1733 lines
195 KiB
XML
1733 lines
195 KiB
XML
<document ID-DOI="10.5281/zenodo.3374526" ID-GBIF-Dataset="2a40cf95-9ffb-4f3c-894b-cf53b8788a0a" ID-Zenodo-Dep="3374526" approvalRequired="109" approvalRequired_for_document="1" approvalRequired_for_illustrations="43" approvalRequired_for_originalDoi="1" approvalRequired_for_taxonomicNames="38" approvalRequired_for_textStreams="1" approvalRequired_for_treatments="25" checkinTime="1566477704804" checkinUser="jeremy" docAuthor="Holtz, TR jr." docDate="2004" docId="012B87EDFF9AD81B4F6D2639DC99B431" docLanguage="en" docName="Holtz2004Tyrannosauroidea.pdf" docOrigin="The Dinosauria, University of California Press" docStyle="DocumentStyle{}" docTitle="Tyrannosauroidea" docType="treatment" docVersion="19" lastPageNumber="7" masterDocId="FD12FF95FF9AD80B4D3D2775DD29B37C" masterDocTitle="Tyrannosauroidea" masterLastPageNumber="136" masterPageNumber="111" pageNumber="1" updateTime="1656386983924" updateUser="felipe">
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<mods:titleInfo>
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<mods:title>Tyrannosauroidea</mods:title>
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<mods:name type="personal">
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Holtz, TR jr.</mods:namePart>
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<mods:dateIssued>2004</mods:dateIssued>
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<mods:publisher>University of California Press</mods:publisher>
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<mods:roleTerm>Editor</mods:roleTerm>
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<mods:namePart>Weishampel DB</mods:namePart>
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<mods:namePart>Dodson P</mods:namePart>
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<mods:namePart>Osmolska H</mods:namePart>
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<mods:titleInfo>
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<mods:title>The Dinosauria</mods:title>
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<mods:identifier type="DOI">10.5281/zenodo.3374526</mods:identifier>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.3483196" ID-GBIF-Taxon="163642024" ID-Zenodo-Dep="3483196" LSID="urn:lsid:plazi:treatment:012B87EDFF9AD81B4F6D2639DC99B431" httpUri="http://treatment.plazi.org/id/012B87EDFF9AD81B4F6D2639DC99B431" lastPageId="16" lastPageNumber="7" pageId="0" pageNumber="1">
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<subSubSection box="[592,870,332,367]" pageId="0" pageNumber="1" type="nomenclature">
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<paragraph blockId="0.[592,870,332,367]" box="[592,870,332,367]" pageId="0" pageNumber="1">
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<heading box="[592,870,332,367]" centered="true" class="Reptilia" fontSize="14" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" level="0" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="superfamily" reason="1" superfamily="Tyrannosauroidea">
|
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<taxonomicName box="[592,870,332,367]" class="Reptilia" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="superFamily" superFamily="Tyrannosauroidea">
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<docTitle box="[592,870,332,367]" pageId="0" pageNumber="1">Tyrannosauroidea</docTitle>
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</taxonomicName>
|
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</heading>
|
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</paragraph>
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</subSubSection>
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<subSubSection lastPageId="16" lastPageNumber="7" pageId="0" pageNumber="1" type="nomenclature">
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<paragraph blockId="0.[601,861,423,443]" box="[601,861,423,443]" pageId="0" pageNumber="1">THOMAS R. HOLTZ JR.</paragraph>
|
||
<paragraph blockId="0.[99,713,731,1926]" pageId="0" pageNumber="1">
|
||
Tyrannosauroids are among the most distinctive and best known of Mesozoic theropods (
|
||
<tableCitation box="[343,433,759,779]" pageId="0" pageNumber="1" startId="1.[685,741,104,117]" targetBox="[99,1357,196,1687]" targetIsTable="true" targetPageId="1">table 5.1</tableCitation>
|
||
). They are characterized by large skulls with a specialized, heterodont dentition, a derived squamosal-quadratojugal flange, and a highly pneumatic basicranium; greatly reduced forelimbs (both in size and in digit count); and elongate hindlimbs bearing a pinched third metatarsal (arctometatarsus).
|
||
</paragraph>
|
||
<paragraph blockId="0.[99,713,731,1926]" pageId="0" pageNumber="1">
|
||
In recent years new discoveries and the use of modern phylogenetic analytical techniques have revealed the presence of taxa outside
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[174,336,989,1008]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
<emphasis box="[340,448,990,1008]" italics="true" pageId="0" pageNumber="1">sensu stricto</emphasis>
|
||
but closer to this clade than to all other theropod groups. These include the newly described Early Cretaceous
|
||
<taxonomicName authorityName="Hutt, Naish, Martill, Barker, et Newberry" authorityYear="2001" box="[269,423,1047,1066]" class="Reptilia" family="Not" genus="Eotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="lengi">
|
||
<emphasis box="[269,423,1047,1066]" italics="true" pageId="0" pageNumber="1">Eotyrannus lengi</emphasis>
|
||
</taxonomicName>
|
||
(Hutt et al. 2001; Naish et al. 2001), the Late Jurassic
|
||
<taxonomicName authorityName="Madsen" authorityYear="1974" box="[331,458,1075,1094]" class="Reptilia" family="Allosauridae" genus="Stokesosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="genus">
|
||
<emphasis box="[331,458,1075,1094]" italics="true" pageId="0" pageNumber="1">Stokesosaurus</emphasis>
|
||
</taxonomicName>
|
||
(Madsen 1974; Foster and Chure 2000; Rauhut 2000a), and the Late Cretaceous
|
||
<taxonomicName baseAuthorityName="Cope" baseAuthorityYear="1866" class="Reptilia" family="Not" genus="Dryptosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="aquilunguis">
|
||
<emphasis italics="true" pageId="0" pageNumber="1">Dryptosaurus aquilunguis</emphasis>
|
||
</taxonomicName>
|
||
(Cope 1866; Carpenter et al. 1997; Carr et al., in press). The latter is historically significant, for it was the taxon that revealed that carnivorous dinosaurs were obligate bipeds rather than quadrupeds (Cope 1866). Together with
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
, these aforementioned taxa are members of Tyrannosauroidea, a clade of basal coelurosaurs. As recognized here there are at least six, and possibly nine, genera within
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
, as well as three genera for which a basal tyrannosauroid position is well supported. Based on more questionable evidence, basal tyrannosauroids may be additional taxa.
|
||
</paragraph>
|
||
<paragraph blockId="0.[99,713,731,1926]" pageId="0" pageNumber="1">
|
||
Whereas most other taxa of large-bodied theropods (abelisaurids, spinosauroids, carnosaurs) are represented by only a few, often incomplete skeletons, several species of tyrannosaurids are represented by multiple complete skulls and postcrania
|
||
<emphasis italics="true" pageId="0" pageNumber="1">
|
||
(
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[176,357,1534,1553]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
|
||
,
|
||
<taxonomicName baseAuthorityName="Maleev" baseAuthorityYear="1955" box="[368,555,1534,1553]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="bataar">Tarbosaurus bataar</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="torosus">Daspletosaurus torosus</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[178,428,1563,1582]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="sarcophagus">Albertosaurus sarcophagus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and
|
||
<emphasis box="[484,684,1563,1582]" italics="true" pageId="0" pageNumber="1">
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[484,677,1563,1582]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="libratus">Gorgosaurus libratus</taxonomicName>
|
||
)
|
||
</emphasis>
|
||
of various ontogenetic stages. Among multi-tonne theropods only
|
||
<taxonomicName baseAuthorityName="Marsh" baseAuthorityYear="1877" box="[149,315,1620,1639]" class="Reptilia" family="Allosauridae" genus="Allosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="fragilis">
|
||
<emphasis box="[149,315,1620,1639]" italics="true" pageId="0" pageNumber="1">Allosaurus fragilis</emphasis>
|
||
</taxonomicName>
|
||
is better represented in the fossil record. Furthermore, tyrannosaurids are characterized by copious synapomorphies relative to other well-known theropod clades; basal tyrannosauroids reveal some of the sequence of acquisition of these features.
|
||
</paragraph>
|
||
<paragraph blockId="0.[99,713,731,1926]" lastBlockId="0.[751,1364,731,1925]" pageId="0" pageNumber="1">
|
||
Diagnostic skeletal material for
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[445,611,1763,1782]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
is at present limited to the last part of the Late Cretaceous (Campanian and Maastrichtian) of eastern and central Asia (China and Mongolia) and North America; the more inclusive Tyrannosauroidea includes taxa from the Late Jurassic of Europe and North America and the Early Cretaceous of Europe and Asia (and possibly South America). Abundant bones of tyrannosauroids (likely tyrannosaurids) are present throughout the Late Cretaceous of western North America and eastern and central Asia (Weishampel et al., this vol.); however, due to their incompleteness they cannot be definitively referred to
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1070,1236,846,865]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
as that taxon is defined here. The oldest confirmed tyrannosauroid is
|
||
<taxonomicName authorityName="Madsen" authorityYear="1974" class="Reptilia" family="Allosauridae" genus="Stokesosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="genus">
|
||
<emphasis italics="true" pageId="0" pageNumber="1">Stokesosaurus</emphasis>
|
||
</taxonomicName>
|
||
from the Kimmeridgian of the western United States (the same taxon or a related form is present in deposits of the same age in Portugal [Rauhut 2000b]). Based on phylogenetic inference, however, if
|
||
<taxonomicName authorityName="Xu, Zhao & Clark" authorityYear="2001" box="[926,1054,990,1009]" class="Reptilia" family="Therizinosauridae" genus="Eshanosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="genus">
|
||
<emphasis box="[926,1054,990,1009]" italics="true" pageId="0" pageNumber="1">Eshanosaurus</emphasis>
|
||
</taxonomicName>
|
||
from the Hettangian of China (Xu et al. 2001a) proves to be a member of Therizinosauroidea (implying a split between tyrannosauroids and maniraptoriforms, including therizinosauroids, had already occurred by the Early Jurassic), then tyrannosauroids were present from the Early Jurassic. The latest Maastrichtian western North American
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis italics="true" pageId="0" pageNumber="1">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
(and
|
||
<emphasis box="[963,1193,1162,1181]" italics="true" pageId="0" pageNumber="1">
|
||
<taxonomicName baseAuthorityName="Gilmore" baseAuthorityYear="1946" box="[963,1190,1162,1181]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="lancensis">Nanotyrannus lancensis</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
if it is a distinct taxon) is the latest-known tyrannosaurid and among the latestknown nonavian theropods.
|
||
</paragraph>
|
||
<paragraph blockId="0.[751,1364,731,1925]" pageId="0" pageNumber="1">Tyrannosauroids make up the sole large-bodied carnivorous theropods known from the Campanian–Maastrichtian of central and eastern Asia and North America; presumably these taxa were the top predators of those communities. Before that, however, tyrannosauroids were of only moderate body size and lived in faunas in which ceratosaurs, spinosauroids, and/or carnosaurs represented the largest carnivorous theropods (Foster et al. 2001; Farlow and Holtz 2002).</paragraph>
|
||
<paragraph blockId="0.[751,1364,731,1925]" lastBlockId="3.[99,716,960,1237]" lastPageId="3" lastPageNumber="4" pageId="0" pageNumber="1">
|
||
Tyrannosaurids include some of the largest known theropod taxa (up to 13 m long and weighing six or more tonnes). The five most completely known species
|
||
<emphasis italics="true" pageId="0" pageNumber="1">
|
||
(
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1070,1242,1535,1554]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
|
||
,
|
||
<taxonomicName baseAuthorityName="Maleev" baseAuthorityYear="1955" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="bataar">Tarbosaurus bataar</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[819,1027,1563,1582]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="torosus">Daspletosaurus torosus</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1034,1275,1563,1582]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="sarcophagus">Albertosaurus sarcophagus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and
|
||
<emphasis italics="true" pageId="0" pageNumber="1">
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="libratus">Gorgosaurus libratus</taxonomicName>
|
||
)
|
||
</emphasis>
|
||
are all represented by individuals with femora more than 100 cm in length, reaching 138 cm in the largest
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[751,929,1650,1668]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[751,929,1650,1668]" italics="true" pageId="0" pageNumber="1">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
(Brochu 2002). The basal tyrannosauroid
|
||
<taxonomicName authorityName="Marsh" authorityYear="1877" box="[751,875,1678,1696]" class="Reptilia" family="Dryptosauridae" genus="Dryptosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="genus">
|
||
<emphasis box="[751,875,1678,1696]" italics="true" pageId="0" pageNumber="1">Dryptosaurus</emphasis>
|
||
</taxonomicName>
|
||
does not reach this size, but with a femoral length of more than 77 cm it was still a large-bodied theropod (Carpenter et al. 1997). Skulls exceeding 1 m in length are known for
|
||
<emphasis box="[786,1056,1764,1783]" italics="true" pageId="0" pageNumber="1">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[786,926,1764,1782]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Maleev" authorityYear="1955" box="[936,1052,1764,1783]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="genus">Tarbosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and
|
||
<emphasis box="[1106,1325,1764,1783]" italics="true" pageId="0" pageNumber="1">
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[1106,1318,1764,1783]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="torosus">Daspletosaurus torosus</taxonomicName>
|
||
;
|
||
</emphasis>
|
||
the premaxilla-occipital condyle length of the largest
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1255,1307,1793,1811]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1255,1307,1793,1811]" italics="true" pageId="0" pageNumber="1">T. rex</emphasis>
|
||
</taxonomicName>
|
||
skull (153 cm) exceeds the same measurement of the skull of
|
||
<taxonomicName authorityName="Coria & Salgado" authorityYear="1995" class="Reptilia" family="Carcharodontosauridae" genus="Giganotosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="genus">
|
||
<emphasis italics="true" pageId="0" pageNumber="1">Giganotosaurus</emphasis>
|
||
</taxonomicName>
|
||
(148 cm). However, despite being among the largest known theropods, tyrannosaurids are also characterized by proportionately elongate and slender hindlimbs. Tyrannosaurids are markedly heterodont for theropods, and the D- or U-shaped premaxillary teeth of all tyrannosaurids and incrassate (thickened) “lateral” (i.e., maxillary and dentary) teeth of tyrannosaurids are easy to distinguish from those of nontyrannosaurid taxa.
