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<mods:title id="259A0CA21765C8C1A97729ADE1334172">Reassessment of the enigmatic Late Cretaceous theropod dinosaur, Bagaraatan ostromi</mods:title>
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<mods:namePart id="372731EFCC1E8B41EAA44C5867BB53D3">Słowiak, Justyna</mods:namePart>
<mods:affiliation id="7579D2B425769A1307A6DDC41F00E411">Institute of Paleobiology, Polish Academy of Sciences, Warsaw, Poland</mods:affiliation>
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<mods:namePart id="4886FFAF9FF248C9799CF785921B1C85">Brusatte, Stephen L.</mods:namePart>
<mods:affiliation id="CB8864AD6FFF3667EB54CCEBDC3EE4B8">School of GeoSciences, University of Edinburgh, Edinburgh, UK</mods:affiliation>
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<mods:namePart id="6887DACA1660F33C402D7D980604E4F6">Szczygielski, Tomasz</mods:namePart>
<mods:affiliation id="C32395F873CFA599DE9877AC33E1EAFA">Institute of Paleobiology, Polish Academy of Sciences, Warsaw, Poland</mods:affiliation>
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<mods:date id="964FDFB66C3AE40FF08B425EFA61308C">2024</mods:date>
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<subSubSection id="C33082DA994DFFDFB90003AAFDADFF91" box="[318,588,144,170]" pageId="32" pageNumber="33" type="nomenclature">
<paragraph id="8B95D151994DFFDFB90003AAFDADFF91" blockId="32.[318,588,144,170]" box="[318,588,144,170]" pageId="32" pageNumber="33">
<heading id="D0DD663D994DFFDFB90003AAFDADFF91" box="[318,588,144,170]" centered="true" fontSize="9" level="2" pageId="32" pageNumber="33" reason="2">
<emphasis id="B95E0D43994DFFDFB90003AAFDADFF91" bold="true" box="[318,588,144,170]" pageId="32" pageNumber="33">
Is
<taxonomicName id="4C2AAAD2994DFFDFB96B03AAFE5BFF91" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[341,442,144,170]" family="Tyrannosauridae" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994DFFDFB96B03AAFE5BFF91" bold="true" box="[341,442,144,170]" italics="true" pageId="32" pageNumber="33">B. ostromi</emphasis>
</taxonomicName>
a valid taxon?
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C33082DA994DFFDDB8BF038CFED1FFFC" lastPageId="34" lastPageNumber="35" pageId="32" pageNumber="33" type="discussion">
<paragraph id="8B95D151994DFFDFB8BF038CFD34FDBE" blockId="32.[129,778,182,833]" pageId="32" pageNumber="33">
Osmólska (1996) listed eight diagnostic features for
<taxonomicName id="4C2AAAD2994DFFDFBAA1038DFCE3FFF5" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[671,770,183,206]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994DFFDFBAA1038DFCE3FFF5" box="[671,770,183,206]" italics="true" pageId="32" pageNumber="33">B. ostromi</emphasis>
</taxonomicName>
: (i) two surangular foramina [also considered by Holtz (2004) as an autapomorphy of
<taxonomicName id="4C2AAAD2994DFFDFB9BE03CCFE07FE36" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[384,486,246,269]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994DFFDFB9BE03CCFE07FE36" box="[384,486,246,269]" italics="true" pageId="32" pageNumber="33">B. ostromi</emphasis>
</taxonomicName>
]; (ii) articular with an oblique posterior surface and a short retroarticular surface; (iii) caudal vertebrae with thin-walled centra; (iv) hyposphenehypantrum articulations in at least the first 16 caudal vertebrae [also listed by Holtz (2004) as an autapomorphy of
<taxonomicName id="4C2AAAD2994DFFDFBAAB0249FD16FEB1" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[661,759,371,394]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994DFFDFBAAB0249FD16FEB1" box="[661,759,371,394]" italics="true" pageId="32" pageNumber="33">B. ostromi</emphasis>
</taxonomicName>
]; (v) prezygapophyses in proximal caudal vertebrae with ridges on the lateral surfaces; (vi) ilium with two deep depressions; (vii) femur with the anterior trochanter (anterior crest
<emphasis id="B95E0D43994DFFDFBAE802EBFCE8FED3" box="[726,777,465,488]" italics="true" pageId="32" pageNumber="33">sensu</emphasis>
Osmólska); and (viii) tibia and fibula fused distally. The status of those features is briefly discussed below. Given that we have now re-identified the hindlimb bones as belonging to other non-tyrannosaurid taxa, those features regarding the hindlimb were discussed above, hence they will be omitted in this section.
</paragraph>
<paragraph id="8B95D151994DFFDFB8A201B6FBABFFDD" blockId="32.[129,778,182,833]" lastBlockId="32.[825,1475,144,826]" pageId="32" pageNumber="33">
The ilium with a distinct ridge on the lateral surface of the postacetabular process, demarcated anteriorly, medially, and posteriorly by depressions, is striking (
<figureCitation id="1311CDD4994DFFDFBA2B01F1FDBDFDD8" box="[533,604,715,739]" captionStart="Figure 16" captionStartId="20.[129,194,1516,1540]" captionTargetBox="[131,1471,146,1486]" captionTargetId="figure-232@20.[129,1473,144,1488]" captionTargetPageId="20" captionText="Figure 16. Ilium of Bagaraatan ostromi ZPAL MgD-I/108. A, B, fragment of left preacetabular process in lateral (A) and medial (B) view. CF, left postacetabular process in lateral (C), medial (D), dorsal (E), and ventral (F) view.GJ, right postacetabular process in dorsal (G), ventral (H), lateral (I), and medial (J) view.KN, two fragments of ilium blade in lateral (K, M) and medial (L, N) view." figureDoi="http://doi.org/10.5281/zenodo.14284263" httpUri="https://zenodo.org/record/14284263/files/figure.png" pageId="32" pageNumber="33">Fig. 16</figureCitation>
). It occurs symmetrically on both ilia, and better preserved on the left, which is less compressed. Such ridges are not found in other theropods, to our knowledge, and are not present in juvenile
<emphasis id="B95E0D43994DFFDFBAD40010FCEBFC7A" box="[746,778,810,833]" italics="true" pageId="32" pageNumber="33">Ta.</emphasis>
<taxonomicName id="4C2AAAD2994DFFDFBB0703AAFC9BFF93" box="[825,890,144,168]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994DFFDFBB0703AAFC9BFF93" box="[825,890,144,168]" italics="true" pageId="32" pageNumber="33">bataar</emphasis>
</taxonomicName>
(MPC-D 107/7) nor the other tyrannosaurid juvenile,
<emphasis id="B95E0D43994DFFDFBD9203ABFA22FF93" box="[1452,1475,145,168]" italics="true" pageId="32" pageNumber="33">R.</emphasis>
<taxonomicName id="4C2AAAD2994DFFDFBB070395FB8EFFFC" authority="(Sereno et al. 2009)" baseAuthorityName="Sereno" baseAuthorityYear="2009" box="[825,1135,175,199]" class="Reptilia" family="Tyrannosauridae" genus="Raptorex" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="kriegsteini">
<emphasis id="B95E0D43994DFFDFBB070395FC7AFFFC" box="[825,923,175,199]" italics="true" pageId="32" pageNumber="33">kriegsteini</emphasis>
(Sereno
<emphasis id="B95E0D43994DFFDFBBC4038AFBC8FFFC" box="[1018,1065,175,199]" italics="true" pageId="32" pageNumber="33">et al.</emphasis>
2009)
</taxonomicName>
. Therefore, they might be a diagnostic feature of
<taxonomicName id="4C2AAAD2994DFFDFBBDC03F5FBA2FFDD" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[994,1091,207,230]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994DFFDFBBDC03F5FBA2FFDD" box="[994,1091,207,230]" italics="true" pageId="32" pageNumber="33">B. ostromi</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8B95D151994DFFDEBB6B03D4FEACFE3D" blockId="32.[825,1475,144,826]" lastBlockId="33.[113,766,144,1985]" lastPageId="33" lastPageNumber="34" pageId="32" pageNumber="33">
The ridges on the lateral surfaces of the prezygapophyses are found also in the proximal caudal vertebrae of
<emphasis id="B95E0D43994DFFDFBD780234FA6EFE1E" box="[1350,1423,270,293]" italics="true" pageId="32" pageNumber="33">
<taxonomicName id="4C2AAAD2994DFFDFBD780234FA63FE1E" baseAuthorityName="Carr" baseAuthorityYear="2020" box="[1350,1410,270,293]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="rex">Ty. rex</taxonomicName>
&amp;
</emphasis>
running from the anterior margin of the transverse process to the prezygapopysis (
<bibRefCitation id="EFBBACA0994DFFDFBC1F0276FB51FE5F" author="Brochu CA" box="[1057,1200,332,356]" pageId="32" pageNumber="33" pagination="1 - 138" refId="ref25263" refString="Brochu CA. Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull. Journal of Vertebrate Paleonotology 2003; 7: 1 - 138. https: // doi. org / 10.2307 / 3889334" year="2003">Brochu 2003</bibRefCitation>
). Similar ridges on the prezygapophyses of anterior caudal vertebrae are also present in
<taxonomicName id="4C2AAAD2994DFFDFBBB702B1FC10FE99" box="[905,1009,394,418]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994DFFDFBBB702B1FC10FE99" box="[905,1009,394,418]" italics="true" pageId="32" pageNumber="33">Ta. bataar</emphasis>
</taxonomicName>
(ZPAL MgD-I/176). Osmólska (1996) did not quantify how thin-walled the caudal vertebrae centra of
<taxonomicName id="4C2AAAD2994DFFDFBD900290FC60FEDA" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994DFFDFBD900290FC60FEDA" italics="true" pageId="32" pageNumber="33">B. ostromi</emphasis>
</taxonomicName>
are in comparison to other theropods. We do not recognize any clear difference between the centrum thickness of
<taxonomicName id="4C2AAAD2994DFFDFBD9002D3FC60FD24" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994DFFDFBD9002D3FC60FD24" italics="true" pageId="32" pageNumber="33">B. ostromi</emphasis>
</taxonomicName>
and other theropods. The stout hyposphenehypantrum articulations in at least the first 16 caudal vertebrae were considered an autapomorphy of
<taxonomicName id="4C2AAAD2994DFFDFBC62017DFB5CFD65" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[1116,1213,583,606]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994DFFDFBC62017DFB5CFD65" box="[1116,1213,583,606]" italics="true" pageId="32" pageNumber="33">B. ostromi</emphasis>
</taxonomicName>
by Osmólska (1996) and Holtz (2004). The presence of the hyposphenehypantrum articulation is seen in many archosaurs, is strongly correlated with body size, and is often already present at a young age, before the articulation is necessary to support the large mass of the fully grown animal (Stefanic and
<bibRefCitation id="EFBBACA0994DFFDFBC5B01D9FB0CFDC0" author="Nesbitt S &amp; Loewen M" box="[1125,1261,739,763]" pageId="32" pageNumber="33" pagination="892 - 9" refId="ref28234" refString="Nesbitt S, Denton RK, Loewen M et al. A mid-Cretaceous tyrannosauroid and the origin of North American end-Cretaceous dinosaur assemblages. Nature Ecology &amp; Evolution 2019; 3: 892 - 9." year="2019">Nesbitt 2019</bibRefCitation>
). The hyposphenehypantrum articulations are common in theropods, and for instance, are present in the caudal vertebrae of medium-sized
<emphasis id="B95E0D43994CFFDEB84F03ABFF3EFF93" box="[113,223,144,168]" italics="true" pageId="33" pageNumber="34">Ta. bataar</emphasis>
(ZPAL MgD-I/176). The oblique posterior surface and short retroarticular surface of the articular, also listed by Osmólska (1996), are tyrannosauroid synapomorphies (
<bibRefCitation id="EFBBACA0994CFFDEB84203D4FEDCFE3D" author="Brusatte SL &amp; Lloyd GT &amp; Wang SC" box="[124,317,238,262]" pageId="33" pageNumber="34" pagination="2386 - 92" refId="ref25593" refString="Brusatte SL, Lloyd GT, Wang SC et al. Gradual assembly of avian body plan culminated in rapid rates of evolution across the dinosaur-bird transition. Current Biology: CB 2014; 24: 2386 - 92. https: // doi. org / 10.1016 / j. cub. 2014.08.034" year="2014">
Brusatte
<emphasis id="B95E0D43994CFFDEB8E803D5FEE4FE3D" box="[214,261,238,262]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
2014
</bibRefCitation>
).
</paragraph>
<caption id="DF5581D9994DFFDFB8BF04BEFDB4F883" ID-DOI="http://doi.org/10.5281/zenodo.14284283" ID-Zenodo-Dep="14284283" httpUri="https://zenodo.org/record/14284283/files/figure.png" pageId="32" pageNumber="33" startId="32.[129,194,1924,1948]" targetBox="[136,1471,887,1868]" targetPageId="32">
<paragraph id="8B95D151994DFFDFB8BF04BEFDB4F883" blockId="32.[129,1409,1924,1976]" pageId="32" pageNumber="33">
<emphasis id="B95E0D43994DFFDFB8BF04BEFF04F8A7" bold="true" box="[129,229,1924,1948]" pageId="32" pageNumber="33">Figure 25.</emphasis>
Reconstruction of
<emphasis id="B95E0D43994DFFDFB9A404BFFD1BF8A6" box="[410,762,1925,1949]" italics="true" pageId="32" pageNumber="33">Bagaraatan ostromi ZPAL MgD-I/108</emphasis>
including only tyrannosaurid bones found in the assemblage and life reconstruction of the dinosaur by Jakub Zalewski.
</paragraph>
</caption>
<paragraph id="8B95D151994CFFDEB8B30234FD49FC62" blockId="33.[113,766,144,1985]" pageId="33" pageNumber="34">
The presence of two surangular foramina and the ridge on the lateral surface of the postacetabular process of ilium seem to be the only two features listed by Osmólska (1996) that distinguish
<taxonomicName id="4C2AAAD2994CFFDEB8910256FEF3FEB8" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[175,274,364,387]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994CFFDEB8910256FEF3FEB8" box="[175,274,364,387]" italics="true" pageId="33" pageNumber="34">B. ostromi</emphasis>
</taxonomicName>
from other tyrannosaurids. The two surangular foramina were also later listed by Holtz (2004) as unique for
<taxonomicName id="4C2AAAD2994CFFDEBAD802B1FF58FEF9" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994CFFDEBAD802B1FF58FEF9" italics="true" pageId="33" pageNumber="34">B. ostromi</emphasis>
</taxonomicName>
in comparison to other theropods (
<bibRefCitation id="EFBBACA0994CFFDEBA0D0290FD08FEF9" author="Currie PJ &amp; Hurum JH &amp; Sabath K" box="[563,745,426,450]" pageId="33" pageNumber="34" pagination="227 - 34" refId="ref26465" refString="Currie PJ, Hurum JH, Sabath K. Skull structure and evolution in tyrannosaurid dinosaurs. Acta Palaeontologica Polonica 2003; 48: 227 - 34." year="2003">
Currie
<emphasis id="B95E0D43994CFFDEBA400291FD4EFEF9" box="[638,687,426,450]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
2003
</bibRefCitation>
). There is some confusion in the literature about the size of the surangular foramen in tyrannosauroids and its phylogenetic significance and ontogenetic and individual variation. In their phylogenetic dataset of tyrannosauroids,
<bibRefCitation id="EFBBACA0994CFFDEBA0B0112FF5DFD65" author="Brusatte SL &amp; Carr TD" pageId="33" pageNumber="34" refId="ref25346" refString="Brusatte SL, Carr TD. The phylogeny and evolutionary history of tyrannosauroid dinosaurs. Scientific Reports 2016; 6: 20252." year="2016">Brusatte and Carr (2016)</bibRefCitation>
used a character that simply divided the size of the foramen into two states: those with a dorsoventral depth &lt;30% or&gt; 30% of the depth of the posterior end of the surangular. This was based on earlier characters used by Sereno
<emphasis id="B95E0D43994CFFDEBA78019FFD95FD87" box="[582,628,676,700]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
(2009), Carr and Williamson (2010), and
<bibRefCitation id="EFBBACA0994CFFDEB9A901FEFD8CFDE7" author="Brusatte SL &amp; Norell MA &amp; Carr TD" box="[407,621,708,732]" pageId="33" pageNumber="34" pagination="1481 - 5" refId="ref25425" refString="Brusatte SL, Norell MA, Carr TD et al. Tyrannosaur paleobiology: new research on ancient exemplar organisms. Science 2010; 329: 1481 - 5. https: // doi. org / 10.1126 / science. 1193304" year="2010">
Brusatte
<emphasis id="B95E0D43994CFFDEB9CE01FFFDFFFDE7" box="[496,542,708,732]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
(2010)
</bibRefCitation>
. The enlarged condition was found to be synapomorphic of a clade consisting of
<taxonomicName id="4C2AAAD2994CFFDEB8A40039FEFEFC21" authorityName="Marsh" authorityYear="1877" box="[154,287,771,794]" class="Reptilia" family="Dryptosauridae" genus="Dryptosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
<emphasis id="B95E0D43994CFFDEB8A40039FEFEFC21" box="[154,287,771,794]" italics="true" pageId="33" pageNumber="34">Dryptosaurus</emphasis>
</taxonomicName>
+ Tyrannosauridae, whereas the primitive smaller foramen is seen in more basal tyrannosauroids, such as
<taxonomicName id="4C2AAAD2994CFFDEB84F007BFF1BFC62" box="[113,250,833,857]" genus="Suskityrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
<emphasis id="B95E0D43994CFFDEB84F007BFF1BFC62" box="[113,250,833,857]" italics="true" pageId="33" pageNumber="34">Suskityrannus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C2AAAD2994CFFDEB93B0078FE94FC62" authorityName="Hutt, Naish, Martill, Barker &amp; Newbery" authorityYear="2001" box="[261,373,834,857]" class="Reptilia" genus="Eotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
<emphasis id="B95E0D43994CFFDEB93B0078FE94FC62" box="[261,373,834,857]" italics="true" pageId="33" pageNumber="34">Eotyrannus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C2AAAD2994CFFDEB941007BFE20FC62" box="[383,449,833,857]" class="Reptilia" family="Tyrannosauridae" genus="Dilong" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
<emphasis id="B95E0D43994CFFDEB941007BFE20FC62" box="[383,449,833,857]" italics="true" pageId="33" pageNumber="34">Dilong</emphasis>
</taxonomicName>
, and proceratosaurids.
</paragraph>
<paragraph id="8B95D151994CFFDEB8B3005BFD86FA55" blockId="33.[113,766,144,1985]" pageId="33" pageNumber="34">
Other authors, however, have considered the foramen differently. The size of the surangular foramen in tyrannosaurids was divided into moderate (
<taxonomicName id="4C2AAAD2994CFFDEB98C009AFDCCFC8C" box="[434,557,927,951]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="libratus">
<emphasis id="B95E0D43994CFFDEB98C009AFDCCFC8C" box="[434,557,927,951]" italics="true" pageId="33" pageNumber="34">Go. libratus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C2AAAD2994CFFDEBA7E00A5FE88FCED" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="sarcophagus">
<emphasis id="B95E0D43994CFFDEBA7E00A5FF2BFCEC" italics="true" pageId="33" pageNumber="34">Albertosaurus sarcophagus</emphasis>
<bibRefCitation id="EFBBACA0994CFFDEB8E80085FE88FCED" author="Osborn HF" box="[214,361,958,983]" pageId="33" pageNumber="34" pagination="259 - 65" refId="ref28311" refString="Osborn HF. Tyrannosaurus and other Cretaceous carnivorous dinosaurs. Bulletin of the American Museum of Natural History 1905; 21: 259 - 65." year="1905">Osborn, 1905</bibRefCitation>
</taxonomicName>
,
<taxonomicName id="4C2AAAD2994CFFDEB9470085FE5EFCED" baseAuthorityName="Carr" baseAuthorityYear="2020" box="[377,447,959,982]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="rex">
<emphasis id="B95E0D43994CFFDEB9470085FE5EFCED" box="[377,447,959,982]" italics="true" pageId="33" pageNumber="34">Ty. rex</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C2AAAD2994CFFDEBA3F0085FD8DFCEC" box="[513,620,959,983]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994CFFDEBA3F0085FD8DFCEC" box="[513,620,959,983]" italics="true" pageId="33" pageNumber="34">Ta. bataar</emphasis>
</taxonomicName>
) or enlarged (
<taxonomicName id="4C2AAAD2994CFFDEB84200E5FF07FCCD" box="[124,230,990,1014]" class="Reptilia" genus="Bistahieversor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="sealeyi">
<emphasis id="B95E0D43994CFFDEB84200E5FF07FCCD" box="[124,230,990,1014]" italics="true" pageId="33" pageNumber="34">Bi. sealeyi</emphasis>
</taxonomicName>
,
<taxonomicName id="4C2AAAD2994CFFDEB8C400E4FE28FCCE" box="[250,457,990,1014]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B95E0D43994CFFDEB8C400E4FE6FFCCD" box="[250,398,990,1014]" italics="true" pageId="33" pageNumber="34">Daspletosaurus</emphasis>
spp.
</taxonomicName>
,
<emphasis id="B95E0D43994CFFDEB9E500E5FE18FCCD" box="[475,505,991,1014]" italics="true" pageId="33" pageNumber="34">Te.</emphasis>
<taxonomicName id="4C2AAAD2994CFFDEBA3900E5FD1BFCCD" authority=", Nanatotheristes" authorityName="Nanatotheristes" box="[519,762,990,1014]" class="Reptilia" genus="Teratophoneus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Avetheropoda" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="curriei">
<emphasis id="B95E0D43994CFFDEBA3900E5FDA4FCCD" box="[519,581,991,1014]" italics="true" pageId="33" pageNumber="34">curriei</emphasis>
,
<emphasis id="B95E0D43994CFFDEBA6400E4FD1BFCCD" box="[602,762,990,1014]" italics="true" pageId="33" pageNumber="34">Ŋanatotheristes</emphasis>
</taxonomicName>
<emphasis id="B95E0D43994CFFDEB84F00C7FF0CFB2E" box="[113,237,1021,1045]" italics="true" pageId="33" pageNumber="34">degrootorum</emphasis>
Voris
<emphasis id="B95E0D43994CFFDEB90D00C4FE84FB2E" box="[307,357,1021,1045]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
, 2020,
<taxonomicName id="4C2AAAD2994CFFDEB98D00C4FDFBFB2E" box="[435,538,1022,1045]" family="Tyrannosauridae" genus="Qianzhousaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="sinensis">
<emphasis id="B95E0D43994CFFDEB98D00C4FDFBFB2E" box="[435,538,1022,1045]" italics="true" pageId="33" pageNumber="34">Q. sinensis</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C2AAAD2994CFFDEBA6800C7FD0EFB2E" box="[598,751,1021,1045]" class="Reptilia" family="Tyrannosauridae" genus="Alioramus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="altai">
<emphasis id="B95E0D43994CFFDEBA6800C7FD0EFB2E" box="[598,751,1021,1045]" italics="true" pageId="33" pageNumber="34">Alioramus altai</emphasis>
</taxonomicName>
) states by Voris
<emphasis id="B95E0D43994CFFDEB92E0727FEA0FB0E" box="[272,321,1053,1077]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
(2021). However, the surangular foramen in
<taxonomicName id="4C2AAAD2994CFFDEB8B50707FF2AFB6F" baseAuthorityName="Carr" baseAuthorityYear="2020" box="[139,203,1085,1108]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="rex">
<emphasis id="B95E0D43994CFFDEB8B50707FF2AFB6F" box="[139,203,1085,1108]" italics="true" pageId="33" pageNumber="34">Ty.rex</emphasis>
</taxonomicName>
and
<taxonomicName id="4C2AAAD2994CFFDEB8C20706FE1FFB6F" authorityName="Osborn" authorityYear="1905" box="[252,510,1084,1108]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="sarcophagus">
<emphasis id="B95E0D43994CFFDEB8C20706FE1FFB6F" box="[252,510,1084,1108]" italics="true" pageId="33" pageNumber="34">Albertosaurus sarcophagus</emphasis>
</taxonomicName>
also used to be classified as smaller than in other tyrannosaurids (
<bibRefCitation id="EFBBACA0994CFFDEBA210761FF44FBA8" author="Carr TD &amp; Williamson TE" pageId="33" pageNumber="34" pagination="479 - 523" refId="ref25757" refString="Carr TD, Williamson TE. Diversity of late Maastrichtian Tyrannosauridae (Dinosauria: Theropoda) from western North America. Zoological Journal of the Linnean Society 2004; 142: 479 - 523. https: // doi. org / 10.1111 / j. 1096 - 3642.2004.00130. x" year="2004">Carr and Williamson 2004</bibRefCitation>
). Other authors reported that the surangular foramen in
<taxonomicName id="4C2AAAD2994CFFDEB84F07A1FF39FB89" box="[113,216,1178,1202]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994CFFDEB84F07A1FF39FB89" box="[113,216,1178,1202]" italics="true" pageId="33" pageNumber="34">Ta. bataar</emphasis>
</taxonomicName>
is smaller than in other tyrannosaurids and invariant during ontogeny (Tsuihiji
<emphasis id="B95E0D43994CFFDEB9420780FE4BFBEA" box="[380,426,1209,1233]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
2011, Voris
<emphasis id="B95E0D43994CFFDEBA180780FDB5FBEA" box="[550,596,1209,1233]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
2021). Also, the surangular foramen in
<taxonomicName id="4C2AAAD2994CFFDEB95307E2FD25FBCB" authorityName="' Dong" authorityYear="1977" box="[365,708,1240,1264]" class="Reptilia" family="Tyrannosauridae" genus="Shanshanosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="huoyanshanensis">
<emphasis id="B95E0D43994CFFDEB95307E2FD25FBCB" box="[365,708,1240,1264]" italics="true" pageId="33" pageNumber="34">Shanshanosaurus huoyanshanensis</emphasis>
</taxonomicName>
was described as large (
<bibRefCitation id="EFBBACA0994CFFDEB90007C2FDCFFA2B" author="Currie PJ &amp; Dong ZM" box="[318,558,1272,1296]" pageId="33" pageNumber="34" pagination="1729 - 37" refId="ref26336" refString="Currie PJ, Dong ZM. New information on Shanshanosaurus huoyanshanensis, a juvenile tyronnsaurid (Theropoda, Dinosauria) from the Late Cretaceous of China. Canadian Journal of Earth Sciences 2001; 38: 1729 - 37." year="2001">Currie and Dong 2001</bibRefCitation>
), but later as small (Tsuihiji
<emphasis id="B95E0D43994CFFDEB8EE0622FEE0FA14" box="[208,257,1303,1327]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
2011). For
<taxonomicName id="4C2AAAD2994CFFDEB9430622FE5EFA14" baseAuthorityName="Carr" baseAuthorityYear="2020" box="[381,447,1304,1327]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="rex">
<emphasis id="B95E0D43994CFFDEB9430622FE5EFA14" box="[381,447,1304,1327]" italics="true" pageId="33" pageNumber="34">Ty. rex</emphasis>
</taxonomicName>
, the size of the surangular foramen was first reported as increasing (
<bibRefCitation id="EFBBACA0994CFFDEBA39060DFD93FA74" author="Carr TD" box="[519,626,1335,1359]" pageId="33" pageNumber="34" pagination="497 - 520" refId="ref25647" refString="Carr TD. Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria). Journal of Vertebrate Paleontology 1999; 19: 497 - 520. https: // doi. org / 10.1080 / 02724634.1999.10011161" year="1999">Carr 1999</bibRefCitation>
) but later as decreasing in size through ontogeny (
<bibRefCitation id="EFBBACA0994CFFDEB9D0066CFDB7FA55" author="Carr TD" box="[494,598,1366,1390]" pageId="33" pageNumber="34" refId="ref25682" refString="Carr TD. A high-resolution growth series of Tyrannosaurus rex obtained from multiple lines of evidence. PeerJ 2020; 8: e 9192. https: // doi. org / 10.7717 / peerj. 9192" year="2020">Carr 2020</bibRefCitation>
).
