="Longipterygidae" genus="Rapaxavis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Longipterygiformes" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="pani">
<emphasis box="[138,266,742,762]" italics="true" pageId="7" pageNumber="8">Rapaxavis pani</emphasis>
</taxonomicName>
(DNHM 2522)
</th>
<td box="[580,622,742,762]" gridcol="1" gridrow="15" pageId="7" pageNumber="8">22.7</td>
<td box="[712,753,742,762]" gridcol="2" gridrow="15" pageId="7" pageNumber="8">22.9</td>
<td box="[850,892,742,762]" gridcol="3" gridrow="15" pageId="7" pageNumber="8">11.8</td>
<td box="[954,995,742,762]" gridcol="4" gridrow="15" pageId="7" pageNumber="8">19.3</td>
<td box="[1091,1133,742,762]" gridcol="5" gridrow="15" pageId="7" pageNumber="8">23.2</td>
<td box="[1184,1225,742,762]" gridcol="6" gridrow="15" pageId="7" pageNumber="8">13.0</td>
<td box="[1288,1449,742,762]" gridcol="7" gridrow="15" pageId="7" pageNumber="8">1.03</td>
</tr>
<tr box="[138,1449,779,799]" gridrow="16" pageId="7" pageNumber="8">
<th box="[138,537,779,799]" gridcol="0" gridrow="16" pageId="7" pageNumber="8">
<taxonomicName box="[138,354,779,799]" class="Aves" family="Longipterygidae" genus="Shanweiniao" higherTaxonomySource="GBIF" kingdom="Animalia" order="Longipterygiformes" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="cooperorum">
<emphasis box="[138,354,779,799]" italics="true" pageId="7" pageNumber="8">Shanweiniao cooperorum</emphasis>
</taxonomicName>
(DNHM 1878/1/2)
</th>
<td box="[580,622,779,799]" gridcol="1" gridrow="16" pageId="7" pageNumber="8">22.4</td>
<td box="[712,753,779,799]" gridcol="2" gridrow="16" pageId="7" pageNumber="8">23.4</td>
<td box="[850,892,779,799]" gridcol="3" gridrow="16" pageId="7" pageNumber="8"></td>
<td box="[954,995,779,799]" gridcol="4" gridrow="16" pageId="7" pageNumber="8">17.6</td>
<td box="[1091,1133,779,799]" gridcol="5" gridrow="16" pageId="7" pageNumber="8">22.5</td>
<td box="[1184,1225,779,799]" gridcol="6" gridrow="16" pageId="7" pageNumber="8">11.8</td>
<td box="[1288,1449,779,799]" gridcol="7" gridrow="16" pageId="7" pageNumber="8">NA</td>
</tr>
<tr box="[138,1449,817,837]" gridrow="17" pageId="7" pageNumber="8">
<th box="[138,537,817,837]" gridcol="0" gridrow="17" pageId="7" pageNumber="8">
<taxonomicName box="[138,293,817,837]" class="Aves" family="Bohaiornithidae" genus="Shenqiornis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="mengi">
<emphasis box="[138,293,817,837]" italics="true" pageId="7" pageNumber="8">Shenqiornis mengi</emphasis>
</taxonomicName>
(DNHM 2950)
</th>
<td box="[580,622,817,837]" gridcol="1" gridrow="17" pageId="7" pageNumber="8">46.6</td>
<td box="[712,753,817,837]" gridcol="2" gridrow="17" pageId="7" pageNumber="8">46.8</td>
<td box="[850,892,817,837]" gridcol="3" gridrow="17" pageId="7" pageNumber="8">25.5</td>
<td box="[954,995,817,837]" gridcol="4" gridrow="17" pageId="7" pageNumber="8">38.8</td>
<td box="[1091,1133,817,837]" gridcol="5" gridrow="17" pageId="7" pageNumber="8"></td>
<td box="[1184,1225,817,837]" gridcol="6" gridrow="17" pageId="7" pageNumber="8">25.0</td>
<td box="[1288,1449,817,837]" gridcol="7" gridrow="17" pageId="7" pageNumber="8">NA</td>
</tr>
<tr box="[138,1449,854,874]" gridrow="18" pageId="7" pageNumber="8">
<th box="[138,537,854,874]" gridcol="0" gridrow="18" pageId="7" pageNumber="8">
<taxonomicName box="[138,288,854,874]" pageId="7" pageNumber="8">
<emphasis box="[138,288,854,874]" italics="true" pageId="7" pageNumber="8">Sinornis santensis</emphasis>
</taxonomicName>
(BVP 538)
</th>
<td box="[580,622,854,874]" gridcol="1" gridrow="18" pageId="7" pageNumber="8">24.0</td>
<td box="[712,753,854,874]" gridcol="2" gridrow="18" pageId="7" pageNumber="8"></td>
<td box="[850,892,854,874]" gridcol="3" gridrow="18" pageId="7" pageNumber="8">10.8</td>
<td box="[954,995,854,874]" gridcol="4" gridrow="18" pageId="7" pageNumber="8"></td>
<td box="[1091,1133,854,874]" gridcol="5" gridrow="18" pageId="7" pageNumber="8">26.4</td>
<td box="[1184,1225,854,874]" gridcol="6" gridrow="18" pageId="7" pageNumber="8">14.6</td>
<td box="[1288,1449,854,874]" gridcol="7" gridrow="18" pageId="7" pageNumber="8">NA</td>
</tr>
<tr box="[138,1449,892,912]" gridrow="19" pageId="7" pageNumber="8">
<th box="[138,537,892,912]" gridcol="0" gridrow="19" pageId="7" pageNumber="8">
<taxonomicName box="[138,312,892,912]" class="Aves" genus="Vescornis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="hebeiensis">
<emphasis box="[138,312,892,912]" italics="true" pageId="7" pageNumber="8">Vescornis hebeiensis</emphasis>