|
||
</paragraph>
|
||
<paragraph blockId="1.[648,815,100,150]" pageId="1" pageNumber="112">TABLE 5.1 Tyrannosauroidea</paragraph>
|
||
<paragraph box="[611,1154,196,215]" pageId="1" pageNumber="112">
|
||
<tableNote box="[611,1154,196,215]" pageId="1" pageNumber="112">
|
||
<emphasis box="[611,711,196,215]" italics="true" pageId="1" pageNumber="112">Occurrence</emphasis>
|
||
<emphasis box="[899,932,196,215]" italics="true" pageId="1" pageNumber="112">Age</emphasis>
|
||
<emphasis box="[1075,1154,196,215]" italics="true" pageId="1" pageNumber="112">Material</emphasis>
|
||
</tableNote>
|
||
</paragraph>
|
||
<paragraph blockId="2.[611,1154,164,183]" box="[611,1154,164,183]" pageId="2" pageNumber="113">
|
||
<tableNote box="[611,1154,164,183]" pageId="2" pageNumber="113">
|
||
<emphasis box="[611,711,164,183]" italics="true" pageId="2" pageNumber="113">Occurrence</emphasis>
|
||
<emphasis box="[899,932,164,183]" italics="true" pageId="2" pageNumber="113">Age</emphasis>
|
||
<emphasis box="[1075,1154,164,183]" italics="true" pageId="2" pageNumber="113">Material</emphasis>
|
||
</tableNote>
|
||
</paragraph>
|
||
<paragraph blockId="2.[1265,1363,1908,1927]" box="[1265,1363,1908,1927]" pageId="2" pageNumber="113">
|
||
<emphasis box="[1265,1363,1908,1927]" italics="true" pageId="2" pageNumber="113">
|
||
<tableNote box="[1265,1363,1908,1927]" pageId="2" pageNumber="113">(continued)</tableNote>
|
||
</emphasis>
|
||
</paragraph>
|
||
<paragraph blockId="3.[611,1154,164,183]" box="[611,1154,164,183]" pageId="3" pageNumber="4">
|
||
<tableNote box="[611,1154,164,183]" pageId="3" pageNumber="4">
|
||
<emphasis box="[611,711,164,183]" italics="true" pageId="3" pageNumber="4">Occurrence</emphasis>
|
||
<emphasis box="[899,932,164,183]" italics="true" pageId="3" pageNumber="4">Age</emphasis>
|
||
<emphasis box="[1075,1154,164,183]" italics="true" pageId="3" pageNumber="4">Material</emphasis>
|
||
</tableNote>
|
||
</paragraph>
|
||
<paragraph blockId="3.[119,992,527,845]" box="[119,773,527,546]" pageId="3" pageNumber="4">
|
||
<emphasis box="[119,251,527,546]" italics="true" pageId="3" pageNumber="4">Nomina dubia</emphasis>
|
||
<emphasis box="[694,773,527,546]" italics="true" pageId="3" pageNumber="4">Material</emphasis>
|
||
</paragraph>
|
||
<paragraph blockId="3.[119,992,527,845]" box="[119,749,586,605]" pageId="3" pageNumber="4">
|
||
<treatmentCitation author="Marsh" box="[119,406,586,605]" httpUri="http://treatment.plazi.org/id/03B58786-FFCE-FFC1-48A6-F911FC2DD183 " pageId="3" pageNumber="4" rank="species" treatmentCitationLabel="nomenDubium" year="1892">
|
||
<taxonomicName authorityName="Marsh" authorityYear="1892" box="[119,406,586,605]" class="Reptilia" family="Tyrannosauridae" genus="Aublysodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="amplus" status="nomen dubium">
|
||
<emphasis box="[119,288,586,605]" italics="true" pageId="3" pageNumber="4">Aublysodon amplus</emphasis>
|
||
Marsh, 1892
|
||
</taxonomicName>
|
||
</treatmentCitation>
|
||
Tooth
|
||
</paragraph>
|
||
<paragraph blockId="3.[119,992,527,845]" box="[119,749,613,632]" pageId="3" pageNumber="4">
|
||
<treatmentCitation author="Marsh" box="[119,416,613,632]" httpUri="http://treatment.plazi.org/id/03B58786-FFCE-FFC6-48A6-F7C6FB32D794 " pageId="3" pageNumber="4" rank="species" treatmentCitationLabel="nomenDubium" year="1892">
|
||
<taxonomicName authorityName="Marsh" authorityYear="1892" box="[119,416,613,632]" class="Reptilia" family="Tyrannosauridae" genus="Aublysodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="cristatus" status="nomen dubium">
|
||
<emphasis box="[119,298,613,632]" italics="true" pageId="3" pageNumber="4">Aublysodon cristatus</emphasis>
|
||
Marsh, 1892
|
||
</taxonomicName>
|
||
</treatmentCitation>
|
||
Tooth
|
||
</paragraph>
|
||
<paragraph blockId="3.[119,992,527,845]" box="[119,745,639,658]" pageId="3" pageNumber="4">
|
||
<treatmentCitation author="Cope" box="[119,409,639,658]" httpUri="http://treatment.plazi.org/id/03BCF91D-FFDE-2E6A-FE5E-FCADB2D7F85F " pageId="3" pageNumber="4" rank="species" treatmentCitationLabel="nomenDubium" year="1876">
|
||
<taxonomicName authorityName="Cope" authorityYear="1876" box="[119,409,639,658]" class="Reptilia" family="Tyrannosauridae" genus="Aublysodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="lateralis" status="nomen dubium">
|
||
<emphasis box="[119,294,639,658]" italics="true" pageId="3" pageNumber="4">Aublysodon lateralis</emphasis>
|
||
Cope, 1876a
|
||
</taxonomicName>
|
||
</treatmentCitation>
|
||
Teeth
|
||
</paragraph>
|
||
<paragraph blockId="3.[119,992,527,845]" box="[119,864,666,685]" pageId="3" pageNumber="4">
|
||
<treatmentCitation author="Leidy" box="[119,415,666,685]" httpUri="http://treatment.plazi.org/id/BB1087B9-FFBC-FFC1-C482-8274FB1A9E19 " pageId="3" pageNumber="4" rank="species" treatmentCitationLabel="nomenDubium" year="1868">
|
||
<taxonomicName authorityName="Leidy" authorityYear="1868" box="[119,415,666,685]" class="Reptilia" family="Tyrannosauridae" genus="Aublysodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="mirandus" status="nomen dubium">
|
||
<emphasis box="[119,307,666,685]" italics="true" pageId="3" pageNumber="4">Aublysodon mirandus</emphasis>
|
||
Leidy, 1868
|
||
</taxonomicName>
|
||
</treatmentCitation>
|
||
Premaxillary teeth
|
||
</paragraph>
|
||
<paragraph blockId="3.[119,992,527,845]" box="[119,745,693,712]" pageId="3" pageNumber="4">
|
||
<treatmentCitation author="Leidy" box="[119,388,693,712]" httpUri="http://treatment.plazi.org/id/03CE87F5-FF86-725D-FF13-DDAC88A2FD3A " pageId="3" pageNumber="4" rank="species" treatmentCitationLabel="nomenDubium" year="1856">
|
||
<taxonomicName authorityName="Leidy" authorityYear="1856" box="[119,388,693,712]" class="Reptilia" family="Tyrannosauridae" genus="Deinodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="horridus" status="nomen dubium">
|
||
<emphasis box="[119,279,693,712]" italics="true" pageId="3" pageNumber="4">Deinodon horridus</emphasis>
|
||
Leidy, 1856
|
||
</taxonomicName>
|
||
</treatmentCitation>
|
||
Teeth
|
||
</paragraph>
|
||
<paragraph blockId="3.[119,992,527,845]" box="[119,992,719,738]" pageId="3" pageNumber="4">
|
||
<treatmentCitation author="Cope" box="[119,380,719,738]" httpUri="http://treatment.plazi.org/id/03BD87E2-D529-FFF5-FF7D-D7351A93F777 " pageId="3" pageNumber="4" rank="species" treatmentCitationLabel="nomenDubium" year="1868">
|
||
<taxonomicName authority="Cope, 1868" authorityName="Cope" authorityYear="1868" box="[119,380,719,738]" class="Arachnida" family="Ologamasidae" genus="Laelaps" higherTaxonomySource="CoL" kingdom="Animalia" order="Mesostigmata" pageId="3" pageNumber="4" phylum="Arthropoda" rank="species" species="macropus" status="nomen dubium">
|
||
<emphasis box="[119,272,719,738]" italics="true" pageId="3" pageNumber="4">Laelaps macropus</emphasis>
|
||
Cope, 1868
|
||
</taxonomicName>
|
||
</treatmentCitation>
|
||
Fragmentary hindlimb elements
|
||
</paragraph>
|
||
<paragraph blockId="3.[119,992,527,845]" box="[119,749,746,765]" pageId="3" pageNumber="4">
|
||
<taxonomicName authority="Leidy, 1865" authorityName="Leidy" authorityYear="1865" box="[119,396,746,765]" class="Reptilia" family="Colubridae" genus="Tomodon" kingdom="Animalia" order="Squamata" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="horrificus" status="nomen dubium">
|
||
<emphasis box="[119,287,746,765]" italics="true" pageId="3" pageNumber="4">Tomodon horrificus</emphasis>
|
||
Leidy, 1865
|
||
</taxonomicName>
|
||
Tooth
|
||
</paragraph>
|
||
<paragraph blockId="3.[119,992,527,845]" box="[139,463,773,792]" pageId="3" pageNumber="4">
|
||
(type of
|
||
<taxonomicName authority="Leidy, 1868 a" authorityName="Leidy" authorityYear="1868" box="[214,457,773,792]" class="Reptilia" family="Dryptosauridae" genus="Diplotomodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis box="[214,338,773,792]" italics="true" pageId="3" pageNumber="4">Diplotomodon</emphasis>
|
||
Leidy, 1868a
|
||
</taxonomicName>
|
||
)
|
||
</paragraph>
|
||
<paragraph blockId="3.[119,992,527,845]" box="[119,745,799,818]" pageId="3" pageNumber="4">
|
||
<taxonomicName authority="Yeh, 1975" authorityName="Yeh" authorityYear="1975" box="[119,451,799,818]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="lanpingensis" status="nomen dubium">
|
||
<emphasis box="[119,360,799,818]" italics="true" pageId="3" pageNumber="4">Tyrannosaurus lanpingensis</emphasis>
|
||
Yeh, 1975
|
||
</taxonomicName>
|
||
Teeth
|
||
</paragraph>
|
||
<paragraph blockId="3.[119,992,527,845]" box="[119,930,826,845]" pageId="3" pageNumber="4">
|
||
<taxonomicName authority="Zhai, Zheng, et Tong, 1978" authorityName="Zhai, Zheng, et Tong" authorityYear="1978" box="[119,598,826,845]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="turpanensis" status="nomen dubium">
|
||
<emphasis box="[119,352,826,845]" italics="true" pageId="3" pageNumber="4">Tyrannosaurus turpanensis</emphasis>
|
||
Zhai, Zheng, et Tong, 1978
|
||
</taxonomicName>
|
||
Teeth and sacral vertebrae
|
||
</paragraph>
|
||
<paragraph blockId="3.[99,716,960,1237]" pageId="3" pageNumber="4">
|
||
Most modern authors have followed D. A. Russell (1970a) in preferring Osborn’s (1906) name
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[423,591,1132,1151]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
, while many earlier workers (e.g., Matthew and Brown 1922, 1923; Huene 1923, 1926c; Maleev 1974) used
|
||
<taxonomicName authority="Cope, 1866" authorityName="Cope" authorityYear="1866" box="[410,556,1189,1208]" class="Reptilia" family="Deinodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="family">Deinodontidae</taxonomicName>
|
||
Brown, 1914, itself an emendation of
|
||
<taxonomicName authority="Cope, 1866" authorityName="Cope" authorityYear="1866" box="[317,573,1218,1237]" class="Reptilia" family="Dinodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="family">Dinodontidae Cope, 1866</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<paragraph blockId="3.[99,388,1306,1330]" box="[99,388,1306,1330]" pageId="3" pageNumber="4">
|
||
<heading bold="true" box="[99,388,1306,1330]" fontSize="10" level="1" pageId="3" pageNumber="4" reason="0">
|
||
<emphasis bold="true" box="[99,388,1306,1330]" pageId="3" pageNumber="4">Definition and Diagnosis</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="3.[99,716,1362,1926]" lastBlockId="3.[747,1364,960,1381]" pageId="3" pageNumber="4">
|
||
Tyrannosauroidea as used herein is defined as the clade composed of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[191,363,1392,1410]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[191,363,1392,1410]" italics="true" pageId="3" pageNumber="4">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
and all taxa sharing a more recent common ancestor with
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[344,397,1420,1438]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[344,397,1420,1438]" italics="true" pageId="3" pageNumber="4">T. rex</emphasis>
|
||
</taxonomicName>
|
||
than with
|
||
<emphasis italics="true" pageId="3" pageNumber="4">
|
||
<taxonomicName authorityName="Marsh" authorityYear="1890" box="[518,711,1420,1439]" class="Reptilia" family="Ornithomimidae" genus="Ornithomimus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="velox">Ornithomimus velox</taxonomicName>
|
||
,
|
||
<taxonomicName baseAuthorityName="Ostrom" baseAuthorityYear="1969" box="[99,328,1448,1467]" class="Reptilia" family="Dromaeosauridae" genus="Deinonychus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="antirrhopus">Deinonychus antirrhopus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
or
|
||
<emphasis box="[362,531,1448,1467]" italics="true" pageId="3" pageNumber="4">
|
||
<taxonomicName baseAuthorityName="Marsh" baseAuthorityYear="1877" box="[362,527,1448,1467]" class="Reptilia" family="Allosauridae" genus="Allosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="fragilis">Allosaurus fragilis</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Tyrannosauroidea, or at least a phylogenetically secure subclade therein comprising
|
||
<emphasis box="[99,490,1506,1525]" italics="true" pageId="3" pageNumber="4">
|
||
<taxonomicName authorityName="Madsen" authorityYear="1974" box="[99,229,1506,1525]" class="Reptilia" family="Allosauridae" genus="Stokesosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">Stokesosaurus</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Marsh" authorityYear="1877" box="[240,367,1506,1524]" class="Reptilia" family="Dryptosauridae" genus="Dryptosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">Dryptosaurus</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Hutt, Naish, Martill, Barker, et Newberry" authorityYear="2001" box="[379,486,1506,1524]" class="Reptilia" family="Not" genus="Eotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">Eotyrannus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[544,712,1505,1524]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
, may be diagnosed by the following synapomorphies: the main body of the premaxilla taller dorsoventrally than it is long rostrocaudally (also in
|
||
<taxonomicName authorityName="Bonaparte & Novas" authorityYear="1985" box="[284,424,1591,1610]" class="Reptilia" family="Abelisauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="family">Abelisauridae</taxonomicName>
|
||
); a prominent and laterally extending horizontal shelf on the lateral surface of the surangular, rostral and ventral to the mandibular condyle; a large opening for the caudal surangular foramen; the retroarticular process reduced; the dorsal surface of the iliac blades converging closely on the midline (also found in ornithomimosaurs and mononykine alvarezsaurids); the dorsal portion of the cranial margin of the preacetabular blade of the ilium concave cranially and the ventral portion convex; an accessory broad, ventral hooklike projection from the preacetabular blade of ilium; a straight supracetabular crest on the lateral margin of the ilium in dorsal view (also found in various maniraptorans); a median vertical ridge on the external surface of the ilium; a ventral flange on the puboischial peduncle; the cranial margin of the pubic peduncle concave; the pubis shaft having a marked concave curvature cranially (convergent with some oviraptorosaurs); the shaft of the ischium much more slender than the pubis shaft; a semicircular scar on the caudolateral surface of ischium, just distal to the iliac process (also in ornithomimosaurs); a sheet of bone extending from the obturator process continuing down at least half the length of the ischium; the fibular cranial tubercle distal to the cranial expansion composed of two longitudinal ridges; elongate metatarsals relative to those of other theropods having femora of the same length (also in
|
||
<emphasis box="[747,886,1304,1323]" italics="true" pageId="3" pageNumber="4">
|
||
<taxonomicName authorityName="Janensch" authorityYear="1920" box="[747,881,1304,1323]" class="Reptilia" family="Ornithomimidae" genus="Elaphrosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">Elaphrosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
ornithomimosaurs, and troodontids); and the proximal surface of metatarsal III crescentic and limited to the plantar half of the proximal surface of the metatarsus.