</paragraph>
<paragraph id="8B95D151994CFFDEB8B3064CFC8EFC62" blockId="33.[113,766,144,1985]" lastBlockId="33.[810,1460,144,1986]" pageId="33" pageNumber="34">
Because of this confusion, we built a dataset to examine the size of foramina in a quantitative context. In tyrannosaurids, growth of the mandible, skull, and femur is isometric and related to the body size of the individual (
<bibRefCitation id="EFBBACA0994CFFDEBA2006E9FD4BFAD0" author="Currie PJ" box="[542,682,1491,1515]" pageId="33" pageNumber="34" pagination="651 - 65" refId="ref26287" refString="Currie PJ. Allometric growth in tyrannosaurids (Dinosauria: Theropoda) from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences 2003 b; 40: 651 - 65. https: // doi. org / 10.1139 / e 02 - 083" year="2003">Currie 2003b</bibRefCitation>
). Thus, we assessed the relationship between the size of the surangular foramen and skull length (as a proxy for body size). Our results (
<figureCitation id="1311CDD4994CFFDEB842050BFF20F972" box="[124,193,1585,1609]" captionStart="Figure 22" captionStartId="29.[113,178,818,842]" captionTargetBox="[117,1455,147,788]" captionTargetId="figure-606@29.[114,1458,144,791]" captionTargetPageId="29" captionText="Figure 22. Plots showing the relationship between the anteroposterior length of the surangular foramen and the skull length (as a proxy of body size) in tyrannosaurids.Bagaraatan ostromi was measured for the single (posterior only) or double (posterior + anterior) foramina. A, regression analysis including all specimens shows a negative slope and a high correlation coefficient. B, regression analysis for specific taxa shows the same trend. No distinction between small, medium, or large size of the surangular foramen can be noticed in the tyrannosaurids." figureDoi="http://doi.org/10.5281/zenodo.14284277" httpUri="https://zenodo.org/record/14284277/files/figure.png" pageId="33" pageNumber="34">Fig. 22</figureCitation>
) show that in all taxa the size of the surangular foramen decreases during ontogeny (negative allometry) and is strongly correlated with the length of the skull. Thus, e.g.
<taxonomicName id="4C2AAAD2994CFFDEBAA8054AFF7CF99C" class="Reptilia" family="Tyrannosauridae" genus="Alioramus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="altai">
<emphasis id="B95E0D43994CFFDEBAA8054AFF7CF99C" italics="true" pageId="33" pageNumber="34">Alioramus altai</emphasis>
</taxonomicName>
(IGM 100/1844) and the similar-sized
<taxonomicName id="4C2AAAD2994CFFDEBA0005B5FD50F99C" box="[574,689,1679,1703]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="libratus">
<emphasis id="B95E0D43994CFFDEBA0005B5FD50F99C" box="[574,689,1679,1703]" italics="true" pageId="33" pageNumber="34">Go. libratus</emphasis>
</taxonomicName>
(TMP 1991.36.500) have surangular foramina of proportionally the same size. Although the surangular foramenskull length correlation is statistically significant, variability in surangular foramen size is also apparent, especially in
<taxonomicName id="4C2AAAD2994CFFDEB9F70437FDDAF81F" box="[457,571,1804,1828]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="libratus">
<emphasis id="B95E0D43994CFFDEB9F70437FDDAF81F" box="[457,571,1804,1828]" italics="true" pageId="33" pageNumber="34">Go. libratus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C2AAAD2994CFFDEBA520437FD33F81F" box="[620,722,1804,1828]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994CFFDEBA520437FD33F81F" box="[620,722,1804,1828]" italics="true" pageId="33" pageNumber="34">Ta. bataar</emphasis>
</taxonomicName>
, for which the data are less fitted to the trend than for the other species (
<emphasis id="B95E0D43994CFFDEB8990476FF56F858" box="[167,183,1868,1891]" italics="true" pageId="33" pageNumber="34">
<collectionCode id="ED3B4994994CFFDEB8990476FF56F858" box="[167,183,1868,1891]" country="Chile" name="Departamento de Geologia, Universidad de Chile" pageId="33" pageNumber="34">R</collectionCode>
</emphasis>
<superScript id="7C5F7C19994CFFDEB8890470FF5EF863" attach="left" box="[183,191,1866,1880]" fontSize="6" pageId="33" pageNumber="34">2</superScript>
=.76.78, vs.
<emphasis id="B95E0D43994CFFDEB96C0476FE81F858" box="[338,352,1868,1891]" italics="true" pageId="33" pageNumber="34">
<collectionCode id="ED3B4994994CFFDEB96C0476FE81F858" box="[338,352,1868,1891]" country="Chile" name="Departamento de Geologia, Universidad de Chile" pageId="33" pageNumber="34">R</collectionCode>
</emphasis>
<superScript id="7C5F7C19994CFFDEB95E0470FE89F863" attach="left" box="[352,360,1866,1880]" fontSize="6" pageId="33" pageNumber="34">2</superScript>
&gt;.88.
<quantity id="4CD27CB4994CFFDEB9880471FE0AF858" box="[438,491,1867,1891]" metricMagnitude="0" metricUnit="m" metricValue="2.3876" pageId="33" pageNumber="34" unit="in" value="94.0">94 in</quantity>
<taxonomicName id="4C2AAAD2994CFFDEB9D10471FD50F858" box="[495,689,1867,1891]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B95E0D43994CFFDEB9D10471FD62F858" box="[495,643,1867,1891]" italics="true" pageId="33" pageNumber="34">Daspletosaurus</emphasis>
spp.
</taxonomicName>
and
<taxonomicName id="4C2AAAD2994CFFDEBAE10476FF6EF8B9" baseAuthorityName="Carr" baseAuthorityYear="2020" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="rex">
<emphasis id="B95E0D43994CFFDEBAE10476FF6EF8B9" italics="true" pageId="33" pageNumber="34">Ty. rex</emphasis>
</taxonomicName>
;
<figureCitation id="1311CDD4994CFFDEB8A00451FF05F8B8" box="[158,228,1899,1923]" captionStart="Figure 22" captionStartId="29.[113,178,818,842]" captionTargetBox="[117,1455,147,788]" captionTargetId="figure-606@29.[114,1458,144,791]" captionTargetPageId="29" captionText="Figure 22. Plots showing the relationship between the anteroposterior length of the surangular foramen and the skull length (as a proxy of body size) in tyrannosaurids.Bagaraatan ostromi was measured for the single (posterior only) or double (posterior + anterior) foramina. A, regression analysis including all specimens shows a negative slope and a high correlation coefficient. B, regression analysis for specific taxa shows the same trend. No distinction between small, medium, or large size of the surangular foramen can be noticed in the tyrannosaurids." figureDoi="http://doi.org/10.5281/zenodo.14284277" httpUri="https://zenodo.org/record/14284277/files/figure.png" pageId="33" pageNumber="34">Fig. 22</figureCitation>
). Indeed, although the surangular foramen is rather enlarged in
<taxonomicName id="4C2AAAD2994CFFDEB8D704B0FE83F899" authorityName="Maleev" authorityYear="1955" box="[233,354,1930,1954]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
<emphasis id="B95E0D43994CFFDEB8D704B0FE83F899" box="[233,354,1930,1954]" italics="true" pageId="33" pageNumber="34">Tarbosaurus</emphasis>
</taxonomicName>
individuals (as in other tyrannosaurids; Fig. 22) bigger than MPC-D 100/66 (skull length:
<quantity id="4CD27CB4994CFFDEBABF0493FD25F8FA" box="[641,708,1961,1985]" metricMagnitude="-1" metricUnit="m" metricValue="4.5" pageId="33" pageNumber="34" unit="cm" value="45.0">45 cm</quantity>
), few exceptions were found within the hypodigm. The surangular foramen of the medium-sized specimen MPC-D 107/14 is exceptionally small in comparison to other
<taxonomicName id="4C2AAAD2994CFFDEBCEA03F5FADAFFDC" box="[1236,1339,207,231]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994CFFDEBCEA03F5FADAFFDC" box="[1236,1339,207,231]" italics="true" pageId="33" pageNumber="34">Ta. bataar</emphasis>
</taxonomicName>
individuals of similar size (e.g. ZPAL MgD-I/3 and MPC-D 107/5;
<figureCitation id="1311CDD4994CFFDEBD5403D4FA43FE3D" box="[1386,1442,238,262]" captionStart="Figure 5" captionStartId="8.[129,194,1539,1563]" captionTargetBox="[131,1471,146,1509]" captionTargetId="figure-205@8.[129,1473,144,1511]" captionTargetPageId="8" captionText="Figure 5. Comparison of Bagaraatan ostromi (A) and Tarbosaurus bataar (BF) distal end of the mandible in lateral view.A, ZPAL MgD-I/108. B, ZPAL MgD-I/31. C, ZPAL MgD-I/3. D, MPC-D 107/14. E, ZPAL MgD-I/4. F, ZPAL MgD-I/5. Note the exceptionally small surangular foramen in the medium-sized individual MPC-D 107/14 (D) in comparison to similar-sized ZPAL MgD-I/3 (C). Asterisks indicate the posterior surangular foramen.Note that the placement of the double surangular foramina in B.ostromi is similar to the enlarged posterior surangular foramen in Ta.bataar. During ontogeny, the surangular shelf faces more laterally and the glenoid becomes longer anteroposteriorly." figureDoi="http://doi.org/10.5281/zenodo.14284239" httpUri="https://zenodo.org/record/14284239/files/figure.png" pageId="33" pageNumber="34">Fig. 5</figureCitation>
). Moreover, a specimen larger than those listed above, MPC-D 100/60, shows asymmetrical surangular foramina: the left one is smaller (anteroposterior length:
<quantity id="4CD27CB4994CFFDEBCA40276FB09FE5F" box="[1178,1256,332,356]" metricMagnitude="-2" metricUnit="m" metricValue="2.3" pageId="33" pageNumber="34" unit="mm" value="23.0">23 mm</quantity>
) and the right one larger (anteroposterior length:
<quantity id="4CD27CB4994CFFDEBC570251FB56FEB8" box="[1129,1207,363,387]" metricMagnitude="-2" metricUnit="m" metricValue="4.0" pageId="33" pageNumber="34" unit="mm" value="40.0">40 mm</quantity>
). The smaller surangular foramen of the left mandible can be noticed as an outlier in the
<figureCitation id="1311CDD4994CFFDEBB6D0290FC56FEF9" box="[851,951,426,450]" captionStart="Figure 22" captionStartId="29.[113,178,818,842]" captionTargetBox="[117,1455,147,788]" captionTargetId="figure-606@29.[114,1458,144,791]" captionTargetPageId="29" captionText="Figure 22. Plots showing the relationship between the anteroposterior length of the surangular foramen and the skull length (as a proxy of body size) in tyrannosaurids.Bagaraatan ostromi was measured for the single (posterior only) or double (posterior + anterior) foramina. A, regression analysis including all specimens shows a negative slope and a high correlation coefficient. B, regression analysis for specific taxa shows the same trend. No distinction between small, medium, or large size of the surangular foramen can be noticed in the tyrannosaurids." figureDoi="http://doi.org/10.5281/zenodo.14284277" httpUri="https://zenodo.org/record/14284277/files/figure.png" pageId="33" pageNumber="34">Figure 22</figureCitation>
. It would appear that there was some variability in the timing of the surangular foramen enlargement, at least in
<taxonomicName id="4C2AAAD2994CFFDEBB1402D3FC70FD3B" box="[810,913,488,512]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994CFFDEBB1402D3FC70FD3B" box="[810,913,488,512]" italics="true" pageId="33" pageNumber="34">Ta. bataar</emphasis>
</taxonomicName>
. The size of the surangular foramen in
<taxonomicName id="4C2AAAD2994CFFDEBD1902D3FA6BFD3B" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[1319,1418,489,512]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994CFFDEBD1902D3FA6BFD3B" box="[1319,1418,489,512]" italics="true" pageId="33" pageNumber="34">B. ostromi</emphasis>
</taxonomicName>
, regardless of whether it is measured for the single (posterior only) or double (posterior + anterior) foramina, falls into the overall variability of surangular size in the tyrannosaurids generally and
<taxonomicName id="4C2AAAD2994CFFDEBB14015DFC6EFD45" box="[810,911,614,638]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994CFFDEBB14015DFC6EFD45" box="[810,911,614,638]" italics="true" pageId="33" pageNumber="34">Ta. bataar</emphasis>
</taxonomicName>
specifically. The position of the surangular foramen in
<taxonomicName id="4C2AAAD2994CFFDEBB0E01BFFC3BFDA6" authorityName="Dong" authorityYear="1977" box="[816,986,645,669]" class="Reptilia" family="Tyrannosauridae" genus="Shanshanosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
<emphasis id="B95E0D43994CFFDEBB0E01BFFC3BFDA6" box="[816,986,645,669]" italics="true" pageId="33" pageNumber="34">Shanshanosaurus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C2AAAD2994CFFDEBC3101BCFB94FDA6" box="[1039,1141,645,669]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994CFFDEBC3101BCFB94FDA6" box="[1039,1141,645,669]" italics="true" pageId="33" pageNumber="34">Ta. bataar</emphasis>
</taxonomicName>
MPC-D 107/7 is similar to the position of the posterior opening in the surangular of
<taxonomicName id="4C2AAAD2994CFFDEBD05019FFA4FFD87" authorityName="Osmolska" authorityYear="1996" box="[1339,1454,677,700]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
<emphasis id="B95E0D43994CFFDEBD05019FFA4FFD87" box="[1339,1454,677,700]" italics="true" pageId="33" pageNumber="34">Bagaraatan</emphasis>
</taxonomicName>
, and those individuals cluster together on the plot. In turn, if the length of the area of both surangular foramina is measured for
<taxonomicName id="4C2AAAD2994CFFDEBB140039FC6CFC21" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[810,909,771,794]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994CFFDEBB140039FC6CFC21" box="[810,909,771,794]" italics="true" pageId="33" pageNumber="34">B. ostromi</emphasis>
</taxonomicName>
, it clusters with
<taxonomicName id="4C2AAAD2994CFFDEBC060039FB56FC20" box="[1080,1207,771,795]" class="Reptilia" family="Tyrannosauridae" genus="Raptorex" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="kriegsteini">
<emphasis id="B95E0D43994CFFDEBC060039FB56FC20" box="[1080,1207,771,795]" italics="true" pageId="33" pageNumber="34">R. kriegsteini</emphasis>
</taxonomicName>
, the surangular foramen of which was previously reported to be enlarged (
<bibRefCitation id="EFBBACA0994CFFDEBD080018FCBFFC62" author="Fowler DW &amp; Woodward HN &amp; Freedman EA" pageId="33" pageNumber="34" refId="ref26728" refString="Fowler DW, Woodward HN, Freedman EA et al. Reanalysis of &quot; Raptorex kriegsteini &quot;: a juvenile tyrannosaurid dinosaur from Mongolia. PLoS One 2011; 6: e 21376. https: // doi. org / 10.1371 / journal. pone. 0021376" year="2011">
Fowler
<emphasis id="B95E0D43994CFFDEBDBA0019FA55FC01" box="[1412,1460,802,826]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
2011
</bibRefCitation>
).
</paragraph>
<paragraph id="8B95D151994CFFDDBB7B005BFED1FFFC" blockId="33.[810,1460,144,1986]" lastBlockId="34.[129,777,144,199]" lastPageId="34" lastPageNumber="35" pageId="33" pageNumber="34">
What might explain the strange double set of foramina in
<taxonomicName id="4C2AAAD2994CFFDEBB1400BBFC7CFCA3" authority="Ne" authorityName="Osmolska" authorityYear="1996" box="[810,925,897,920]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
<emphasis id="B95E0D43994CFFDEBB1400BBFC7CFCA3" box="[810,925,897,920]" italics="true" pageId="33" pageNumber="34">Bagaraatan</emphasis>
</taxonomicName>
? The bone between the anterior and posterior surangular foramina in
<taxonomicName id="4C2AAAD2994CFFDEBC20009AFB62FC8C" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[1054,1155,928,951]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994CFFDEBC20009AFB62FC8C" box="[1054,1155,928,951]" italics="true" pageId="33" pageNumber="34">B. ostromi</emphasis>
</taxonomicName>
is very thin, and the relative position of this area and both foramina matches the surangular foramen of
<taxonomicName id="4C2AAAD2994CFFDEBB9E00E5FBFFFCCD" box="[928,1054,990,1014]" class="Reptilia" family="Tyrannosauridae" genus="Raptorex" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="kriegsteini">
<emphasis id="B95E0D43994CFFDEBB9E00E5FBFFFCCD" box="[928,1054,990,1014]" italics="true" pageId="33" pageNumber="34">R. kriegsteini</emphasis>
</taxonomicName>
and
<taxonomicName id="4C2AAAD2994CFFDEBC7100E5FB55FCCD" box="[1103,1204,990,1014]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994CFFDEBC7100E5FB55FCCD" box="[1103,1204,990,1014]" italics="true" pageId="33" pageNumber="34">Ta. bataar</emphasis>
</taxonomicName>
(ZPAL MgD-I/31). The surangular in tyrannosaurids during the early years of life was invaded by a pneumatic diverticulum (
<bibRefCitation id="EFBBACA0994CFFDEBC870726FABCFB0E" author="Gold MEL &amp; Brusatte SL &amp; Norell MA" box="[1209,1373,1052,1077]" pageId="33" pageNumber="34" pagination="1 - 46" refId="ref27220" refString="Gold MEL, Brusatte SL, Norell MA. The cranial pneumatic sinuses of the tyrannosaurid Alioramus (Dinosauria: Theropoda) and the evolution of cranial pneumaticity in theropod dinosaurs. American Museum Novitates 2013; 3790: 1 - 46. https: // doi. org / 10.1206 / 3790.1" year="2013">
Gold
<emphasis id="B95E0D43994CFFDEBCCA0727FAC5FB0F" box="[1268,1316,1052,1076]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
2013
</bibRefCitation>
), which pneumatized the bone and formed the enlarged surangular foramen, bordered by a pneumatic pocket posterodorsal to it. Given that more basal tyrannosauroids have a small foramen without a pneumatic pocket, it is not clear whether there was any pneumatic diverticulum in this region in these species. Owing to the fact that pneumatic diverticula induce bone resorption when they contact bone (
<bibRefCitation id="EFBBACA0994CFFDEBBC607C2FB61FA2B" author="Bremer JL" box="[1016,1152,1272,1296]" pageId="33" pageNumber="34" pagination="197 - 211" refId="ref25223" refString="Bremer JL. The pneumatization of the humerus in common fowl and the associated activity of theelin. Anatomical Record 1940; 77: 197 - 211. https: // doi. org / 10.1002 / ar. 1090770209" year="1940">Bremer 1940</bibRefCitation>
, Witmer 1997,
<bibRefCitation id="EFBBACA0994CFFDEBD1807C2FA43FA2B" author="Wedel M" box="[1318,1442,1272,1296]" pageId="33" pageNumber="34" refId="ref29166" refString="Wedel M. Postcranial pneumaticity in dinosaurs and the origin of the avian lung. Unpublished PhD Thesis, University of California, 2007." type="book" year="2007">Wedel 2007</bibRefCitation>
), we propose that the mandible of
<taxonomicName id="4C2AAAD2994CFFDEBCBA0622FB09FA14" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[1156,1256,1304,1327]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994CFFDEBCBA0622FB09FA14" box="[1156,1256,1304,1327]" italics="true" pageId="33" pageNumber="34">B. ostromi</emphasis>
</taxonomicName>
exhibits local bone resorption, induced by the pneumatic diverticula, that would explain the extremely thin bone between the anterior and posterior foramen. We hypothesize that if the pneumatization process continued slightly longer, the two foramina might have merged into a single large foramen, which is the common condition in
<taxonomicName id="4C2AAAD2994CFFDEBBAF06EEFBF7FAD0" authorityName="Marsh" authorityYear="1877" box="[913,1046,1492,1515]" class="Reptilia" family="Dryptosauridae" genus="Dryptosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
<emphasis id="B95E0D43994CFFDEBBAF06EEFBF7FAD0" box="[913,1046,1492,1515]" italics="true" pageId="33" pageNumber="34">Dryptosaurus</emphasis>
</taxonomicName>
+ Tyrannosauridae (
<bibRefCitation id="EFBBACA0994CFFDEBCCB06EEFCBFF930" author="Brusatte SL &amp; Carr TD" pageId="33" pageNumber="34" refId="ref25346" refString="Brusatte SL, Carr TD. The phylogeny and evolutionary history of tyrannosauroid dinosaurs. Scientific Reports 2016; 6: 20252." year="2016">Brusatte and Carr 2016</bibRefCitation>
). This proposal is supported by the fact that the posterior surangular foramen in
<taxonomicName id="4C2AAAD2994CFFDEBC260529FB9CF911" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[1048,1149,1555,1578]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994CFFDEBC260529FB9CF911" box="[1048,1149,1555,1578]" italics="true" pageId="33" pageNumber="34">B. ostromi</emphasis>
</taxonomicName>
is similar in length and positioned in a similar place as in the smaller
<taxonomicName id="4C2AAAD2994CFFDEBCE9050BFA60F972" authorityName="Dong" authorityYear="1977" box="[1239,1409,1585,1609]" class="Reptilia" family="Tyrannosauridae" genus="Shanshanosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
<emphasis id="B95E0D43994CFFDEBCE9050BFA60F972" box="[1239,1409,1585,1609]" italics="true" pageId="33" pageNumber="34">Shanshanosaurus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C2AAAD2994CFFDEBB14056BFC74F952" box="[810,917,1617,1641]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994CFFDEBB14056BFC74F952" box="[810,917,1617,1641]" italics="true" pageId="33" pageNumber="34">Ta. bataar</emphasis>
</taxonomicName>
MPC-D 107/7 (skull length ~
<quantity id="4CD27CB4994CFFDEBCD6056BFAD0F952" box="[1256,1329,1617,1641]" metricMagnitude="-1" metricUnit="m" metricValue="2.9" pageId="33" pageNumber="34" unit="cm" value="29.0">29 cm</quantity>
) specimens. Furthermore, the area of the surangular containing the posterior and anterior surangular foramina and the thinned bone between them matches the length and position of the surangular foramen in
<taxonomicName id="4C2AAAD2994CFFDEBBB905F5FC00F9DD" box="[903,993,1743,1766]" class="Reptilia" family="Tyrannosauridae" genus="Raptorex" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
<emphasis id="B95E0D43994CFFDEBBB905F5FC00F9DD" box="[903,993,1743,1766]" italics="true" pageId="33" pageNumber="34">Raptorex</emphasis>
</taxonomicName>
(skull length ~
<quantity id="4CD27CB4994CFFDEBCBD05F4FB2BF9DD" box="[1155,1226,1742,1766]" metricMagnitude="-1" metricUnit="m" metricValue="3.0" pageId="33" pageNumber="34" unit="cm" value="30.0">30 cm</quantity>
). Therefore,
<taxonomicName id="4C2AAAD2994CFFDEBD7005F5FA52F9DD" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[1358,1459,1743,1766]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994CFFDEBD7005F5FA52F9DD" box="[1358,1459,1743,1766]" italics="true" pageId="33" pageNumber="34">B. ostromi</emphasis>
</taxonomicName>
(skull also ~
<quantity id="4CD27CB4994CFFDEBB9905D7FC08F83E" box="[935,1001,1773,1797]" metricMagnitude="-1" metricUnit="m" metricValue="3.0" pageId="33" pageNumber="34" unit="cm" value="30.0">30 cm</quantity>
long) possibly captures the precise moment of ongoing bone resorption and perforation attributable to the pneumatic diverticulum. This process probably occurred early in ontogeny, in specimens
<quantity id="4CD27CB4994CFFDEBC1A0471FB89F858" box="[1060,1128,1867,1891]" metricMagnitude="0" metricUnit="m" metricValue="2.5" metricValueMax="3.0" metricValueMin="2.0" pageId="33" pageNumber="34" unit="m" value="2.5" valueMax="3.0" valueMin="2.0">23 m</quantity>
long, which were probably 23 years old at the time of death (as indicated for
<taxonomicName id="4C2AAAD2994CFFDEBCC80451FABAF8B9" box="[1270,1371,1898,1922]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994CFFDEBCC80451FABAF8B9" box="[1270,1371,1898,1922]" italics="true" pageId="33" pageNumber="34">Ta. bataar</emphasis>
</taxonomicName>
MPC-D 107/7 by Tsuihiji
<emphasis id="B95E0D43994CFFDEBBD604B1FBFBF899" box="[1000,1050,1930,1954]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
2011). Apparently, around this growth stage the pneumatic diverticulum invaded the bone, and thus variability in the size, shape, and even the number of foramina is to be expected.