</taxonomicName>
(IVPP130722)
</th>
<td box="[580,622,892,912]" gridcol="1" gridrow="19" pageId="7" pageNumber="8">22.6</td>
<td box="[712,753,892,912]" gridcol="2" gridrow="19" pageId="7" pageNumber="8">16.1</td>
<td box="[850,892,892,912]" gridcol="3" gridrow="19" pageId="7" pageNumber="8">11.6</td>
<td box="[954,995,892,912]" gridcol="4" gridrow="19" pageId="7" pageNumber="8">24.1</td>
<td box="[1091,1133,892,912]" gridcol="5" gridrow="19" pageId="7" pageNumber="8">29.8</td>
<td box="[1184,1225,892,912]" gridcol="6" gridrow="19" pageId="7" pageNumber="8">16.5</td>
<td box="[1288,1449,892,912]" gridcol="7" gridrow="19" pageId="7" pageNumber="8">0.71</td>
</tr>
<tr box="[138,1449,929,949]" gridrow="20" pageId="7" pageNumber="8">
<th box="[138,537,929,949]" gridcol="0" gridrow="20" pageId="7" pageNumber="8">
<taxonomicName box="[138,265,929,949]" class="Aves" family="Bohaiornithidae" genus="Zhouornis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="hani">
<emphasis box="[138,265,929,949]" italics="true" pageId="7" pageNumber="8">Zhouornis hani</emphasis>
</taxonomicName>
(CNU VB0903)
</th>
<td box="[580,622,929,949]" gridcol="1" gridrow="20" pageId="7" pageNumber="8">50.6</td>
<td box="[712,753,929,949]" gridcol="2" gridrow="20" pageId="7" pageNumber="8">53.8</td>
<td box="[850,892,929,949]" gridcol="3" gridrow="20" pageId="7" pageNumber="8">24.0</td>
<td box="[954,995,929,949]" gridcol="4" gridrow="20" pageId="7" pageNumber="8">44.0</td>
<td box="[1091,1133,929,949]" gridcol="5" gridrow="20" pageId="7" pageNumber="8">51.6</td>
<td box="[1184,1225,929,949]" gridcol="6" gridrow="20" pageId="7" pageNumber="8">25.9</td>
<td box="[1288,1449,929,949]" gridcol="7" gridrow="20" pageId="7" pageNumber="8">1.06</td>
</tr>
</table>
</paragraph>
<paragraph blockId="7.[122,1444,973,1049]" pageId="7" pageNumber="8">
<tableNote pageId="7" pageNumber="8" targetBox="[138,1449,175,949]" targetPageId="7">Anatomical abbreviations: H--humerus, U--ulna, Cmc--carpometacarpus, Tbt--tibiotarsus, and Tmt--tarsometatarsus. Institution abbreviations: DNHM- Dalian Natural History Museum, BMNHC/BVP-Beijing Museum of Natural History, China,CNU- Capital Normal University, and STM Shandong Tianyu Museum.</tableNote>
</paragraph>
<paragraph blockId="7.[122,770,1112,1964]" lastBlockId="7.[818,1467,1112,1964]" pageId="7" pageNumber="8">
The known ecological diversity of the Enantiornithes is largely inferred on basis of variation in the shape of the rostrum and dental morphologies (
<bibRefCitation author="Li, Z. &amp; Zhou, Z. &amp; Wang, M. &amp; Clarke, J. A." box="[340,482,1355,1375]" journalOrPublisher="Journal of Paleontology" pageId="7" pageNumber="8" pagination="99 - 108" part="88" refId="ref7568" refString="Li, Z., Zhou, Z., Wang, M. &amp; Clarke, J. A. (2014) A new specimen of largebodied basal enantiornithine Bohaiornis from the Early Cretaceous of China and the inference of feeding ecology in Mesozoic birds. Journal of Paleontology, 88, 99 - 108." title="A new specimen of largebodied basal enantiornithine Bohaiornis from the Early Cretaceous of China and the inference of feeding ecology in Mesozoic birds" type="journal article" year="2014">Li et al., 2014</bibRefCitation>
,
<bibRefCitation author="Li, Z. &amp; Wang, C. - C. &amp; Wang, M. &amp; Chiang, C. - C. &amp; Wang, Y. &amp; Zheng, X." box="[492,546,1355,1375]" journalOrPublisher="BMC Evolutionary Biology" pageId="7" pageNumber="8" pagination="1 - 8" part="20" refId="ref7462" refString="Li, Z., Wang, C. - C., Wang, M., Chiang, C. - C., Wang, Y., Zheng, X. et al. (2020) Ultramicrostructural reductions in teeth: implications for dietary transition from non-avian dinosaurs to birds. BMC Evolutionary Biology, 20, 1 - 8." title="Ultramicrostructural reductions in teeth: implications for dietary transition from non-avian dinosaurs to birds" type="journal article" year="2020">2020</bibRefCitation>
). An inferred raptorial feeding ecology and fish consumption have been proposed for the long-rostrumed
<taxonomicName baseAuthorityName="Stidham &amp; O'Connor" baseAuthorityYear="2021" box="[280,390,1424,1444]" class="Aves" family="Longipterygidae" genus="Longipteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Longipterygiformes" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
<emphasis box="[280,390,1424,1444]" italics="true" pageId="7" pageNumber="8">Longipteryx</emphasis>
</taxonomicName>
and
<emphasis box="[436,568,1424,1444]" italics="true" pageId="7" pageNumber="8">Changzuiornis</emphasis>