|
||
</paragraph>
|
||
<paragraph blockId="3.[747,851,1450,1474]" box="[747,851,1450,1474]" pageId="3" pageNumber="4">
|
||
<heading bold="true" box="[747,851,1450,1474]" fontSize="10" level="1" pageId="3" pageNumber="4" reason="0">
|
||
<emphasis bold="true" box="[747,851,1450,1474]" pageId="3" pageNumber="4">Anatomy</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="3.[747,1364,1505,1754]" pageId="3" pageNumber="4">Tyrannosauroid anatomy has been described by Lambe (1904a, 1917a), Osborn (1905, 1906, 1912a, 1917), Parks (1933), Gilmore (1946a), Russell (1970a), Madsen (1974), Maleev (1974), Bakker et al. (1988), Molnar et al. (1990), Molnar (1991), Carpenter et al. (1997), Chure and Madsen (1998), Carr (1999), Carr and Williamson (2000, in press), Foster and Chure (2000), Carpenter and Smith (2001), Hutt et al. (2001), Naish et al. (2001), Brochu (2002), Carr et al. (in press), Currie (2003b), and Hurum and Sabath (2003).</paragraph>
|
||
<paragraph blockId="3.[747,932,1826,1845]" box="[747,932,1826,1845]" pageId="3" pageNumber="4">
|
||
<heading bold="true" box="[747,932,1826,1845]" fontSize="8" level="2" pageId="3" pageNumber="4" reason="2">
|
||
<emphasis bold="true" box="[747,932,1826,1845]" pageId="3" pageNumber="4">Skull and Mandible</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="3.[747,1364,1879,1927]" lastBlockId="4.[99,716,1018,1927]" lastPageId="4" lastPageNumber="5" pageId="3" pageNumber="4">
|
||
Complete or near-complete skulls are known for
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1226,1364,1880,1898]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1226,1364,1880,1898]" italics="true" pageId="3" pageNumber="4">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
(Osborn 1912a; Gilmore 1946a; Bakker et al. 1988; Molnar 1991; Carr 1999; Brochu 2002; Carr and Williamson, in press),
|
||
<taxonomicName authorityName="Maleev" authorityYear="1955" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis italics="true" pageId="4" pageNumber="5">Tarbosaurus</emphasis>
|
||
</taxonomicName>
|
||
(Maleev 1974; Hurum and Sabath 2003),
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[574,715,1047,1066]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis box="[574,715,1047,1066]" italics="true" pageId="4" pageNumber="5">Daspletosaurus</emphasis>
|
||
</taxonomicName>
|
||
(Russell 1970a; Currie 2003b),
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[388,574,1076,1095]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="libratus">
|
||
<emphasis box="[388,574,1076,1095]" italics="true" pageId="4" pageNumber="5">Gorgosaurus libratus</emphasis>
|
||
</taxonomicName>
|
||
(Lambe 1917a; Carr 1999; Currie 2003b), and
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[390,514,1105,1124]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis box="[390,514,1105,1124]" italics="true" pageId="4" pageNumber="5">Albertosaurus</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation box="[525,642,1104,1124]" pageId="4" pageNumber="5" startId-0="4.[99,166,871,886]" startId-1="4.[747,815,1782,1797]" startId-2="5.[99,167,726,741]" targetBox-0="[103,707,99,847]" targetBox-1="[751,1359,99,1757]" targetBox-2="[103,710,100,700]" targetPageId-0="4" targetPageId-1="4" targetPageId-2="5">figs. 5.1–5.3</figureCitation>
|
||
; Lambe 1904a; Currie 2003b). Partial skulls are known for
|
||
<taxonomicName authorityName="Kurzanov" authorityYear="1976" class="Reptilia" family="Tyrannosauridae" genus="Alioramus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="remotus">
|
||
<emphasis italics="true" pageId="4" pageNumber="5">Alioramus remotus</emphasis>
|
||
</taxonomicName>
|
||
(Kurzanov 1976a; Currie 2003b) and the unnamed Alabama tyrannosaurid (Carr et al., in press). Cranial material is presently known for
|
||
<taxonomicName authorityName="Gilmore" authorityYear="1933" box="[302,484,1219,1238]" class="Reptilia" family="Tyrannosauridae" genus="Alectrosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="olseni">
|
||
<emphasis box="[302,484,1219,1238]" italics="true" pageId="4" pageNumber="5">Alectrosaurus olseni</emphasis>
|
||
</taxonomicName>
|
||
(Gilmore 1933a; Mader and Bradley 1989), although more recently discovered skulls may be referable to this taxon (Perle 1977). Only fragmentary cranial material is known for
|
||
<taxonomicName authorityName="Hutt, Naish, Martill, Barker, et Newberry" authorityYear="2001" box="[383,487,1306,1324]" class="Reptilia" family="Not" genus="Eotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis box="[383,487,1306,1324]" italics="true" pageId="4" pageNumber="5">Eotyrannus</emphasis>
|
||
</taxonomicName>
|
||
(Hutt et al. 2001; Naish et al. 2001),
|
||
<taxonomicName authorityName="Marsh" authorityYear="1877" box="[213,337,1335,1353]" class="Reptilia" family="Dryptosauridae" genus="Dryptosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis box="[213,337,1335,1353]" italics="true" pageId="4" pageNumber="5">Dryptosaurus</emphasis>
|
||
</taxonomicName>
|
||
(Cope 1866; Carpenter et al. 1997), and
|
||
<taxonomicName authority="(Osmolska 1996)" authorityName="Osmolska" authorityYear="1996" baseAuthorityName="Osmolska" baseAuthorityYear="1996" box="[99,380,1363,1382]" class="Reptilia" family="Not" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis box="[99,206,1363,1381]" italics="true" pageId="4" pageNumber="5">Bagaraatan</emphasis>
|
||
(Osmólska 1996)
|
||
</taxonomicName>
|
||
; cranial material referred to
|
||
<taxonomicName authorityName="Madsen" authorityYear="1974" class="Reptilia" family="Allosauridae" genus="Stokesosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis italics="true" pageId="4" pageNumber="5">Stokesosaurus</emphasis>
|
||
</taxonomicName>
|
||
(Madsen 1974; Chure and Madsen 1998) cannot be definitely referred to this taxon.
|
||
</paragraph>
|
||
<caption httpUri="https://zenodo.org/record/3374528/files/figure.png" pageId="4" pageNumber="5" startId="4.[99,166,871,886]" targetBox="[103,707,99,847]" targetPageId="4">
|
||
<paragraph blockId="4.[99,700,869,911]" pageId="4" pageNumber="5">
|
||
FIGURE 5.1. Skull of subadult
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[358,524,870,887]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="libratus">
|
||
<emphasis box="[358,524,870,887]" italics="true" pageId="4" pageNumber="5">Gorgosaurus libratus</emphasis>
|
||
</taxonomicName>
|
||
in dorsal
|
||
<emphasis box="[606,663,870,886]" italics="true" pageId="4" pageNumber="5">(upper)</emphasis>
|
||
and right lateral
|
||
<emphasis box="[203,259,894,911]" italics="true" pageId="4" pageNumber="5">(lower)</emphasis>
|
||
views.Scale = 10 cm. (After Carr 1999.)
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="4.[99,716,1018,1927]" lastBlockId="5.[99,716,874,1926]" lastPageId="5" lastPageNumber="6" pageId="4" pageNumber="5">
|
||
Tyrannosauroid skulls are long and low among small individuals (adults of small-bodied taxa and juveniles of larger species), deepening dorsoventrally in the adults of the largebodied taxa (
|
||
<figureCitation box="[230,350,1534,1554]" pageId="4" pageNumber="5" startId-0="4.[99,166,871,886]" startId-1="4.[747,815,1782,1797]" startId-2="5.[99,167,726,741]" targetBox-0="[103,707,99,847]" targetBox-1="[751,1359,99,1757]" targetBox-2="[103,710,100,700]" targetPageId-0="4" targetPageId-1="4" targetPageId-2="5">figs. 5.1–5.3</figureCitation>
|
||
; Carr 1999; Carr and Williamson, in press; Currie 2003b). In general form the skulls of most tyrannosaurids resemble those of other large carnivorous theropods and nondinosaurian archosaurs (e.g., poposaurids and rauisuchids) in being considerably deeper dorsoventrally than wide transversely; however,
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[280,457,1679,1697]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[280,457,1679,1697]" italics="true" pageId="4" pageNumber="5">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
has a more rounded rostrum, proportionately wider at the base than in more primitive tyrannosaurids. Where the skulls are complete, tyrannosaurids have a proportionately wider rostrum than do carnosaurs of similar size. In
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[242,290,1794,1812]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[242,290,1794,1812]" italics="true" pageId="4" pageNumber="5">T. rex</emphasis>
|
||
</taxonomicName>
|
||
this width is greatly increased postorbitally, so that the maximum skull width is approximately two-thirds of the premaxilla-occipital condyle length of the skull (compared with
|
||
<emphasis box="[150,255,1879,1898]" italics="true" pageId="4" pageNumber="5">
|
||
<taxonomicName box="[150,251,1879,1898]" class="Reptilia" family="Allosauridae" genus="Allosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">Allosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
for example, in which the comparable figure is only one-third of the skull base length, or the tyrannosaurid
|
||
<emphasis box="[99,247,875,894]" italics="true" pageId="5" pageNumber="6">
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[99,243,875,894]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Daspletosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
in which it is slightly less than half of the skull base length in). The cranial and some postdentary bones have extensive internal chambers. Tyrannosaurids have variable cranial ornamentation, including rugosities on the nasals, postorbitals, and jugals and hornlets on the lacrimals.
|
||
</paragraph>
|
||
<caption httpUri="https://zenodo.org/record/3374530/files/figure.png" pageId="4" pageNumber="5" startId="4.[747,815,1782,1797]" targetBox="[751,1359,99,1757]" targetPageId="4">
|
||
<paragraph blockId="4.[747,1292,1780,1846]" pageId="4" pageNumber="5">
|
||
FIGURE 5.2. Skull of adult tyrannosaurids: A,
|
||
<emphasis box="[1138,1292,1781,1797]" italics="true" pageId="4" pageNumber="5">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1138,1288,1781,1797]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
|
||
;
|
||
</emphasis>
|
||
B,
|
||
<emphasis box="[768,958,1805,1822]" italics="true" pageId="4" pageNumber="5">
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[768,953,1805,1822]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="torosus">Daspletosaurus torosus</taxonomicName>
|
||
;
|
||
</emphasis>
|
||
C,
|
||
<emphasis box="[985,1143,1805,1822]" italics="true" pageId="4" pageNumber="5">
|
||
<taxonomicName authorityName="Kurzanov" authorityYear="1976" box="[985,1139,1805,1822]" class="Reptilia" family="Tyrannosauridae" genus="Alioramus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="remotus">Alioramus remotus</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Scale = 50 cm. (Courtesy of T. L. Ford.)
|
||
</paragraph>
|
||
</caption>
|
||
<caption httpUri="https://zenodo.org/record/3374532/files/figure.png" pageId="5" pageNumber="6" startId="5.[99,167,726,741]" targetBox="[103,710,100,700]" targetPageId="5">
|
||
<paragraph blockId="5.[99,706,724,790]" pageId="5" pageNumber="6">
|
||
FIGURE 5.3. Skulls of juvenile tyrannosaurids: A,
|
||
<taxonomicName baseAuthorityName="Gilmore" baseAuthorityYear="1946" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="lancensis">
|
||
<emphasis italics="true" pageId="5" pageNumber="6">Nanotyrannus lancensis</emphasis>
|
||
</taxonomicName>
|
||
(possibly a juvenile
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[292,441,749,765]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[292,441,749,765]" italics="true" pageId="5" pageNumber="6">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
); B,
|
||
<emphasis italics="true" pageId="5" pageNumber="6">
|
||
“
|
||
<taxonomicName authorityName="Dong" authorityYear="1977" class="Reptilia" family="Tyrannosauridae" genus="Shanshanosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="houyanshanensis">Shanshanosaurus houyanshanensis</taxonomicName>
|
||
.”
|
||
</emphasis>
|
||
Scale = 5 cm. (After Carr 1999; and Currie and Dong 2001b.)