</paragraph>
</subSubSection>
</treatment>
</document>

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@ -0,0 +1,436 @@
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<subSubSection id="C33082DA994FFFDDB92303DDFD8CFE3A" box="[285,621,231,257]" pageId="34" pageNumber="35" type="nomenclature">
<paragraph id="8B95D151994FFFDDB92303DDFD8CFE3A" blockId="34.[128,778,231,1985]" box="[285,621,231,257]" pageId="34" pageNumber="35">
<heading id="D0DD663D994FFFDDB92303DDFD8CFE3A" box="[285,621,231,257]" centered="true" fontSize="9" level="2" pageId="34" pageNumber="35" reason="3">
<emphasis id="B95E0D43994FFFDDB92303DDFD8CFE3A" bold="true" box="[285,621,231,257]" pageId="34" pageNumber="35">
Juvenile
<taxonomicName id="4C2AAAD2994FFFDDB94903DDFDCDFE3A" authorityName="Osborn" authorityYear="1905" box="[375,556,231,257]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
indet.
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C33082DA994FFFDCB8BF0237FF13FD45" lastPageId="35" lastPageNumber="36" pageId="34" pageNumber="35" type="discussion">
<paragraph id="8B95D151994FFFDDB8BF0237FD87FD04" blockId="34.[128,778,231,1985]" pageId="34" pageNumber="35">
Owing to its small body size and similarity to other juvenile tyrannosaurid specimens from the Nemegt, it is likely that
<taxonomicName id="4C2AAAD2994FFFDDBACB0217FF28FE5F" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994FFFDDBACB0217FF28FE5F" italics="true" pageId="34" pageNumber="35">B. ostromi</emphasis>
</taxonomicName>
is a juvenile tyrannosaurid. We tested this hypothesis further by determining whether
<taxonomicName id="4C2AAAD2994FFFDDB9E80256FDDBFEB8" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[470,570,364,387]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994FFFDDB9E80256FDDBFEB8" box="[470,570,364,387]" italics="true" pageId="34" pageNumber="35">B. ostromi</emphasis>
</taxonomicName>
shows juvenile features that have been well documented in
<taxonomicName id="4C2AAAD2994FFFDDBA0B02B1FD98FE99" baseAuthorityName="Carr" baseAuthorityYear="2020" box="[565,633,395,418]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="rex">
<emphasis id="B95E0D43994FFFDDBA0B02B1FD98FE99" box="[565,633,395,418]" italics="true" pageId="34" pageNumber="35">Ty. rex</emphasis>
</taxonomicName>
, whose ontogenetic osteological changes have been chronicled in detail by
<bibRefCitation id="EFBBACA0994FFFDDB8BF02F3FEE3FEDA" author="Carr TD" box="[129,258,457,481]" pageId="34" pageNumber="35" refId="ref25682" refString="Carr TD. A high-resolution growth series of Tyrannosaurus rex obtained from multiple lines of evidence. PeerJ 2020; 8: e 9192. https: // doi. org / 10.7717 / peerj. 9192" year="2020">Carr (2020)</bibRefCitation>
. We recognized that
<taxonomicName id="4C2AAAD2994FFFDDB9DC02F0FDA7FEDA" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[482,582,458,481]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994FFFDDB9DC02F0FDA7FEDA" box="[482,582,458,481]" italics="true" pageId="34" pageNumber="35">B. ostromi</emphasis>
</taxonomicName>
shows
<specimenCount id="9D2C1AD8994FFFDDBAA802F3FCE8FEDA" box="[662,777,457,481]" count="22" pageId="34" pageNumber="35" type="juvenile">22 juvenile</specimenCount>
and only
<specimenCount id="9D2C1AD8994FFFDDB8DA02D3FEA5FD3A" box="[228,324,489,513]" count="5" pageId="34" pageNumber="35" type="adult">five adult</specimenCount>
mandible features found in
<taxonomicName id="4C2AAAD2994FFFDDBA5002D3FD50FD3B" baseAuthorityName="Carr" baseAuthorityYear="2020" box="[622,689,489,512]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="rex">
<emphasis id="B95E0D43994FFFDDBA5002D3FD50FD3B" box="[622,689,489,512]" italics="true" pageId="34" pageNumber="35">Ty. rex</emphasis>
</taxonomicName>
by
<bibRefCitation id="EFBBACA0994FFFDDBAE402D3FF2AFD1B" author="Carr TD" pageId="34" pageNumber="35" refId="ref25682" refString="Carr TD. A high-resolution growth series of Tyrannosaurus rex obtained from multiple lines of evidence. PeerJ 2020; 8: e 9192. https: // doi. org / 10.7717 / peerj. 9192" year="2020">Carr (2020)</bibRefCitation>
. Four of the
<specimenCount id="9D2C1AD8994FFFDDB9750132FE5AFD1B" box="[331,443,520,544]" count="22" pageId="34" pageNumber="35" type="juvenile">22 juvenile</specimenCount>
features were found only in early juveniles, and the remaining
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late juveniles.
</paragraph>
<paragraph id="8B95D151994FFFDDB8A2017DFF24FA14" blockId="34.[128,778,231,1985]" pageId="34" pageNumber="35">
The mandible characters recognized both in
<taxonomicName id="4C2AAAD2994FFFDDBA62017DFD5CFD65" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[604,701,583,606]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994FFFDDBA62017DFD5CFD65" box="[604,701,583,606]" italics="true" pageId="34" pageNumber="35">B. ostromi</emphasis>
</taxonomicName>
and juveniles of
<taxonomicName id="4C2AAAD2994FFFDDB8D7015DFECAFD45" baseAuthorityName="Carr" baseAuthorityYear="2020" box="[233,299,615,638]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="rex">
<emphasis id="B95E0D43994FFFDDB8D7015DFECAFD45" box="[233,299,615,638]" italics="true" pageId="34" pageNumber="35">Ty. rex</emphasis>
</taxonomicName>
are as follows: (i) size of the first three alveoli of the dentary increasing posteriorly; (ii) shallow dentary in lateral view; (iii) shallow coronoid region of the surangular; (iv) no ridge delimiting the caudoventral fossa of the angular caudal process; (v) first two dentary alveoli much smaller than the latter alveoli; (vi) eighth tooth is the mesiodistally longest in the dentary; (vii) the alveoli decreasing posteriorly in mesiodistal length from the sixth to seventh alveolus; (viii) single large pit medial to the Meckelian fossa; (ix) low angle of the chin; (x) lightly textured chin region; (xi) distance of the ventralmost dentary foramen from the dorsal margin of the dentary to the total height of the bone&gt; 40%; (xii) the lateral extension of the surangular shelf horizontal; (xiii) surangular shelf not slanted; (xiv) small surangular foramen; (xv) caudal extent of the coronoid process declining before it reaches the glenoid; (xvi) presence of an embayment on the caudal margin of the surangular foramen; (xvii) cleft between the caudal glenoid process dorsoventrally short and shallow; (xvii) caudal end of the surangular shelf fading below the glenoid region; (xix) lateral scar on the surangular present; (xx) caudal glenoid process as tall as the rostral process; (xxi) lateral scar on the surangular rugose and shallow; and (xxii) dorsal orientation of the anterior glenoid foramen (
<bibRefCitation id="EFBBACA0994FFFDDBAE407C2FF54FA14" author="Carr TD" pageId="34" pageNumber="35" refId="ref25682" refString="Carr TD. A high-resolution growth series of Tyrannosaurus rex obtained from multiple lines of evidence. PeerJ 2020; 8: e 9192. https: // doi. org / 10.7717 / peerj. 9192" year="2020">Carr 2020</bibRefCitation>
).
</paragraph>
<paragraph id="8B95D151994FFFDDB8A2060DFB42FD04" blockId="34.[128,778,231,1985]" lastBlockId="34.[825,1475,143,1985]" pageId="34" pageNumber="35">
The prevalence of features shared by
<taxonomicName id="4C2AAAD2994FFFDDBA21060DFD62FA75" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[543,643,1335,1358]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994FFFDDBA21060DFD62FA75" box="[543,643,1335,1358]" italics="true" pageId="34" pageNumber="35">B. ostromi</emphasis>
</taxonomicName>
and juvenile
<taxonomicName id="4C2AAAD2994FFFDDB8BF066CFF29FA56" baseAuthorityName="Carr" baseAuthorityYear="2020" box="[129,200,1366,1389]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="rex">
<emphasis id="B95E0D43994FFFDDB8BF066CFF29FA56" box="[129,200,1366,1389]" italics="true" pageId="34" pageNumber="35">Ty. rex</emphasis>
</taxonomicName>
supports the identification of
<taxonomicName id="4C2AAAD2994FFFDDBA23066CFD64FA56" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[541,645,1366,1389]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994FFFDDBA23066CFD64FA56" box="[541,645,1366,1389]" italics="true" pageId="34" pageNumber="35">B. ostromi</emphasis>
</taxonomicName>
as a young tyrannosaurid. The less numerous adult
<taxonomicName id="4C2AAAD2994FFFDDBA14064CFD8FFAB6" baseAuthorityName="Carr" baseAuthorityYear="2020" box="[554,622,1398,1421]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="rex">
<emphasis id="B95E0D43994FFFDDBA14064CFD8FFAB6" box="[554,622,1398,1421]" italics="true" pageId="34" pageNumber="35">Ty. rex</emphasis>
</taxonomicName>
features (
<bibRefCitation id="EFBBACA0994FFFDDBAE4064FFF55FA97" author="Carr TD" pageId="34" pageNumber="35" refId="ref25682" refString="Carr TD. A high-resolution growth series of Tyrannosaurus rex obtained from multiple lines of evidence. PeerJ 2020; 8: e 9192. https: // doi. org / 10.7717 / peerj. 9192" year="2020">Carr 2020</bibRefCitation>
) found in
<taxonomicName id="4C2AAAD2994FFFDDB91606AFFE6DFA97" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[296,396,1429,1452]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994FFFDDB91606AFFE6DFA97" box="[296,396,1429,1452]" italics="true" pageId="34" pageNumber="35">B. ostromi</emphasis>
</taxonomicName>
are listed below, with comments regarding variability within
<taxonomicName id="4C2AAAD2994FFFDDB9B0068EFE16FAF7" box="[398,503,1460,1484]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994FFFDDB9B0068EFE16FAF7" box="[398,503,1460,1484]" italics="true" pageId="34" pageNumber="35">Ta. bataar</emphasis>
</taxonomicName>
. First, the second dentary tooth is&gt; 75% of the mesiodistal length of the third dentary tooth. The proportions between the first three dentary teeth in
<taxonomicName id="4C2AAAD2994FFFDDB8BE0528FF05F911" box="[128,228,1554,1578]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994FFFDDB8BE0528FF05F911" box="[128,228,1554,1578]" italics="true" pageId="34" pageNumber="35">Ta.bataar</emphasis>
</taxonomicName>
seem to be variable. In the subadult ZPAL MgD-I/175, the second dentary tooth is &lt;75% of the mesiodistal length of the third dentary tooth, and in the adults (ZPAL MgD-I/5) it is between 70 and 78%. Second, the combined mesiodistal lengths of the first two alveoli of the dentary are greater than the mesiodistal length of the third alveolus, as in all examined individuals of
<taxonomicName id="4C2AAAD2994FFFDDB92505F4FE67F9DD" box="[283,390,1742,1766]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994FFFDDB92505F4FE67F9DD" box="[283,390,1742,1766]" italics="true" pageId="34" pageNumber="35">Ta. bataar</emphasis>
</taxonomicName>
(subadults ZPAL MgD-I/45, ZPAL MgD-I/46, and ZPAL MgD-I/175; and adults ZPAL MgD-I/4 and ZPAL MgD-I/5; Table 1). However, the difference in all
<taxonomicName id="4C2AAAD2994FFFDDBAD40437FF20F878" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994FFFDDBAD40437FF20F878" italics="true" pageId="34" pageNumber="35">Ta. bataar</emphasis>
</taxonomicName>
specimens is greater (~
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) than in
<taxonomicName id="4C2AAAD2994FFFDDBA6E0416FD52F878" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[592,691,1836,1859]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994FFFDDBA6E0416FD52F878" box="[592,691,1836,1859]" italics="true" pageId="34" pageNumber="35">B. ostromi</emphasis>
</taxonomicName>
(
<quantity id="4CD27CB4994FFFDDBAF90416FD1EF87F" box="[711,767,1836,1860]" metricMagnitude="-2" metricUnit="m" metricValue="1.0" pageId="34" pageNumber="35" unit="cm" value="1.0">1 cm</quantity>
). Third, there is no deviation in the chin region, which is not recognized in any examined
<taxonomicName id="4C2AAAD2994FFFDDB9980451FDECF8B9" box="[422,525,1898,1922]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994FFFDDB9980451FDECF8B9" box="[422,525,1898,1922]" italics="true" pageId="34" pageNumber="35">Ta. bataar</emphasis>
</taxonomicName>
specimen (ZPAL MgD-I/45, ZPAL MgD-I/46, ZPAL MgD-I/175, ZPAL MgD-I/4, and ZPAL MgD-I/5), nor it has been described in juvenile Ta.
<taxonomicName id="4C2AAAD2994FFFDDBB0703B5FC9BFF9C" box="[825,890,143,167]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994FFFDDBB0703B5FC9BFF9C" box="[825,890,143,167]" italics="true" pageId="34" pageNumber="35">bataar</emphasis>
</taxonomicName>
MPC-D 107/7 (Tsuihiji
<emphasis id="B95E0D43994FFFDDBCBC03AAFB52FF9C" box="[1154,1203,143,167]" italics="true" pageId="34" pageNumber="35">et al.</emphasis>
2011). Fourth, the caudal surangular foramen is positioned far anteriorly to the glenoid, as in all
<taxonomicName id="4C2AAAD2994FFFDDBB4A03F5FC3AFFDD" box="[884,987,206,230]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994FFFDDBB4A03F5FC3AFFDD" box="[884,987,206,230]" italics="true" pageId="34" pageNumber="35">Ta. bataar</emphasis>
</taxonomicName>
individuals (ZPAL MgD-I/4, ZPAL MgD-I/5, and ZPAL MgD-I/31), including the juvenile MPC-D 107/7 (Tsuihiji
<emphasis id="B95E0D43994FFFDDBBA80234FC24FE1E" box="[918,965,269,293]" italics="true" pageId="34" pageNumber="35">et al.</emphasis>
2011). Fifth, the glenoid fossa is short and deep in adult
<taxonomicName id="4C2AAAD2994FFFDDBB4C0217FC52FE7F" baseAuthorityName="Carr" baseAuthorityYear="2020" box="[882,947,301,324]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="rex">
<emphasis id="B95E0D43994FFFDDBB4C0217FC52FE7F" box="[882,947,301,324]" italics="true" pageId="34" pageNumber="35">Ty. rex</emphasis>
</taxonomicName>
and
<taxonomicName id="4C2AAAD2994FFFDDBBDD0217FBA5FE7F" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[995,1092,301,324]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994FFFDDBBDD0217FBA5FE7F" box="[995,1092,301,324]" italics="true" pageId="34" pageNumber="35">B. ostromi</emphasis>
</taxonomicName>
. A shallow and long glenoid fossa can be recognized in
<taxonomicName id="4C2AAAD2994FFFDDBBD10271FAA3FE58" authorityName="' Dong" authorityYear="1977" box="[1007,1346,331,355]" class="Reptilia" family="Tyrannosauridae" genus="Shanshanosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="huoyanshanensis">
<emphasis id="B95E0D43994FFFDDBBD10271FAA3FE58" box="[1007,1346,331,355]" italics="true" pageId="34" pageNumber="35">Shanshanosaurus huoyanshanensis</emphasis>
</taxonomicName>
see (
<bibRefCitation id="EFBBACA0994FFFDDBD410276FC37FEB8" author="Currie PJ &amp; Dong ZM" pageId="34" pageNumber="35" pagination="1729 - 37" refId="ref26336" refString="Currie PJ, Dong ZM. New information on Shanshanosaurus huoyanshanensis, a juvenile tyronnsaurid (Theropoda, Dinosauria) from the Late Cretaceous of China. Canadian Journal of Earth Sciences 2001; 38: 1729 - 37." year="2001">Currie and Dong 2001</bibRefCitation>
), but already in the slightly larger MPC-D 107/7 and
<taxonomicName id="4C2AAAD2994FFFDDBB5A02B1FC25FE99" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[868,964,395,418]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994FFFDDBB5A02B1FC25FE99" box="[868,964,395,418]" italics="true" pageId="34" pageNumber="35">B. ostromi</emphasis>
</taxonomicName>
, as in young and adult
<taxonomicName id="4C2AAAD2994FFFDDBC9802B1FAEAFE98" box="[1190,1291,395,419]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994FFFDDBC9802B1FAEAFE98" box="[1190,1291,395,419]" italics="true" pageId="34" pageNumber="35">Ta. bataar</emphasis>
</taxonomicName>
(ZPAL MgD-I/4, ZPAL MgD-I/5, and ZPAL MgD-I/31), it is narrow and deep. Those features possibly indicate some species-dependent variability, similar to the proportion of the antorbital fenestra, which does not shorten as much during the ontogeny of
<taxonomicName id="4C2AAAD2994FFFDDBD0E0133FA74FD1B" box="[1328,1429,520,544]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994FFFDDBD0E0133FA74FD1B" box="[1328,1429,520,544]" italics="true" pageId="34" pageNumber="35">Ta. bataar</emphasis>
</taxonomicName>
as it does in
<taxonomicName id="4C2AAAD2994FFFDDBBB60112FB7FFD04" authority="(Tsuihiji et al. 2011)" baseAuthorityName="Tsuihiji" baseAuthorityYear="2011" box="[904,1182,551,575]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="rex">
<emphasis id="B95E0D43994FFFDDBBB60112FC29FD04" box="[904,968,552,575]" italics="true" pageId="34" pageNumber="35">Ty. rex</emphasis>
(Tsuihiji
<emphasis id="B95E0D43994FFFDDBC150112FBBBFD04" box="[1067,1114,551,575]" italics="true" pageId="34" pageNumber="35">et al.</emphasis>
2011)
</taxonomicName>
.
</paragraph>
<paragraph id="8B95D151994FFFDDBB6B017DFA9CF9DD" blockId="34.[825,1475,143,1985]" pageId="34" pageNumber="35">
As it is clear that the
<taxonomicName id="4C2AAAD2994FFFDDBC16017DFB6BFD65" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[1064,1162,583,606]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994FFFDDBC16017DFB6BFD65" box="[1064,1162,583,606]" italics="true" pageId="34" pageNumber="35">B. ostromi</emphasis>
</taxonomicName>
holotype belongs to a juvenile tyrannosaurid, the question becomes: can we identify which species it belonged to? We can first make comparisons with the other Nemegt tyrannosaurids:
<taxonomicName id="4C2AAAD2994FFFDDBC4F019FFB37FD86" box="[1137,1238,677,701]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994FFFDDBC4F019FFB37FD86" box="[1137,1238,677,701]" italics="true" pageId="34" pageNumber="35">Ta. bataar</emphasis>
</taxonomicName>
and
<taxonomicName id="4C2AAAD2994FFFDDBD3B019FFA76FD87" box="[1285,1431,677,701]" class="Reptilia" family="Tyrannosauridae" genus="Alioramus" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B95E0D43994FFFDDBD3B019FFA8BFD86" box="[1285,1386,677,701]" italics="true" pageId="34" pageNumber="35">Alioramus</emphasis>
spp.
</taxonomicName>
The mandible of
<taxonomicName id="4C2AAAD2994FFFDDBB8801FFFBF7FDE7" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[950,1046,709,732]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994FFFDDBB8801FFFBF7FDE7" box="[950,1046,709,732]" italics="true" pageId="34" pageNumber="35">B. ostromi</emphasis>
</taxonomicName>
is generally similar to juvenile
<taxonomicName id="4C2AAAD2994FFFDDBD7A01FFFA49FDE7" box="[1348,1448,708,732]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994FFFDDBD7A01FFFA49FDE7" box="[1348,1448,708,732]" italics="true" pageId="34" pageNumber="35">Ta. bataar</emphasis>
</taxonomicName>
or putative juveniles of that species, like
<taxonomicName id="4C2AAAD2994FFFDDBC8201DEFA94FDC0" authorityName="' Dong" authorityYear="1977" box="[1212,1397,739,763]" class="Reptilia" family="Tyrannosauridae" genus="Shanshanosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="huoyanshanensis">
<emphasis id="B95E0D43994FFFDDBC8201DEFA94FDC0" box="[1212,1397,739,763]" italics="true" pageId="34" pageNumber="35">S. huoyanshanensis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C2AAAD2994FFFDDBD9501DEFC7AFC21" class="Reptilia" family="Tyrannosauridae" genus="Raptorex" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="kriegsteini">
<emphasis id="B95E0D43994FFFDDBD9501DEFC7AFC21" italics="true" pageId="34" pageNumber="35">R. kriegsteini</emphasis>
</taxonomicName>
(see
<bibRefCitation id="EFBBACA0994FFFDDBBF10038FB55FC20" author="Currie PJ &amp; Dong ZM" box="[975,1204,770,795]" pageId="34" pageNumber="35" pagination="1729 - 37" refId="ref26336" refString="Currie PJ, Dong ZM. New information on Shanshanosaurus huoyanshanensis, a juvenile tyronnsaurid (Theropoda, Dinosauria) from the Late Cretaceous of China. Canadian Journal of Earth Sciences 2001; 38: 1729 - 37." year="2001">Currie and Dong 2001</bibRefCitation>
, Sereno
<emphasis id="B95E0D43994FFFDDBD350039FAD8FC21" box="[1291,1337,770,794]" italics="true" pageId="34" pageNumber="35">et al.</emphasis>
2009,
<bibRefCitation id="EFBBACA0994FFFDDBD450038FC47FC01" author="Fowler DW &amp; Woodward HN &amp; Freedman EA" pageId="34" pageNumber="35" refId="ref26728" refString="Fowler DW, Woodward HN, Freedman EA et al. Reanalysis of &quot; Raptorex kriegsteini &quot;: a juvenile tyrannosaurid dinosaur from Mongolia. PLoS One 2011; 6: e 21376. https: // doi. org / 10.1371 / journal. pone. 0021376" year="2011">
Fowler
<emphasis id="B95E0D43994FFFDDBB070019FC8AFC01" box="[825,875,802,826]" italics="true" pageId="34" pageNumber="35">et al.</emphasis>
2011
</bibRefCitation>
, Tsuihiji
<emphasis id="B95E0D43994FFFDDBC340019FBDDFC01" box="[1034,1084,802,826]" italics="true" pageId="34" pageNumber="35">et al.</emphasis>
2011). The dentary is straight in the dorsal and ventral view, shallow, slender, and thickens and tapers dorsally at the anterior end.