(
<bibRefCitation author="Huang, J. &amp; Wang, X. &amp; Hu, Y. &amp; Liu, J. &amp; Peteya, J. A. &amp; Clarke, J. A." box="[579,761,1424,1444]" journalOrPublisher="PeerJ" pageId="7" pageNumber="8" pagination="1765" part="4" refId="ref7303" refString="Huang, J., Wang, X., Hu, Y., Liu, J., Peteya, J. A. &amp; Clarke, J. A. (2016) A new ornithurine from the Early Cretaceous of China sheds light on the evolution of early ecological and cranial diversity in birds. PeerJ, 4, e 1765." title="A new ornithurine from the Early Cretaceous of China sheds light on the evolution of early ecological and cranial diversity in birds" type="journal article" year="2016">Huang et al., 2016</bibRefCitation>
). A recent study of the quadrate in
<taxonomicName baseAuthorityName="Stidham &amp; O'Connor" baseAuthorityYear="2021" box="[456,566,1459,1479]" class="Aves" family="Longipterygidae" genus="Longipteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Longipterygiformes" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
<emphasis box="[456,566,1459,1479]" italics="true" pageId="7" pageNumber="8">Longipteryx</emphasis>
</taxonomicName>
suggests potentially a stronger or quicker bite in that bird (
<bibRefCitation author="Stidham, T. A. &amp; O'Connor, J. K." box="[494,760,1494,1514]" journalOrPublisher="Journal of Anatomy" pageId="7" pageNumber="8" pagination="1066 - 1074" part="239" refId="ref8012" refString="Stidham, T. A. &amp; O'Connor, J. K. (2021) The evolutionary and functional implications of the unusual quadrate of Longipteryx chaoyangensis (Avialae: Enantiornithes) from the Cretaceous Jehol Biota of China. Journal of Anatomy, 239, 1066 - 1074." title="The evolutionary and functional implications of the unusual quadrate of Longipteryx chaoyangensis (Avialae: Enantiornithes) from the Cretaceous Jehol Biota of China" type="journal article" year="2021">Stidham &amp; O'Connor, 2021</bibRefCitation>
). The long and delicate rostrum of
<taxonomicName box="[457,587,1528,1548]" class="Aves" family="Longipterygidae" genus="Longirostravis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Longipterygiformes" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
<emphasis box="[457,587,1528,1548]" italics="true" pageId="7" pageNumber="8">Longirostravis</emphasis>
</taxonomicName>
has been hypothesized for filtering invertebrate and other nutrients from muddy sediments (
<bibRefCitation author="Hou, L. &amp; Chiappe, L. M. &amp; Zhang, F. &amp; Chuong, C. - M." box="[234,399,1598,1618]" journalOrPublisher="Die Naturwissenschaften" pageId="7" pageNumber="8" pagination="22 - 25" part="91" refId="ref7252" refString="Hou, L., Chiappe, L. M., Zhang, F. &amp; Chuong, C. - M. (2004) New early Cretaceous fossil from China documents a novel trophic specialization for Mesozoic birds. Die Naturwissenschaften, 91, 22 - 25." title="New early Cretaceous fossil from China documents a novel trophic specialization for Mesozoic birds" type="journal article" year="2004">Hou et al., 2004</bibRefCitation>
). In contrast,
<taxonomicName box="[534,691,1598,1618]" class="Aves" family="Bohaiornithidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="8" phylum="Chordata" rank="family">Bohaiornithidae</taxonomicName>
is characterized by a stout rostrum (
<bibRefCitation author="Li, Z. &amp; Zhou, Z. &amp; Wang, M. &amp; Clarke, J. A." box="[417,556,1632,1652]" journalOrPublisher="Journal of Paleontology" pageId="7" pageNumber="8" pagination="99 - 108" part="88" refId="ref7568" refString="Li, Z., Zhou, Z., Wang, M. &amp; Clarke, J. A. (2014) A new specimen of largebodied basal enantiornithine Bohaiornis from the Early Cretaceous of China and the inference of feeding ecology in Mesozoic birds. Journal of Paleontology, 88, 99 - 108." title="A new specimen of largebodied basal enantiornithine Bohaiornis from the Early Cretaceous of China and the inference of feeding ecology in Mesozoic birds" type="journal article" year="2014">Li et al., 2014</bibRefCitation>
). This wide variety of rostral shapes among enantiornithines suggests that the combination of cranial characters and shapes may have facilitated the reduction of competition among these sympatric taxa, maximizing the exploitation of food resources. The spectrum of rostral variation can be analogous to the diversified beak forms among crown group birds (