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="5.[99,716,874,1926]" pageId="5" pageNumber="6">
|
||
The nares are rostrocaudally elongate and bounded by the premaxillae, the maxillae, and the nasals. The orbits are oval, with the dorsoventral axis becoming proportionately longer as the skull size increases (except in
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[465,657,1104,1124]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="libratus">
|
||
<emphasis box="[465,657,1104,1124]" italics="true" pageId="5" pageNumber="6">Gorgosaurus libratus</emphasis>
|
||
</taxonomicName>
|
||
[Carr 1999; Carr and Williamson, in press]). The eyes of most tyrannosaurids are laterally directed, but those of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[510,680,1162,1180]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[510,680,1162,1180]" italics="true" pageId="5" pageNumber="6">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
are more rostrally directed, reoriented due to the postorbital expansion of the skull.The infratemporal fenestra of tyrannosaurids is distinctive, with a vertical rostral margin and a caudal margin shaped like a sideways
|
||
<emphasis box="[327,353,1277,1295]" italics="true" pageId="5" pageNumber="6">W.</emphasis>
|
||
The central constriction divides the fenestra in half due to a synapomorphic extension of both the squamosal and the quadratojugal, with the suture between the two subhorizontal and down the longest part of the protrusion. This differs from the constriction of the infratemporal fenestrae of basal tetanurans such as
|
||
<emphasis box="[435,716,1420,1439]" italics="true" pageId="5" pageNumber="6">
|
||
<taxonomicName authority="," box="[435,539,1420,1439]" class="Reptilia" family="Allosauridae" genus="Allosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Allosaurus,</taxonomicName>
|
||
<taxonomicName authority="Zhao & Currie 1993" authorityName="Zhao & Currie" authorityYear="1993" box="[545,712,1420,1439]" class="Reptilia" family="Not" genus="Monolophosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Monolophosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and
|
||
<emphasis box="[141,268,1448,1467]" italics="true" pageId="5" pageNumber="6">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1903" box="[141,265,1448,1467]" class="Reptilia" family="Compsognathidae" genus="Ornitholestes" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Ornitholestes</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
in which the squamosal forms the majority of the protrusion. The supratemporal fenestra is smaller than the infratemporal fenestra, situated entirely dorsally, and bounded by the frontal, parietal, supraoccipital, squamosal, and postorbital. The posttemporal fenestrae are greatly reduced. The choanae are located about halfway along the length of the skull and are surrounded by the maxilla, the palatine, and the vomer. The palatal surface is roofed by extensive medial projections from the premaxillae and maxillae.
|
||
</paragraph>
|
||
<paragraph blockId="5.[99,716,874,1926]" lastBlockId="5.[747,1364,101,1926]" pageId="5" pageNumber="6">
|
||
The premaxilla is short and deep in tyrannosauroids; the ventral ramus of the premaxilla is taller dorsoventrally than it is long rostrocaudally, and it is positioned so that it is more obscured by the maxilla in lateral view than in other theropods. There are only four premaxillary teeth. The premaxillary teeth of
|
||
<taxonomicName authorityName="Hutt, Naish, Martill, Barker, et Newberry" authorityYear="2001" box="[123,227,1850,1868]" class="Reptilia" family="Not" genus="Eotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[123,227,1850,1868]" italics="true" pageId="5" pageNumber="6">Eotyrannus</emphasis>
|
||
</taxonomicName>
|
||
and tyrannosaurids are arranged in an arcade that is much more transversely oriented than rostrocaudally oriented, so that the rostralmost surface of the last tooth is rostral to the caudalmost surface of the first tooth (in contrast to the case of most theropods, in which the premaxillary tooth arcade is arranged more closely to the main skull axis).
|
||
</paragraph>
|
||
<paragraph blockId="5.[747,1364,101,1926]" pageId="5" pageNumber="6">
|
||
The maxilla is an elongate, roughly triangular element with a rounded rostral apex and two caudal rami separated by the internal antorbital fenestra. The ventral margin of large tyrannosaurids has a convex curvature, most pronounced in the
|
||
<taxonomicName authorityName="Maleev" authorityYear="1955" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis italics="true" pageId="5" pageNumber="6">Tarbosaurus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<emphasis box="[858,1003,302,320]" italics="true" pageId="5" pageNumber="6">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[858,999,302,320]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
The maxillary antorbital fossa bears both promaxillary and maxillary fenestrae, the former often obscured in lateral view in larger individuals by a shelf on the lateral surface of the maxilla (Carr et al., in press). In tyrannosaurines the promaxillary fenestra rotates to face rostrally, and the rostral margin of the maxillary fenestra contacts the rostral rim of the antorbital fossa. Medially the maxillae of tyrannosaurids house a series of internal chambers (maxillary and promaxillary antra and pneumatic interalveolar recesses); as preserved these chambers are not separated from each other by shelves of bone (similar to the condition in
|
||
<taxonomicName box="[1177,1275,588,607]" class="Reptilia" family="Allosauridae" genus="Allosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1177,1275,588,607]" italics="true" pageId="5" pageNumber="6">Allosaurus</emphasis>
|
||
</taxonomicName>
|
||
[Witmer 1997a]). The number of maxillary teeth in tyrannosaurids varies: a maximum of 18 are reported in skulls referred to
|
||
<taxonomicName authorityName="Gilmore" authorityYear="1933" class="Reptilia" family="Tyrannosauridae" genus="Alectrosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="olseni">
|
||
<emphasis italics="true" pageId="5" pageNumber="6">Alectrosaurus olseni</emphasis>
|
||
</taxonomicName>
|
||
and a minimum of 11 in some
|
||
<emphasis box="[1185,1364,675,693]" italics="true" pageId="5" pageNumber="6">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1185,1361,675,693]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
The number decreases with skull size in
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[1166,1363,703,722]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="libratus">
|
||
<emphasis box="[1166,1363,703,722]" italics="true" pageId="5" pageNumber="6">Gorgosaurus libratus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[790,838,732,750]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[790,838,732,750]" italics="true" pageId="5" pageNumber="6">T. rex</emphasis>
|
||
</taxonomicName>
|
||
(accompanied by a proportional increase in the crosssectional size of the teeth during ontogeny, so that larger teeth exclude the cranialmost tooth position [see Currie 2003b for a dissenting view]). The interdental plates are large and rugose. The palatal shelves of the maxillae of tyrannosaurids are much more extensive then those of carnosaurs or other large theropods (
|
||
<figureCitation box="[808,896,903,923]" pageId="5" pageNumber="6" startId="6.[219,287,843,858]" targetBox="[224,1242,103,817]" targetPageId="6">fig. 5.4A</figureCitation>
|
||
). In most tyrannosaurids these shelves contact the vomers along the midline for the rostral half of the length of the tooth row: they angle back toward the tooth row forming a diagonal contact with the palatines along their caudomedial surface. In
|
||
<emphasis box="[854,999,1019,1037]" italics="true" pageId="5" pageNumber="6">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[854,994,1019,1037]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
however, these projections are much more extensive, contacting the vomers for more than threequarters of the length of the tooth row.
|
||
</paragraph>
|
||
<paragraph blockId="5.[747,1364,101,1926]" pageId="5" pageNumber="6">
|
||
The elongate nasals of
|
||
<taxonomicName authorityName="Hutt, Naish, Martill, Barker, et Newberry" authorityYear="2001" box="[1016,1120,1105,1123]" class="Reptilia" family="Not" genus="Eotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1016,1120,1105,1123]" italics="true" pageId="5" pageNumber="6">Eotyrannus</emphasis>
|
||
</taxonomicName>
|
||
and tyrannosaurids are fused along the midline of the dorsal surface, although the rostralmost and caudalmost ends are not entirely fused and the suture line is visible along the ventral surface. In adult tyrannosaurids the dorsal surface is rugose, and it can have an almost barklike texture in some cases.The nasals are penetrated by many foramina, suggesting that a horny covering was present. In
|
||
<taxonomicName authorityName="Kurzanov" authorityYear="1976" class="Reptilia" family="Tyrannosauridae" genus="Alioramus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis italics="true" pageId="5" pageNumber="6">Alioramus</emphasis>
|
||
</taxonomicName>
|
||
there is a row of five vertical blades along the midline, while in the Alabama tyrannosaurid there are six small bumps.
|
||
</paragraph>
|
||
<paragraph blockId="5.[747,1364,101,1926]" pageId="5" pageNumber="6">
|
||
The lacrimal is shaped like an inverted
|
||
<emphasis box="[1160,1178,1363,1381]" italics="true" pageId="5" pageNumber="6">L;</emphasis>
|
||
the rostral and descending rami of the lacrimals form about half of the dorsal and all of the caudal surface, respectively, of the internal antorbital fenestra. The rostral ramus contains internal pneumatic chambers; in adult
|
||
<taxonomicName authorityName="Maleev" authorityYear="1955" box="[896,1012,1477,1496]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[896,1012,1477,1496]" italics="true" pageId="5" pageNumber="6">Tarbosaurus</emphasis>
|
||
</taxonomicName>
|
||
and especially
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1175,1315,1477,1495]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1175,1315,1477,1495]" italics="true" pageId="5" pageNumber="6">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
this portion of the bone can have an inflated appearance. The dorsal surface bears a triangular cornual process in albertosaurines and
|
||
<emphasis box="[789,933,1563,1582]" italics="true" pageId="5" pageNumber="6">
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[789,928,1563,1582]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Daspletosaurus</taxonomicName>
|
||
;
|
||
</emphasis>
|
||
this prominence is lost in the
|
||
<taxonomicName authorityName="Maleev" authorityYear="1955" box="[747,860,1591,1610]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis italics="true" pageId="5" pageNumber="6">Tyrannosaurus-Tarbosaurus</emphasis>
|
||
</taxonomicName>
|
||
clade.
|
||
</paragraph>
|
||
<paragraph blockId="5.[747,1364,101,1926]" lastBlockId="6.[99,716,990,1926]" lastPageId="6" lastPageNumber="7" pageId="5" pageNumber="6">
|
||
The postorbital is a triradiate bone with longer descending and caudal rami and a shorter medial process. In
|
||
<emphasis italics="true" pageId="5" pageNumber="6">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1219,1359,1649,1667]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Maleev" authorityYear="1955" box="[747,863,1677,1696]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Tarbosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and
|
||
<emphasis box="[918,1051,1677,1696]" italics="true" pageId="5" pageNumber="6">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[918,1048,1677,1696]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Albertosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and to a much lesser degree in
|
||
<emphasis box="[747,895,1706,1725]" italics="true" pageId="5" pageNumber="6">
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[747,891,1706,1725]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Daspletosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
a prong of the postorbital constricts the orbit and demarcates a dorsal (eyeball-bearing) and ventral portion of the orbit; similar constrictions are found in the carnosaurs
|
||
<taxonomicName authorityName="Coria & Salgado" authorityYear="1995" box="[747,891,1793,1812]" class="Reptilia" family="Carcharodontosauridae" genus="Giganotosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[747,891,1793,1812]" italics="true" pageId="5" pageNumber="6">Giganotosaurus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<emphasis box="[944,1148,1792,1811]" italics="true" pageId="5" pageNumber="6">
|
||
<taxonomicName authorityName="Stromer" authorityYear="1931" box="[944,1143,1792,1811]" class="Reptilia" family="Carcharodontosauridae" genus="Carcharodontosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Carcharodontosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
some
|
||
<emphasis box="[1214,1318,1792,1811]" italics="true" pageId="5" pageNumber="6">
|
||
<taxonomicName box="[1214,1315,1792,1811]" class="Reptilia" family="Allosauridae" genus="Allosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Allosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and some abelisaurids (Chure 1998c). In the case of
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[1206,1321,1821,1840]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1206,1321,1821,1840]" italics="true" pageId="5" pageNumber="6">Gorgosaurus</emphasis>
|
||
</taxonomicName>
|
||
this suborbital flange extends to the descending of the ramus of the lacrimal, creating a circular orbit in the largest adults. Smaller individual tyrannosaurids have a slight bump on the thickened dorsal rim above and behind the orbit that becomes greatly expanded in larger individuals. This structure forms a rugose postorbital boss in most adult
|
||
<taxonomicName authorityName="Maleev" authorityYear="1955" box="[362,475,1047,1066]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
||
<emphasis box="[362,475,1047,1066]" italics="true" pageId="6" pageNumber="7">Tarbosaurus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<emphasis box="[527,672,1048,1066]" italics="true" pageId="6" pageNumber="7">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[527,668,1048,1066]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
The enlargement of the dorsal surface of the lacrimal excludes the prefrontals and frontals from view on the lateral aspect; in some
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[99,239,1134,1152]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
||
<emphasis box="[99,239,1134,1152]" italics="true" pageId="6" pageNumber="7">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
this exclusion is completed by intergrowth of the lacrimal and postorbital rugosities.
|
||
</paragraph>
|
||
<caption httpUri="https://zenodo.org/record/3374534/files/figure.png" pageId="6" pageNumber="7" startId="6.[219,287,843,858]" targetBox="[224,1242,103,817]" targetPageId="6">
|
||
<paragraph blockId="6.[219,1231,841,907]" pageId="6" pageNumber="7">
|
||
FIGURE 5.4. Palatal elements:A, palatal view of
|
||
<emphasis box="[630,784,842,858]" italics="true" pageId="6" pageNumber="7">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[630,780,842,858]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
|
||
;
|
||
</emphasis>
|
||
B, left ectopterygoid of
|
||
<taxonomicName baseAuthorityName="Maleev" baseAuthorityYear="1955" box="[987,1144,842,859]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis box="[987,1144,842,859]" italics="true" pageId="6" pageNumber="7">Tarbosaurus bataar</emphasis>
|
||
</taxonomicName>
|
||
in ventral view; C, right palatine of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[434,647,866,883]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="sarcophagus">
|
||
<emphasis box="[434,647,866,883]" italics="true" pageId="6" pageNumber="7">Albertosaurus sarcophagus</emphasis>
|
||
</taxonomicName>
|
||
in lateral view.Scale = 10 cm (A), 3 cm (B), 5 cm (C). (After Molnar 1991; Maleev 1974; and Carr et al., in press.)
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="6.[99,716,990,1926]" pageId="6" pageNumber="7">The squamosal is pneumatized (Witmer 1997a, 1997b) and forms a complex, four-pronged structure. The ventral process forms the dorsal half of the prong that constricts the infratemporal fenestra; the suture with the quadratojugal is nearly as long as the squamosal is tall in most taxa. The quadratojugal is a complex element, wide at the dorsal and ventral surfaces and narrowest dorsal to the juncture of the ascending and caudoventral rami. The dorsal portion of the ascending process is closely appressed medially to the lateral margin of the quadrate.</paragraph>
|
||
<paragraph blockId="6.[99,716,990,1926]" pageId="6" pageNumber="7">
|
||
The jugal is a large triradiate element. The dorsal surface of the rostral prong participates in the internal antorbital fenestra and contains an enlarged rostral foramen. This aperture opens to a complex of internal chambers that also exit via a smaller foramen below the lacrimal articulation in some
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis italics="true" pageId="6" pageNumber="7">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
(Molnar 1991). The lacrimal articulation contributes only slightly to the orbital margin but forms most of the rostral margin of the antorbital fenestra. In tyrannosaurids a low cornual process extends caudoventrally from the middle portion of the ventrolateral surface of the jugal; in large individuals this jugal horn can be rugose.
|
||
</paragraph>
|
||
<paragraph blockId="6.[99,716,990,1926]" lastBlockId="6.[747,1364,990,1927]" pageId="6" pageNumber="7">The prefrontal is greatly reduced in all tyrannosaurids. It is bordered primarily by the lacrimal and the frontal, with a small contribution from the nasal. The frontals of tyrannosaurids vary in shape relative to skull size, starting as triangular bones but expanding caudally to become rectangular in tyrannosaurids having the largest skulls (Currie 2003a). In the largest tyrannosaurids there is a sagittal crest along most of the midline of the paired frontals. In all tyrannosaurids the frontal is firmly sutured medially.</paragraph>
|
||
<paragraph blockId="6.[747,1364,990,1927]" pageId="6" pageNumber="7">
|
||
Tyrannosaurids possess a sagittal crest on the fused parietals (formed along the midline where the supratemporal fossae are confluent). Caudally the parietals bear a large transverse nuchal crest, which rises above the supraoccipital (
|
||
<figureCitation box="[1169,1240,1161,1181]" pageId="6" pageNumber="7" startId="7.[99,167,650,665]" targetBox="[120,690,104,621]" targetPageId="7">fig. 5.5</figureCitation>
|
||
). The crest is more pronounced in tyrannosaurids than in other large-bodied theropods, particularly in
|
||
<emphasis box="[1005,1231,1219,1238]" italics="true" pageId="6" pageNumber="7">
|
||
<taxonomicName authorityName="Kurzanov" authorityYear="1976" box="[1005,1102,1219,1238]" class="Reptilia" family="Tyrannosauridae" genus="Alioramus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">Alioramus</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Maleev" authorityYear="1955" box="[1111,1227,1219,1238]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">Tarbosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and
|
||
<emphasis italics="true" pageId="6" pageNumber="7">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
In these latter taxa the crest is often rugose on the dorsal surface. Fossae for the ligamentum nuchae are present dorsally on the caudal surface of the transverse nuchal crest.