<taxonomicName id="4C2AAAD2994FFFDDBC52005BFAD1FC43" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[1132,1328,865,888]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994FFFDDBC52005BFAD1FC43" box="[1132,1328,865,888]" italics="true" pageId="34" pageNumber="35">Bagaraatan ostromi</emphasis>
</taxonomicName>
, like MPC-D 107/7, but in contrast to
<taxonomicName id="4C2AAAD2994FFFDDBC7400BBFB53FCA3" box="[1098,1202,896,920]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994FFFDDBC7400BBFB53FCA3" box="[1098,1202,896,920]" italics="true" pageId="34" pageNumber="35">Ta. bataar</emphasis>
</taxonomicName>
specimens and
<taxonomicName id="4C2AAAD2994FFFDDBD6300BAFC84FC8C" class="Reptilia" family="Tyrannosauridae" genus="Alioramus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="altai">
<emphasis id="B95E0D43994FFFDDBD6300BAFC84FC8C" italics="true" pageId="34" pageNumber="35">Alioramus altai</emphasis>
</taxonomicName>
, does not show any pneumatic pocket behind the surangular foramen (Tsuihiji
<emphasis id="B95E0D43994FFFDDBBCA0085FBC5FCED" box="[1012,1060,958,982]" italics="true" pageId="34" pageNumber="35">et al.</emphasis>
2011,
<bibRefCitation id="EFBBACA0994FFFDDBC540085FACFFCEC" author="Brusatte SL &amp; Carr TD &amp; Norell MA" box="[1130,1326,958,983]" pageId="34" pageNumber="35" pagination="1 - 197" refId="ref25467" refString="Brusatte SL, Carr TD, Norell MA. The osteology of Alioramus, a gracile and long-snouted tyrannosaurid (Dinosauria: Theropoda) from the Late Cretaceous of Mongolia. Bulletin of the American Museum of Natural History 2012; 366: 1 - 197. https: // doi. org / 10.1206 / 770.1" year="2012">
Brusatte
<emphasis id="B95E0D43994FFFDDBCFB0085FB14FCED" box="[1221,1269,958,982]" italics="true" pageId="34" pageNumber="35">et al.</emphasis>
2012
</bibRefCitation>
). The cervical vertebrae of
<taxonomicName id="4C2AAAD2994FFFDDBBFC00E5FBC9FCCE" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[962,1064,990,1014]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994FFFDDBBFC00E5FBC9FCCE" box="[962,1064,990,1014]" italics="true" pageId="34" pageNumber="35">B. ostromi</emphasis>
</taxonomicName>
strongly resemble the middle or posterior cervical vertebrae of
<taxonomicName id="4C2AAAD2994FFFDDBC6200C7FAF9FB2E" authorityName="' Dong" authorityYear="1977" box="[1116,1304,1021,1045]" class="Reptilia" family="Tyrannosauridae" genus="Shanshanosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="huoyanshanensis">
<emphasis id="B95E0D43994FFFDDBC6200C7FAF9FB2E" box="[1116,1304,1021,1045]" italics="true" pageId="34" pageNumber="35">S. huoyanshanensis</emphasis>
</taxonomicName>
. They share the posterodorsal rather than dorsal inclination of the neural spines, and have less flexed centra than in adult, large tyrannosaurids (
<bibRefCitation id="EFBBACA0994FFFDDBB7A0761FBA8FB48" author="Currie PJ &amp; Zhiming D" box="[836,1097,1115,1139]" pageId="34" pageNumber="35" pagination="1729 - 1737" refId="ref26525" refString="Currie PJ, Zhiming D. New information on Shanshanosaurus huoyanshanensis, a juvenile tyrannosaurid (Theropoda, Dinosauria) from the Late Cretaceous of China. Canadian Journal of Earth Sciences 2001; 38: 1729 - 1737. https: // doi. org / 10.1139 / cjes- 38 - 12 - 1729" year="2001">Currie and Zhiming 2001</bibRefCitation>
). The fusion of some bones occurred early in the ontogeny of tyrannosaurids, e.g. the juvenile
<taxonomicName id="4C2AAAD2994FFFDDBD9D0741FC9BFB89" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994FFFDDBD9D0741FC9BFB89" italics="true" pageId="34" pageNumber="35">Ta. bataar</emphasis>
</taxonomicName>
already has fused nasals (Tsuihiji
<emphasis id="B95E0D43994FFFDDBCE307A0FAEFFB89" box="[1245,1294,1178,1202]" italics="true" pageId="34" pageNumber="35">et al.</emphasis>
2011). However, the articular remains unfused with the surangular in
<taxonomicName id="4C2AAAD2994FFFDDBD670780FA5DFBEA" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[1369,1468,1210,1233]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994FFFDDBD670780FA5DFBEA" box="[1369,1468,1210,1233]" italics="true" pageId="34" pageNumber="35">B. ostromi</emphasis>
</taxonomicName>
, similar to
<taxonomicName id="4C2AAAD2994FFFDDBB8A07E3FB94FBCB" authorityName="' Dong" authorityYear="1977" box="[948,1141,1240,1264]" class="Reptilia" family="Tyrannosauridae" genus="Shanshanosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="huoyanshanensis">
<emphasis id="B95E0D43994FFFDDBB8A07E3FB94FBCB" box="[948,1141,1240,1264]" italics="true" pageId="34" pageNumber="35">S. huoyanshanensis</emphasis>
</taxonomicName>
, juvenile
<taxonomicName id="4C2AAAD2994FFFDDBCD707E3FAB7FBCB" box="[1257,1366,1240,1264]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994FFFDDBCD707E3FAB7FBCB" box="[1257,1366,1240,1264]" italics="true" pageId="34" pageNumber="35">Ta. bataar</emphasis>
</taxonomicName>
(MPC-D 107/7), and
<taxonomicName id="4C2AAAD2994FFFDDBBFF07C2FBBBFA2B" box="[961,1114,1272,1296]" class="Reptilia" family="Tyrannosauridae" genus="Alioramus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="altai">
<emphasis id="B95E0D43994FFFDDBBFF07C2FBBBFA2B" box="[961,1114,1272,1296]" italics="true" pageId="34" pageNumber="35">Alioramus altai</emphasis>
</taxonomicName>
(see
<bibRefCitation id="EFBBACA0994FFFDDBCAA07C2FABFFA2B" author="Brusatte SL &amp; Carr TD &amp; Norell MA" box="[1172,1374,1272,1296]" pageId="34" pageNumber="35" pagination="1 - 197" refId="ref25467" refString="Brusatte SL, Carr TD, Norell MA. The osteology of Alioramus, a gracile and long-snouted tyrannosaurid (Dinosauria: Theropoda) from the Late Cretaceous of Mongolia. Bulletin of the American Museum of Natural History 2012; 366: 1 - 197. https: // doi. org / 10.1206 / 770.1" year="2012">
Brusatte
<emphasis id="B95E0D43994FFFDDBCCF07C2FAC3FA2B" box="[1265,1314,1272,1296]" italics="true" pageId="34" pageNumber="35">et al.</emphasis>
2012
</bibRefCitation>
). In contrast, in
<taxonomicName id="4C2AAAD2994FFFDDBBB10622FC08FA14" box="[911,1001,1304,1327]" class="Reptilia" family="Tyrannosauridae" genus="Raptorex" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="genus">
<emphasis id="B95E0D43994FFFDDBBB10622FC08FA14" box="[911,1001,1304,1327]" italics="true" pageId="34" pageNumber="35">Raptorex</emphasis>
</taxonomicName>
and larger
<taxonomicName id="4C2AAAD2994FFFDDBC600622FB24FA14" box="[1118,1221,1303,1327]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994FFFDDBC600622FB24FA14" box="[1118,1221,1303,1327]" italics="true" pageId="34" pageNumber="35">Ta. bataar</emphasis>
</taxonomicName>
individuals the articular is fused to the surangular. Moreover, early in ontogeny partial fusion of the pelvis was reported in
<taxonomicName id="4C2AAAD2994FFFDDBC8E066CFAEBFA56" box="[1200,1290,1366,1389]" class="Reptilia" family="Tyrannosauridae" genus="Raptorex" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="genus">
<emphasis id="B95E0D43994FFFDDBC8E066CFAEBFA56" box="[1200,1290,1366,1389]" italics="true" pageId="34" pageNumber="35">Raptorex</emphasis>
</taxonomicName>
(see
<bibRefCitation id="EFBBACA0994FFFDDBD7A066CFC8CFAB6" author="Fowler DW &amp; Woodward HN &amp; Freedman EA" pageId="34" pageNumber="35" refId="ref26728" refString="Fowler DW, Woodward HN, Freedman EA et al. Reanalysis of &quot; Raptorex kriegsteini &quot;: a juvenile tyrannosaurid dinosaur from Mongolia. PLoS One 2011; 6: e 21376. https: // doi. org / 10.1371 / journal. pone. 0021376" year="2011">
Fowler
<emphasis id="B95E0D43994FFFDDBDAC066CFA22FA55" box="[1426,1475,1366,1390]" italics="true" pageId="34" pageNumber="35">et al.</emphasis>
2011
</bibRefCitation>
) and young
<taxonomicName id="4C2AAAD2994FFFDDBBCA064CFBD6FAB6" box="[1012,1079,1398,1421]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="rex">
<emphasis id="B95E0D43994FFFDDBBCA064CFBD6FAB6" box="[1012,1079,1398,1421]" italics="true" pageId="34" pageNumber="35">Ty. rex</emphasis>
</taxonomicName>
(BMR P
<date id="FF94F791994FFFDDBCA8064FFB13FAB6" box="[1174,1266,1397,1421]" pageId="34" pageNumber="35" value="2002-04-01">2002.4.1</date>
, Jane;
<bibRefCitation id="EFBBACA0994FFFDDBD7C064FFC8CFA97" author="Parrish JM &amp; Molar RE &amp; Currie PJ &amp; Koppelhus EB" pageId="34" pageNumber="35" refId="ref28561" refString="Parrish JM, Molar RE, Currie PJ, Koppelhus EB. Tyrannosaurid Paleobiology. Bloomington and Indianapolis, Indiana University Press; 2013." type="book" year="2013">
Parrish
<emphasis id="B95E0D43994FFFDDBDAC064CFA22FAB6" box="[1426,1475,1397,1421]" italics="true" pageId="34" pageNumber="35">et al.</emphasis>
2013
</bibRefCitation>
). In contrast, the pelvic bones are unfused in
<taxonomicName id="4C2AAAD2994FFFDDBD0406AFFA7AFA97" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[1338,1435,1429,1452]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994FFFDDBD0406AFFA7AFA97" box="[1338,1435,1429,1452]" italics="true" pageId="34" pageNumber="35">B. ostromi</emphasis>
</taxonomicName>
, juvenile
<taxonomicName id="4C2AAAD2994FFFDDBB42068FFC03FAF7" box="[892,994,1460,1484]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994FFFDDBB42068FFC03FAF7" box="[892,994,1460,1484]" italics="true" pageId="34" pageNumber="35">Ta. bataar</emphasis>
</taxonomicName>
(MPC-D 107/7), and subadult
<taxonomicName id="4C2AAAD2994FFFDDBD12068EFA23FAF7" box="[1324,1474,1460,1484]" class="Reptilia" family="Tyrannosauridae" genus="Alioramus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="altai">
<emphasis id="B95E0D43994FFFDDBD12068EFA23FAF7" box="[1324,1474,1460,1484]" italics="true" pageId="34" pageNumber="35">Alioramus altai</emphasis>
</taxonomicName>
(see
<bibRefCitation id="EFBBACA0994FFFDDBB5706EEFBCAFAD0" author="Brusatte SL &amp; Carr TD &amp; Norell MA" box="[873,1067,1491,1515]" pageId="34" pageNumber="35" pagination="1 - 197" refId="ref25467" refString="Brusatte SL, Carr TD, Norell MA. The osteology of Alioramus, a gracile and long-snouted tyrannosaurid (Dinosauria: Theropoda) from the Late Cretaceous of Mongolia. Bulletin of the American Museum of Natural History 2012; 366: 1 - 197. https: // doi. org / 10.1206 / 770.1" year="2012">
Brusatte
<emphasis id="B95E0D43994FFFDDBBFD06EEFC13FAD0" box="[963,1010,1491,1515]" italics="true" pageId="34" pageNumber="35">et al.</emphasis>
2012
</bibRefCitation>
) However, an early fusion of the cranial sutures might not necessarily be associated with an early fusion of postcrania, because these functional units could be subjected to developmental plasticity or separate evolutionary pressure depending on the ecology and preferred or available diet. The co-ossification of postcranial sutures and fusion between bones among tyranosaurids require further study. However, owing to their high variability, also in juveniles, we do not find them to be an adequate indicator for growth stage in tyrannosaurids.
</paragraph>
<paragraph id="8B95D151994FFFDCBB6B05D4FF13FD45" blockId="34.[825,1475,143,1985]" lastBlockId="35.[113,763,144,638]" lastPageId="35" lastPageNumber="36" pageId="34" pageNumber="35">
We can more thoroughly compare
<taxonomicName id="4C2AAAD2994FFFDDBCEA05D4FADFF83E" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[1236,1342,1774,1797]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994FFFDDBCEA05D4FADFF83E" box="[1236,1342,1774,1797]" italics="true" pageId="34" pageNumber="35">B. ostromi</emphasis>
</taxonomicName>
with young juvenile
<taxonomicName id="4C2AAAD2994FFFDDBBAB0437FBE1F81F" box="[917,1024,1804,1828]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994FFFDDBBAB0437FBE1F81F" box="[917,1024,1804,1828]" italics="true" pageId="34" pageNumber="35">Ta. bataar</emphasis>
</taxonomicName>
, because no young juveniles of
<taxonomicName id="4C2AAAD2994FFFDDBD640436FC85F878" class="Reptilia" family="Tyrannosauridae" genus="Alioramus" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B95E0D43994FFFDDBD640436FA23F81F" box="[1370,1474,1804,1828]" italics="true" pageId="34" pageNumber="35">Alioramus</emphasis>
spp.
</taxonomicName>
are known thus far. Given that both tyrannosaurids occur in the Nemegt Formation and that
<taxonomicName id="4C2AAAD2994FFFDDBC900476FAF5F858" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[1198,1300,1868,1891]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994FFFDDBC900476FAF5F858" box="[1198,1300,1868,1891]" italics="true" pageId="34" pageNumber="35">B. ostromi</emphasis>
</taxonomicName>
lacks diagnostic features of either
<taxonomicName id="4C2AAAD2994FFFDDBBC80450FB71F8B9" box="[1014,1168,1898,1922]" class="Reptilia" family="Tyrannosauridae" genus="Alioramus" kingdom="Animalia" order="Saurischia" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B95E0D43994FFFDDBBC80450FBBFF8B9" box="[1014,1118,1898,1922]" italics="true" pageId="34" pageNumber="35">Alioramus</emphasis>
spp.
</taxonomicName>
(see
<bibRefCitation id="EFBBACA0994FFFDDBCF50451FA79F8B8" author="Brusatte SL &amp; Carr TD &amp; Norell MA" box="[1227,1432,1898,1923]" pageId="34" pageNumber="35" pagination="1 - 197" refId="ref25467" refString="Brusatte SL, Carr TD, Norell MA. The osteology of Alioramus, a gracile and long-snouted tyrannosaurid (Dinosauria: Theropoda) from the Late Cretaceous of Mongolia. Bulletin of the American Museum of Natural History 2012; 366: 1 - 197. https: // doi. org / 10.1206 / 770.1" year="2012">
Brusatte
<emphasis id="B95E0D43994FFFDDBD140451FABDF8B9" box="[1322,1372,1898,1922]" italics="true" pageId="34" pageNumber="35">et al.</emphasis>
2012
</bibRefCitation>
) or
<emphasis id="B95E0D43994FFFDDBB0704B1FC42F899" box="[825,931,1930,1954]" italics="true" pageId="34" pageNumber="35">Ta. bataar</emphasis>
(see
<bibRefCitation id="EFBBACA0994FFFDDBBE204B0FB03F899" author="Hurum JH &amp; Sabath K" box="[988,1250,1930,1954]" pageId="34" pageNumber="35" pagination="161 - 90" refId="ref27424" refString="Hurum JH, Sabath K. Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared. Acta Palaeontologica Polonica 2003; 48: 161 - 90." year="2003">Hurum and Sabath 2003</bibRefCitation>
), which is mostly attributable to the fragmentary nature of the holotype skeleton, we cannot assign
<emphasis id="B95E0D43994EFFDCB90503AAFE1FFF93" box="[315,510,144,168]" italics="true" pageId="35" pageNumber="36">ZPAL MgD-I/108</emphasis>
to any particular species. Some subtle features suggest that
<taxonomicName id="4C2AAAD2994EFFDCBA29038AFD9EFFFC" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[535,639,176,199]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="35" pageNumber="36" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994EFFDCBA29038AFD9EFFFC" box="[535,639,176,199]" italics="true" pageId="35" pageNumber="36">B. ostromi</emphasis>
</taxonomicName>
might be a juvenile of
<taxonomicName id="4C2AAAD2994EFFDCB8D503F5FEB6FFDC" box="[235,343,207,231]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="35" pageNumber="36" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994EFFDCB8D503F5FEB6FFDC" box="[235,343,207,231]" italics="true" pageId="35" pageNumber="36">Ta. bataar</emphasis>
</taxonomicName>
: e.g. (i) already strongly expanded anterior end of the dentary; (ii) chin well demarcated; and (iii) lack of the pneumatic pocket next to the surangular foramen. However, because those features might potentially be a result of intraspecific variability or be more widespread among juvenile tyrannosaurids than currently suspected, we cannot clearly determine whether
<taxonomicName id="4C2AAAD2994EFFDCB9B502B1FE11FE99" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[395,496,395,418]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="35" pageNumber="36" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994EFFDCB9B502B1FE11FE99" box="[395,496,395,418]" italics="true" pageId="35" pageNumber="36">B. ostromi</emphasis>
</taxonomicName>
is a juvenile of
<taxonomicName id="4C2AAAD2994EFFDCBAAF02B1FD1BFE98" box="[657,762,395,419]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="35" pageNumber="36" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43994EFFDCBAAF02B1FD1BFE98" box="[657,762,395,419]" italics="true" pageId="35" pageNumber="36">Ta. bataar</emphasis>
</taxonomicName>
or
<taxonomicName id="4C2AAAD2994EFFDCB8AC0290FECFFEFA" box="[146,302,426,450]" class="Reptilia" family="Tyrannosauridae" genus="Alioramus" kingdom="Animalia" order="Saurischia" pageId="35" pageNumber="36" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B95E0D43994EFFDCB8AC0290FF1BFEF9" box="[146,250,426,450]" italics="true" pageId="35" pageNumber="36">Alioramus</emphasis>
spp.
</taxonomicName>
Thus, we consider
<taxonomicName id="4C2AAAD2994EFFDCBA3B0291FD8EFEF9" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[517,623,427,450]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="35" pageNumber="36" phylum="Chordata" rank="species" species="ostromi">
<emphasis id="B95E0D43994EFFDCBA3B0291FD8EFEF9" box="[517,623,427,450]" italics="true" pageId="35" pageNumber="36">B. ostromi</emphasis>
</taxonomicName>
to be an indeterminate juvenile representative of the Tyrannosauridae. This assessment might be modified in the future, when more juvenile individuals of tyrannosaurid taxa are known (particularly young individuals of
<taxonomicName id="4C2AAAD2994EFFDCB989011DFDB4FD04" box="[439,597,551,575]" class="Reptilia" family="Tyrannosauridae" genus="Alioramus" kingdom="Animalia" order="Saurischia" pageId="35" pageNumber="36" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B95E0D43994EFFDCB989011DFDFEFD04" box="[439,543,551,575]" italics="true" pageId="35" pageNumber="36">Alioramus</emphasis>
spp.
</taxonomicName>
) and when the growth series and variability at early life stages are better understood.
</paragraph>
</subSubSection>
</treatment>
</document>

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<subSubSection id="C33082DA997AFFE8B95D0395FD54FFF2" box="[355,693,175,201]" pageId="23" pageNumber="24" type="nomenclature">
<paragraph id="8B95D151997AFFE8B95D0395FD54FFF2" blockId="23.[166,710,144,201]" box="[355,693,175,201]" pageId="23" pageNumber="24">
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<emphasis id="B95E0D43997AFFE8B95D0395FD54FFF2" bold="true" box="[355,693,175,201]" pageId="23" pageNumber="24">Caenagnathidae Stenberg, 1940</emphasis>
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<subSubSection id="C33082DA997AFFE8B95B03E3FDE6FFC8" box="[357,519,217,243]" pageId="23" pageNumber="24" type="materials_examined">
<paragraph id="8B95D151997AFFE8B95B03E3FDE6FFC8" blockId="23.[357,519,217,243]" box="[357,519,217,243]" pageId="23" pageNumber="24">
<heading id="D0DD663D997AFFE8B95B03E3FDE6FFC8" box="[357,519,217,243]" centered="true" fontSize="9" level="2" pageId="23" pageNumber="24" reason="2">
<emphasis id="B95E0D43997AFFE8B95B03E3FDE6FFC8" box="[357,519,217,243]" italics="true" pageId="23" pageNumber="24">Referred material</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C33082DA997AFFE8B84F023AFE78FD4B" pageId="23" pageNumber="24" type="diagnosis">
<paragraph id="8B95D151997AFFE8B84F023AFD39FE0C" blockId="23.[113,762,256,311]" pageId="23" pageNumber="24">ZPAL MgD-I/108/1: Left manus phalanx II-1, manus ungual I-2, proximal and distal ends of the left femur, tibiotarsus, and rib.</paragraph>
<paragraph id="8B95D151997AFFE8B911026DFDDCFE4B" blockId="23.[303,573,342,368]" box="[303,573,342,368]" pageId="23" pageNumber="24">
<heading id="D0DD663D997AFFE8B911026DFDDCFE4B" box="[303,573,342,368]" centered="true" fontSize="9" level="2" pageId="23" pageNumber="24" reason="2">
<emphasis id="B95E0D43997AFFE8B911026DFDDCFE4B" box="[303,573,342,368]" italics="true" pageId="23" pageNumber="24">Note on diagnostic characters</emphasis>
</heading>
</paragraph>
<paragraph id="8B95D151997AFFE8B84F0247FE78FD4B" blockId="23.[113,763,381,624]" pageId="23" pageNumber="24">
We provide full details below, because we must first describe all the bones of the
<taxonomicName id="4C2AAAD2997AFFE8B91502A7FE7FFE8F" authorityName="Osmolska" authorityYear="1996" box="[299,414,413,436]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="23" pageNumber="24" phylum="Chordata" rank="genus">
<emphasis id="B95E0D43997AFFE8B91502A7FE7FFE8F" box="[299,414,413,436]" italics="true" pageId="23" pageNumber="24">Bagaraatan</emphasis>
</taxonomicName>
original series before untangling which different taxa they belong to. However, we note here that this set of bones can be referred to
<taxonomicName id="4C2AAAD2997AFFE8B9E502E1FD61FEC8" authorityName="Stenberg" authorityYear="1940" box="[475,640,475,499]" class="Reptilia" family="Caenagnathidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="23" pageNumber="24" phylum="Chordata" rank="family">Caenagnathidae</taxonomicName>
because of: (i) the presence of lateral pleurocoels in the proximal caudal centra; (ii) lesser and greater trochanters in contact; (iii) clearly demarcated accessory trochanter; and (iv) gracile and straight shape of the manual phalanx.
</paragraph>
</subSubSection>
<subSubSection id="C33082DA997AFFE8B95701AAFD1AFDF5" pageId="23" pageNumber="24" type="distribution">
<paragraph id="8B95D151997AFFE8B95701AAFDE3FD91" blockId="23.[113,763,656,723]" box="[361,514,656,682]" pageId="23" pageNumber="24">
<heading id="D0DD663D997AFFE8B95701AAFDE3FD91" box="[361,514,656,682]" centered="true" fontSize="9" level="2" pageId="23" pageNumber="24" reason="2">
<emphasis id="B95E0D43997AFFE8B95701AAFDE3FD91" box="[361,514,656,682]" italics="true" pageId="23" pageNumber="24">Locality and age</emphasis>
</heading>
</paragraph>
<paragraph id="8B95D151997AFFE8B84F018CFD1AFDF5" blockId="23.[113,763,656,723]" box="[113,763,694,723]" pageId="23" pageNumber="24">
Northern Sayr, Nemegt,
<collectingRegion id="49EE1FB3997AFFE8B95E0182FE27FDE8" box="[352,454,696,723]" country="Mongolia" name="Omnogovi" pageId="23" pageNumber="24">Ömnögov</collectingRegion>
,
<collectingCountry id="F33D91C1997AFFE8B9F3018CFDD1FDF5" box="[461,560,694,718]" name="Mongolia" pageId="23" pageNumber="24">Mongolia</collectingCountry>
; Nemegt Formation.
</paragraph>
</subSubSection>
<subSubSection id="C33082DA997AFFE3B9BE01D5FABCFB0E" lastPageId="28" lastPageNumber="29" pageId="23" pageNumber="24" type="description">
<paragraph id="8B95D151997AFFE8B9BE01D5FE0DFC33" blockId="23.[113,763,751,1126]" box="[384,492,751,776]" pageId="23" pageNumber="24">
<emphasis id="B95E0D43997AFFE8B9BE01D5FE0DFC33" box="[384,492,751,776]" italics="true" pageId="23" pageNumber="24">Description</emphasis>
</paragraph>
<paragraph id="8B95D151997AFFE8B84F002EFF57FB5D" blockId="23.[113,763,751,1126]" pageId="23" pageNumber="24">
<emphasis id="B95E0D43997AFFE8B84F002EFEC2FC17" box="[113,291,788,812]" italics="true" pageId="23" pageNumber="24">Caudal vertebrae:</emphasis>
The centrum of one caudal vertebra is preserved (
<figureCitation id="1311CDD4997AFFE8B8FA000EFED5FC77" box="[196,308,820,844]" captionStart="Figure 19" captionStartId="24.[130,195,1427,1451]" captionTargetBox="[129,1473,144,1399]" captionTargetId="figure-251@24.[129,1473,144,1399]" captionTargetPageId="24" captionText="Figure 19. Various bones of Caenagnathidae indet. ZPAL MgD-I/108/1. AF, caudal vertebra in anterior (A), left lateral (B), posterior (C), right lateral (D), dorsal (E), and ventral (F) view. G, H, proximal part of dorsal rib. IN, manus ungual II-3 in dorsal (I), medial (J), ventral (K), lateral (L), anterior (M), and posterior (N) view.OT, left manus phalanx II-1 in medial (O), ventral (P), lateral (Q), dorsal (R), anterior (S), and posterior (T) view.UAʹ, left pedal phalanx II-2 in medial (U), ventral (W), lateral (X), dorsal (Y), anterior (Z), and posterior (Aʹ) view." figureDoi="http://doi.org/10.5281/zenodo.14284269" httpUri="https://zenodo.org/record/14284269/files/figure.png" pageId="23" pageNumber="24">Fig. 19AF</figureCitation>
). It is
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long,
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tall, and
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wide (the height to width ratio of the centrum is 0.8). The centrum is oval, only slightly compressed dorsoventrally. Laterally, the centrum bears one pleurocoel (pneumatic foramen) on each side. The presence of lateral pleurocoels in the caudal vertebrae is a synapomorphy of Caenagnathoidea (
<bibRefCitation id="EFBBACA0997AFFE8BA2700EBFD08FCD3" author="Lamanna MC &amp; Sues HD" box="[537,745,976,1000]" pageId="23" pageNumber="24" refId="ref27733" refString="Lamanna MC, Sues HD, Schachner ER et al. A new large-bodied oviraptorosaurian theropod dinosaur from the Latest Cretaceous of Western North America. PLoS One 2014; 9: e 92022. https: // doi. org / 10.1371 / journal. pone. 0092022" year="2014">
Lamanna
<emphasis id="B95E0D43997AFFE8BABE00EBFD51FCD3" box="[640,688,976,1000]" italics="true" pageId="23" pageNumber="24">et al.</emphasis>
2014
</bibRefCitation>
). The centrum is only slightly concave laterally. Ventrally, two parallel ridges extend along the centrum, as in
<taxonomicName id="4C2AAAD2997AFFE8BA6A0735FD1BFB1C" authorityName="Osmolska" authorityYear="1981" box="[596,762,1039,1063]" class="Reptilia" family="Caenagnathidae" genus="Elmisaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="rarus">
<emphasis id="B95E0D43997AFFE8BA6A0735FD1BFB1C" box="[596,762,1039,1063]" italics="true" pageId="23" pageNumber="24">Elmisaurus rarus</emphasis>
</taxonomicName>
(specimen MPC-D 100/119
<taxonomicName id="4C2AAAD2997AFFE8B99C0715FF44FB5D" authority="' Barsbold et al. 2000" authorityName="Barsbold" authorityYear="2000" class="Reptilia" family="Caenagnathidae" genus="Nomingia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="gobiensis">
<emphasis id="B95E0D43997AFFE8B99C0715FDBEFB7D" box="[418,607,1070,1094]" italics="true" pageId="23" pageNumber="24">Nomingia gobiensis</emphasis>
Barsbold
<emphasis id="B95E0D43997AFFE8BAF20715FD1AFB7D" box="[716,763,1070,1094]" italics="true" pageId="23" pageNumber="24">et al.</emphasis>
2000
</taxonomicName>
).