<bibRefCitation author="O'Connor, P. M. &amp; Turner, A. H. &amp; Groenke, J. R. &amp; Felice, R. N. &amp; Rogers, R. R. &amp; Krause, D. W." box="[252,478,1840,1860]" journalOrPublisher="Nature" pageId="7" pageNumber="8" pagination="272 - 276" part="588" refId="ref7690" refString="O'Connor, P. M., Turner, A. H., Groenke, J. R., Felice, R. N., Rogers, R. R., Krause, D. W. et al. (2020) Late Cretaceous bird from Madagascar reveals unique development of beaks. Nature, 588, 272 - 276." title="Late Cretaceous bird from Madagascar reveals unique development of beaks" type="journal article" year="2020">OConnor et al., 2020</bibRefCitation>
) indicative of a high degree of ecological diversification among enantiornithines within the Cretaceous forested ecosystem (
<bibRefCitation author="Zhou, Z. &amp; Barrett, P. M. &amp; Hilton, J." box="[447,622,1910,1930]" journalOrPublisher="Nature" pageId="7" pageNumber="8" pagination="807" part="421" refId="ref8803" refString="Zhou, Z., Barrett, P. M. &amp; Hilton, J. (2003) An exceptionally preserved Lower Cretaceous ecosystem. Nature, 421, 807." title="An exceptionally preserved Lower Cretaceous ecosystem" type="journal article" year="2003">Zhou et al., 2003</bibRefCitation>
). For taxa with a similarly short rostrum, the
<taxonomicName box="[414,570,1944,1964]" class="Aves" family="Bohaiornithidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="8" phylum="Chordata" rank="family">Bohaiornithidae</taxonomicName>
and
<taxonomicName box="[620,769,1944,1964]" class="Aves" family="Pengornithidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="8" phylum="Chordata" rank="family">Pengornithidae</taxonomicName>
are differentiated further by remarkable tooth morphologies.
<taxonomicName box="[818,917,1147,1167]" class="Aves" family="Bohaiornithidae" genus="Bohaiornis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
<emphasis box="[818,917,1147,1167]" italics="true" pageId="7" pageNumber="8">Bohaiornis</emphasis>
</taxonomicName>
has been hypothesized to use their robust teeth for crushing hard materials, while
<taxonomicName box="[1119,1210,1182,1202]" class="Aves" family="Pengornithidae" genus="Pengornis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
<emphasis box="[1119,1210,1182,1202]" italics="true" pageId="7" pageNumber="8">Pengornis</emphasis>
</taxonomicName>
might have used its blunt teeth for grinding (
<bibRefCitation author="Zhou, Z. &amp; Clarke, J. &amp; Zhang, F." box="[1002,1173,1216,1236]" journalOrPublisher="Journal of Anatomy" pageId="7" pageNumber="8" pagination="565 - 577" part="212" refId="ref8835" refString="Zhou, Z., Clarke, J. &amp; Zhang, F. (2008) Insight into diversity, body size and morphological evolution from the largest Early Cretaceous enantiornithine bird. Journal of Anatomy, 212, 565 - 577." title="Insight into diversity, body size and morphological evolution from the largest Early Cretaceous enantiornithine bird" type="journal article" year="2008">Zhou et al., 2008</bibRefCitation>
).
</paragraph>
<paragraph blockId="7.[818,1467,1112,1964]" lastBlockId="8.[122,770,1494,1964]" lastPageId="8" lastPageNumber="9" pageId="7" pageNumber="8">
In addition to the rostrum and dental variation,
<taxonomicName box="[1327,1466,1251,1271]" genus="Brevirostruavis" pageId="7" pageNumber="8" rank="genus">
<emphasis box="[1327,1466,1251,1271]" italics="true" pageId="7" pageNumber="8">Brevirostruavis</emphasis>
</taxonomicName>
adds significant new data related to the known diversity in feeding specializations present in enantiornithines and early diverging stem birds as a whole. The combination of a short rostrum paired with a hyper-elongated hyoid apparatus (i.e., ceratobranchial) is not known among early avialans. The significant elongation and distinct dorsal curvature of the ceratobranchials might have functioned similarly to the elongate epibranchials present in living birds, such as hummingbirds, woodpeckers, and honeyeaters (
<bibRefCitation author="Paton, D. C. &amp; Collins, B. G." journalOrPublisher="Australian Journal of Ecology" pageId="7" pageNumber="8" pagination="473 - 506" part="14" refId="ref7885" refString="Paton, D. C. &amp; Collins, B. G. (1989) Bills and tongues of nectar-feeding birds: a review of morphology, function and performance, with intercontinental comparisons. Australian Journal of Ecology, 14, 473 - 506." title="Bills and tongues of nectar-feeding birds: a review of morphology, function and performance, with intercontinental comparisons" type="journal article" year="1989">Paton &amp; Collins, 1989</bibRefCitation>
). The elongate ceratobranchials in
<taxonomicName box="[1212,1351,1563,1583]" genus="Brevirostruavis" pageId="7" pageNumber="8" rank="genus">