|
||
</paragraph>
|
||
<paragraph blockId="6.[747,1364,990,1927]" lastBlockId="7.[99,716,789,1927]" lastPageId="7" lastPageNumber="8" pageId="6" pageNumber="7">
|
||
The parasphenoid is shaped as a thin rostral prong. It forms the rostral wall of the hypophyseal fossa, is fused with the basisphenoid, and contacts the orbitosphenoid and the laterosphenoid dorsally. The sphenethmoid is known for
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[1249,1364,1420,1439]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1249,1364,1420,1439]" italics="true" pageId="6" pageNumber="7">Gorgosaurus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<emphasis box="[792,939,1449,1467]" italics="true" pageId="6" pageNumber="7">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[792,935,1449,1467]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
in which it is a boxlike structure housing the passages for the olfactory nerves. A median septum separates the two passages. Just behind the sphenethmoid lies the subquadrate orbitosphenoid, bearing the optical foramen. Almost horizontal, it is sutured to the frontal laterally and to the laterosphenoid caudally. The prootic, which is caudal to the laterosphenoid and partly overlaps the opisthotic, is penetrated by the fenestra ovalis and the jugular foramen. The laterosphenoid is tall and forms much of the lateral wall of the braincase. Dorsolaterally it contacts the postorbital by an apparently immobile condyloid joint. The laterosphenoid floors the endocranial cavity and forms the dorsal part of the sella turcica. The oculomotor and trochlear nerves pass through the base of the pila antotica of the laterosphenoid
|
||
<emphasis box="[1143,1295,1822,1841]" italics="true" pageId="6" pageNumber="7">
|
||
(
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1149,1288,1822,1841]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||
)
|
||
</emphasis>
|
||
or the laterosphenoid-orbitosphenoid junction
|
||
<emphasis box="[1150,1272,1850,1869]" italics="true" pageId="6" pageNumber="7">
|
||
(
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[1157,1272,1850,1869]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">Gorgosaurus</taxonomicName>
|
||
</emphasis>
|
||
and
|
||
<emphasis italics="true" pageId="6" pageNumber="7">
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">Daspletosaurus</taxonomicName>
|
||
).
|
||
</emphasis>
|
||
The trigeminal foramen penetrates the laterosphenoid. The opisthotic bears the foramen of the facial nerve. It is extended caudolaterally as a vertical plate and forms, with the exoccipital, the paroccipital process. The paroccipital process of tyrannosaurids is hollow and projects more horizontally and transversely than in carnosaurs and other large tetanurans, in which it is more ventrally and rostrocaudally oriented. The two branches of the vestibulocochlear nerve are separate from the facial nerve. Three tympanic recesses connect to the inner ear, the caudal invading the base of the paroccipital process through a large foramen. The occipital region is caudally oriented in
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[99,214,1048,1067]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
|
||
<emphasis box="[99,214,1048,1067]" italics="true" pageId="7" pageNumber="8">Gorgosaurus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<emphasis box="[263,411,1047,1066]" italics="true" pageId="7" pageNumber="8">
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[263,405,1047,1066]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">Daspletosaurus</taxonomicName>
|
||
;
|
||
</emphasis>
|
||
however, in
|
||
<emphasis italics="true" pageId="7" pageNumber="8">
|
||
<taxonomicName authorityName="Kurzanov" authorityYear="1976" box="[535,632,1047,1066]" class="Reptilia" family="Tyrannosauridae" genus="Alioramus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">Alioramus</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">Albertosaurus</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Maleev" authorityYear="1955" box="[172,288,1076,1095]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">Tarbosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[342,480,1077,1095]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
|
||
<emphasis box="[342,480,1077,1095]" italics="true" pageId="7" pageNumber="8">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
it is more ventrally oriented, suggesting that the skull was oriented with the rostrum directed slightly ventrally in life.
|
||
</paragraph>
|
||
<caption httpUri="https://zenodo.org/record/3374536/files/figure.png" pageId="7" pageNumber="8" startId="7.[99,167,650,665]" targetBox="[120,690,104,621]" targetPageId="7">
|
||
<paragraph blockId="7.[99,641,648,690]" pageId="7" pageNumber="8">
|
||
FIGURE 5.5. Caudal view of the skull of
|
||
<emphasis box="[441,641,649,666]" italics="true" pageId="7" pageNumber="8">
|
||
<taxonomicName baseAuthorityName="Gilmore" baseAuthorityYear="1946" box="[441,637,649,666]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="lancensis">Nanotyrannus lancensis</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Scale = 5 cm.(After Carr 1999.)
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="7.[99,716,789,1927]" pageId="7" pageNumber="8">
|
||
The occipital elements of tyrannosaurids are often extensively fused (
|
||
<figureCitation box="[226,298,1190,1210]" pageId="7" pageNumber="8" startId="7.[99,167,650,665]" targetBox="[120,690,104,621]" targetPageId="7">fig. 5.5</figureCitation>
|
||
). The supraoccipital forms a broad rectangular structure that bears a pair of tablike processes. Internally the supraoccipital houses a tripartite sinus arching over the endocranial cavity (Brochu 2002). The basioccipital forms about half of the occipital condyle and most of the occipital plate below, with extensive contributions from the exoccipitals. Deep subcondylar recesses are situated immediately lateral and slightly ventral to the occipital condyle (Witmer 1997a, 1997b). The basisphenoid region is expanded transversely relative to that in most other large theropods. The basisphenoid floors the braincase. The ventrolaterally directed basipterygoid processes extend from the base of the basisphenoid to contact the pterygoids at well-defined articular surfaces. The basisphenoid is excavated into a deep, ventrally opened recess (Witmer 1997a, 1997b) surrounded by a basipterygoid web (connecting the basipterygoids), a basituberal web (connecting the basal tubera), and webs connecting the basal tubera and the basipterygoid processes on each side (Bakker et al. 1988). The basisphenoid recess is penetrated by a pair of large foramina in tyrannosaurids (Witmer 1997a, 1997b). Although these foramina are reported as a single opening (Russell 1970a) or paired but much reduced (Bakker et al. 1988) in
|
||
<emphasis box="[219,438,1793,1812]" italics="true" pageId="7" pageNumber="8">
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[219,434,1793,1812]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="torosus">Daspletosaurus torosus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
new evidence suggests that they were similarly formed in
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[383,521,1822,1841]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
|
||
<emphasis box="[383,521,1822,1841]" italics="true" pageId="7" pageNumber="8">Daspletosaurus</emphasis>
|
||
</taxonomicName>
|
||
as in other tyrannosaurids (Carr 1999). The basal tubera are plesiomorphically large in
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[128,245,1879,1898]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
|
||
<emphasis box="[128,245,1879,1898]" italics="true" pageId="7" pageNumber="8">Gorgosaurus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[300,444,1879,1898]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
|
||
<emphasis box="[300,444,1879,1898]" italics="true" pageId="7" pageNumber="8">Daspletosaurus</emphasis>
|
||
</taxonomicName>
|
||
but reduced in
|
||
<emphasis italics="true" pageId="7" pageNumber="8">
|
||
<taxonomicName authorityName="Kurzanov" authorityYear="1976" box="[612,712,1879,1898]" class="Reptilia" family="Tyrannosauridae" genus="Alioramus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">Alioramus</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[99,229,1908,1927]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">Albertosaurus</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Maleev" authorityYear="1955" box="[237,353,1908,1927]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">Tarbosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and
|
||
<emphasis box="[404,549,1908,1926]" italics="true" pageId="7" pageNumber="8">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[404,545,1908,1926]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
</paragraph>
|
||
<paragraph blockId="7.[747,1364,101,1926]" pageId="7" pageNumber="8">
|
||
The quadrate of tyrannosaurids and probably
|
||
<taxonomicName authority="(Hutt et al. 2001)" authorityName="Hutt" authorityYear="2001" baseAuthorityName="Hutt" baseAuthorityYear="2001" class="Reptilia" family="Not" genus="Eotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1259,1363,102,120]" italics="true" pageId="7" pageNumber="8">Eotyrannus</emphasis>
|
||
(Hutt et al. 2001)
|
||
</taxonomicName>
|
||
is pneumatic. The jaw articulation of tyrannosaurids lies no farther caudally than the caudalmost point of the occipital condyles and rostral to the caudalmost extent of the paroccipital processes; this is unlike the condition in other large-skulled theropods, in which the jaw articulation is well caudal to the occipital condyle and to the caudalmost point of the paroccipital processes.
|
||
</paragraph>
|
||
<paragraph blockId="7.[747,1364,101,1926]" pageId="7" pageNumber="8">
|
||
The pterygoid has two main components: a horizontal, flat palatal plate and a vertical, thin quadrate ramus. The main body of the ectopterygoid has several interconnecting chambers, which open caudoventrally (
|
||
<figureCitation box="[1035,1122,415,435]" pageId="7" pageNumber="8" startId="6.[219,287,843,858]" targetBox="[224,1242,103,817]" targetPageId="6">fig. 5.4B</figureCitation>
|
||
). Two foramina open to the ventral ectopterygoid sinus in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1074,1198,445,464]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1074,1198,445,464]" italics="true" pageId="7" pageNumber="8">Albertosaurus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<emphasis box="[1245,1362,445,464]" italics="true" pageId="7" pageNumber="8">
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[1245,1358,445,464]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">Gorgosaurus</taxonomicName>
|
||
;
|
||
</emphasis>
|
||
the ectopterygoid chambers of
|
||
<emphasis box="[1050,1322,474,493]" italics="true" pageId="7" pageNumber="8">
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[1050,1194,474,493]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">Daspletosaurus</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Maleev" authorityYear="1955" box="[1202,1318,474,493]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">Tarbosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[747,885,503,521]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
|
||
<emphasis box="[747,885,503,521]" italics="true" pageId="7" pageNumber="8">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
are inflated.
|
||
</paragraph>
|
||
<paragraph blockId="7.[747,1364,101,1926]" pageId="7" pageNumber="8">
|
||
The palatine in most tyrannosaurids
|
||
<emphasis italics="true" pageId="7" pageNumber="8">
|
||
(
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[1161,1279,531,550]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">Gorgosaurus</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">Albertosaurus</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[821,965,560,579]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">Daspletosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and subadult
|
||
<taxonomicName authorityName="Maleev" authorityYear="1955" box="[1113,1226,560,579]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1113,1226,560,579]" italics="true" pageId="7" pageNumber="8">Tarbosaurus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<emphasis italics="true" pageId="7" pageNumber="8">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||
)
|
||
</emphasis>
|
||
is triradiate (
|
||
<figureCitation box="[949,1039,587,607]" pageId="7" pageNumber="8" startId="6.[219,287,843,858]" targetBox="[224,1242,103,817]" targetPageId="6">fig. 5.4C</figureCitation>
|
||
), with maxillary, pterygoid, and vomerine processes but lacking the jugal process of noncoelurosaurian tetanurans. In adult
|
||
<taxonomicName authorityName="Maleev" authorityYear="1955" box="[1111,1224,646,665]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1111,1224,646,665]" italics="true" pageId="7" pageNumber="8">Tarbosaurus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
|
||
<emphasis italics="true" pageId="7" pageNumber="8">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
the palatine is a pneumatic trapezoidal element with a flat ventral surface. The long maxillary process (or the long side of the trapezoid in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[972,1110,732,750]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
|
||
<emphasis box="[972,1110,732,750]" italics="true" pageId="7" pageNumber="8">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
) contacts the palatal shelf of the maxilla, and the maxillary and vomerine processes form the caudal margin of the choana. The lateral surface of the tyrannosaurid palatine bears a recess, and a large foramen or foramina (one in the Alabama tyrannosaurid,
|
||
<emphasis box="[1216,1364,846,865]" italics="true" pageId="7" pageNumber="8">
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[1216,1360,846,865]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">Daspletosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and some
|
||
<emphasis box="[845,1017,876,894]" italics="true" pageId="7" pageNumber="8">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[845,1011,876,894]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
|
||
;
|
||
</emphasis>
|
||
two or more in
|
||
<emphasis italics="true" pageId="7" pageNumber="8">
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[1168,1283,875,894]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">Gorgosaurus</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">Albertosaurus</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Maleev" authorityYear="1955" box="[821,937,904,923]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">Tarbosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and other
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1055,1107,904,922]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1055,1107,904,922]" italics="true" pageId="7" pageNumber="8">T. rex</emphasis>
|
||
</taxonomicName>
|
||
) lead to a strutted cavity within the bone (Witmer 1997a, 1997b).
|
||
</paragraph>
|
||
<paragraph blockId="7.[747,1364,101,1926]" pageId="7" pageNumber="8">The vomer comprises two portions, a rostral rhomboid plate and a caudal laterally compressed stem (Molnar 1999). The rostral plate is expanded in tyrannosaurids, unlike in most other theropods, and is either trapezoidal or diamond-shaped. The vomer has extensive contact with the palatal shelves of the maxilla, and the rostral end of the plate may contact the ventral surface of the palatal shelves of the premaxillae.</paragraph>
|
||
<paragraph blockId="7.[747,1364,101,1926]" pageId="7" pageNumber="8">
|
||
To date, the epipterygoid has been described only in
|
||
<emphasis italics="true" pageId="7" pageNumber="8">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">Albertosaurus</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Kurzanov" authorityYear="1976" box="[874,976,1190,1209]" class="Reptilia" family="Tyrannosauridae" genus="Alioramus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">Alioramus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and
|
||
<emphasis box="[1037,1188,1191,1209]" italics="true" pageId="7" pageNumber="8">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1037,1184,1191,1209]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
It is a teardropshaped bone linking the quadrate process of the pterygoid to the laterosphenid.
|
||
</paragraph>
|
||
<paragraph blockId="7.[747,1364,101,1926]" pageId="7" pageNumber="8">
|
||
The endocranial cavity is larger in tyrannosaurids than in carnosaurs of equivalent skull size (Larsson et al. 2000; Larsson 2001). The use of computer tomography (CT) on various tyrannosaurids (Brochu 2000, 2002) has greatly enhanced previous knowledge, which was drawn mostly from endocranial molds (Osborn 1912a; Maleev 1965; Hopson 1979). The cavity is short but tall. There is a transversely expanded rostral region followed by a transversely constricted but dorsoventrally expanded region that tapers toward the foramen magnum. The rostral region presumably corresponds to the forebrain, while the dorsoventrally expanded region housed the mid- and hindbrains.The floor of the cavity is flexed upward between the jugular and trigeminal foramina. Just rostral to the trigeminal foramen, the cavity is widened to accommodate the optic lobes or the cerebrum. Brochu (2000, 2002) has demonstrated the greatly enlarged olfactory lobes of
|
||
<emphasis box="[1023,1205,1707,1725]" italics="true" pageId="7" pageNumber="8">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1023,1200,1707,1725]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
However, without a comparative series of this region in theropods of intermediate size it is not certain whether these are apomorphically enlarged in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[868,920,1793,1811]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[868,920,1793,1811]" italics="true" pageId="7" pageNumber="8">T. rex</emphasis>
|
||
</taxonomicName>
|
||
(or
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[965,1132,1792,1811]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
) relative to other nonavian theropods or instead are the size expected by allometry; hence, any paleobiological interpretation for this taxon based on the absolute size of its olfactory lobe remains spurious at present.
|
||
</paragraph>
|
||
<caption httpUri="https://zenodo.org/record/3374538/files/figure.png" pageId="8" pageNumber="9" startId="8.[99,166,750,765]" targetBox="[104,708,102,723]" targetPageId="8">
|
||
<paragraph blockId="8.[99,702,748,814]" pageId="8" pageNumber="9">
|
||
FIGURE 5.6. Mandible of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[319,467,749,765]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[319,467,749,765]" italics="true" pageId="8" pageNumber="9">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
in A, lateral view, B, medial view, and C, medial view with splenial removed.Scale = 10 cm. (After Hurum and Currie 2000.)