</paragraph>
<paragraph id="8B95D151997AFFE8B84F07B7FDAFFAE5" blockId="23.[113,763,1165,1502]" pageId="23" pageNumber="24">
<emphasis id="B95E0D43997AFFE8B84F07B7FF42FB9E" box="[113,163,1165,1189]" italics="true" pageId="23" pageNumber="24">Ribs:</emphasis>
Only a proximal part of a dorsal rib is preserved; the rib is broken at the tuberculum (
<figureCitation id="1311CDD4997AFFE8B9910796FDCFFBFF" box="[431,558,1196,1220]" captionStart="Figure 19" captionStartId="24.[130,195,1427,1451]" captionTargetBox="[129,1473,144,1399]" captionTargetId="figure-251@24.[129,1473,144,1399]" captionTargetPageId="24" captionText="Figure 19. Various bones of Caenagnathidae indet. ZPAL MgD-I/108/1. AF, caudal vertebra in anterior (A), left lateral (B), posterior (C), right lateral (D), dorsal (E), and ventral (F) view. G, H, proximal part of dorsal rib. IN, manus ungual II-3 in dorsal (I), medial (J), ventral (K), lateral (L), anterior (M), and posterior (N) view.OT, left manus phalanx II-1 in medial (O), ventral (P), lateral (Q), dorsal (R), anterior (S), and posterior (T) view.UAʹ, left pedal phalanx II-2 in medial (U), ventral (W), lateral (X), dorsal (Y), anterior (Z), and posterior (Aʹ) view." figureDoi="http://doi.org/10.5281/zenodo.14284269" httpUri="https://zenodo.org/record/14284269/files/figure.png" pageId="23" pageNumber="24">Fig. 19G, H</figureCitation>
). The capitulum is bulbous. Behind the slightly convex articular surface, no depression is present, in contrast to tyrannosaurids (
<taxonomicName id="4C2AAAD2997AFFE8BA5E07D6FD2BFA38" box="[608,714,1259,1283]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43997AFFE8BA5E07D6FD2BFA38" box="[608,714,1259,1283]" italics="true" pageId="23" pageNumber="24">Ta. bataar</emphasis>
</taxonomicName>
, e.g. ZPAL MgD-I/3, ZPAL MgD-I/4, and ZPAL MgD-I/175, and
<taxonomicName id="4C2AAAD2997AFFE8B84F0613FEE9FA7A" box="[113,264,1321,1345]" class="Reptilia" family="Tyrannosauridae" genus="Alioramus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="altai">
<emphasis id="B95E0D43997AFFE8B84F0613FEE9FA7A" box="[113,264,1321,1345]" italics="true" pageId="23" pageNumber="24">Alioramus altai</emphasis>
</taxonomicName>
; see
<bibRefCitation id="EFBBACA0997AFFE8B9050610FE1CFA79" author="Brusatte SL &amp; Carr TD &amp; Norell MA" box="[315,509,1321,1346]" pageId="23" pageNumber="24" pagination="1 - 197" refId="ref25467" refString="Brusatte SL, Carr TD, Norell MA. The osteology of Alioramus, a gracile and long-snouted tyrannosaurid (Dinosauria: Theropoda) from the Late Cretaceous of Mongolia. Bulletin of the American Museum of Natural History 2012; 366: 1 - 197. https: // doi. org / 10.1206 / 770.1" year="2012">
Brusatte
<emphasis id="B95E0D43997AFFE8B9AB0610FE25FA7A" box="[405,452,1321,1345]" italics="true" pageId="23" pageNumber="24">et al.</emphasis>
2012
</bibRefCitation>
). Also, in contrast to the latter, the tuberculum is enlarged. Because the capitulum and tuberculum are at a similar level, the rib is likely to come from the posterior part of the ribcage. The overall shape of the preserved part of the rib corresponds to the morphology in caenagnathids (e.g.
<taxonomicName id="4C2AAAD2997AFFE8B89C06FCFEA6FAE5" authorityName="Stenberg" authorityYear="1940" box="[162,327,1478,1502]" class="Reptilia" family="Caenagnathidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="23" pageNumber="24" phylum="Chordata" rank="family">Caenagnathidae</taxonomicName>
indet. ZPAL MgD-I/99).
</paragraph>
<paragraph id="8B95D151997AFFE8B84F0533FD92F801" blockId="23.[113,763,1544,1976]" pageId="23" pageNumber="24">
<emphasis id="B95E0D43997AFFE8B84F0533FEA0F91B" box="[113,321,1544,1568]" italics="true" pageId="23" pageNumber="24">Manus phalanx II-1:</emphasis>
The left phalanx is straight and elongated, measuring
<quantity id="4CD27CB4997AFFE8B8DD051DFEA0F904" box="[227,321,1575,1599]" metricMagnitude="-2" metricUnit="m" metricValue="7.659999999999999" pageId="23" pageNumber="24" unit="mm" value="76.6">76.6 mm</quantity>
(
<figureCitation id="1311CDD4997AFFE8B96D051DFE2CF97B" box="[339,461,1575,1600]" captionStart="Figure 19" captionStartId="24.[130,195,1427,1451]" captionTargetBox="[129,1473,144,1399]" captionTargetId="figure-251@24.[129,1473,144,1399]" captionTargetPageId="24" captionText="Figure 19. Various bones of Caenagnathidae indet. ZPAL MgD-I/108/1. AF, caudal vertebra in anterior (A), left lateral (B), posterior (C), right lateral (D), dorsal (E), and ventral (F) view. G, H, proximal part of dorsal rib. IN, manus ungual II-3 in dorsal (I), medial (J), ventral (K), lateral (L), anterior (M), and posterior (N) view.OT, left manus phalanx II-1 in medial (O), ventral (P), lateral (Q), dorsal (R), anterior (S), and posterior (T) view.UAʹ, left pedal phalanx II-2 in medial (U), ventral (W), lateral (X), dorsal (Y), anterior (Z), and posterior (Aʹ) view." figureDoi="http://doi.org/10.5281/zenodo.14284269" httpUri="https://zenodo.org/record/14284269/files/figure.png" pageId="23" pageNumber="24">Fig. 19OT</figureCitation>
). The proximal articular surface is taller (
<quantity id="4CD27CB4997AFFE8B8C7057DFEA6F964" box="[249,327,1607,1631]" metricMagnitude="-2" metricUnit="m" metricValue="1.8" pageId="23" pageNumber="24" unit="mm" value="18.0">18 mm</quantity>
) than wide (
<quantity id="4CD27CB4997AFFE8B9F7057DFDF6F964" box="[457,535,1607,1631]" metricMagnitude="-2" metricUnit="m" metricValue="1.6" pageId="23" pageNumber="24" unit="mm" value="16.0">16 mm</quantity>
) and divided by a low ridge, which is narrow dorsally and wide ventrally. On both sides of the ridge, the articular surfaces are teardrop-shaped and strongly concave. The distal medial condyle (
<quantity id="4CD27CB4997AFFE8BA7A059FFD45F987" box="[580,676,1701,1725]" metricMagnitude="-2" metricUnit="m" metricValue="1.35" pageId="23" pageNumber="24" unit="mm" value="13.5">13.5 mm</quantity>
high) is smaller than the lateral one (
<quantity id="4CD27CB4997AFFE8B9A605FEFE16F9E7" box="[408,503,1732,1756]" metricMagnitude="-2" metricUnit="m" metricValue="1.54" pageId="23" pageNumber="24" unit="mm" value="15.4">15.4 mm</quantity>
high) and separated by a deep and narrow furrow. The medial ligament pit is shallower than the lateral ligament pit. The width of the distal end is
<quantity id="4CD27CB4997AFFE8BADE0439FF7CF801" metricMagnitude="-2" metricUnit="m" metricValue="1.5" pageId="23" pageNumber="24" unit="mm" value="15.0">15 mm</quantity>
; the length to width ratio of the phalanx is 4.7.
</paragraph>
<paragraph id="8B95D151997AFFE8B8B30478FAA5FD04" blockId="23.[113,763,1544,1976]" lastBlockId="23.[810,1459,144,575]" pageId="23" pageNumber="24">
The gracile and straight shape of the manus phalanx II-1 of ZPAL MgD-I/108/1 is the same as in
<taxonomicName id="4C2AAAD2997AFFE8BA32045BFD1AF842" authority="ZPAL" authorityName="ZPAL" box="[524,763,1889,1913]" class="Reptilia" family="Caenagnathidae" genus="Elmisaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="rarus">
<emphasis id="B95E0D43997AFFE8BA32045BFD54F843" box="[524,693,1889,1913]" italics="true" pageId="23" pageNumber="24">Elmisaurus rarus</emphasis>
ZPAL
</taxonomicName>
MgD-I/98, although the phalanx of ZPAL MgD-I/108/1 is larger. The length of manus phalanx II-1 of ZPAL MgD-I/98 is 66 mm, the proximal width 14 mm, and the distal width 12 mm (the length to width ratio is 4.7, same as for ZPAL MgDI/108/1). The manus phalanx II-1 of ZPAL MgD-I/108/1 shares also with
<taxonomicName id="4C2AAAD2997AFFE8BBDA03D4FB71FE3E" box="[996,1168,238,262]" class="Reptilia" family="Caenagnathidae" genus="Elmisaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="rarus">
<emphasis id="B95E0D43997AFFE8BBDA03D4FB71FE3E" box="[996,1168,238,262]" italics="true" pageId="23" pageNumber="24">Elmisaurus rarus</emphasis>
</taxonomicName>
slightly downturned distal condyles and expanded articular surfaces of the distal condyles. Other theropods known from the Nemegt Formation, i.e. tyrannosaurids (
<taxonomicName id="4C2AAAD2997AFFE8BBC40276FB83FE5F" box="[1018,1122,332,356]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43997AFFE8BBC40276FB83FE5F" box="[1018,1122,332,356]" italics="true" pageId="23" pageNumber="24">Ta. bataar</emphasis>
</taxonomicName>
, e.g. ZPAL MgD-I/3 and ZPAL MgD-I/4), ornithomimids (
<taxonomicName id="4C2AAAD2997AFFE8BC580251FB0BFEB8" box="[1126,1258,363,387]" class="Reptilia" family="Ornithomimidae" genus="Gallimimus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="bullatus">
<emphasis id="B95E0D43997AFFE8BC580251FB0BFEB8" box="[1126,1258,363,387]" italics="true" pageId="23" pageNumber="24">Ga. bullatus</emphasis>
</taxonomicName>
, cast of
<emphasis id="B95E0D43997AFFE8BD5D0251FC96FE99" italics="true" pageId="23" pageNumber="24">MPC-D 100/11</emphasis>
;
<taxonomicName id="4C2AAAD2997AFFE8BBB702B0FB34FE99" authority="Osmolska" authorityName="Osmolska" box="[905,1237,394,418]" class="Reptilia" family="Deinocheiridae" genus="Deinocheirus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="mirificus">
<emphasis id="B95E0D43997AFFE8BBB702B0FB85FE99" box="[905,1124,394,418]" italics="true" pageId="23" pageNumber="24">Deinocheirus mirificus</emphasis>
Osmólska
</taxonomicName>
&amp; Roniewicz, 1970, cast of
<emphasis id="B95E0D43997AFFE8BB4F0290FBF5FEF9" box="[881,1044,426,450]" italics="true" pageId="23" pageNumber="24">MPC-D 100/18</emphasis>
), avimimids [alvarezsaurids (
<taxonomicName id="4C2AAAD2997AFFE8BD7C0290FBD2FEDA" authority="Perle et al., 1993" authorityName="Perle" authorityYear="1993" class="Reptilia" family="Alvarezsauridae" genus="Mononykus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="olecranus">
<emphasis id="B95E0D43997AFFE8BD7C0290FC65FEDA" italics="true" pageId="23" pageNumber="24">Mononykus olecranus</emphasis>
Perle
<emphasis id="B95E0D43997AFFE8BBFA02F0FC15FEDA" box="[964,1012,457,481]" italics="true" pageId="23" pageNumber="24">et al.</emphasis>
, 1993
</taxonomicName>
(Perle
<emphasis id="B95E0D43997AFFE8BC4102F0FB4FFEDA" box="[1151,1198,457,481]" italics="true" pageId="23" pageNumber="24">et al.</emphasis>
1994)], and oviraptorids [e.g.
<taxonomicName id="4C2AAAD2997AFFE8BB5A02D2FAF7FD3B" authority="(Funston et al. 2020 a)" baseAuthorityName="Funston" baseAuthorityYear="2020" box="[868,1302,488,512]" family="Oviraptoridae" genus="Oksoko" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="avarsan">
<emphasis id="B95E0D43997AFFE8BB5A02D2FBE8FD3B" box="[868,1033,488,512]" italics="true" pageId="23" pageNumber="24">Oksoko avarsan</emphasis>
(
<bibRefCitation id="EFBBACA0997AFFE8BC1D02D3FAEDFD3B" author="Funston GF &amp; Tsogtbaatar C &amp; Tsogtbaatar K" box="[1059,1292,488,512]" pageId="23" pageNumber="24" refId="ref27006" refString="Funston GF, Tsogtbaatar C, Tsogtbaatar K et al. A new two-fingered dinosaur sheds light on the radiation of Oviraptorosauria. Royal Society Open Science 2020 a; 7: 201184." year="2020">
Funston
<emphasis id="B95E0D43997AFFE8BCB902D3FB5EFD3B" box="[1159,1215,488,512]" italics="true" pageId="23" pageNumber="24">et al.</emphasis>
2020a
</bibRefCitation>
)
</taxonomicName>
,
<taxonomicName id="4C2AAAD2997AFFE8BD1502D3FBFBFD1B" authority="Lu et al., 2005" authorityName="Lu" authorityYear="2005" class="Reptilia" family="Oviraptoridae" genus="Nemegtomaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="barsboldi">
<emphasis id="B95E0D43997AFFE8BD1502D3FC65FD1B" italics="true" pageId="23" pageNumber="24">Nemegtomaia barsboldi</emphasis>
<bibRefCitation id="EFBBACA0997AFFE8BBB40132FBFBFD1B" author="Lu J &amp; Tomida Y &amp; Azuma Y" box="[906,1050,520,544]" pageId="23" pageNumber="24" refId="ref27973" refString="Lu J, Tomida Y, Azuma Y et al. Nemegtomaia gen. nov., a replacement name for the oviraptorosaurian dinosaur Nemegtia Lu et al., 2004, a preoccupied name. Bulletin of the National Science Museum &amp; Tokyo &amp; Series C 2005; 31: 51." year="2005">
Lu
<emphasis id="B95E0D43997AFFE8BB920133FC3AFD1B" box="[940,987,520,544]" italics="true" pageId="23" pageNumber="24">et al.</emphasis>
, 2005
</bibRefCitation>
</taxonomicName>
(Fanti
<emphasis id="B95E0D43997AFFE8BC5B0133FB75FD1B" box="[1125,1172,520,544]" italics="true" pageId="23" pageNumber="24">et al.</emphasis>
2012)] do not have manual phalanges that are so straight, slender, and elongated.
</paragraph>
<paragraph id="8B95D151997AFFE8BB14015BFA7AFB2B" blockId="23.[810,1460,608,1666]" pageId="23" pageNumber="24">
<emphasis id="B95E0D43997AFFE8BB14015BFC0AFD42" box="[810,1003,609,633]" italics="true" pageId="23" pageNumber="24">Manus ungual II-3:</emphasis>
The ungual is elongated (
<quantity id="4CD27CB4997AFFE8BCCB015AFAA0FD43" box="[1269,1345,608,632]" metricMagnitude="-2" metricUnit="m" metricValue="5.4" pageId="23" pageNumber="24" unit="mm" value="54.0">54 mm</quantity>
in length), curved, and very narrow, and the proximal articular surface is
<quantity id="4CD27CB4997AFFE8BB1401A5FC95FD8C" box="[810,884,671,695]" metricMagnitude="-2" metricUnit="m" metricValue="1.3" pageId="23" pageNumber="24" unit="mm" value="13.0">13 mm</quantity>
wide (
<figureCitation id="1311CDD4997AFFE8BB8001A5FBCFFD8C" box="[958,1070,671,695]" captionStart="Figure 19" captionStartId="24.[130,195,1427,1451]" captionTargetBox="[129,1473,144,1399]" captionTargetId="figure-251@24.[129,1473,144,1399]" captionTargetPageId="24" captionText="Figure 19. Various bones of Caenagnathidae indet. ZPAL MgD-I/108/1. AF, caudal vertebra in anterior (A), left lateral (B), posterior (C), right lateral (D), dorsal (E), and ventral (F) view. G, H, proximal part of dorsal rib. IN, manus ungual II-3 in dorsal (I), medial (J), ventral (K), lateral (L), anterior (M), and posterior (N) view.OT, left manus phalanx II-1 in medial (O), ventral (P), lateral (Q), dorsal (R), anterior (S), and posterior (T) view.UAʹ, left pedal phalanx II-2 in medial (U), ventral (W), lateral (X), dorsal (Y), anterior (Z), and posterior (Aʹ) view." figureDoi="http://doi.org/10.5281/zenodo.14284269" httpUri="https://zenodo.org/record/14284269/files/figure.png" pageId="23" pageNumber="24">Fig. 19IN</figureCitation>
). The ungual lacks only the distal tip. The proximal articular surface is oval (longer dorsoventrally than mediolaterally). A vertical ridge, which is dorsally and ventrally expanded but constricted in the middle section, extends across the middle of the articular surface. The articular surfaces on both sides of the ridge are strongly concave. The dorsal edge of the articular surface forms a robust dorsal lip, surrounded by a depression. A ventral process is present on the ventral edge of the articular surface. The articular surface is separated by a notch from the ventrodistally located enlarged flexor tubercle. Laterally and medially, the collateral groove extends along the entire ungual, starting from the area above the flexor tubercle.
</paragraph>
<paragraph id="8B95D151997AFFE8BB7B072DFC6FF9B9" blockId="23.[810,1460,608,1666]" pageId="23" pageNumber="24">
The manus ungual II-3 is not known in
<taxonomicName id="4C2AAAD2997AFFE8BCF1072DFA95FB14" box="[1231,1396,1047,1071]" class="Reptilia" family="Caenagnathidae" genus="Elmisaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="rarus">
<emphasis id="B95E0D43997AFFE8BCF1072DFA95FB14" box="[1231,1396,1047,1071]" italics="true" pageId="23" pageNumber="24">Elmisaurus rarus</emphasis>
</taxonomicName>
; however, the presence of the distinctive dorsal lip indicates that the ungual corresponds to the manual unguals of
<taxonomicName id="4C2AAAD2997AFFE8BD37076CFA51FB55" authorityName="Stenberg" authorityYear="1940" box="[1289,1456,1110,1134]" class="Reptilia" family="Caenagnathidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="23" pageNumber="24" phylum="Chordata" rank="family">Caenagnathidae</taxonomicName>
. In comparison to the manual unguals of an North American caenagnathid,
<taxonomicName id="4C2AAAD2997AFFE8BBFA07AEFA84FB96" authority="Gilmore, 1924" authorityName="Gilmore" authorityYear="1924" box="[964,1381,1172,1197]" class="Reptilia" family="Caenagnathidae" genus="Chirostenotes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="pergracillis">
<emphasis id="B95E0D43997AFFE8BBFA07AEFB5DFB97" box="[964,1212,1172,1196]" italics="true" pageId="23" pageNumber="24">Chirostenotes pergracillis</emphasis>
<bibRefCitation id="EFBBACA0997AFFE8BCF707AEFA84FB96" author="Gilmore CW" box="[1225,1381,1172,1197]" pageId="23" pageNumber="24" pagination="1 - 12" refId="ref27186" refString="Gilmore CW. A new coelurid dinosaur from the Belly River Cretaceous of Alberta. Canada Department of Mines Geological Survey Bulletin (Geological Series) 1924; 38: 1 - 12." year="1924">Gilmore, 1924</bibRefCitation>
</taxonomicName>
,
<emphasis id="B95E0D43997AFFE8BD4607AFFCBFFBF7" italics="true" pageId="23" pageNumber="24">CMN 2367</emphasis>
(
<bibRefCitation id="EFBBACA0997AFFE8BB4D078EFBE6FBF7" author="Funston GF" box="[883,1031,1204,1228]" pageId="23" pageNumber="24" pagination="105 - 53" refId="ref26776" refString="Funston GF. Caenagnathids of the Dinosaur Park Formation (Campanian) of Alberta, Canada: anatomy, osteo-histology, taxonomy, and evolution. Vertebrate Anatomy Morphology Palaeontology 2020; 8: 105 - 53. https: // doi. org / 10.18435 / vamp 29362" year="2020">Funston 2020</bibRefCitation>
), the ungual of ZPAL MgD-I/108/1 is less curved than the phalanges I-2 and III-4, but more straight, similar to II-3. Moreover, the proximal articulation is offset, and the flexor tubercle is distally positioned and smaller in contrast to unguals I-2 and III-4, which further supports its identification as II-3 of a caenagnathid. Other theropods known from the Nemegt Formation, i.e. tyrannosaurids (
<taxonomicName id="4C2AAAD2997AFFE8BD7D064AFA4CFAB3" box="[1347,1453,1392,1416]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D43997AFFE8BD7D064AFA4CFAB3" box="[1347,1453,1392,1416]" italics="true" pageId="23" pageNumber="24">Ta. bataar</emphasis>
</taxonomicName>
, e.g. ZPAL MgD-I/3 and ZPAL MgD-I/4), ornithomimids (
<taxonomicName id="4C2AAAD2997AFFE8BDB106B5FC99FAFD" class="Reptilia" family="Ornithomimidae" genus="Gallimimus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="bullatus">
<emphasis id="B95E0D43997AFFE8BDB106B5FC99FAFD" italics="true" pageId="23" pageNumber="24">Ga. bullatus</emphasis>
</taxonomicName>
, cast of
<emphasis id="B95E0D43997AFFE8BBEA0695FB9BFAFC" box="[980,1146,1455,1479]" italics="true" pageId="23" pageNumber="24">MPC-D 100/11</emphasis>
;
<emphasis id="B95E0D43997AFFE8BCB40695FAE6FAFD" box="[1162,1287,1454,1478]" italics="true" pageId="23" pageNumber="24">
De.
<taxonomicName id="4C2AAAD2997AFFE8BC8A0694FAE6FAFD" box="[1204,1287,1454,1478]" class="Reptilia" family="Deinocheiridae" genus="Deinocheirus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="mirificus">mirificus</taxonomicName>
</emphasis>
, cast of
<emphasis id="B95E0D43997AFFE8BD5D0695FC99FADD" italics="true" pageId="23" pageNumber="24">MPC-D 100/18</emphasis>
), alvarezsaurids [
<taxonomicName id="4C2AAAD2997AFFE8BC0F06F4FA94FADD" authority="(Perle et al. 1994)" baseAuthorityName="Perle" baseAuthorityYear="1994" box="[1073,1397,1486,1510]" class="Reptilia" family="Alvarezsauridae" genus="Mononykus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="olecranus">
<emphasis id="B95E0D43997AFFE8BC0F06F4FB4EFADD" box="[1073,1199,1486,1510]" italics="true" pageId="23" pageNumber="24">M. olecranus</emphasis>
(Perle
<emphasis id="B95E0D43997AFFE8BCC306F4FACFFADD" box="[1277,1326,1486,1510]" italics="true" pageId="23" pageNumber="24">et al.</emphasis>
1994)
</taxonomicName>
], and oviraptorids [
<taxonomicName id="4C2AAAD2997AFFE8BB8906D7FAEBF93E" authority="(Funston et al. 2020 a)" baseAuthorityName="Funston" baseAuthorityYear="2020" box="[951,1290,1517,1541]" family="Oviraptoridae" genus="Oksoko" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="avarsan">
<emphasis id="B95E0D43997AFFE8BB8906D7FBC1F93E" box="[951,1056,1517,1541]" italics="true" pageId="23" pageNumber="24">O. avarsan</emphasis>
(
<bibRefCitation id="EFBBACA0997AFFE8BC0F06D4FB1EF93E" author="Funston GF &amp; Tsogtbaatar C &amp; Tsogtbaatar K" box="[1073,1279,1517,1541]" pageId="23" pageNumber="24" refId="ref27006" refString="Funston GF, Tsogtbaatar C, Tsogtbaatar K et al. A new two-fingered dinosaur sheds light on the radiation of Oviraptorosauria. Royal Society Open Science 2020 a; 7: 201184." year="2020">
Funston
<emphasis id="B95E0D43997AFFE8BCB206D4FB5DF93E" box="[1164,1212,1517,1541]" italics="true" pageId="23" pageNumber="24">et al.</emphasis>
2020a
</bibRefCitation>
)
</taxonomicName>
and
<taxonomicName id="4C2AAAD2997AFFE8BD0206D4FC3EF91E" authority="(Fanti et al. 2012)" baseAuthorityName="Fanti" baseAuthorityYear="2012" class="Reptilia" family="Oviraptoridae" genus="Nemegtomaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="barsboldi">
<emphasis id="B95E0D43997AFFE8BD0206D4FA52F93E" box="[1340,1459,1517,1541]" italics="true" pageId="23" pageNumber="24">N. barsboldi</emphasis>
(Fanti
<emphasis id="B95E0D43997AFFE8BB500537FC7DF91F" box="[878,924,1548,1572]" italics="true" pageId="23" pageNumber="24">et al.</emphasis>
2012)
</taxonomicName>
] do not have such enlarged, curved, and transversely narrow manual unguals with an enlarged flexor tubercle distinctly separated from the ventral process and distinctive dorsal lip.