<emphasis box="[1212,1351,1563,1583]" italics="true" pageId="7" pageNumber="8">Brevirostruavis</emphasis>
</taxonomicName>
might have compensated for the absence of ossified epibranchials in early birds who required a long hyobranchial for food manipulation. Similar elongation of branchial elements appears to have evolved independently in
<taxonomicName authorityName="Zhou &amp; Zhang" authorityYear="2002" box="[996,1089,1702,1722]" class="Aves" family="Jeholornithidae" genus="Jeholornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Jeholornithiformes" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
<emphasis box="[996,1089,1702,1722]" italics="true" pageId="7" pageNumber="8">Jeholornis</emphasis>
</taxonomicName>
and
<taxonomicName box="[1144,1218,1702,1722]" class="Aves" family="Bohaiornithidae" genus="Sulcavis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
<emphasis box="[1144,1218,1702,1722]" italics="true" pageId="7" pageNumber="8">Sulcavis</emphasis>
</taxonomicName>
with their longer rostra (
<figureCitation box="[823,904,1736,1756]" captionStart="FIGURE 6" captionStartId="9.[122,214,1478,1499]" captionTargetBox="[122,1465,57,1436]" captionTargetId="figure-275@9.[132,1455,123,1438]" captionText="FIGURE 6 Proposed hyoid apparatus character evolution in birds including ceratobranchial elongation, mineralization of basihyale, and epibranchial evolution (modified from Li et al., 2018). The fleshy part of tongue is elaborated in extant taxa" figureDoi="http://doi.org/10.5281/zenodo.5808234" httpUri="https://zenodo.org/record/5808234/files/figure.png" pageId="7" pageNumber="8">Figure 6</figureCitation>
). The long hyoid apparatus and associated tongue appendage (i.e. fleshy tongue and cartilaginous paraglossum) are known to be good indicators of dietary adaptations in crown birds (
<bibRefCitation author="Erdogan, S. &amp; Iwasaki, S. - I." journalOrPublisher="Annals of Anatomy-Anatomischer Anzeiger" pageId="7" pageNumber="8" pagination="75 - 87" part="196" refId="ref7102" refString="Erdogan, S. &amp; Iwasaki, S. - I. (2014) Function-related morphological char- acteristics and specialized structures of the avian tongue. Annals of Anatomy-Anatomischer Anzeiger, 196, 75 - 87." title="Function-related morphological char- acteristics and specialized structures of the avian tongue" type="journal article" year="2014">Erdoğan &amp; Iwasaki, 2014</bibRefCitation>
). For instance, grazing-, filtering-feeding, and piscivorous anatids maintain different shapes of their fleshy and bony tongues, particularly related to their different feeding adaptation. The simplified and rudimentary shaped muscular tongue is present in
<taxonomicName authorityName="Brisson" authorityYear="1760" box="[146,244,1494,1514]" class="Aves" family="Anatidae" genus="Merganser" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anseriformes" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
<emphasis box="[146,244,1494,1514]" italics="true" pageId="8" pageNumber="9">Merganser</emphasis>
</taxonomicName>
and other piscivorous ducks. By contrast, the muscular shape is strikingly different, being much wider and flattened, in grazing and filtering-feeding anatids (
<bibRefCitation author="Erdogan, S. &amp; Iwasaki, S. - I." box="[480,723,1563,1583]" journalOrPublisher="Annals of Anatomy-Anatomischer Anzeiger" pageId="8" pageNumber="9" pagination="75 - 87" part="196" refId="ref7102" refString="Erdogan, S. &amp; Iwasaki, S. - I. (2014) Function-related morphological char- acteristics and specialized structures of the avian tongue. Annals of Anatomy-Anatomischer Anzeiger, 196, 75 - 87." title="Function-related morphological char- acteristics and specialized structures of the avian tongue" type="journal article" year="2014">Erdoğan &amp; Iwasaki, 2014</bibRefCitation>
;
<bibRefCitation author="Li, Z. &amp; Clarke, J. A." journalOrPublisher="Evolutionary Biology" pageId="8" pageNumber="9" pagination="12 - 25" part="43" refId="ref7412" refString="Li, Z. &amp; Clarke, J. A. (2016) The craniolingual morphology of waterfowl (Aves, Anseriformes) and its relationship with feeding mode revealed through contrast-enhanced X-ray computed tomography and 2 D morphometrics. Evolutionary Biology, 43, 12 - 25." title="The craniolingual morphology of waterfowl (Aves, Anseriformes) and its relationship with feeding mode revealed through contrast-enhanced X-ray computed tomography and 2 D morphometrics" type="journal article" year="2016">Li &amp; Clarke, 2016</bibRefCitation>
).