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="8.[99,717,874,1925]" pageId="8" pageNumber="9">
|
||
The mandibles of primitive tyrannosauroids, as well as small individuals of tyrannosaurids, tend to be slender (
|
||
<figureCitation box="[616,705,902,922]" pageId="8" pageNumber="9" startId="5.[99,167,726,741]" targetBox="[103,710,100,700]" targetPageId="5">fig. 5.3B</figureCitation>
|
||
), while those of adult tyrannosaurids (and adult and juvenile
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[99,237,961,979]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
||
<emphasis box="[99,237,961,979]" italics="true" pageId="8" pageNumber="9">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
) are more robust and deeper dorsoventrally than in more primitive taxa (
|
||
<figureCitation box="[336,408,988,1008]" pageId="8" pageNumber="9" startId="8.[99,166,750,765]" targetBox="[104,708,102,723]" targetPageId="8">fig. 5.6</figureCitation>
|
||
; Carr 1999; Currie 2003b). The mandibular fenestra is small, while the caudal surangular foramen is large in most tyrannosaurids, although it is secondarily reduced in
|
||
<emphasis box="[209,395,1075,1094]" italics="true" pageId="8" pageNumber="9">
|
||
<taxonomicName baseAuthorityName="Maleev" baseAuthorityYear="1955" box="[209,391,1075,1094]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="bataar">Tarbosaurus bataar</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
The dentary bears as many as 18 teeth
|
||
<emphasis box="[157,354,1104,1123]" italics="true" pageId="8" pageNumber="9">
|
||
(
|
||
<taxonomicName authorityName="Gilmore" authorityYear="1933" box="[163,348,1104,1123]" class="Reptilia" family="Tyrannosauridae" genus="Alectrosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="olseni">Alectrosaurus olseni</taxonomicName>
|
||
)
|
||
</emphasis>
|
||
and as few as 11
|
||
<emphasis box="[524,717,1104,1123]" italics="true" pageId="8" pageNumber="9">
|
||
(
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[530,706,1104,1123]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
|
||
).
|
||
</emphasis>
|
||
The interdental plates of tyrannosaurids are large, as in the maxilla; in
|
||
<taxonomicName box="[218,324,1162,1180]" class="Reptilia" family="Not" genus="Eotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
||
<emphasis box="[218,324,1162,1180]" italics="true" pageId="8" pageNumber="9">Eotyrannus</emphasis>
|
||
</taxonomicName>
|
||
they are small spikes. The caudal end of the dentary flares dorsally and ventrally in adult tyrannosaurids, marking the deepening of the postdentary region compared with that in more primitive forms. The supradentary of tyrannosaurids is fused to the coronoid (
|
||
<figureCitation box="[502,589,1275,1295]" pageId="8" pageNumber="9" startId="8.[99,166,750,765]" targetBox="[104,708,102,723]" targetPageId="8">fig. 5.6C</figureCitation>
|
||
; Hurum and Currie 2000). Indeed, it is not at all certain that these represent two separate bones in any theropod (Currie 2003b).The splenial is large, flat, and triangular and bears a large rostral mylohyoid foramen and a caudal notch forming the rostral margin for the internal mandibular fenestra.The prearticular forms a large, flat crescent (the curvature that is greatest in those taxa with the deepest postdentary region) and forms the caudal, ventral, and rostroventral margins of the adductor fossa; the small triangular coronoid forms the rostrodorsal margin. The surangular is a large, thin, curved vertical plate that is dorsoventrally deep on the rostral margin. The rostral external surangular foramen is small, while the caudal surangular foramen is large (except in
|
||
<taxonomicName authorityName="Maleev" authorityYear="1955" box="[99,212,1649,1668]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
||
<emphasis box="[99,212,1649,1668]" italics="true" pageId="8" pageNumber="9">Tarbosaurus</emphasis>
|
||
</taxonomicName>
|
||
). Dorsal to the caudal surangular foramen is a horizontal shelf dorsal to which is a flat facet, presumably for the attachment of the superficial mandibular adductor musculature. The articular is roughly tetrahedral and bears a large foramen aerosum and a large central sinus. Tyrannosaurids do not possess a retroarticular process; instead, a shallow concavity occupies the whole of the caudal surface of the articular, presumably for the insertion of the mandibular depressor muscle. The shape of the glenoid in tyrannosaurids closely conforms to that of the mandibular condyles of the quadrate.
|
||
</paragraph>
|
||
<paragraph blockId="8.[747,1364,100,1468]" pageId="8" pageNumber="9">
|
||
Tyrannosauroid premaxillary (
|
||
<figureCitation box="[1079,1163,100,120]" pageId="8" pageNumber="9" startId="9.[267,335,620,635]" targetBox="[267,1191,99,599]" targetPageId="9">fig. 5.7A</figureCitation>
|
||
) and “lateral” (maxillary and dentary) teeth (
|
||
<figureCitation box="[997,1078,129,149]" pageId="8" pageNumber="9" startId="9.[267,335,620,635]" targetBox="[267,1191,99,599]" targetPageId="9">fig. 5.7B</figureCitation>
|
||
) are distinct from each other, more so than in most theropod taxa; in fact, Marsh (1892) considered isolated premaxillary teeth of tyrannosaurids to be mammalian. The incisiform premaxillary teeth of
|
||
<taxonomicName box="[1259,1363,216,234]" class="Reptilia" family="Not" genus="Eotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1259,1363,216,234]" italics="true" pageId="8" pageNumber="9">Eotyrannus</emphasis>
|
||
</taxonomicName>
|
||
and tyrannosaurids have a D- or U-shaped basal cross section that is mesiodistally as deep as or deeper than it is labiolingually (
|
||
<figureCitation box="[754,839,301,321]" pageId="8" pageNumber="9" startId="9.[267,335,620,635]" targetBox="[267,1191,99,599]" targetPageId="9">fig. 5.7A</figureCitation>
|
||
).Both carinae lie along the caudal surface of the tooth, on a plane perpendicular to the main axis of the skull. These teeth often have a strongly developed vertical ridge on their distal surface. Some tyrannosaurid premaxillary teeth lack serrations on their carinae; however, it is not at all certain that these
|
||
<emphasis box="[807,934,445,464]" italics="true" pageId="8" pageNumber="9">
|
||
“
|
||
<taxonomicName authorityName="Leidy" authorityYear="1868" box="[813,927,445,464]" class="Reptilia" family="Tyrannosauridae" genus="Aublysodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">Aublysodon</taxonomicName>
|
||
”
|
||
</emphasis>
|
||
teeth (Molnar and Carpenter 1989; Currie et al. 1990; Lehman and Carpenter 1990; Holtz 2001b) are not simply the product of wear, digestion, or postmortem erosion (Brochu 2002; Carr and Williamson, in press). In some tyrannosaurid teeth the carinae are bifurcated, perhaps due to trauma or aberrant tooth replacement, but this is also consistent with genetic factors (Erickson 1995). In some tyrannosaurids the rostralmost maxillary teeth are incisiform or subconical. Otherwise, the lateral teeth of tyrannosaurids are larger than the premaxillary teeth, and the carinae are slightly offset from the rostral and caudal margins of the tooth (more so in more robust teeth than in slender forms). In primitive tyrannosauroids
|
||
<emphasis box="[747,883,789,808]" italics="true" pageId="8" pageNumber="9">
|
||
(
|
||
<taxonomicName authorityName="Marsh" authorityYear="1877" box="[754,883,789,808]" class="Reptilia" family="Dryptosauridae" genus="Dryptosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">Dryptosaurus</taxonomicName>
|
||
</emphasis>
|
||
and
|
||
<emphasis box="[941,1056,789,808]" italics="true" pageId="8" pageNumber="9">
|
||
<taxonomicName box="[941,1049,789,808]" class="Reptilia" family="Not" genus="Eotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">Eotyrannus</taxonomicName>
|
||
)
|
||
</emphasis>
|
||
and in juveniles of tyrannosaurids the teeth are ziphodont (bladelike), although the crosssectional diameter is greater labiolingually than found in the teeth of other theropods of the same crown height. In larger advanced tyrannosaurid taxa, however, the teeth are incrassate, so that the labiolingual width at the tooth base sometimes is equal to or exceeds the mesiodistal length. The tooth root can constitute almost two-thirds of the total length in some tyrannosaurids. Denticles on the lateral teeth are wider labiolingually than they are tall proximodistally. In
|
||
<taxonomicName box="[1148,1252,1048,1066]" class="Reptilia" family="Not" genus="Eotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1148,1252,1048,1066]" italics="true" pageId="8" pageNumber="9">Eotyrannus</emphasis>
|
||
</taxonomicName>
|
||
and some tyrannosaurids the denticulations extend over the tip of the tooth. In some of the largest tyrannosaurid teeth, wrinkles in the enamel extend from the carinae.
|
||
</paragraph>
|
||
<paragraph blockId="8.[747,1364,100,1468]" pageId="8" pageNumber="9">Tyrannosaurid teeth show three kinds of wear: slight rounding of the tip, development of flat, oblique facets adjacent to the top both labially and lingually, and abrasion of the serrations both mesially and distally (Molnar et al. 1990; Abler 1992; Farlow and Brinkman 1994). The facets do not correspond in the upper and lower dentitions, so it is unlikely that they were formed by tooth-to-tooth contact.</paragraph>
|
||
<paragraph blockId="8.[747,1364,100,1468]" pageId="8" pageNumber="9">
|
||
To date, ceratobranchials have only been reported in a probable juvenile
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[885,1060,1392,1410]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[885,1060,1392,1410]" italics="true" pageId="8" pageNumber="9">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
(Gilmore 1946a; Bakker et al. 1988). These bones are elongate rods with a curved segment near the middle of the shaft.
|
||
</paragraph>
|
||
<paragraph blockId="8.[747,946,1535,1554]" box="[747,946,1535,1554]" pageId="8" pageNumber="9">
|
||
<heading bold="true" box="[747,946,1535,1554]" fontSize="8" level="2" pageId="8" pageNumber="9" reason="2">
|
||
<emphasis bold="true" box="[747,946,1535,1554]" pageId="8" pageNumber="9">Postcranial Skeleton</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="8.[747,1364,1588,1807]" pageId="8" pageNumber="9">
|
||
The majority of the postcranium of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1087,1225,1589,1607]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1087,1225,1589,1607]" italics="true" pageId="8" pageNumber="9">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
(Osborn 1917; Brochu 2002),
|
||
<taxonomicName authority="(Maleev 1974)" authorityName="Maleev" authorityYear="1974" baseAuthorityName="Maleev" baseAuthorityYear="1974" box="[895,1156,1616,1636]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
||
<emphasis box="[895,1008,1617,1636]" italics="true" pageId="8" pageNumber="9">Tarbosaurus</emphasis>
|
||
(Maleev 1974)
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName authority="(Russell 1970 a)" authorityName="Russell" authorityYear="1970" baseAuthorityName="Russell" baseAuthorityYear="1970" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1166,1307,1617,1636]" italics="true" pageId="8" pageNumber="9">Daspletosaurus</emphasis>
|
||
(Russell 1970a)
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[860,1123,1645,1665]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
||
<emphasis box="[860,975,1646,1665]" italics="true" pageId="8" pageNumber="9">Gorgosaurus</emphasis>
|
||
(Lambe 1917a)
|
||
</taxonomicName>
|
||
, and
|
||
<taxonomicName authority="(Parks 1933)" authorityName="Parks" authorityYear="1933" baseAuthorityName="Parks" baseAuthorityYear="1933" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1173,1300,1645,1664]" italics="true" pageId="8" pageNumber="9">Albertosaurus</emphasis>
|
||
(Parks 1933)
|
||
</taxonomicName>
|
||
is known (
|
||
<figureCitation box="[909,980,1673,1693]" pageId="8" pageNumber="9" startId="9.[99,167,1887,1902]" targetBox="[103,1363,778,1863]" targetPageId="9">fig. 5.8</figureCitation>
|
||
). More fragmentary remains have been recovered for the Alabama tyrannosaurid (Carr et al., in press),
|
||
<taxonomicName authority="(Kurzanov 1976 a)" authorityName="Kurzanov" authorityYear="1976" baseAuthorityName="Kurzanov" baseAuthorityYear="1976" box="[747,1025,1731,1750]" class="Reptilia" family="Tyrannosauridae" genus="Alioramus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
||
<emphasis box="[747,842,1731,1750]" italics="true" pageId="8" pageNumber="9">Alioramus</emphasis>
|
||
(Kurzanov 1976a)
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Gilmore" authorityYear="1933" box="[1035,1159,1731,1750]" class="Reptilia" family="Tyrannosauridae" genus="Alectrosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1035,1159,1731,1750]" italics="true" pageId="8" pageNumber="9">Alectrosaurus</emphasis>
|
||
</taxonomicName>
|
||
(Mader and Bradley 1 989),
|
||
<taxonomicName box="[815,919,1760,1778]" class="Reptilia" family="Not" genus="Eotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
||
<emphasis box="[815,919,1760,1778]" italics="true" pageId="8" pageNumber="9">Eotyrannus</emphasis>
|
||
</taxonomicName>
|
||
(Hutt et al. 2001; Naish et al. 2001),
|
||
<taxonomicName authority="(Carpenter et al. 1997)" authorityName="Carpenter" authorityYear="1997" baseAuthorityName="Carpenter" baseAuthorityYear="1997" class="Reptilia" family="Not" genus="Dryptosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
||
<emphasis italics="true" pageId="8" pageNumber="9">Dryptosaurus</emphasis>
|
||
(Carpenter et al. 1997)
|
||
</taxonomicName>
|
||
, and
|
||
<taxonomicName authority="(Osmolska 1996)" authorityName="Osmolska" authorityYear="1996" baseAuthorityName="Osmolska" baseAuthorityYear="1996" box="[1082,1360,1788,1807]" class="Reptilia" family="Not" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1082,1189,1789,1807]" italics="true" pageId="8" pageNumber="9">Bagaraatan</emphasis>
|
||
(Osmólska 1996)
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<paragraph blockId="8.[747,925,1832,1849]" box="[747,925,1832,1849]" pageId="8" pageNumber="9">
|
||
<heading allCaps="true" bold="true" box="[747,925,1832,1849]" fontSize="7" level="4" pageId="8" pageNumber="9" reason="2">
|
||
<emphasis bold="true" box="[747,925,1832,1849]" pageId="8" pageNumber="9">AXIAL SKELETON</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="8.[747,1364,1878,1926]" lastBlockId="10.[99,716,1017,1898]" lastPageId="10" lastPageNumber="1" pageId="8" pageNumber="9">
|
||
In tyrannosaurids there are 10 cervicals, 13 dorsals, 5 sacrals (although the caudalmost dorsal is slightly sacralized), and 35–44 caudals. Cervicals, dorsals, and some sacrals of tyrannosaurids (and, as far as can be ascertained,
|
||
<taxonomicName box="[501,605,1048,1066]" class="Reptilia" family="Not" genus="Eotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="10" pageNumber="1" phylum="Chordata" rank="genus">
|
||
<emphasis box="[501,605,1048,1066]" italics="true" pageId="10" pageNumber="1">Eotyrannus</emphasis>
|
||
</taxonomicName>
|
||
) bear pleurocoels, and the centra have a complex camellate structure (Britt 1993, 1997). The spinous processes of tyrannosaurids are also highly pneumatized (Brochu 2002).