</paragraph>
<paragraph id="8B95D151997AFFE8BB14059EFA61F863" blockId="23.[810,1460,1700,1975]" pageId="23" pageNumber="24">
<emphasis id="B95E0D43997AFFE8BB14059EFC91F980" box="[810,880,1700,1723]" italics="true" pageId="23" pageNumber="24">Femur:</emphasis>
Two parts of the left femur are preserved: the proximal and distal end; most of the shaft is missing, hence the length of the femur is unknown (
<figureCitation id="1311CDD4997AFFE8BC2005D8FB85F9C1" box="[1054,1124,1762,1786]" captionStart="Figure 20" captionStartId="26.[129,194,1836,1860]" captionTargetBox="[175,1427,146,1806]" captionTargetId="figure-6@26.[173,1429,144,1808]" captionTargetPageId="26" captionText="Figure 20. Left femur of Caenagnathidae indet. ZPAL MgD-I/108/1. AE, proximal end of the femur in anterior (A), lateral (B), posterior (C), medial (D), and dorsal (E) view.FJ, distal end of the femur in anterior (F), lateral (G), posterior (H), medial (I), and ventral (J) view.K, L, thin section of the femoral cortex under polarized light. Red arrows indicate secondary osteons; green arrows point to resorption cavities; yellow arrows indicate lamellar bone; and purple arrows point to parallel-fibred bone." figureDoi="http://doi.org/10.5281/zenodo.14284271" httpUri="https://zenodo.org/record/14284271/files/figure.png" pageId="23" pageNumber="24">Fig. 20</figureCitation>
). The circumference of the shaft portions preserved with the distal and proximal parts is
<quantity id="4CD27CB4997AFFE8BD67043BFA51F822" box="[1369,1456,1793,1817]" metricMagnitude="-1" metricUnit="m" metricValue="1.05" pageId="23" pageNumber="24" unit="mm" value="105.0">105 mm</quantity>
. Osmólska (1996) hypothesized that ~
<quantity id="4CD27CB4997AFFE8BC8C041BFAC5F803" box="[1202,1316,1825,1849]" metricMagnitude="-2" metricUnit="m" metricValue="8.5" metricValueMax="9.0" metricValueMin="8.0" pageId="23" pageNumber="24" unit="mm" value="85.0" valueMax="90.0" valueMin="80.0">8090 mm</quantity>
of the shaft is missing, adding up to a total femur length of
<quantity id="4CD27CB4997AFFE8BCCE047AFA9DF863" box="[1264,1404,1856,1880]" metricMagnitude="-1" metricUnit="m" metricValue="3.15" metricValueMax="3.2" metricValueMin="3.1" pageId="23" pageNumber="24" unit="mm" value="315.0" valueMax="320.0" valueMin="310.0">310320 mm</quantity>
.
</paragraph>
<paragraph id="8B95D151997AFFE6BB7B045AFEF2FFFC" blockId="23.[810,1460,1700,1975]" lastBlockId="25.[113,763,144,1734]" lastPageId="25" lastPageNumber="26" pageId="23" pageNumber="24">
The proximal part of the femur (
<figureCitation id="1311CDD4997AFFE8BC91045AFAC7F843" box="[1199,1318,1888,1912]" captionStart="Figure 20" captionStartId="26.[129,194,1836,1860]" captionTargetBox="[175,1427,146,1806]" captionTargetId="figure-6@26.[173,1429,144,1808]" captionTargetPageId="26" captionText="Figure 20. Left femur of Caenagnathidae indet. ZPAL MgD-I/108/1. AE, proximal end of the femur in anterior (A), lateral (B), posterior (C), medial (D), and dorsal (E) view.FJ, distal end of the femur in anterior (F), lateral (G), posterior (H), medial (I), and ventral (J) view.K, L, thin section of the femoral cortex under polarized light. Red arrows indicate secondary osteons; green arrows point to resorption cavities; yellow arrows indicate lamellar bone; and purple arrows point to parallel-fibred bone." figureDoi="http://doi.org/10.5281/zenodo.14284271" httpUri="https://zenodo.org/record/14284271/files/figure.png" pageId="23" pageNumber="24">Fig. 20AE</figureCitation>
) is narrower lateromedially than longer anteroposteriorly. In dorsal view, the femur is L-shaped. The posterior part of the greater trochanter is connected to the femoral head that projects mediodistally, and the anterior part of the greater trochanter is widened anteriorly. In anterior view, the femoral head is positioned higher than the greater trochanter, and they are separated by a broad, shallow depression. The surface of the rounded femoral head is rugose. In posterior view, a wide groove for the capital ligament is present on the femoral head. In medial view, the femoral head is ovoid, and its posterodorsal margin is wider than the anteroventral end. The neck is narrower anteroposteriorly than the head; and the ventral margin of the head is directed downwards before connecting to the neck. The neck extends upwards from the greater trochanter, which is wider lateromedially than the lesser trochanter. The lesser trochanter is almond-shaped in anterior view. The dorsal margin of the femoral trochanters in lateral view is arched; the small, anteriorly positioned lesser trochanter is separated by a shallow groove from the much anteroposteriorly longer greater trochanter. On the lateral surface of the proximal part of the femur, the separation between the lesser and greater trochanter is marked by a shallow and short groove. Below the lesser trochanter, the accessory trochanter (anterior crest
<emphasis id="B95E0D439975FFE7BDB1041DFA23F805" box="[1423,1474,1831,1854]" italics="true" pageId="24" pageNumber="25">sensu</emphasis>
Osmólska 1996) is present. It is slightly expanded anteriorly and extends along the preserved part of the proximal shaft. The accessory trochanter keeps a consistent lateromedial width along the preserved proximal part of the shaft. A posterior tubercle is present below the greater trochanter, well visible in anterior and posterior views.
</paragraph>
<caption id="DF5581D99975FFE7B8BC06A9FEDCF920" ID-DOI="http://doi.org/10.5281/zenodo.14284269" ID-Zenodo-Dep="14284269" httpUri="https://zenodo.org/record/14284269/files/figure.png" pageId="24" pageNumber="25" startId="24.[130,195,1427,1451]" targetBox="[129,1473,144,1399]" targetPageId="24" targetType="figure">
<paragraph id="8B95D1519975FFE7B8BC06A9FEDCF920" blockId="24.[129,1451,1427,1563]" pageId="24" pageNumber="25">
<emphasis id="B95E0D439975FFE7B8BC06A9FF06FA90" bold="true" box="[130,231,1427,1451]" pageId="24" pageNumber="25">Figure 19.</emphasis>
Various bones of
<taxonomicName id="4C2AAAD29975FFE7B9AE06A9FDCBFA90" authorityName="Stenberg" authorityYear="1940" box="[400,554,1427,1451]" class="Reptilia" family="Caenagnathidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="24" pageNumber="25" phylum="Chordata" rank="family">Caenagnathidae</taxonomicName>
indet. ZPAL MgD-I/108/1. AF, caudal vertebra in anterior (A), left lateral (B), posterior (C), right lateral (D), dorsal (E), and ventral (F) view. G, H, proximal part of dorsal rib. IN, manus ungual II-
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dorsal (I), medial (J), ventral (K), lateral (L), anterior (M), and posterior (N) view. OT, left manus phalanx II-
<quantity id="4CD27CB49975FFE7BBE906F1FC16FAD8" box="[983,1015,1483,1507]" metricMagnitude="-2" metricUnit="m" metricValue="2.54" pageId="24" pageNumber="25" unit="in" value="1.0">1 in</quantity>
medial (O), ventral (P), lateral (Q), dorsal (R), anterior (S), and posterior (T) view. UA
<emphasis id="B95E0D439975FFE7BA0206DCFDA0FAC4" box="[572,577,1510,1535]" italics="true" pageId="24" pageNumber="25">ʹ</emphasis>
, left pedal phalanx II-
<quantity id="4CD27CB49975FFE7BB2E06DDFCD5FAC4" box="[784,820,1511,1535]" metricMagnitude="-2" metricUnit="m" metricValue="5.08" pageId="24" pageNumber="25" unit="in" value="2.0">2 in</quantity>
medial (U), ventral (W), lateral (X), dorsal (Y), anterior (Z), and posterior (A
<emphasis id="B95E0D439975FFE7B8C90538FF1DF920" box="[247,252,1538,1563]" italics="true" pageId="24" pageNumber="25">ʹ</emphasis>
) view.
</paragraph>
</caption>
<paragraph id="8B95D1519974FFE6B8B303F5FD1AFD45" blockId="25.[113,763,144,1734]" pageId="25" pageNumber="26">
The distal end of the femur is now longitudinally shorter than described by Osmólska (1996), because it has since been thin sectioned. At that time, it measured
<quantity id="4CD27CB49974FFE6B9C20237FDB8FE1E" box="[508,601,269,293]" metricMagnitude="-1" metricUnit="m" metricValue="1.05" pageId="25" pageNumber="26" unit="mm" value="105.0">105 mm</quantity>
; now, only the distalmost part of the femur including both condyles is present, measuring
<quantity id="4CD27CB49974FFE6B8DB0276FED1FE58" box="[229,304,332,356]" metricMagnitude="-2" metricUnit="m" metricValue="5.2" pageId="25" pageNumber="26" unit="mm" value="52.0">52 mm</quantity>
(
<figureCitation id="1311CDD49974FFE6B97A0276FE50FE5F" box="[324,433,332,356]" captionStart="Figure 20" captionStartId="26.[129,194,1836,1860]" captionTargetBox="[175,1427,146,1806]" captionTargetId="figure-6@26.[173,1429,144,1808]" captionTargetPageId="26" captionText="Figure 20. Left femur of Caenagnathidae indet. ZPAL MgD-I/108/1. AE, proximal end of the femur in anterior (A), lateral (B), posterior (C), medial (D), and dorsal (E) view.FJ, distal end of the femur in anterior (F), lateral (G), posterior (H), medial (I), and ventral (J) view.K, L, thin section of the femoral cortex under polarized light. Red arrows indicate secondary osteons; green arrows point to resorption cavities; yellow arrows indicate lamellar bone; and purple arrows point to parallel-fibred bone." figureDoi="http://doi.org/10.5281/zenodo.14284271" httpUri="https://zenodo.org/record/14284271/files/figure.png" pageId="25" pageNumber="26">Fig. 20FJ</figureCitation>
). The medial condyle is bigger than the lateral condyle, but the lateral condyle extends further distally than the medial condyle. The condyles are distally separated by a deep but narrow notch (the popliteal fossa). Anteriorly and distally, the condyles are separated by shallower and wider depressions (the extensor grooves). The medial condyle is convex, with a slightly rugose surface. The lateral condyle bears an elevation on its distal surface. The tibiofibular crest extends posteromedially. In lateral view, the tibiofibular crest is axeshaped and projects further posteriorly than the medial condyle.
</paragraph>
<paragraph id="8B95D1519974FFE6B8B301BFFF3DF9FD" blockId="25.[113,763,144,1734]" pageId="25" pageNumber="26">
The accessory trochanter appeared in Tetanurae as a branch of the distal base of the lesser trochanter, and it was reduced in Eumaniraptora. The accessory trochanter is smaller in basal Tetanurae, Carnosauria, basal Coelurosauria, Tyrannosauridae, and
<taxonomicName id="4C2AAAD29974FFE6B8A00038FE8DFC21" baseAuthorityName="Barsbold" baseAuthorityYear="1976" box="[158,364,770,794]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithomimosauria" pageId="25" pageNumber="26" phylum="Chordata" rank="order">Ornithomimosauria</taxonomicName>
than in
<taxonomicName id="4C2AAAD29974FFE6B9FD0038FD8FFC21" box="[451,622,770,794]" class="Reptilia" family="Caudipterygidae" genus="Caudipteryx" kingdom="Animalia" order="Dinosauria" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B95E0D439974FFE6B9FD0038FDDCFC21" box="[451,573,770,794]" italics="true" pageId="25" pageNumber="26">Caudipteryx</emphasis>
spp.
</taxonomicName>
,
<taxonomicName id="4C2AAAD29974FFE6BA460039FED9FC01" authority="Ostrom, 1970" authorityName="Ostrom" authorityYear="1970" class="Reptilia" family="Caenagnathidae" genus="Microvenator" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="celer">
<emphasis id="B95E0D439974FFE6BA460039FF7DFC01" italics="true" pageId="25" pageNumber="26">Microvenator celer</emphasis>
Ostrom, 1970
</taxonomicName>
,
<taxonomicName id="4C2AAAD29974FFE6B9790018FE0CFC01" authorityName="Stenberg" authorityYear="1940" box="[327,493,802,826]" class="Reptilia" family="Caenagnathidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="25" pageNumber="26" phylum="Chordata" rank="family">Caenagnathidae</taxonomicName>
, and some
<taxonomicName id="4C2AAAD29974FFE6BA570018FEDFFC62" authority="(Hutchinson 2001)" baseAuthorityName="Hutchinson" baseAuthorityYear="2001" class="Reptilia" family="Oviraptoridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="25" pageNumber="26" phylum="Chordata" rank="family">
Oviraptoridae (
<bibRefCitation id="EFBBACA09974FFE6B842007BFED3FC62" author="Hutchinson" box="[124,306,833,857]" pageId="25" pageNumber="26" pagination="169 - 97" refId="ref27459" refString="Hutchinson JR. The evolution of femoral osteology and soft tissues on the line to extant birds (Neornithes). Zoological Journal of the Linnean Society 2001; 131: 169 - 97. https: // doi. org / 10.1111 / j. 1096 - 3642.2001. tb 01314. x" year="2001">Hutchinson 2001</bibRefCitation>
)
</taxonomicName>
. The accessory trochanter of the femur of ZPAL MgD-I/108/1 is clearly demarcated from the lesser trochanter and forms a dorsoventral flange, comparable to that seen in
<taxonomicName id="4C2AAAD29974FFE6B8B500A5FED0FC8C" authorityName="Stenberg" authorityYear="1940" box="[139,305,927,951]" class="Reptilia" family="Caenagnathidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="25" pageNumber="26" phylum="Chordata" rank="family">Caenagnathidae</taxonomicName>
, e.g.
<taxonomicName id="4C2AAAD29974FFE6B95E00A5FDA9FC8C" authority="ZPAL" authorityName="ZPAL" box="[352,584,927,951]" class="Reptilia" family="Caenagnathidae" genus="Elmisaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="rarus">
<emphasis id="B95E0D439974FFE6B95E00A5FDE4FC8C" box="[352,517,927,951]" italics="true" pageId="25" pageNumber="26">Elmisaurus rarus</emphasis>
ZPAL
</taxonomicName>
MgD-I/98,
<taxonomicName id="4C2AAAD29974FFE6BAFB00A5FE64FCEC" authority="Lamanna et al., 2014" authorityName="Lamanna" authorityYear="2014" class="Reptilia" family="Caenagnathidae" genus="Anzu" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="oyliei">
<emphasis id="B95E0D439974FFE6BAFB00A5FF48FCED" italics="true" pageId="25" pageNumber="26">Anzu oyliei</emphasis>
<bibRefCitation id="EFBBACA09974FFE6B88E0085FE64FCEC" author="Lamanna MC &amp; Sues HD" box="[176,389,958,983]" pageId="25" pageNumber="26" refId="ref27733" refString="Lamanna MC, Sues HD, Schachner ER et al. A new large-bodied oviraptorosaurian theropod dinosaur from the Latest Cretaceous of Western North America. PLoS One 2014; 9: e 92022. https: // doi. org / 10.1371 / journal. pone. 0092022" year="2014">
Lamanna
<emphasis id="B95E0D439974FFE6B9280085FEA7FCED" box="[278,326,958,982]" italics="true" pageId="25" pageNumber="26">et al.</emphasis>
, 2014
</bibRefCitation>
</taxonomicName>
, or
<taxonomicName id="4C2AAAD29974FFE6B9900084FF44FCCD" authority="Gilmore, 1924" authorityName="Gilmore" authorityYear="1924" class="Reptilia" family="Caenagnathidae" genus="Chirostenotes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="pergracilis">
<emphasis id="B95E0D439974FFE6B9900084FD79FCED" box="[430,664,958,982]" italics="true" pageId="25" pageNumber="26">Chirostenotes pergracilis</emphasis>
<bibRefCitation id="EFBBACA09974FFE6BAA10084FF44FCCD" author="Gilmore CW" pageId="25" pageNumber="26" pagination="1 - 12" refId="ref27186" refString="Gilmore CW. A new coelurid dinosaur from the Belly River Cretaceous of Alberta. Canada Department of Mines Geological Survey Bulletin (Geological Series) 1924; 38: 1 - 12." year="1924">Gilmore, 1924</bibRefCitation>
</taxonomicName>
(Currie and Russell 1987). The lesser and greater trochanters are in contact, as in all Caenagnathoidea (
<bibRefCitation id="EFBBACA09974FFE6BABC00C4FF22FB0E" author="Lamanna MC &amp; Sues HD" pageId="25" pageNumber="26" refId="ref27733" refString="Lamanna MC, Sues HD, Schachner ER et al. A new large-bodied oviraptorosaurian theropod dinosaur from the Latest Cretaceous of Western North America. PLoS One 2014; 9: e 92022. https: // doi. org / 10.1371 / journal. pone. 0092022" year="2014">
Lamanna
<emphasis id="B95E0D439974FFE6BAD700C4FF6BFB0F" italics="true" pageId="25" pageNumber="26">et al.</emphasis>
2014
</bibRefCitation>
). The proximal end of the femur further resembles the femur of
<taxonomicName id="4C2AAAD29974FFE6B8EC0706FE5FFB6F" authority="ZPAL" authorityName="ZPAL" box="[210,446,1084,1108]" class="Reptilia" family="Caenagnathidae" genus="Elmisaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="rarus">
<emphasis id="B95E0D439974FFE6B8EC0706FE98FB68" box="[210,377,1084,1108]" italics="true" pageId="25" pageNumber="26">Elmisaurus rarus</emphasis>
ZPAL
</taxonomicName>
MgD-I/
<quantity id="4CD27CB49974FFE6BA270706FDAFFB6F" box="[537,590,1084,1108]" metricMagnitude="0" metricUnit="m" metricValue="2.4892" pageId="25" pageNumber="26" unit="in" value="98.0">98 in</quantity>
possessing a cylindrical head positioned higher than the greater trochanter and separated by a depression, which is wider in the larger (ontogenetically older, as indicated by the difference in size between those specimens) ZPAL MgD-I/108/1. Such an embayment is also present in other
<taxonomicName id="4C2AAAD29974FFE6B92407E2FE21FBCB" authorityName="Stenberg" authorityYear="1940" box="[282,448,1240,1264]" class="Reptilia" family="Caenagnathidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="25" pageNumber="26" phylum="Chordata" rank="family">Caenagnathidae</taxonomicName>
, e.g.
<taxonomicName id="4C2AAAD29974FFE6B9CF07E3FD85FBCB" box="[497,612,1240,1264]" class="Reptilia" family="Caenagnathidae" genus="Anzu" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="oyliei">
<emphasis id="B95E0D439974FFE6B9CF07E3FD85FBCB" box="[497,612,1240,1264]" italics="true" pageId="25" pageNumber="26">Anzu oyliei</emphasis>
</taxonomicName>
(see
<bibRefCitation id="EFBBACA09974FFE6BAA407E3FF3DFA2B" author="Lamanna MC &amp; Sues HD" pageId="25" pageNumber="26" refId="ref27733" refString="Lamanna MC, Sues HD, Schachner ER et al. A new large-bodied oviraptorosaurian theropod dinosaur from the Latest Cretaceous of Western North America. PLoS One 2014; 9: e 92022. https: // doi. org / 10.1371 / journal. pone. 0092022" year="2014">
Lamanna
<emphasis id="B95E0D439974FFE6B84F07C2FF43FA2B" box="[113,162,1272,1296]" italics="true" pageId="25" pageNumber="26">et al.</emphasis>
2014
</bibRefCitation>
),
<taxonomicName id="4C2AAAD29974FFE6B8CA07C2FE7DFA34" box="[244,412,1272,1296]" class="Reptilia" family="Caenagnathidae" genus="Elmisaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="rarus">
<emphasis id="B95E0D439974FFE6B8CA07C2FE7DFA34" box="[244,412,1272,1296]" italics="true" pageId="25" pageNumber="26">Elmisaurus rarus</emphasis>
</taxonomicName>
(see Barsbold
<emphasis id="B95E0D439974FFE6BA0907C2FD89FA2B" box="[567,616,1272,1296]" italics="true" pageId="25" pageNumber="26">et al.</emphasis>
2000), or
<taxonomicName id="4C2AAAD29974FFE6BAE907C2FF32FA14" authorityName="Gilmore" authorityYear="1924" class="Reptilia" family="Caenagnathidae" genus="Chirostenotes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="pergracilis">
<emphasis id="B95E0D439974FFE6BAE907C2FF32FA14" italics="true" pageId="25" pageNumber="26">Ch. pergracilis</emphasis>
</taxonomicName>
(see Currie and Russell 1987). A wide groove on the posterior surface of the femoral head for the capital ligament is present in both
<taxonomicName id="4C2AAAD29974FFE6B923066CFDECFA55" authority="ZPAL" authorityName="ZPAL" box="[285,525,1366,1390]" class="Reptilia" family="Caenagnathidae" genus="Elmisaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="rarus">
<emphasis id="B95E0D439974FFE6B923066CFE27FA56" box="[285,454,1366,1390]" italics="true" pageId="25" pageNumber="26">Elmisaurus rarus</emphasis>
ZPAL
</taxonomicName>
MgD-I/98 and MgDI/108/1. Also, similar to
<taxonomicName id="4C2AAAD29974FFE6B94D064FFDF8FAB6" authorityName="Stenberg" authorityYear="1940" box="[371,537,1397,1421]" class="Reptilia" family="Caenagnathidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="25" pageNumber="26" phylum="Chordata" rank="family">Caenagnathidae</taxonomicName>
, the lateral condyle of the femur ZPAL MgD-I/108/1 is positioned more distally than the medial condyle, and the tibiofibular crest is well demarcated [
<taxonomicName id="4C2AAAD29974FFE6B84206E9FF13FAD0" box="[124,242,1491,1515]" class="Reptilia" family="Caenagnathidae" genus="Anzu" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="oyliei">
<emphasis id="B95E0D439974FFE6B84206E9FF13FAD0" box="[124,242,1491,1515]" italics="true" pageId="25" pageNumber="26">Anzu oyliei</emphasis>
</taxonomicName>
(see
<bibRefCitation id="EFBBACA09974FFE6B91006E9FDE4FAD0" author="Lamanna MC &amp; Sues HD" box="[302,517,1491,1515]" pageId="25" pageNumber="26" refId="ref27733" refString="Lamanna MC, Sues HD, Schachner ER et al. A new large-bodied oviraptorosaurian theropod dinosaur from the Latest Cretaceous of Western North America. PLoS One 2014; 9: e 92022. https: // doi. org / 10.1371 / journal. pone. 0092022" year="2014">
Lamanna
<emphasis id="B95E0D439974FFE6B9A906EEFE28FAD0" box="[407,457,1491,1515]" italics="true" pageId="25" pageNumber="26">et al.</emphasis>
2014
</bibRefCitation>
),
<taxonomicName id="4C2AAAD29974FFE6BA2006E9FD26FAD0" box="[542,711,1491,1515]" class="Reptilia" family="Caenagnathidae" genus="Elmisaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="rarus">
<emphasis id="B95E0D439974FFE6BA2006E9FD26FAD0" box="[542,711,1491,1515]" italics="true" pageId="25" pageNumber="26">Elmisaurus rarus</emphasis>
</taxonomicName>
(see Barsbold
<emphasis id="B95E0D439974FFE6B8EA06C9FEE4F931" box="[212,261,1522,1546]" italics="true" pageId="25" pageNumber="26">et al.</emphasis>
2000), or
<taxonomicName id="4C2AAAD29974FFE6B94A06C8FDE1F931" authorityName="Gilmore" authorityYear="1924" box="[372,512,1522,1546]" class="Reptilia" family="Caenagnathidae" genus="Chirostenotes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="pergracilis">
<emphasis id="B95E0D439974FFE6B94A06C8FDE1F931" box="[372,512,1522,1546]" italics="true" pageId="25" pageNumber="26">Ch. pergracilis</emphasis>
</taxonomicName>
(see Currie and Russell 1987)]. The extensor groove is distinct, but shallow, consistent with
<taxonomicName id="4C2AAAD29974FFE6B899050BFEAEF972" box="[167,335,1585,1609]" class="Reptilia" family="Caenagnathidae" genus="Elmisaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="rarus">
<emphasis id="B95E0D439974FFE6B899050BFEAEF972" box="[167,335,1585,1609]" italics="true" pageId="25" pageNumber="26">Elmisaurus rarus</emphasis>
</taxonomicName>
(see Barsbold
<emphasis id="B95E0D439974FFE6B9D20508FDFCF972" box="[492,541,1585,1609]" italics="true" pageId="25" pageNumber="26">et al.</emphasis>
2000). The proximal and distal ends of the femur ZPAL MgD-I/108/1 are of similar size to the measurements in
<taxonomicName id="4C2AAAD29974FFE6B9AF0555FDD6F9BC" box="[401,567,1647,1671]" class="Reptilia" family="Caenagnathidae" genus="Elmisaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="rarus">
<emphasis id="B95E0D439974FFE6B9AF0555FDD6F9BC" box="[401,567,1647,1671]" italics="true" pageId="25" pageNumber="26">Elmisaurus rarus</emphasis>
</taxonomicName>
(see Barsbold
<emphasis id="B95E0D439974FFE6BAF2054AFD1AF9BC" box="[716,763,1647,1671]" italics="true" pageId="25" pageNumber="26">et al.</emphasis>
2000), hence the probable length of the whole bone was similar, ~
<quantity id="4CD27CB49974FFE6B8410594FF38F9FD" box="[127,217,1710,1734]" metricMagnitude="-1" metricUnit="m" metricValue="2.85" pageId="25" pageNumber="26" unit="mm" value="285.0">285 mm</quantity>
.
</paragraph>
<paragraph id="8B95D1519974FFE6B84F05D2FA4CFE5F" blockId="25.[113,763,1767,1980]" lastBlockId="25.[810,1460,144,1045]" pageId="25" pageNumber="26">
<emphasis id="B95E0D439974FFE6B84F05D2FE74F83B" box="[113,405,1768,1792]" italics="true" pageId="25" pageNumber="26">Bone microstructure of femur:</emphasis>
A histological section of the distal part of the shaft, above the condyles, shows a large marrow cavity and thin (~
<quantity id="4CD27CB49974FFE6B8DB041CFEC4F805" box="[229,293,1830,1854]" metricMagnitude="-3" metricUnit="m" metricValue="2.0" pageId="25" pageNumber="26" unit="mm" value="2.0">2 mm</quantity>
) cortex (
<figureCitation id="1311CDD49974FFE6B9BC041DFE15F805" box="[386,500,1831,1855]" captionStart="Figure 20" captionStartId="26.[129,194,1836,1860]" captionTargetBox="[175,1427,146,1806]" captionTargetId="figure-6@26.[173,1429,144,1808]" captionTargetPageId="26" captionText="Figure 20. Left femur of Caenagnathidae indet. ZPAL MgD-I/108/1. AE, proximal end of the femur in anterior (A), lateral (B), posterior (C), medial (D), and dorsal (E) view.FJ, distal end of the femur in anterior (F), lateral (G), posterior (H), medial (I), and ventral (J) view.K, L, thin section of the femoral cortex under polarized light. Red arrows indicate secondary osteons; green arrows point to resorption cavities; yellow arrows indicate lamellar bone; and purple arrows point to parallel-fibred bone." figureDoi="http://doi.org/10.5281/zenodo.14284271" httpUri="https://zenodo.org/record/14284271/files/figure.png" pageId="25" pageNumber="26">Fig. 20K, L</figureCitation>
). The external part of the cortex (half of its thickness) is built of parallel-fibred bone, with scattered secondary osteons. The vascularization is laminar, and growth marks are absent. In the section, no definite primary osteons were seen, although the external cortex is poorly preserved, possibly obscuring their presence. The inner cortex is sharply demarcated from the external cortex and built of densely packed secondary osteons: up to four generations are present. Close to the marrow cavity, resorption cavities are present, surrounded by a thick layer of lamellar bone (≤
<quantity id="4CD27CB49974FFE6BD1A0237FA98FE1E" box="[1316,1401,269,293]" metricMagnitude="-4" metricUnit="m" metricValue="3.0" pageId="25" pageNumber="26" unit="mm" value="0.3">0.3 mm</quantity>
). The marrow cavity is surrounded by a thinner layer of lamellar bone (
<quantity id="4CD27CB49974FFE6BB0C0276FC73FE5F" box="[818,914,332,356]" metricMagnitude="-4" metricUnit="m" metricValue="1.5" pageId="25" pageNumber="26" unit="mm" value="0.15">0.15 mm</quantity>
) and filled by slender and elongated bony trabeculae.