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.5808232" ID-Zenodo-Dep="5808232" httpUri="https://zenodo.org/record/5808232/files/figure.png" pageId="8" pageNumber="9" startId="8.[123,215,1343,1364]" targetBox="[122,1466,57,1301]" targetPageId="8">
<paragraph blockId="8.[122,1452,1343,1422]" pageId="8" pageNumber="9">
<emphasis bold="true" box="[123,239,1343,1364]" pageId="8" pageNumber="9">FIGURE 5</emphasis>
Phylogenetic position of
<taxonomicName box="[495,774,1344,1364]" genus="Brevirostruavis" pageId="8" pageNumber="9" rank="species" species="macrohyoideus">
<emphasis box="[495,774,1344,1364]" italics="true" pageId="8" pageNumber="9">Brevirostruavis macrohyoideus</emphasis>
</taxonomicName>
(IVPP V 13266) in Avialae. (a) Strict consensus tree from current phylogenetic analysis; (b) reduced consensus tree with ten most unstable taxa removed. The bootstrap and absolute Bremer support values are denoted in normal and bold Italic fonts
</paragraph>
</caption>
<paragraph blockId="8.[122,770,1494,1964]" lastBlockId="9.[122,770,1632,1964]" lastPageId="9" lastPageNumber="10" pageId="8" pageNumber="9">
The elaboration and ossification of different hyoid bony elements appeared early in avialan evolution as demonstrated by this Early Cretaceous indication of hyolinugal diversity. The mineralization (or ossification) of the basihyal is known in the pygostylian
<taxonomicName box="[200,338,1771,1791]" class="Aves" family="Confuciusornithidae" genus="Confuciusornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Confuciusornithiformes" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
<emphasis box="[200,338,1771,1791]" italics="true" pageId="8" pageNumber="9">Confuciusornis</emphasis>
</taxonomicName>
, the ornithuromorph
<taxonomicName authorityName="Zhou &amp; Zhang" authorityYear="2005" box="[575,714,1771,1791]" class="Aves" family="Hongshanornithidae" genus="Hongshanornis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
<emphasis box="[575,714,1771,1791]" italics="true" pageId="8" pageNumber="9">Hongshanornis</emphasis>
</taxonomicName>
, and even in the non-avialan
<taxonomicName authority="(Li et al., 2018)" baseAuthorityName="Li" baseAuthorityYear="2018" box="[365,637,1806,1826]" class="Reptilia" family="Dromaeosauridae" genus="Microraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
<emphasis box="[365,476,1806,1826]" italics="true" pageId="8" pageNumber="9">Microraptor</emphasis>
(
<bibRefCitation author="Li, Z. &amp; Zhou, Z. &amp; Clarke, J. A." box="[490,632,1806,1826]" journalOrPublisher="PLoS One" pageId="8" pageNumber="9" pagination="0198078" part="13" refId="ref7528" refString="Li, Z., Zhou, Z. &amp; Clarke, J. A. (2018) Convergent evolution of a mobile bony tongue in flighted dinosaurs and pterosaurs. PLoS One, 13, e 0198078." title="Convergent evolution of a mobile bony tongue in flighted dinosaurs and pterosaurs" type="journal article" year="2018">Li et al., 2018</bibRefCitation>
)
</taxonomicName>
. The ossified epibranchial and proposed mobile or protractible tongue evolved first in
<taxonomicName authorityName="Zhou &amp; Zhang" authorityYear="2005" box="[196,335,1875,1895]" class="Aves" family="Hongshanornithidae" genus="Hongshanornis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
<emphasis box="[196,335,1875,1895]" italics="true" pageId="8" pageNumber="9">Hongshanornis</emphasis>
</taxonomicName>
(
<bibRefCitation author="Li, Z. &amp; Zhou, Z. &amp; Clarke, J. A." box="[350,492,1875,1895]" journalOrPublisher="PLoS One" pageId="8" pageNumber="9" pagination="0198078" part="13" refId="ref7528" refString="Li, Z., Zhou, Z. &amp; Clarke, J. A. (2018) Convergent evolution of a mobile bony tongue in flighted dinosaurs and pterosaurs. PLoS One, 13, e 0198078." title="Convergent evolution of a mobile bony tongue in flighted dinosaurs and pterosaurs" type="journal article" year="2018">Li et al., 2018</bibRefCitation>
;
<bibRefCitation author="Zhou, Z. &amp; Zhang, F." box="[503,715,1875,1895]" pageId="8" pageNumber="9" pagination="18998 - 19002" refId="ref8919" refString="Zhou, Z. &amp; Zhang, F. (2005) Discovery of an ornithurine bird and its implication for Early Cretaceous avian radiation. Proceedings of the National Academy of Sciences of the United States of America, 102, 18998 - 19002." type="journal article" year="2005">Zhou &amp; Zhang, 2005</bibRefCitation>
). The separate ossification center of the epibranchial represents a novel feature present in all living birds (
<bibRefCitation author="Li, Z. &amp; Zhou, Z. &amp; Clarke, J. A." box="[452,589,1944,1964]" journalOrPublisher="PLoS One" pageId="8" pageNumber="9" pagination="0198078" part="13" refId="ref7528" refString="Li, Z., Zhou, Z. &amp; Clarke, J. A. (2018) Convergent evolution of a mobile bony tongue in flighted dinosaurs and pterosaurs. PLoS One, 13, e 0198078." title="Convergent evolution of a mobile bony tongue in flighted dinosaurs and pterosaurs" type="journal article" year="2018">Li et al., 2018</bibRefCitation>
). We propose that the elongation of the hyoid apparatus evolved independently via different patterns in bird evolution (
<figureCitation box="[1196,1284,1528,1548]" captionStart="FIGURE 6" captionStartId="9.[122,214,1478,1499]" captionTargetBox="[122,1465,57,1436]" captionTargetId="figure-275@9.[132,1455,123,1438]" captionText="FIGURE 6 Proposed hyoid apparatus character evolution in birds including ceratobranchial elongation, mineralization of basihyale, and epibranchial evolution (modified from Li et al., 2018). The fleshy part of tongue is elaborated in extant taxa" figureDoi="http://doi.org/10.5281/zenodo.5808234" httpUri="https://zenodo.org/record/5808234/files/figure.png" pageId="8" pageNumber="9">Figure 6</figureCitation>
). Within avialans, slightly longer ceratobranchials are present in
<taxonomicName authorityName="Zhou &amp; Zhang" authorityYear="2002" box="[1285,1379,1563,1583]" class="Aves" family="Jeholornithidae" genus="Jeholornis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Jeholornithiformes" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