|
||
</paragraph>
|
||
<caption httpUri="https://zenodo.org/record/3374540/files/figure.png" pageId="9" pageNumber="0" startId="9.[267,335,620,635]" targetBox="[267,1191,99,599]" targetPageId="9">
|
||
<paragraph blockId="9.[267,1172,618,708]" pageId="9" pageNumber="0">
|
||
FIGURE 5.7. Dentition of tyrannosaurids: A, premaxillary tooth of
|
||
<taxonomicName baseAuthorityName="Russell" baseAuthorityYear="1970" box="[838,962,619,636]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="9" pageNumber="0" phylum="Chordata" rank="genus">
|
||
<emphasis box="[838,962,619,636]" italics="true" pageId="9" pageNumber="0">Daspletosaurus</emphasis>
|
||
</taxonomicName>
|
||
sp. in lateral and lingual views, with cross sections at indicated levels (labial end to right); B, dentary tooth of
|
||
<taxonomicName baseAuthorityName="Maleev" baseAuthorityYear="1955" box="[988,1144,643,660]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="9" pageNumber="0" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis box="[988,1144,643,660]" italics="true" pageId="9" pageNumber="0">Tarbosaurus bataar</emphasis>
|
||
</taxonomicName>
|
||
in lateral view
|
||
<emphasis box="[369,426,667,683]" italics="true" pageId="9" pageNumber="0">(upper)</emphasis>
|
||
and basal cross section
|
||
<emphasis box="[627,689,667,684]" italics="true" pageId="9" pageNumber="0">(lower).</emphasis>
|
||
Scale = 1 cm. (After Lehman and Carpenter 1990; and Maleev 1974.)
|
||
</paragraph>
|
||
</caption>
|
||
<caption httpUri="https://zenodo.org/record/3374542/files/figure.png" pageId="9" pageNumber="0" startId="9.[99,167,1887,1902]" targetBox="[103,1363,778,1863]" targetPageId="9">
|
||
<paragraph blockId="9.[99,1280,1885,1927]" pageId="9" pageNumber="0">
|
||
FIGURE 5.8. Skeletons of tyrannosauroids:A,
|
||
<taxonomicName authorityName="Hutt, Naish, Martill, Barker, et Newberry" authorityYear="2001" box="[490,623,1886,1903]" class="Reptilia" family="Not" genus="Eotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="9" pageNumber="0" phylum="Chordata" rank="species" species="lengi">
|
||
<emphasis box="[490,623,1886,1903]" italics="true" pageId="9" pageNumber="0">Eotyrannus lengi</emphasis>
|
||
</taxonomicName>
|
||
(shaded elements represent those recovered at present); B,
|
||
<emphasis box="[1125,1280,1886,1902]" italics="true" pageId="9" pageNumber="0">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1125,1277,1886,1902]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="9" pageNumber="0" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Scale = 1 m. (A courtesy of D. Naish; B courtesy of G. S. Paul.)
|
||
</paragraph>
|
||
</caption>
|
||
<caption httpUri="https://zenodo.org/record/3374544/files/figure.png" pageId="10" pageNumber="1" startId="10.[99,166,844,859]" targetBox="[152,662,100,820]" targetPageId="10">
|
||
<paragraph blockId="10.[99,669,842,956]" pageId="10" pageNumber="1">
|
||
FIGURE 5.9. Cervical vertebrae: A, atlas of juvenile tyrannosaurid
|
||
<emphasis box="[99,414,867,884]" italics="true" pageId="10" pageNumber="1">
|
||
(“
|
||
<taxonomicName authorityName="Dong" authorityYear="1977" box="[111,402,867,884]" class="Reptilia" family="Tyrannosauridae" genus="Shanshanosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="10" pageNumber="1" phylum="Chordata" rank="species" species="houyanshanensis">Shanshanosaurus houyanshanensis</taxonomicName>
|
||
”)
|
||
</emphasis>
|
||
in left lateral and right lateral views; B, cervical series of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[320,469,892,908]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="10" pageNumber="1" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[320,469,892,908]" italics="true" pageId="10" pageNumber="1">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
in right lateral view. Scale = 1 cm (A), 10 cm (B). (After Currie and Dong 2001b; and Osborn 1917.)
|
||
</paragraph>
|
||
</caption>
|
||
<caption httpUri="https://zenodo.org/record/3374546/files/figure.png" pageId="10" pageNumber="1" startId="10.[747,815,1034,1049]" targetBox="[788,1290,103,1005]" targetPageId="10">
|
||
<paragraph blockId="10.[747,1318,1032,1098]" pageId="10" pageNumber="1">
|
||
FIGURE 5.10. Dorsal vertebrae: A, dorsals 1–4 of
|
||
<taxonomicName baseAuthorityName="Maleev" baseAuthorityYear="1955" box="[1162,1318,1033,1050]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="10" pageNumber="1" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis box="[1162,1318,1033,1050]" italics="true" pageId="10" pageNumber="1">Tarbosaurus bataar</emphasis>
|
||
</taxonomicName>
|
||
in right lateral view; B, caudal dorsal in cranial and caudal views. Scale = 10 cm. (After Maleev 1974.)
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="10.[99,716,1017,1898]" pageId="10" pageNumber="1">
|
||
As in most theropods, the atlas is composed of only the neurapophyses, the intercentrum, and the odontoid (
|
||
<figureCitation box="[590,662,1189,1209]" pageId="10" pageNumber="1" startId="10.[99,166,844,859]" targetBox="[152,662,100,820]" targetPageId="10">fig. 5.9</figureCitation>
|
||
). The crescentic intercentrum is axially compressed, twice as long as it is high. The axial pleurocentrum is pleurocoelous, generally having only a single pneumatopore. The axial spinous process has only a low vertical component projecting dorsally. Pre- and postzygapophyses are present, and the latter bear epipophyses.
|
||
</paragraph>
|
||
<paragraph blockId="10.[99,716,1017,1898]" pageId="10" pageNumber="1">
|
||
Postaxial cervical centra are amphiplatyan, amphicoelous, or at most weakly opisthocoelous (
|
||
<figureCitation box="[449,536,1390,1410]" pageId="10" pageNumber="1" startId="10.[99,166,844,859]" targetBox="[152,662,100,820]" targetPageId="10">fig. 5.9B</figureCitation>
|
||
); this differs from the condition in other large-bodied theropods, in which the centra are markedly opisthocoelous. The spinous processes, zygapophyses, and rib articulations are well developed. Facets for the zygapophyses are dorsally (rather than craniodorsally) oriented in
|
||
<taxonomicName box="[186,290,1535,1553]" class="Reptilia" family="Not" genus="Eotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="10" pageNumber="1" phylum="Chordata" rank="genus">
|
||
<emphasis box="[186,290,1535,1553]" italics="true" pageId="10" pageNumber="1">Eotyrannus</emphasis>
|
||
</taxonomicName>
|
||
and tyrannosaurids. The cervical central of adult tyrannosaurines are craniocaudally compressed, only about half as long as they are tall; however, those of juveniles and of other tyrannosauroids are as long as or longer than they are tall. Although tyrannosaurids are often characterized as being short-necked (cf. Steel 1970), this description is only applicable to
|
||
<taxonomicName authorityName="Matthew et Brown" authorityYear="1922" box="[126,296,1706,1725]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="10" pageNumber="1" phylum="Chordata" rank="subFamily" subFamily="Tyrannosaurinae">Tyrannosaurinae</taxonomicName>
|
||
(Holtz 2001b; Currie 2003b); the cervical series of
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[183,298,1736,1755]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="10" pageNumber="1" phylum="Chordata" rank="genus">
|
||
<emphasis box="[183,298,1736,1755]" italics="true" pageId="10" pageNumber="1">Gorgosaurus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName baseAuthorityName="Parks" baseAuthorityYear="1933" box="[345,472,1735,1754]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="10" pageNumber="1" phylum="Chordata" rank="genus">
|
||
<emphasis box="[345,472,1735,1754]" italics="true" pageId="10" pageNumber="1">Albertosaurus</emphasis>
|
||
</taxonomicName>
|
||
are almost as long as the dorsal series. Tyrannosaurids’ appearing to be short-necked may be a result of their enlarged skulls rather than the proportions of the sections of the vertebral column.
|
||
</paragraph>
|
||
<paragraph blockId="10.[99,716,1017,1898]" lastBlockId="10.[747,1364,1161,1898]" pageId="10" pageNumber="1">
|
||
Dorsal vertebrae have powerfully developed spinous processes and transverse processes, the latter being horizontally directed (
|
||
<figureCitation box="[754,838,1161,1181]" pageId="10" pageNumber="1" startId="10.[747,815,1034,1049]" targetBox="[788,1290,103,1005]" targetPageId="10">fig. 5.10</figureCitation>
|
||
). The centra are amphicoelous or amphiplatyan and become longer caudally in the series.All dorsals have hyposphenehypantrum articulations. Dorsal ribs are well developed and broadly curved cranially, suggesting a barrel-shaped chest region. The trunk becomes shallower and narrower in front of the sacrum.
|
||
</paragraph>
|
||
<paragraph blockId="10.[747,1364,1161,1898]" pageId="10" pageNumber="1">
|
||
The sacrum proper consists of five vertebrae (
|
||
<figureCitation box="[1219,1298,1333,1353]" pageId="10" pageNumber="1" startId="11.[99,167,454,469]" targetBox="[186,630,102,428]" targetPageId="11">fig. 5.11</figureCitation>
|
||
); however the caudalmost dorsal is completely surrounded laterally by the ilia, with which it has broad contact via the transverse processes, and is thus functionally sacralized. In at least some material the sacrum is compressed, so that the centra are much deeper dorsoventrally than they are wide transversely, although this may be due in part to postmortem distortion. The sacral spinous processes are fused to form a continuous lamina. The sacral ribs and transverse processes are robust and have broad contact with the ilia.
|
||
</paragraph>
|
||
<paragraph blockId="10.[747,1364,1161,1898]" pageId="10" pageNumber="1">
|
||
The transition point between the proximal and distal parts of the caudal series (as marked by the presence or absence of transverse processes and spinous processes, respectively) is located at about caudal 15. The distal centra are more elongate in form than the proximal (
|
||
<figureCitation box="[997,1095,1734,1754]" pageId="10" pageNumber="1" startId="11.[99,167,925,940]" targetBox="[106,705,565,898]" targetPageId="11">fig. 5.12A</figureCitation>
|
||
); the caudal centra are amphiplatyan or amphicoelous (
|
||
<figureCitation box="[1027,1119,1763,1783]" pageId="10" pageNumber="1" startId="11.[99,167,925,940]" targetBox="[106,705,565,898]" targetPageId="11">fig.5.12B</figureCitation>
|
||
).The transverse processes are broad and horizontal. The distal spinous processes are axially elongate. The cranial zygapophyses of the distal series are elongate, sometimes extending more than half of the length of the preceding centrum.
|
||
</paragraph>
|
||
<caption httpUri="https://zenodo.org/record/3374548/files/figure.png" pageId="11" pageNumber="2" startId="11.[99,167,454,469]" targetBox="[186,630,102,428]" targetPageId="11">
|
||
<paragraph blockId="11.[99,637,452,494]" pageId="11" pageNumber="2">
|
||
FIGURE 5.11. Sacrum of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[312,460,453,469]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="2" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[312,460,453,469]" italics="true" pageId="11" pageNumber="2">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
in right lateral view. Scale = 10 cm. (After Osborn 1917.)
|
||
</paragraph>
|
||
</caption>
|
||
<caption httpUri="https://zenodo.org/record/3374550/files/figure.png" pageId="11" pageNumber="2" startId="11.[747,815,702,717]" targetBox="[803,1309,103,675]" targetPageId="11">
|
||
<paragraph blockId="11.[747,1361,700,814]" pageId="11" pageNumber="2">
|
||
FIGURE 5.13. Haemal arches of
|
||
<emphasis box="[1021,1193,701,718]" italics="true" pageId="11" pageNumber="2">
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[1021,1187,701,718]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="2" phylum="Chordata" rank="species" species="libratus">Gorgosaurus libratus</taxonomicName>
|
||
:
|
||
</emphasis>
|
||
A, twentieth hemal arch in right lateral, caudal, and dorsal views; B, fifteenth hemal arch in right lateral, caudal, and dorsal views; C, seventh hemal arch in right lateral and caudal views; D, second hemal arch in right lateral and caudal views. Scale = 5 cm. (After Lambe 1917a.)
|
||
</paragraph>
|
||
</caption>
|
||
<caption httpUri="https://zenodo.org/record/3374552/files/figure.png" pageId="11" pageNumber="2" startId="11.[99,167,925,940]" targetBox="[106,705,565,898]" targetPageId="11">
|
||
<paragraph blockId="11.[99,694,923,1013]" pageId="11" pageNumber="2">
|
||
FIGURE 5.12. Caudal vertebrae: A, caudal 28 of
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[506,673,924,941]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="2" phylum="Chordata" rank="species" species="libratus">
|
||
<emphasis box="[506,673,924,941]" italics="true" pageId="11" pageNumber="2">Gorgosaurus libratus</emphasis>
|
||
</taxonomicName>
|
||
in right lateral, dorsal, ventral, and cranial views; B, midseries caudal of
|
||
<taxonomicName baseAuthorityName="Maleev" baseAuthorityYear="1955" box="[99,255,972,989]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="2" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis box="[99,255,972,989]" italics="true" pageId="11" pageNumber="2">Tarbosaurus bataar</emphasis>
|
||
</taxonomicName>
|
||
in cranial and caudal view. Scale = 1 cm (A), 10 cm (B). (After Lambe 1917a; and Maleev 1974.)