</paragraph>
<paragraph id="8B95D1519974FFE6BB7B0251FC4FFD3B" blockId="25.[810,1460,144,1045]" pageId="25" pageNumber="26">The section shows features typical for the metaphyses of long bones: extensive secondary remodelling, lack of growth marks, and numerous resorption cavities. Thus, owing to the lack of any growth record in the section, it is not possible to estimate the growth ratio.</paragraph>
<paragraph id="8B95D1519974FFE6BB7B0132FA78FB2E" blockId="25.[810,1460,144,1045]" pageId="25" pageNumber="26">
The bone microstructure of the femur in caenagnathids is unknown. Thus far, bone histology of the tibiae of cf.
<taxonomicName id="4C2AAAD29974FFE6BD7E011DFA52FD04" box="[1344,1459,551,575]" class="Reptilia" family="Caenagnathidae" genus="Anzu" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="oyliei">
<emphasis id="B95E0D439974FFE6BD7E011DFA52FD04" box="[1344,1459,551,575]" italics="true" pageId="25" pageNumber="26">Anzu oyliei</emphasis>
</taxonomicName>
(ROM 65884) and
<taxonomicName id="4C2AAAD29974FFE6BBCF017CFB71FD65" authorityName="Stenberg" authorityYear="1940" box="[1009,1168,582,606]" class="Reptilia" family="Caenagnathidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="25" pageNumber="26" phylum="Chordata" rank="family">Caenagnathidae</taxonomicName>
indet. (UALVP 57349) have been described (
<bibRefCitation id="EFBBACA09974FFE6BBE9015CFB3DFD45" author="Funston GF &amp; Currie PJ" box="[983,1244,614,638]" pageId="25" pageNumber="26" pagination="220 - 30" refId="ref26868" refString="Funston GF, Currie PJ. A small caenagnathid tibia from the Horseshoe Canyon Formation (Maastrichtian): implications for growth and lifestyle in oviraptorosaurs. Cretaceous Research 2018; 92: 220 - 30. https: // doi. org / 10.1016 / j. cretres. 2018.08.020" year="2018">Funston and Currie 2018</bibRefCitation>
,
<bibRefCitation id="EFBBACA09974FFE6BCD5015CFA43FD45" author="Cullen TM &amp; Simon DJ &amp; Benner EKC" box="[1259,1442,614,638]" pageId="25" pageNumber="26" pagination="751 - 67" refId="ref26173" refString="Cullen TM, Simon DJ, Benner EKC et al. Morphology and osteohistology of a large-bodied caenagnathid (Theropoda, Oviraptorosauria) from the Hell Creek Formation (Montana): implications for size-based classifications and growth reconstruction in theropods. Papers in Palaeontology 2021; 7: 751 - 67. https: // doi. org / 10.1002 / spp 2.1302" year="2021">
Cullen
<emphasis id="B95E0D439974FFE6BD09015DFA89FD45" box="[1335,1384,614,638]" italics="true" pageId="25" pageNumber="26">et al.</emphasis>
2021
</bibRefCitation>
). Both, however, represent young individuals, as indicated by their predominant fibrolamellar bone, high vascularity, and limited secondary remodelling (none in UALVP 57349 and ≤ 30% of cortex in OMVP 65884). The predominance of fibrolamellar bone and high vascularity are also seen in the cortices of the femora and fibulae of the oviraptorid
<taxonomicName id="4C2AAAD29974FFE6BC120018FB75FC02" box="[1068,1172,802,825]" family="Oviraptoridae" genus="Oksoko" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="avarsan">
<emphasis id="B95E0D439974FFE6BC120018FB75FC02" box="[1068,1172,802,825]" italics="true" pageId="25" pageNumber="26">O. avarsan</emphasis>
</taxonomicName>
, regardless of their ontogenetic age (
<bibRefCitation id="EFBBACA09974FFE6BBB80078FBAEFC62" author="Funston GF &amp; Tsogtbaatar C &amp; Tsogtbaatar K" box="[902,1103,833,857]" pageId="25" pageNumber="26" refId="ref27006" refString="Funston GF, Tsogtbaatar C, Tsogtbaatar K et al. A new two-fingered dinosaur sheds light on the radiation of Oviraptorosauria. Royal Society Open Science 2020 a; 7: 201184." year="2020">
Funston
<emphasis id="B95E0D439974FFE6BBE10078FBEFFC62" box="[991,1038,833,857]" italics="true" pageId="25" pageNumber="26">et al.</emphasis>
2020a
</bibRefCitation>
). Even in the large-bodied cf.
<taxonomicName id="4C2AAAD29974FFE6BD41007BFC81FC43" class="Reptilia" family="Caenagnathidae" genus="Anzu" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="oyliei">
<emphasis id="B95E0D439974FFE6BD41007BFC81FC43" italics="true" pageId="25" pageNumber="26">Anzu oyliei</emphasis>
</taxonomicName>
(ROM 65884), the section from the tibia revealed a predominately primary tissue, with generally high vascularity and limited secondary remodelling (
<bibRefCitation id="EFBBACA09974FFE6BC1C00A5FB38FC8C" author="Cullen TM &amp; Simon DJ &amp; Benner EKC" box="[1058,1241,927,951]" pageId="25" pageNumber="26" pagination="751 - 67" refId="ref26173" refString="Cullen TM, Simon DJ, Benner EKC et al. Morphology and osteohistology of a large-bodied caenagnathid (Theropoda, Oviraptorosauria) from the Hell Creek Formation (Montana): implications for size-based classifications and growth reconstruction in theropods. Papers in Palaeontology 2021; 7: 751 - 67. https: // doi. org / 10.1002 / spp 2.1302" year="2021">
Cullen
<emphasis id="B95E0D439974FFE6BC50009AFB7BFC8C" box="[1134,1178,927,951]" italics="true" pageId="25" pageNumber="26">et al</emphasis>
. 2021
</bibRefCitation>
). As can be noticed, the section taken from ZPAL MgD-I/108/1 is different from the Caenagnathoidea described before, which is a result of its sectioning at the metaphysis, and not the diaphysis as usually done.
</paragraph>
<paragraph id="8B95D1519974FFE6BB14070CFADFFABC" blockId="25.[810,1459,1078,1979]" pageId="25" pageNumber="26">
<emphasis id="B95E0D439974FFE6BB14070CFC7EFB75" box="[810,927,1078,1102]" italics="true" pageId="25" pageNumber="26">Tibiotarsus:</emphasis>
The left tibiotarsus is complete and measures
<quantity id="4CD27CB49974FFE6BDB2070CFCB2FB56" metricMagnitude="-1" metricUnit="m" metricValue="3.8" pageId="25" pageNumber="26" unit="mm" value="380.0">380 mm</quantity>
(
<figureCitation id="1311CDD49974FFE6BB5B076CFC36FB55" box="[869,983,1110,1134]" captionStart="Figure 21" captionStartId="27.[113,178,1572,1596]" captionTargetBox="[116,1456,146,1542]" captionTargetId="figure-215@27.[114,1458,144,1544]" captionTargetPageId="27" captionText="Figure 21. Left tibiotarsus of Caenagnathidae indet. ZPAL MgD-I/108/1. AF, tibiotarsus in anterior (A), lateral (B), posterior (C), medial (D), dorsal (E), and ventral (F) view. GK, proximal fibula in dorsal (G), anterior (H), lateral (I), posterior (J), and medial (K) view." figureDoi="http://doi.org/10.5281/zenodo.14284275" httpUri="https://zenodo.org/record/14284275/files/figure.png" pageId="25" pageNumber="26">Fig. 21AF</figureCitation>
). The bone is slender, slightly bowed laterally (possibly taphonomically exaggerated), and the distal fibula is fused to the distal tibia and calcaneum (
<figureCitation id="1311CDD49974FFE6BCF907AEFADFFB97" box="[1223,1342,1172,1196]" captionStart="Figure 21" captionStartId="27.[113,178,1572,1596]" captionTargetBox="[116,1456,146,1542]" captionTargetId="figure-215@27.[114,1458,144,1544]" captionTargetPageId="27" captionText="Figure 21. Left tibiotarsus of Caenagnathidae indet. ZPAL MgD-I/108/1. AF, tibiotarsus in anterior (A), lateral (B), posterior (C), medial (D), dorsal (E), and ventral (F) view. GK, proximal fibula in dorsal (G), anterior (H), lateral (I), posterior (J), and medial (K) view." figureDoi="http://doi.org/10.5281/zenodo.14284275" httpUri="https://zenodo.org/record/14284275/files/figure.png" pageId="25" pageNumber="26">Fig. 21AC</figureCitation>
). The shaft is elliptical in cross-section (circumference
<quantity id="4CD27CB49974FFE6BCDA0789FACEFBF7" box="[1252,1327,1203,1228]" metricMagnitude="-2" metricUnit="m" metricValue="9.5" pageId="25" pageNumber="26" unit="mm" value="95.0">95 mm</quantity>
) and is compressed anteroposteriorly, possibly as an effect of taphonomical crushing. Proximally, the tibia expands anteriorly and slightly mediolaterally; its anteroposterior depth is
<quantity id="4CD27CB49974FFE6BD29062BFA9DFA12" box="[1303,1404,1297,1321]" metricMagnitude="-2" metricUnit="m" metricValue="4.75" pageId="25" pageNumber="26" unit="mm" value="47.5">47.5 mm</quantity>
and mediolateral width
<quantity id="4CD27CB49974FFE6BBC8060BFBB6FA73" box="[1014,1111,1329,1353]" metricMagnitude="-2" metricUnit="m" metricValue="5.92" pageId="25" pageNumber="26" unit="mm" value="59.2">59.2 mm</quantity>
. Distally, where the tibia is fused with the astragalocalcaneum distally and the fibula laterally, the tibia expands mediolaterally and measures
<quantity id="4CD27CB49974FFE6BCE30655FADAFABC" box="[1245,1339,1391,1415]" metricMagnitude="-2" metricUnit="m" metricValue="5.7299999999999995" pageId="25" pageNumber="26" unit="mm" value="57.3">57.3 mm</quantity>
.
</paragraph>
<paragraph id="8B95D1519974FFE6BB7806B5FC59F880" blockId="25.[810,1459,1078,1979]" pageId="25" pageNumber="26">
The cnemial crest condyle (cranial cnemial crest
<emphasis id="B95E0D439974FFE6BDBE06AAFA52FA9C" box="[1408,1459,1424,1447]" italics="true" pageId="25" pageNumber="26">sensu</emphasis>
Osmólska 1996) is robust, laterally deflected, and short in anterior view, comprising only ~15% of the maximum proximodistal tibiotarsus length. The fibular condyle (lateral cnemial crest
<emphasis id="B95E0D439974FFE6BBFA0537FC16F91F" box="[964,1015,1549,1572]" italics="true" pageId="25" pageNumber="26">sensu</emphasis>
Osmólska 1996) is also robust, slightly curved anteriorly, and shorter mediolaterally and dorsoventrally than the cnemial crest. Between the cnemial crest and fibular condyle, a deep and posteriorly curved incisura tibialis is present (
<figureCitation id="1311CDD49974FFE6BB0B05B3FC4BF999" box="[821,938,1673,1698]" captionStart="Figure 21" captionStartId="27.[113,178,1572,1596]" captionTargetBox="[116,1456,146,1542]" captionTargetId="figure-215@27.[114,1458,144,1544]" captionTargetPageId="27" captionText="Figure 21. Left tibiotarsus of Caenagnathidae indet. ZPAL MgD-I/108/1. AF, tibiotarsus in anterior (A), lateral (B), posterior (C), medial (D), dorsal (E), and ventral (F) view. GK, proximal fibula in dorsal (G), anterior (H), lateral (I), posterior (J), and medial (K) view." figureDoi="http://doi.org/10.5281/zenodo.14284275" httpUri="https://zenodo.org/record/14284275/files/figure.png" pageId="25" pageNumber="26">Fig. 21AE</figureCitation>
). The medial proximal condyle of the tibiotarsus is long anteroposteriorly; anteriorly, it is smoothly connected with the cnemial crest; posteriorly, it is separated from the fibular condyle by a triangular cleft. The posteromedial edge of the medial proximal condyle of the tibiotarsus is posteriorly extended. Below the fibular condyle, the fibular crest is present. It is tall dorsoventrally, ~20% of the tibiotarsus length. The crest becomes wider mediolaterally and deflects anteriorly; however, the crest is not strongly pronounced. The distal end of the crest is rectangular.
</paragraph>
<caption id="DF5581D99977FFE5B8BF0416FC42F8A3" ID-DOI="http://doi.org/10.5281/zenodo.14284271" ID-Zenodo-Dep="14284271" httpUri="https://zenodo.org/record/14284271/files/figure.png" pageId="26" pageNumber="27" startId="26.[129,194,1836,1860]" targetBox="[175,1427,146,1806]" targetPageId="26" targetType="figure">
<paragraph id="8B95D1519977FFE5B8BF0416FC42F8A3" blockId="26.[129,1470,1836,1944]" pageId="26" pageNumber="27">
<emphasis id="B95E0D439977FFE5B8BF0416FF04F87F" bold="true" box="[129,229,1836,1860]" pageId="26" pageNumber="27">Figure 20.</emphasis>
Left femur of
<taxonomicName id="4C2AAAD29977FFE5B9540416FDE2F87F" authorityName="Stenberg" authorityYear="1940" box="[362,515,1836,1860]" class="Reptilia" family="Caenagnathidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="26" pageNumber="27" phylum="Chordata" rank="family">Caenagnathidae</taxonomicName>
indet. ZPAL MgD-I/108/1. AE, proximal end of the femur in anterior (A), lateral (B), posterior (C), medial (D), and dorsal (E) view. FJ, distal end of the femur in anterior (F), lateral (G), posterior (H), medial (I), and ventral (J) view. K, L, thin section of the femoral cortex under polarized light. Red arrows indicate secondary osteons; green arrows point to resorption cavities; yellow arrows indicate lamellar bone; and purple arrows point to parallel-fibred bone.
</paragraph>
</caption>
<caption id="DF5581D99976FFE4B84F051EFAB9F963" ID-DOI="http://doi.org/10.5281/zenodo.14284275" ID-Zenodo-Dep="14284275" httpUri="https://zenodo.org/record/14284275/files/figure.png" pageId="27" pageNumber="28" startId="27.[113,178,1572,1596]" targetBox="[116,1456,146,1542]" targetPageId="27" targetType="figure">
<paragraph id="8B95D1519976FFE4B84F051EFAB9F963" blockId="27.[113,1427,1572,1625]" pageId="27" pageNumber="28">
<emphasis id="B95E0D439976FFE4B84F051EFF37F907" bold="true" box="[113,214,1572,1596]" pageId="27" pageNumber="28">Figure 21.</emphasis>
Left tibiotarsus of
<taxonomicName id="4C2AAAD29976FFE4B9B8051EFDFEF907" authorityName="Stenberg" authorityYear="1940" box="[390,543,1572,1596]" class="Reptilia" family="Caenagnathidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="27" pageNumber="28" phylum="Chordata" rank="family">Caenagnathidae</taxonomicName>
indet. ZPAL MgD-I/108/1. AF, tibiotarsus in anterior (A), lateral (B), posterior (C), medial (D), dorsal (E), and ventral (F) view. GK, proximal fibula in dorsal (G), anterior (H), lateral (I), posterior (J), and medial (K) view.
</paragraph>
</caption>
<paragraph id="8B95D1519976FFE4B8B305B2FAD5F806" blockId="27.[113,763,1672,1978]" lastBlockId="27.[810,1459,1672,1979]" pageId="27" pageNumber="28">
The fibula is fused to the lateral side of the distal end of the tibia, along the distal ~ 23% of the tibiotarsus length (
<figureCitation id="1311CDD49976FFE4BAEA0592FF5DF9E4" captionStart="Figure 21" captionStartId="27.[113,178,1572,1596]" captionTargetBox="[116,1456,146,1542]" captionTargetId="figure-215@27.[114,1458,144,1544]" captionTargetPageId="27" captionText="Figure 21. Left tibiotarsus of Caenagnathidae indet. ZPAL MgD-I/108/1. AF, tibiotarsus in anterior (A), lateral (B), posterior (C), medial (D), dorsal (E), and ventral (F) view. GK, proximal fibula in dorsal (G), anterior (H), lateral (I), posterior (J), and medial (K) view." figureDoi="http://doi.org/10.5281/zenodo.14284275" httpUri="https://zenodo.org/record/14284275/files/figure.png" pageId="27" pageNumber="28">Fig. 21AC</figureCitation>
). The distal end of the fibula is partly fused with the calcaneum. The outline of the distal part of the fibula is marked and distinguishable against the remaining bones. The suture between the astragalocalcaneum and distal tibia are clearer in anterior than posterior view; however, that might be a matter of preservation. No suture is visible between the astragalus and calcaneum. The calcaneum shows a lateral depression (lateral epicondylar depression) below the suture with the fibula. The preserved, incomplete ascending process of the astragalus extends along 7.5% of the length of the tibiotarsus. In anterior view, it has subtriangular, pointed medial and lateral processes, separated by a deep depression. At the base of the ascending process, a shallow median depression is present, above which a low, mediolaterally extended ridge is located (
<figureCitation id="1311CDD49976FFE4BCF3041FFAC2F806" box="[1229,1315,1829,1853]" captionStart="Figure 21" captionStartId="27.[113,178,1572,1596]" captionTargetBox="[116,1456,146,1542]" captionTargetId="figure-215@27.[114,1458,144,1544]" captionTargetPageId="27" captionText="Figure 21. Left tibiotarsus of Caenagnathidae indet. ZPAL MgD-I/108/1. AF, tibiotarsus in anterior (A), lateral (B), posterior (C), medial (D), dorsal (E), and ventral (F) view. GK, proximal fibula in dorsal (G), anterior (H), lateral (I), posterior (J), and medial (K) view." figureDoi="http://doi.org/10.5281/zenodo.14284275" httpUri="https://zenodo.org/record/14284275/files/figure.png" pageId="27" pageNumber="28">Fig. 21A</figureCitation>
).
</paragraph>
<paragraph id="8B95D1519976FFE3BB7B047EFF19FD86" blockId="27.[810,1459,1672,1979]" lastBlockId="28.[129,778,144,1170]" lastPageId="28" lastPageNumber="29" pageId="27" pageNumber="28">
A proper tibiotarsus, in which the tibia is fused to the proximal tarsals, is recognized in three non-avian maniraptoran taxa: Alvarezsauridae, Troodontidae, and Avimimidae. Both alvarezsaurids known from the Nemegt Formation (M. olecranus and
<taxonomicName id="4C2AAAD29971FFE3B89303AAFDF1FF93" authority="Lee et al., 2019" authorityName="Lee" authorityYear="2019" box="[173,528,144,168]" family="Alvarezsauridae" genus="Nemegtonykus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="citus">
<emphasis id="B95E0D439971FFE3B89303AAFE8FFF93" box="[173,366,144,168]" italics="true" pageId="28" pageNumber="29">Nemegtonykus citus</emphasis>
<bibRefCitation id="EFBBACA09971FFE3B94A03AAFDF1FF93" author="Lee S &amp; Park J &amp; Lee Y" box="[372,528,144,168]" pageId="28" pageNumber="29" refId="ref27832" refString="Lee S, Park J, Lee Y et al. A new alvarezsaurid dinosaur from the Nemegt Formation of Mongolia. Scientific Reports 2019; 9: 15493. https: // doi. org / 10.1038 / s 41598 - 019 - 52021 - y" year="2019">
Lee
<emphasis id="B95E0D439971FFE3B99F03ABFE30FF93" box="[417,465,144,168]" italics="true" pageId="28" pageNumber="29">et al.</emphasis>
, 2019
</bibRefCitation>
</taxonomicName>
) have a proximodistally short fibula, which does not reach even the midshaft of the tibiotarsus (Perle
<emphasis id="B95E0D439971FFE3B93B03F5FED2FFDC" box="[261,307,207,231]" italics="true" pageId="28" pageNumber="29">et al.</emphasis>
1994,
<bibRefCitation id="EFBBACA09971FFE3B94B03F5FDE6FFDC" author="Lee S &amp; Park J &amp; Lee Y" box="[373,519,207,231]" pageId="28" pageNumber="29" refId="ref27832" refString="Lee S, Park J, Lee Y et al. A new alvarezsaurid dinosaur from the Nemegt Formation of Mongolia. Scientific Reports 2019; 9: 15493. https: // doi. org / 10.1038 / s 41598 - 019 - 52021 - y" year="2019">
Lee
<emphasis id="B95E0D439971FFE3B99E03F5FE2EFFDC" box="[416,463,207,231]" italics="true" pageId="28" pageNumber="29">et al.</emphasis>
2019
</bibRefCitation>
). The presence of a tibiotarsus in the troodontids known from the Nemegt Formation is variable. In the larger species
<taxonomicName id="4C2AAAD29971FFE3B9DE0237FF21FE7E" authority="(Barsbold, 1974)" baseAuthorityName="Barsbold" baseAuthorityYear="1974" class="Reptilia" family="Troodontidae" genus="Zanabazar" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="junior">
<emphasis id="B95E0D439971FFE3B9DE0237FD72FE1E" box="[480,659,269,293]" italics="true" pageId="28" pageNumber="29">Zanabazar junior</emphasis>
(
<bibRefCitation id="EFBBACA09971FFE3BA960237FF54FE7E" author="Barsbold" pageId="28" pageNumber="29" pagination="5 - 22" refId="ref25112" refString="Barsbold R. Saurornithoididae, a new family of small theropod dinosaurs from Central Asia and North America. Palaeontologia Polonica 1974; 30: 5 - 22." year="1974">Barsbold, 1974</bibRefCitation>
)
</taxonomicName>
, the astagalocalcaneum is not fused to the tibia (
<bibRefCitation id="EFBBACA09971FFE3BAF60217FF09FE5F" author="Norell MA &amp; Makovicky PJ &amp; Bever GS" pageId="28" pageNumber="29" pagination="1 - 63" refId="ref28270" refString="Norell MA, Makovicky PJ, Bever GS et al. A review of the Mongolian Cretaceous dinosaur Saurornithoides. American Museum Novitates 2009; 3654: 1 - 63. https: // doi. org / 10.1206 / 648.1" year="2009">
Norell
<emphasis id="B95E0D439971FFE3B8BF0277FF4EFE5F" box="[129,175,332,356]" italics="true" pageId="28" pageNumber="29">et al.</emphasis>
2009
</bibRefCitation>
), whereas in the smaller
<taxonomicName id="4C2AAAD29971FFE3B9DB0277FD8CFE5F" authorityName="Osmolska" authorityYear="1987" box="[485,621,332,356]" class="Reptilia" family="Troodontidae" genus="Borogovia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="gracilicrus">
<emphasis id="B95E0D439971FFE3B9DB0277FD8CFE5F" box="[485,621,332,356]" italics="true" pageId="28" pageNumber="29">Bo. gracilicrus</emphasis>
</taxonomicName>
, a tibiotarsus is present (Osmólska 1987,
<bibRefCitation id="EFBBACA09971FFE3B9B90251FD89FEB8" author="Cau A &amp; Madzia D" box="[391,616,363,387]" pageId="28" pageNumber="29" refId="ref25973" refString="Cau A, Madzia D. The phylogenetic affinities and morphological peculiarities of the bird- like dinosaur Borogovia gracilicrus from the Upper Cretaceous of Mongolia. PeerJ 2021; 9: e 12640. https: // doi. org / 10.7717 / peerj. 12640" year="2021">Cau and Madzia 2021</bibRefCitation>
). The hindlimb is unknown in the third troodontid from the Nemegt Formation,
<taxonomicName id="4C2AAAD29971FFE3B8BF0290FE3EFEFA" authority="Kurzanov" authorityName="Kurzanov" box="[129,479,426,450]" class="Reptilia" family="Troodontidae" genus="Tochisaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="nemegtensis">
<emphasis id="B95E0D439971FFE3B8BF0290FE90FEF9" box="[129,369,426,450]" italics="true" pageId="28" pageNumber="29">Tochisaurus nemegtensis</emphasis>
<bibRefCitation id="EFBBACA09971FFE3B9450290FD50FEF9" author="Kurzanov SM &amp; Osmolska H" box="[379,689,426,450]" pageId="28" pageNumber="29" pagination="69 - 76" refId="ref27695" refString="Kurzanov SM, Osmolska H. Tochisaurus nemegtensis gen. et sp. n., a new troodontid (Dinosauria, Theropoda) from Mongolia. Acta Palaeontologica Polonica 1991; 36: 69 - 76." year="1991">Kurzanov &amp; Osmólska, 1991</bibRefCitation>
</taxonomicName>
. Only a fragment of proximal right fibula of
<taxonomicName id="4C2AAAD29971FFE3B9C902F0FD60FEDA" authorityName="Osmolska" authorityYear="1987" box="[503,641,457,481]" class="Reptilia" family="Troodontidae" genus="Borogovia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="gracilicrus">
<emphasis id="B95E0D439971FFE3B9C902F0FD60FEDA" box="[503,641,457,481]" italics="true" pageId="28" pageNumber="29">Bo. gracilicrus</emphasis>
</taxonomicName>
is preserved, but the distal end of the tibiotarsus does not show any signs of fusion with the distal fibula, as in other
<taxonomicName id="4C2AAAD29971FFE3BA270132FD47FD1B" authorityName="Gilmore" authorityYear="1924" box="[537,678,520,544]" class="Reptilia" family="Troodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="family">Troodontidae</taxonomicName>
(e.g.