<emphasis box="[1285,1379,1563,1583]" italics="true" pageId="8" pageNumber="9">Jeholornis</emphasis>
</taxonomicName>
, a seedeating basal bird, and another enantiornithine (
<emphasis box="[1301,1454,1598,1618]" italics="true" pageId="8" pageNumber="9">Sulcvis geeorum</emphasis>
). These taxa share relatively longer ceratobranchial elements with the new bird described here, and contrast with the states present in other more derived ornithuromorphan birds. It is reasonable to hypothesize that the evolutionary pathway of hyolingual feeding in Mesozoic stem birds was achieved through the (initial) lengthening of the ceratobranchials in enantiornithines, and via the ossification of epibranchials in ornithuromorphs (including crown birds). The absence of cartilaginous epibranchials in the hyolingual apparatus in this enantiornithine is supported by the morphology of the caudal end of the ceratobranchials with a flattened tapered end that differs significantly from the bulbous and rounded knob-like end which articulates with the epibranchial in living birds (
<figureCitation box="[642,722,1667,1687]" captionStart="FIGURE 6" captionStartId="9.[122,214,1478,1499]" captionTargetBox="[122,1465,57,1436]" captionTargetId="figure-275@9.[132,1455,123,1438]" captionText="FIGURE 6 Proposed hyoid apparatus character evolution in birds including ceratobranchial elongation, mineralization of basihyale, and epibranchial evolution (modified from Li et al., 2018). The fleshy part of tongue is elaborated in extant taxa" figureDoi="http://doi.org/10.5281/zenodo.5808234" httpUri="https://zenodo.org/record/5808234/files/figure.png" pageId="9" pageNumber="10">Figure 6</figureCitation>
) and
<taxonomicName authorityName="Zhou &amp; Zhang" authorityYear="2005" box="[122,261,1702,1722]" class="Aves" family="Hongshanornithidae" genus="Hongshanornis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="9" pageNumber="10" phylum="Chordata" rank="genus">
<emphasis box="[122,261,1702,1722]" italics="true" pageId="9" pageNumber="10">Hongshanornis</emphasis>
</taxonomicName>
.
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.5808234" ID-Zenodo-Dep="5808234" httpUri="https://zenodo.org/record/5808234/files/figure.png" pageId="9" pageNumber="10" startId="9.[122,214,1478,1499]" targetBox="[122,1465,57,1436]" targetPageId="9">
<paragraph blockId="9.[122,1437,1478,1528]" pageId="9" pageNumber="10">
<emphasis bold="true" box="[122,237,1478,1499]" pageId="9" pageNumber="10">FIGURE 6</emphasis>
Proposed hyoid apparatus character evolution in birds including ceratobranchial elongation, mineralization of basihyale, and epibranchial evolution (modified from
<bibRefCitation author="Li, Z. &amp; Zhou, Z. &amp; Clarke, J. A." box="[487,618,1508,1528]" journalOrPublisher="PLoS One" pageId="9" pageNumber="10" pagination="0198078" part="13" refId="ref7528" refString="Li, Z., Zhou, Z. &amp; Clarke, J. A. (2018) Convergent evolution of a mobile bony tongue in flighted dinosaurs and pterosaurs. PLoS One, 13, e 0198078." title="Convergent evolution of a mobile bony tongue in flighted dinosaurs and pterosaurs" type="journal article" year="2018">Li et al., 2018</bibRefCitation>
). The fleshy part of tongue is elaborated in extant taxa
</paragraph>
</caption>
<paragraph blockId="9.[122,770,1632,1964]" lastBlockId="10.[122,770,142,1375]" lastPageId="10" lastPageNumber="11" pageId="9" pageNumber="10">
Being absent in enantiornithines or more stem-ward avialans, the presence of a ossified epibranchial is optimized to have evolved among early diverging ornithuromorphs (
<bibRefCitation author="Li, Z. &amp; Zhou, Z. &amp; Clarke, J. A." box="[534,679,1806,1826]" journalOrPublisher="PLoS One" pageId="9" pageNumber="10" pagination="0198078" part="13" refId="ref7528" refString="Li, Z., Zhou, Z. &amp; Clarke, J. A. (2018) Convergent evolution of a mobile bony tongue in flighted dinosaurs and pterosaurs. PLoS One, 13, e 0198078." title="Convergent evolution of a mobile bony tongue in flighted dinosaurs and pterosaurs" type="journal article" year="2018">Li et al., 2018</bibRefCitation>
). Dietary associated traits in enantiornithines and other Early Cretaceous stem birds (like the presence of a crop and gastroliths) are strongly indicative of a regime where herbivory and seed-eating played a key role in driving the early radiation of several lineages in Avialae (
<bibRefCitation author="Zanno, L. E. &amp; Makovicky, P. J." box="[823,1088,1632,1652]" journalOrPublisher="Proceedings of the National Academy of Sciences" pageId="9" pageNumber="10" pagination="232 - 237" part="108" refId="ref8505" refString="Zanno, L. E. &amp; Makovicky, P. J. (2011) Herbivorous ecomorphology and specialization patterns in theropod dinosaur evolution. Proceedings of the National Academy of Sciences, 108, 232 - 237." title="Herbivorous ecomorphology and specialization patterns in theropod dinosaur evolution" type="journal article" year="2011">Zanno &amp; Makovicky, 2011</bibRefCitation>
). A major shift from carnivory in nonavialan theropods to the herbivorous diet present among basal birds seems to drive cranial evolution leading to the origin of ornithurine birds, including all extant birds. The effective use of available food resources and the reduction in competition with other contemporary non-avialan theropods and pterosaurs might have contributed to their major radiation in the Mesozoic. The elongation of the ceratobranchial bone in
<taxonomicName box="[1189,1329,1875,1895]" genus="Brevirostruavis" pageId="9" pageNumber="10" rank="genus">
<emphasis box="[1189,1329,1875,1895]" italics="true" pageId="9" pageNumber="10">Brevirostruavis</emphasis>
</taxonomicName>
represents a novel evolutionary path that is unknown among extant birds and differs from their Mesozoic relatives. That elongation represents a change in function, an increase in feeding efficiency, or an unknown specialization necessary to exploit certain food resources. In particular, this hyoid apparatus likely helped the bird to achieve protrusion of the tongue beyond the rostrum as a component of foraging and feeding.
</paragraph>
<paragraph blockId="10.[122,770,142,1375]" pageId="10" pageNumber="11">
The linkage between hyolingual feeding with the morphology of the rostrum and beak seems to be a synapomorphy of crown birds (
<bibRefCitation author="Bongo-Tomlinson, C. &amp; Schwenk, K." box="[181,537,384,404]" editor="Schwenk, K." journalOrPublisher="San Diego: Academic Press" pageId="10" pageNumber="11" pagination="359 - 394" refId="ref6836" refString="Bongo-Tomlinson, C. &amp; Schwenk, K. (2000) Feeding in paleognathous birds. In: Schwenk, K. (Ed.) Feeding: form, function, and evolution in tetrapod vertebrates. San Diego: Academic Press, pp. 359 - 394." title="Feeding in paleognathous birds" type="book chapter" volumeTitle="Feeding: form, function, and evolution in tetrapod vertebrates" year="2000">Bongo-Tomlinson &amp; Schwenk, 2000</bibRefCitation>
). However, such a close coupling or correlation between the hyoid and rostrum morphology might be derived from their functional coordination of the two components that is closely tied to feeding specialization, and thus, not linked to a particular clade. This novel linkage between the rostrum and hyolingual apparatus might be associated with the origin of avian cranial kinesis in ornithuromorph birds as well, and requires further testing.
</paragraph>
<paragraph blockId="10.[122,770,142,1375]" pageId="10" pageNumber="11">
It is interesting to note that enantiornithine birds likely lacked a kinetic skull as in crown birds, and even retained static dinosaurian aspects to their skull and palate (
<bibRefCitation author="Wang, M. &amp; Stidham, T. A. &amp; Li, Z. &amp; Xu, X. &amp; Zhou, Z." box="[480,653,731,751]" journalOrPublisher="Nature Communications" pageId="10" pageNumber="11" pagination="1 - 9" part="12" refId="ref8243" refString="Wang, M., Stidham, T. A., Li, Z., Xu, X. &amp; Zhou, Z. (2021) Cretaceous bird with dinosaur skull sheds light on avian cranial evolution. Nature Communications, 12, 1 - 9." title="Cretaceous bird with dinosaur skull sheds light on avian cranial evolution" type="journal article" year="2021">Wang et al., 2021</bibRefCitation>
). Given that framework in enantiornithines along with the absence of epibranchials, the only evolutionary pathway open to them may have been to elongate the ceratobranchials as a functional mechanism to utilize or exploit available dietary resources. That difference also is indicated by the long ceratobranchials of
<taxonomicName box="[427,562,904,924]" genus="Brevirostruavis" pageId="10" pageNumber="11" rank="genus">
<emphasis box="[427,562,904,924]" italics="true" pageId="10" pageNumber="11">Brevirostruavis</emphasis>
</taxonomicName>
contrasting strikingly with its abbreviated rostrum, suggesting an absence of the functional coordination between the rostrum and hyoid in early diverging avialans. Clearly, the combination of a short rostrum and a long tongue evolved in a relationship to obtain some dietary component, and that requirement may have necessitated extending the tongue beyond the rostrum as in some extant birds like woodpeckers. The ancient forests of northeastern China had abundant and various food resources, including diverse insects and even nutritious plant resources such as nectar and related reproductive structures (
<bibRefCitation author="Ren, D. &amp; Shih, C. &amp; Labandeira, C. C." box="[614,764,1216,1236]" journalOrPublisher="ZooKeys" pageId="10" pageNumber="11" pagination="17 - 28" part="129" refId="ref7968" refString="Ren, D., Shih, C. &amp; Labandeira, C. C. (2011) A well-preserved aneuretopsychid from the Jehol Biota of China (Insecta, Mecoptera, Aneuretopsychidae). ZooKeys, 129, 17 - 28." title="A well-preserved aneuretopsychid from the Jehol Biota of China (Insecta, Mecoptera, Aneuretopsychidae)" type="journal article" year="2011">Ren et al., 2011</bibRefCitation>
) that could have been consumed by birds. The combination of a short rostrum and elongated tongue clearly increased the fitness of this enantiornithine bird, and allowed it to utilize a resource that other known birds did not.
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</subSubSection>
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