<bibRefCitation id="EFBBACA09971FFE3BAE00132FF06FD04" author="Gao C &amp; Morschhauser EM &amp; Varricchio DJ" pageId="28" pageNumber="29" refId="ref27130" refString="Gao C, Morschhauser EM, Varricchio DJ et al. A second soundly sleeping dragon: new anatomical details of the Chinese troodontid Mei long with implications for phylogeny and taphonomy. PLoS One 2012; 7: e 45203. https: // doi. org / 10.1371 / journal. pone. 0045203" year="2012">
Gao
<emphasis id="B95E0D439971FFE3B8BF0112FF4EFD04" box="[129,175,551,575]" italics="true" pageId="28" pageNumber="29">et al.</emphasis>
2012
</bibRefCitation>
).
<taxonomicName id="4C2AAAD29971FFE3B8C2011DFE6CFD04" baseAuthorityName="Hutchinson" baseAuthorityYear="2001" box="[252,397,551,575]" class="Reptilia" family="Oviraptoridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="family">Oviraptoridae</taxonomicName>
and
<taxonomicName id="4C2AAAD29971FFE3B983011DFD83FD04" authorityName="Stenberg" authorityYear="1940" box="[445,610,551,575]" class="Reptilia" family="Caenagnathidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="family">Caenagnathidae</taxonomicName>
show a fused astragalus and calcaneum, but not to the tibia (
<bibRefCitation id="EFBBACA09971FFE3BA77017CFD18FD65" author="Currie PJ &amp; Funston GF &amp; Osmolska H" box="[585,761,582,606]" pageId="28" pageNumber="29" pagination="143 - 57" refId="ref26493" refString="Currie PJ, Funston GF, Osmolska H. New specimens of the crested theropod dinosaur Elmisaurus rarus from Mongolia. Acta Palaeontologica Polonica 2016; 61: 143 - 57." year="2016">
Currie
<emphasis id="B95E0D439971FFE3BAAF017DFD21FD65" box="[657,704,582,606]" italics="true" pageId="28" pageNumber="29">et al.</emphasis>
2016
</bibRefCitation>
). Finally, a fused tibiotarsus including the distal end of the fibula is an autapomorphy of
<taxonomicName id="4C2AAAD29971FFE3B95201BCFE1DFDA6" box="[364,508,646,669]" class="Reptilia" family="Avimimidae" genus="Avimimus" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B95E0D439971FFE3B95201BCFE2FFDA6" box="[364,462,646,669]" italics="true" pageId="28" pageNumber="29">Avimimus</emphasis>
spp.
</taxonomicName>
(
<bibRefCitation id="EFBBACA09971FFE3BA3201BCFD48FDA6" author="Kurzanov SM" box="[524,681,645,669]" pageId="28" pageNumber="29" pagination="39 - 49" refId="ref27634" refString="Kurzanov SM. An unusual theropod from the Upper Cretaceous of Mongolia. In: Fossil Vertebrates of Mongolia. Joint Soviet-Mongolian Paleontological Expedition. Moscow: Nauka; 1981, 39 - 49." year="1981">Kurzanov 1981</bibRefCitation>
,
<bibRefCitation id="EFBBACA09971FFE3BA8A01BCFF06FD86" author="Funston GF &amp; Currie PJ" pageId="28" pageNumber="29" pagination="220 - 30" refId="ref26868" refString="Funston GF, Currie PJ. A small caenagnathid tibia from the Horseshoe Canyon Formation (Maastrichtian): implications for growth and lifestyle in oviraptorosaurs. Cretaceous Research 2018; 92: 220 - 30. https: // doi. org / 10.1016 / j. cretres. 2018.08.020" year="2018">
Funston
<emphasis id="B95E0D439971FFE3B8BF019FFF4EFD87" box="[129,175,676,700]" italics="true" pageId="28" pageNumber="29">et al.</emphasis>
2018
</bibRefCitation>
).
</paragraph>
<paragraph id="8B95D1519971FFE3B8A201FEFD38FBA9" blockId="28.[129,778,144,1170]" pageId="28" pageNumber="29">
However, ZPAL MgD-I/108/1 would be an exceptionally large representative of
<taxonomicName id="4C2AAAD29971FFE3B95A01D9FE27FDC1" box="[356,454,739,762]" class="Reptilia" family="Avimimidae" genus="Avimimus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="genus">
<emphasis id="B95E0D439971FFE3B95A01D9FE27FDC1" box="[356,454,739,762]" italics="true" pageId="28" pageNumber="29">Avimimus</emphasis>
</taxonomicName>
; the largest reported tibiotarsus of
<taxonomicName id="4C2AAAD29971FFE3B8A70039FD6CFC21" authority="Funston, Mendonca, Currie" authorityName="Funston, Mendonca, Currie" box="[153,653,770,794]" class="Reptilia" family="Avimimidae" genus="Avimimus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="nemegtensis">
<emphasis id="B95E0D439971FFE3B8A70039FE91FC21" box="[153,368,771,794]" italics="true" pageId="28" pageNumber="29">Avimimus nemegtensis</emphasis>
Funston, Mendonca, Currie
</taxonomicName>
&amp; Barsbold, 2018 MPC-D 102/92 measures
<quantity id="4CD27CB49971FFE3B9F30018FDC5FC02" box="[461,548,802,826]" metricMagnitude="-1" metricUnit="m" metricValue="2.82" pageId="28" pageNumber="29" unit="mm" value="282.0">282 mm</quantity>
(
<bibRefCitation id="EFBBACA09971FFE3BA0B0018FD18FC01" author="Funston GF &amp; Currie PJ &amp; Eberth DA" box="[565,761,802,826]" pageId="28" pageNumber="29" refId="ref26919" refString="Funston GF, Currie PJ, Eberth DA et al. The first oviraptorosaur (Dinosauria: Theropoda) bonebed: evidence of gregarious behaviour in a maniraptoran theropod. Scientific Reports 2016; 6: 35782." year="2016">
Funston
<emphasis id="B95E0D439971FFE3BAAE0019FD5EFC01" box="[656,703,802,826]" italics="true" pageId="28" pageNumber="29">et al.</emphasis>
2016
</bibRefCitation>
), and in
<taxonomicName id="4C2AAAD29971FFE3B8FC007BFD89FC62" authority="Kurzanov, 1981 PIN" authorityName="Kurzanov" authorityYear="1981" box="[194,616,833,857]" class="Reptilia" family="Avimimidae" genus="Avimimus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="portentosus">
<emphasis id="B95E0D439971FFE3B8FC007BFE77FC62" box="[194,406,833,857]" italics="true" pageId="28" pageNumber="29">Avimimus portentosus</emphasis>
<bibRefCitation id="EFBBACA09971FFE3B9A40078FDD8FC62" author="Kurzanov SM" box="[410,569,833,857]" pageId="28" pageNumber="29" pagination="39 - 49" refId="ref27634" refString="Kurzanov SM. An unusual theropod from the Upper Cretaceous of Mongolia. In: Fossil Vertebrates of Mongolia. Joint Soviet-Mongolian Paleontological Expedition. Moscow: Nauka; 1981, 39 - 49." year="1981">Kurzanov, 1981</bibRefCitation>
<emphasis id="B95E0D439971FFE3BA030078FD89FC62" box="[573,616,834,857]" italics="true" pageId="28" pageNumber="29">PIN</emphasis>
</taxonomicName>
<emphasis id="B95E0D439971FFE3BA55007BFD58FC62" box="[619,697,833,857]" italics="true" pageId="28" pageNumber="29">3907/1</emphasis>
it is
<quantity id="4CD27CB49971FFE3BADC007BFF4BFC43" metricMagnitude="-1" metricUnit="m" metricValue="2.57" pageId="28" pageNumber="29" unit="mm" value="257.0">257 mm</quantity>
long (
<bibRefCitation id="EFBBACA09971FFE3B8D4005BFE67FC42" author="Kurzanov SM" box="[234,390,865,889]" pageId="28" pageNumber="29" pagination="39 - 49" refId="ref27634" refString="Kurzanov SM. An unusual theropod from the Upper Cretaceous of Mongolia. In: Fossil Vertebrates of Mongolia. Joint Soviet-Mongolian Paleontological Expedition. Moscow: Nauka; 1981, 39 - 49." year="1981">Kurzanov 1981</bibRefCitation>
), whereas ZPAL MgD-I/108/1 measures
<quantity id="4CD27CB49971FFE3B88C00BAFEEAFCAC" box="[178,267,896,920]" metricMagnitude="-1" metricUnit="m" metricValue="3.8" pageId="28" pageNumber="29" unit="mm" value="380.0">380 mm</quantity>
, similar to
<taxonomicName id="4C2AAAD29971FFE3B94000BAFDC4FCA3" box="[382,549,896,920]" class="Reptilia" family="Caenagnathidae" genus="Elmisaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="rarus">
<emphasis id="B95E0D439971FFE3B94000BAFDC4FCA3" box="[382,549,896,920]" italics="true" pageId="28" pageNumber="29">Elmisaurus rarus</emphasis>
</taxonomicName>
MPC-D 100/119, i.e.
<quantity id="4CD27CB49971FFE3B8BF00A5FF38FC8C" box="[129,217,927,951]" metricMagnitude="-1" metricUnit="m" metricValue="3.55" pageId="28" pageNumber="29" unit="mm" value="355.0">355 mm</quantity>
. The histological sections of the Iren Dabasu avimimids revealed that the largest sectioned specimens were already adults (Funston
<emphasis id="B95E0D439971FFE3B8D600E4FEF8FCCD" box="[232,281,990,1014]" italics="true" pageId="28" pageNumber="29">et al.</emphasis>
2019). Moreover, three features of the tibia are shared by ZPAL MgD-I/108/1 and
<taxonomicName id="4C2AAAD29971FFE3BA3900C7FD53FB2E" box="[519,690,1021,1045]" class="Reptilia" family="Caenagnathidae" genus="Elmisaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="rarus">
<emphasis id="B95E0D439971FFE3BA3900C7FD53FB2E" box="[519,690,1021,1045]" italics="true" pageId="28" pageNumber="29">Elmisaurus rarus</emphasis>
</taxonomicName>
: (i) the medial proximal condyle is more protruded dorsally than in
<taxonomicName id="4C2AAAD29971FFE3B8BF0706FEF5FB68" box="[129,276,1084,1107]" class="Reptilia" family="Avimimidae" genus="Avimimus" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B95E0D439971FFE3B8BF0706FF02FB68" box="[129,227,1084,1107]" italics="true" pageId="28" pageNumber="29">Avimimus</emphasis>
spp.
</taxonomicName>
; (ii) the fibular crest is longer, and its distal end is rectangular, not arcuate as in
<taxonomicName id="4C2AAAD29971FFE3B9960761FDDAFB48" box="[424,571,1115,1139]" class="Reptilia" family="Avimimidae" genus="Avimimus" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B95E0D439971FFE3B9960761FDEBFB49" box="[424,522,1115,1138]" italics="true" pageId="28" pageNumber="29">Avimimus</emphasis>
spp.
</taxonomicName>
; and (iii) the medial malleolus protrudes further medially than in
<taxonomicName id="4C2AAAD29971FFE3BA770741FD38FBA9" box="[585,729,1147,1170]" class="Reptilia" family="Avimimidae" genus="Avimimus" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B95E0D439971FFE3BA770741FD4AFBA9" box="[585,683,1147,1170]" italics="true" pageId="28" pageNumber="29">Avimimus</emphasis>
spp.
</taxonomicName>
</paragraph>
<paragraph id="8B95D1519971FFE3B8BF078DFDDDF9FE" blockId="28.[128,778,1207,1858]" pageId="28" pageNumber="29">
<emphasis id="B95E0D439971FFE3B8BF078DFF27FBF4" box="[129,198,1207,1231]" italics="true" pageId="28" pageNumber="29">Fibula:</emphasis>
The left fibula is complete, measuring
<quantity id="4CD27CB49971FFE3BA79078DFD7EFBF4" box="[583,671,1207,1231]" metricMagnitude="-1" metricUnit="m" metricValue="3.4" pageId="28" pageNumber="29" unit="mm" value="340.0">340 mm</quantity>
. The distal end is partly fused to the calcaneum and laterally fused to the tibia, along ~33% of the length of the fibula. It is similar to
<taxonomicName id="4C2AAAD29971FFE3BA9907CCFF0EFA16" authorityName="Kurzanov" authorityYear="1981" class="Reptilia" family="Avimimidae" genus="Avimimus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="portentosus">
<emphasis id="B95E0D439971FFE3BA9907CCFF0EFA16" italics="true" pageId="28" pageNumber="29">Avimimus portentosus</emphasis>
</taxonomicName>
, in which the fibula is fused to the tibia along onethird of its length (
<bibRefCitation id="EFBBACA09971FFE3B970060FFE0FFA77" author="Kurzanov SM" box="[334,494,1332,1356]" pageId="28" pageNumber="29" pagination="39 - 49" refId="ref27634" refString="Kurzanov SM. An unusual theropod from the Upper Cretaceous of Mongolia. In: Fossil Vertebrates of Mongolia. Joint Soviet-Mongolian Paleontological Expedition. Moscow: Nauka; 1981, 39 - 49." year="1981">Kurzanov 1981</bibRefCitation>
). The proximal anteroposterior length of the fibula is
<quantity id="4CD27CB49971FFE3B99B066EFDE5FA50" box="[421,516,1364,1388]" metricMagnitude="-2" metricUnit="m" metricValue="4.71" pageId="28" pageNumber="29" unit="mm" value="47.1">47.1 mm</quantity>
. The proximal end of the fibula is triangular in the lateromedial aspect, only slightly concave medially in dorsal view (
<figureCitation id="1311CDD49971FFE3B99506A8FDC3FA91" box="[427,546,1426,1450]" captionStart="Figure 21" captionStartId="27.[113,178,1572,1596]" captionTargetBox="[116,1456,146,1542]" captionTargetId="figure-215@27.[114,1458,144,1544]" captionTargetPageId="27" captionText="Figure 21. Left tibiotarsus of Caenagnathidae indet. ZPAL MgD-I/108/1. AF, tibiotarsus in anterior (A), lateral (B), posterior (C), medial (D), dorsal (E), and ventral (F) view. GK, proximal fibula in dorsal (G), anterior (H), lateral (I), posterior (J), and medial (K) view." figureDoi="http://doi.org/10.5281/zenodo.14284275" httpUri="https://zenodo.org/record/14284275/files/figure.png" pageId="28" pageNumber="29">Fig. 21GK</figureCitation>
), similar to
<taxonomicName id="4C2AAAD29971FFE3BAA206A8FF55FAF2" class="Reptilia" family="Caenagnathidae" genus="Elmisaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="rarus">
<emphasis id="B95E0D439971FFE3BAA206A8FF55FAF2" italics="true" pageId="28" pageNumber="29">Elmisaurus rarus</emphasis>
</taxonomicName>
(MPC-D 100/119; Barsbold
<emphasis id="B95E0D439971FFE3B9DC0688FDF0FAF1" box="[482,529,1458,1482]" italics="true" pageId="28" pageNumber="29">et al.</emphasis>
2000) and in contrast to the rectangular proximal end in
<taxonomicName id="4C2AAAD29971FFE3B9F806E8FDB6FAD2" box="[454,599,1490,1513]" class="Reptilia" family="Avimimidae" genus="Avimimus" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B95E0D439971FFE3B9F806E8FDC9FAD2" box="[454,552,1490,1513]" italics="true" pageId="28" pageNumber="29">Avimimus</emphasis>
spp.
</taxonomicName>
(
<bibRefCitation id="EFBBACA09971FFE3BA5806E8FCE5FAD2" author="Kurzanov SM" box="[614,772,1489,1513]" pageId="28" pageNumber="29" pagination="39 - 49" refId="ref27634" refString="Kurzanov SM. An unusual theropod from the Upper Cretaceous of Mongolia. In: Fossil Vertebrates of Mongolia. Joint Soviet-Mongolian Paleontological Expedition. Moscow: Nauka; 1981, 39 - 49." year="1981">Kurzanov 1981</bibRefCitation>
,
<bibRefCitation id="EFBBACA09971FFE3B8BE06CBFEA9F932" author="Funston GF &amp; Currie PJ &amp; Eberth DA" box="[128,328,1521,1545]" pageId="28" pageNumber="29" refId="ref26919" refString="Funston GF, Currie PJ, Eberth DA et al. The first oviraptorosaur (Dinosauria: Theropoda) bonebed: evidence of gregarious behaviour in a maniraptoran theropod. Scientific Reports 2016; 6: 35782." year="2016">
Funston
<emphasis id="B95E0D439971FFE3B8E306CBFEECF932" box="[221,269,1521,1545]" italics="true" pageId="28" pageNumber="29">et al.</emphasis>
2016
</bibRefCitation>
). Distal to the expanded proximal end, the fibula narrows anteroposteriorly. On the medial surface is a long (
<quantity id="4CD27CB49971FFE3B8B60515FF0DF97C" box="[136,236,1583,1607]" metricMagnitude="-2" metricUnit="m" metricValue="7.42" pageId="28" pageNumber="29" unit="mm" value="74.2">74.2 mm</quantity>
, ~22% of total length) fusiform attachment for the fibular crest of the tibia. At this level, on the anterior surface of the fibula, an elliptical iliofibularis tubercle is present. Distally, the fibula strongly narrows anteroposteriorly and has a triangular cross-section along ~65% of its total length.
</paragraph>
<paragraph id="8B95D1519971FFE3B8A205F6FDC6F87A" blockId="28.[128,778,1207,1858]" pageId="28" pageNumber="29">
Despite the distal part of fibula being fused with the tibia in a similar manner to
<taxonomicName id="4C2AAAD29971FFE3B90205D6FE2EF838" box="[316,463,1772,1795]" class="Reptilia" family="Avimimidae" genus="Avimimus" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B95E0D439971FFE3B90205D6FE7FF838" box="[316,414,1772,1795]" italics="true" pageId="28" pageNumber="29">Avimimus</emphasis>
spp.
</taxonomicName>
, the proximal end of the fibula is more triangular, as in MPC-D 100/119 (Barsbold
<emphasis id="B95E0D439971FFE3BAAC0431FD20F818" box="[658,705,1803,1827]" italics="true" pageId="28" pageNumber="29">et al.</emphasis>
2000), than rectangular, as seen in
<taxonomicName id="4C2AAAD29971FFE3B9A80410FDC6F87A" box="[406,551,1834,1857]" class="Reptilia" family="Avimimidae" genus="Avimimus" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B95E0D439971FFE3B9A80410FE19F87A" box="[406,504,1834,1857]" italics="true" pageId="28" pageNumber="29">Avimimus</emphasis>
spp.
</taxonomicName>
</paragraph>
<paragraph id="8B95D1519971FFE3B8BE0456FA97FD04" blockId="28.[128,777,1898,1986]" lastBlockId="28.[825,1475,144,1077]" pageId="28" pageNumber="29">
<emphasis id="B95E0D439971FFE3B8BE0456FEF2F8BF" box="[128,275,1900,1924]" italics="true" pageId="28" pageNumber="29">Pedal phalanx:</emphasis>
The left phalanx II-2 is
<quantity id="4CD27CB49971FFE3B9C50456FDA2F8B8" box="[507,579,1900,1924]" metricMagnitude="-2" metricUnit="m" metricValue="3.5" pageId="28" pageNumber="29" unit="mm" value="35.0">35 mm</quantity>
long (
<figureCitation id="1311CDD49971FFE3BABD0456FD24F8BF" box="[643,709,1900,1924]" captionStart="Figure 19" captionStartId="24.[130,195,1427,1451]" captionTargetBox="[129,1473,144,1399]" captionTargetId="figure-251@24.[129,1473,144,1399]" captionTargetPageId="24" captionText="Figure 19. Various bones of Caenagnathidae indet. ZPAL MgD-I/108/1. AF, caudal vertebra in anterior (A), left lateral (B), posterior (C), right lateral (D), dorsal (E), and ventral (F) view. G, H, proximal part of dorsal rib. IN, manus ungual II-3 in dorsal (I), medial (J), ventral (K), lateral (L), anterior (M), and posterior (N) view.OT, left manus phalanx II-1 in medial (O), ventral (P), lateral (Q), dorsal (R), anterior (S), and posterior (T) view.UAʹ, left pedal phalanx II-2 in medial (U), ventral (W), lateral (X), dorsal (Y), anterior (Z), and posterior (Aʹ) view." figureDoi="http://doi.org/10.5281/zenodo.14284269" httpUri="https://zenodo.org/record/14284269/files/figure.png" pageId="28" pageNumber="29">Fig. 19</figureCitation>
U-A
<emphasis id="B95E0D439971FFE3BACD0450FD18F8B8" box="[755,761,1898,1923]" italics="true" pageId="28" pageNumber="29">ʹ</emphasis>
), and its length to width ratio is two. The proximal articular surface is triangular in posterior view and can be divided into lateral and medial teardrop-shaped concave articular surfaces, separated from each other by a smooth ridge running in the middle of the proximal articular surface. In the lateral and medial views, the proximal articular surface is strongly concave, the plantar margin extends backwards, and the lip-shaped dorsal margin (extensor turbecle) is elevated dorsally and directed posteriorly. The distal articular surface is composed of the lateral condyle and slightly shorter plantodorsally medial condyle, which are separated by a concavity that is shallow dorsally but becomes deeper along the articular surface to its end on the plantar side. In anterior view, the lateral condyle extends further downwards than the medial condyle. The ligament pits are well marked on the both sides of the phalanx; the lateral ligament pit is elongated anteroposteriorly, and the medial ligament pit is circular.
</paragraph>
<paragraph id="8B95D1519971FFE3BB6B017DFABCFB0E" blockId="28.[825,1475,144,1077]" pageId="28" pageNumber="29">
The phalanx II-2 is similar to the corresponding phalanx of
<taxonomicName id="4C2AAAD29971FFE3BB07015CFC05FD46" box="[825,996,614,638]" class="Reptilia" family="Caenagnathidae" genus="Elmisaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="rarus">
<emphasis id="B95E0D439971FFE3BB07015CFC05FD46" box="[825,996,614,638]" italics="true" pageId="28" pageNumber="29">Elmisaurus rarus</emphasis>
</taxonomicName>
(ZPAL MgD-I/98; length to width ratio of 2.1), especially in the structure of the proximal articular surface, i.e. two teardrop-shaped surfaces separated by a ridge, and the lip-like extensor tubercle. The phalanx II-
<quantity id="4CD27CB49971FFE3BD2D01FEFADAFDE7" box="[1299,1339,708,732]" metricMagnitude="-2" metricUnit="m" metricValue="5.08" pageId="28" pageNumber="29" unit="in" value="2.0">2 in</quantity>
other theropods from the Nemegt formation differs from the one of ZPAL MgD-I/108/1. This phalanx in
<taxonomicName id="4C2AAAD29971FFE3BC420039FB15FC21" box="[1148,1268,770,794]" class="Reptilia" family="Ornithomimidae" genus="Gallimimus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="bullatus">
<emphasis id="B95E0D439971FFE3BC420039FB15FC21" box="[1148,1268,770,794]" italics="true" pageId="28" pageNumber="29">Ga. bullatus</emphasis>
</taxonomicName>
(ZPAL MgD-I/94) is compressed (the length to width ratio is 1.4), and the extensor tubercle is less pronounced than in ZPAL MgD-I/108/1. The phalanx II-2 is even more compressed in
<taxonomicName id="4C2AAAD29971FFE3BCD6005BFA93FC43" authorityName="Osmolska" authorityYear="1987" box="[1256,1394,864,888]" class="Reptilia" family="Troodontidae" genus="Borogovia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="gracilicrus">
<emphasis id="B95E0D439971FFE3BCD6005BFA93FC43" box="[1256,1394,864,888]" italics="true" pageId="28" pageNumber="29">Bo. gracilicrus</emphasis>
</taxonomicName>
(ZPAL MgD-I/174; the length to width ratio is 1.2). The proximal articular surface in
<taxonomicName id="4C2AAAD29971FFE3BBD2009AFBB2FC8C" box="[1004,1107,927,951]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D439971FFE3BBD2009AFBB2FC8C" box="[1004,1107,927,951]" italics="true" pageId="28" pageNumber="29">Ta. bataar</emphasis>
</taxonomicName>
(ZPAL MgD-I/3, ZPAL MgD-I/4, ZPAL MgD-I/29, and ZPAL MgD-I/175) is wider than long, in contrast to
<taxonomicName id="4C2AAAD29971FFE3BBF100E4FB99FCCE" box="[975,1144,990,1014]" class="Reptilia" family="Caenagnathidae" genus="Elmisaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="rarus">
<emphasis id="B95E0D439971FFE3BBF100E4FB99FCCE" box="[975,1144,990,1014]" italics="true" pageId="28" pageNumber="29">Elmisaurus rarus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C2AAAD29971FFE3BCB800E4FAE1FCCD" box="[1158,1280,990,1014]" class="Reptilia" family="Ornithomimidae" genus="Gallimimus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="bullatus">
<emphasis id="B95E0D439971FFE3BCB800E4FAE1FCCD" box="[1158,1280,990,1014]" italics="true" pageId="28" pageNumber="29">Ga. bullatus</emphasis>
</taxonomicName>
, and ZPAL MgDI/108/1. The length to width ratio of the phalanx II-
<quantity id="4CD27CB49971FFE3BD4E00C7FA78FB2E" box="[1392,1433,1021,1045]" metricMagnitude="-2" metricUnit="m" metricValue="5.08" pageId="28" pageNumber="29" unit="in" value="2.0">2 in</quantity>
<taxonomicName id="4C2AAAD29971FFE3BD9D00C4FC9BFB0F" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="bataar">
<emphasis id="B95E0D439971FFE3BD9D00C4FC9BFB0F" italics="true" pageId="28" pageNumber="29">Ta. bataar</emphasis>
</taxonomicName>
is 1.51.6, depending on the ontogenetical age.
</paragraph>
</subSubSection>
</treatment>
</document>