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="11.[99,716,1075,1898]" pageId="11" pageNumber="2">
|
||
The hemal processes in tyrannosaurids show a transition from proximal L-shaped forms through caudal 10 (
|
||
<figureCitation box="[584,704,1103,1123]" pageId="11" pageNumber="2" startId="11.[747,815,702,717]" targetBox="[803,1309,103,675]" targetPageId="11">fig.5.13C, D</figureCitation>
|
||
); hatchet-shaped forms, with the distal portion longer craniocaudally than the proximal portion and a ventrally convex margin, from caudal 11 to 15 (
|
||
<figureCitation box="[366,463,1189,1209]" pageId="11" pageNumber="2" startId="11.[747,815,702,717]" targetBox="[803,1309,103,675]" targetPageId="11">fig. 5.13B</figureCitation>
|
||
); and boat-shaped hemal processes with cranial and caudal projections, and more than twice as long craniocaudally as tall dorsoventrally, from caudal 15 onward (
|
||
<figureCitation box="[215,312,1275,1295]" pageId="11" pageNumber="2" startId="11.[747,815,702,717]" targetBox="[803,1309,103,675]" targetPageId="11">fig. 5.13A</figureCitation>
|
||
). Osborn (1917) miscalculated the length and transitional sequence of the (then incompletely known) tail of
|
||
<emphasis box="[124,301,1334,1352]" italics="true" pageId="11" pageNumber="2">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[124,297,1334,1352]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="2" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
He thus restored this tyrannosaurid, using
|
||
<emphasis box="[99,204,1362,1381]" italics="true" pageId="11" pageNumber="2">
|
||
<taxonomicName box="[99,200,1362,1381]" class="Reptilia" family="Allosauridae" genus="Allosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="2" phylum="Chordata" rank="genus">Allosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
with 53 caudals, as a model. He placed the only discovered hatchet-shaped hemal process between caudals 21 and 23 (ten or more vertebrae distal to the actual position) and interpolated a gradual sequence from the elongate proximal hemal processes. This incorrect model of the tyrannosaurid tail has been used to restore other incomplete skeletons since and may have been the source for some misinterpretation of the actual caudal anatomy of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[361,530,1563,1582]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="2" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
(e.g., Forster et al. 1998).
|
||
</paragraph>
|
||
<caption httpUri="https://zenodo.org/record/3374554/files/figure.png" pageId="11" pageNumber="2" startId="11.[747,815,1390,1405]" targetBox="[768,1342,870,1363]" targetPageId="11">
|
||
<paragraph blockId="11.[747,1340,1388,1430]" pageId="11" pageNumber="2">
|
||
FIGURE 5.14. Cranial portion of gastralia series of
|
||
<emphasis box="[1177,1340,1389,1406]" italics="true" pageId="11" pageNumber="2">
|
||
<taxonomicName baseAuthorityName="Maleev" baseAuthorityYear="1955" box="[1177,1336,1389,1406]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="2" phylum="Chordata" rank="species" species="bataar">Tarbosaurus bataar</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Scale = 15 cm. (After Maleev 1974.)
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="11.[99,716,1075,1898]" lastBlockId="11.[747,1363,1501,1549]" pageId="11" pageNumber="2">
|
||
Gastralia have saddle-shaped facets between the medial and lateral segments and between interlocking portions of each sequential medial segment (
|
||
<figureCitation box="[364,446,1677,1697]" pageId="11" pageNumber="2" startId="11.[747,815,1390,1405]" targetBox="[768,1342,870,1363]" targetPageId="11">fig. 5.14</figureCitation>
|
||
; Lambe 1917a; Parks 1933; Maleev 1974; Brochu 2002). The median segment of each gastralium is longer than the lateral segment. In
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[559,676,1736,1755]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="2" phylum="Chordata" rank="genus">
|
||
<emphasis box="[559,676,1736,1755]" italics="true" pageId="11" pageNumber="2">Gorgosaurus</emphasis>
|
||
</taxonomicName>
|
||
the series had 19 segments (Lambe 1917a); Brochu (2002) estimates that 18 were present in
|
||
<emphasis box="[348,497,1793,1811]" italics="true" pageId="11" pageNumber="2">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[348,493,1793,1811]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="2" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
The cranialmost two segments of the gastral basket of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[429,567,1822,1840]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="2" phylum="Chordata" rank="genus">
|
||
<emphasis box="[429,567,1822,1840]" italics="true" pageId="11" pageNumber="2">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
(Brochu 2002) and
|
||
<taxonomicName authority="(Maleev 1974)" authorityName="Maleev" authorityYear="1974" baseAuthorityName="Maleev" baseAuthorityYear="1974" box="[144,401,1850,1869]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="2" phylum="Chordata" rank="genus">
|
||
<emphasis box="[144,257,1850,1869]" italics="true" pageId="11" pageNumber="2">Tarbosaurus</emphasis>
|
||
(Maleev 1974)
|
||
</taxonomicName>
|
||
are fused into a platelike mass; given the similarity of this element to the supposed sternum of
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[747,1012,1501,1520]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="2" phylum="Chordata" rank="genus">
|
||
<emphasis box="[747,862,1501,1520]" italics="true" pageId="11" pageNumber="2">Gorgosaurus</emphasis>
|
||
(Lambe 1917a)
|
||
</taxonomicName>
|
||
, that latter structure is more simply explained as also being a fused gastral plate.
|
||
</paragraph>
|
||
<paragraph blockId="11.[747,1029,1573,1590]" box="[747,1029,1573,1590]" pageId="11" pageNumber="2">
|
||
<heading allCaps="true" bold="true" box="[747,1029,1573,1590]" fontSize="7" level="4" pageId="11" pageNumber="2" reason="2">
|
||
<emphasis bold="true" box="[747,1029,1573,1590]" pageId="11" pageNumber="2">APPENDICULAR SKELETON</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="11.[747,1364,1620,1897]" pageId="11" pageNumber="2">
|
||
The scapula is long and straplike. The caudal end is only slightly expanded in
|
||
<emphasis box="[872,1259,1649,1668]" italics="true" pageId="11" pageNumber="2">
|
||
<taxonomicName box="[872,979,1649,1667]" class="Reptilia" family="Not" genus="Eotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="2" phylum="Chordata" rank="genus">Eotyrannus</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[986,1103,1649,1668]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="2" phylum="Chordata" rank="genus">Gorgosaurus</taxonomicName>
|
||
,
|
||
<taxonomicName baseAuthorityName="Russell" baseAuthorityYear="1970" box="[1111,1255,1649,1668]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="2" phylum="Chordata" rank="genus">Daspletosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and
|
||
<taxonomicName baseAuthorityName="Maleev" baseAuthorityYear="1974" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="2" phylum="Chordata" rank="genus">
|
||
<emphasis italics="true" pageId="11" pageNumber="2">Tarbosaurus</emphasis>
|
||
</taxonomicName>
|
||
but much more so in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1036,1176,1678,1696]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="2" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1036,1176,1678,1696]" italics="true" pageId="11" pageNumber="2">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<emphasis box="[1228,1364,1677,1696]" italics="true" pageId="11" pageNumber="2">
|
||
<taxonomicName baseAuthorityName="Parks" baseAuthorityYear="1933" box="[1228,1361,1677,1696]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="2" phylum="Chordata" rank="genus">Albertosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
the latter having more expansion dorsally than ventrally. The acromial expansion is proportionately large. The scapula contributes more of the glenoid than does the coracoid. The coracoid is an oval plate approximately one-quarter of the scapular length and has a well-developed caudoventral process. The coracoid foramen is large. The biceps tubercle is present and moderately well developed.
|
||
</paragraph>
|
||
<caption httpUri="https://zenodo.org/record/3374556/files/figure.png" pageId="12" pageNumber="3" startId="12.[99,166,883,898]" targetBox="[105,1354,102,856]" targetPageId="12">
|
||
<paragraph blockId="12.[99,1354,881,947]" pageId="12" pageNumber="3">
|
||
FIGURE 5.15. Humeri: A, right humerus of
|
||
<taxonomicName authorityName="Hutt, Naish, Martill, Barker, et Newberry" authorityYear="2001" box="[469,603,882,899]" class="Reptilia" family="Not" genus="Eotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="species" species="lengi">
|
||
<emphasis box="[469,603,882,899]" italics="true" pageId="12" pageNumber="3">Eotyrannus lengi</emphasis>
|
||
</taxonomicName>
|
||
in cranial and lateral views (
|
||
<taxonomicName box="[846,937,882,898]" class="Reptilia" family="Not" genus="Eotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[846,937,882,898]" italics="true" pageId="12" pageNumber="3">Eotyrannus</emphasis>
|
||
</taxonomicName>
|
||
lateral tooth and
|
||
<taxonomicName authorityName="Galton" authorityYear="1977" box="[1088,1188,882,899]" class="Reptilia" family="Dryosauridae" genus="Valdosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1088,1188,882,899]" italics="true" pageId="12" pageNumber="3">Valdosaurus</emphasis>
|
||
</taxonomicName>
|
||
tooth root still attached by matrix); B, left humerus of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[418,566,906,922]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[418,566,906,922]" italics="true" pageId="12" pageNumber="3">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
in proximal, distal, cranial, lateral, caudal, and medial views. Scale = 5 cm. (After Naish et al. 2001; and Carpenter and Smith 2001.)
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="12.[99,716,1018,1897]" pageId="12" pageNumber="3">
|
||
Furculae are known for
|
||
<emphasis box="[363,623,1018,1037]" italics="true" pageId="12" pageNumber="3">
|
||
<taxonomicName baseAuthorityName="Parks" baseAuthorityYear="1933" box="[363,493,1018,1037]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="genus">Albertosaurus</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[502,619,1018,1037]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="genus">Gorgosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and
|
||
<taxonomicName authority="Russell, 1970" baseAuthorityName="Russell" baseAuthorityYear="1970" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis italics="true" pageId="12" pageNumber="3">Daspletosaurus</emphasis>
|
||
</taxonomicName>
|
||
(Makovicky and Currie 1998). Tyrannosaurid furculae have a broad curvature. They have a short hypocleideum, and larger individuals have expanded epicleidial facets. Furculae articulate with the acromial processes, and the apex lies cranial and ventral to the cranial margins of the coracoids. Tyrannosaurid furculae can be distinguished from the fused cranialmost medial gastral segments of some theropods in that the latter are more massive, have more gradually tapering and straight (rather than sigmoid) rami, and do not have epicleidial facets (Currie and Makovicky 1998).
|
||
</paragraph>
|
||
<paragraph blockId="12.[99,716,1018,1897]" pageId="12" pageNumber="3">
|
||
The sternum was reported for
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[440,712,1333,1353]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[440,556,1334,1353]" italics="true" pageId="12" pageNumber="3">Gorgosaurus</emphasis>
|
||
(Lambe 1917a)
|
||
</taxonomicName>
|
||
, but based on new observations from
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[486,626,1363,1381]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[486,626,1363,1381]" italics="true" pageId="12" pageNumber="3">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
(Brochu 2002) that element is more likely a fused gastral plate. Thus, no ossified sternum has been confirmed for any member of Tyrannosauroidea.
|
||
</paragraph>
|
||
<paragraph blockId="12.[99,716,1018,1897]" pageId="12" pageNumber="3">
|
||
The tyrannosaurid forelimb is greatly reduced relative to the hindlimb and the scapula. Ratios of femoral to humeral length range from approximately 2.8 in
|
||
<taxonomicName baseAuthorityName="Russell" baseAuthorityYear="1970" box="[422,563,1534,1553]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[422,563,1534,1553]" italics="true" pageId="12" pageNumber="3">Daspletosaurus</emphasis>
|
||
</taxonomicName>
|
||
to 4.0 in
|
||
<emphasis italics="true" pageId="12" pageNumber="3">
|
||
<taxonomicName baseAuthorityName="Maleev" baseAuthorityYear="1974" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="genus">Tarbosaurus</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Ratios of scapular to humeral length range from 2.2 in
|
||
<taxonomicName baseAuthorityName="Russell" baseAuthorityYear="1970" box="[99,237,1592,1611]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[99,237,1592,1611]" italics="true" pageId="12" pageNumber="3">Daspletosaurus</emphasis>
|
||
</taxonomicName>
|
||
to 2.9 in
|
||
<emphasis box="[324,442,1592,1611]" italics="true" pageId="12" pageNumber="3">
|
||
<taxonomicName baseAuthorityName="Maleev" baseAuthorityYear="1974" box="[324,438,1592,1611]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="genus">Tarbosaurus</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
The antebrachium is shorter than the humerus, while the manus is intermediate in length between the two. The forelimb of
|
||
<taxonomicName box="[435,559,1650,1668]" class="Reptilia" family="Not" genus="Dryptosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[435,559,1650,1668]" italics="true" pageId="12" pageNumber="3">Dryptosaurus</emphasis>
|
||
</taxonomicName>
|
||
is also reduced, but that of
|
||
<taxonomicName box="[208,312,1678,1696]" class="Reptilia" family="Not" genus="Eotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[208,312,1678,1696]" italics="true" pageId="12" pageNumber="3">Eotyrannus</emphasis>
|
||
</taxonomicName>
|
||
is plesiomorphically elongate.
|
||
</paragraph>
|
||
<paragraph blockId="12.[99,716,1018,1897]" lastBlockId="12.[747,1364,1018,1898]" pageId="12" pageNumber="3">
|
||
The humerus is slender, and the deltopectoral crest is weakly developed in all tyrannosauroids except
|
||
<taxonomicName box="[504,608,1736,1754]" class="Reptilia" family="Not" genus="Eotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[504,608,1736,1754]" italics="true" pageId="12" pageNumber="3">Eotyrannus</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation box="[622,705,1734,1754]" pageId="12" pageNumber="3" startId="12.[99,166,883,898]" targetBox="[105,1354,102,856]" targetPageId="12">fig. 5.15</figureCitation>
|
||
). The crest is largest in
|
||
<emphasis box="[312,582,1764,1783]" italics="true" pageId="12" pageNumber="3">
|
||
<taxonomicName box="[312,439,1764,1782]" class="Reptilia" family="Not" genus="Dryptosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="genus">Dryptosaurus</taxonomicName>
|
||
,
|
||
<taxonomicName baseAuthorityName="Parks" baseAuthorityYear="1933" box="[448,578,1764,1783]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="genus">Albertosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and
|
||
<emphasis italics="true" pageId="12" pageNumber="3">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
The humeral head is not offset from the lateral tuberosity by a cleft. The shaft is straighter than in most large tetanurans, although within
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[266,435,1850,1868]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[266,435,1850,1868]" italics="true" pageId="12" pageNumber="3">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
two different forms are present, one with a more pronounced medial curvature and one with a much straighter shaft; Carpenter and Smith (2001) suggest that this difference might be a product of sexual dimorphism. The distal end is weakly expanded. The radius is a simple cylinder (
|
||
<figureCitation box="[843,941,1103,1123]" pageId="12" pageNumber="3" startId="13.[99,167,855,870]" targetBox="[107,710,99,834]" targetPageId="13">fig. 5.16B</figureCitation>
|
||
). The length of the ulna is approximately 60% of the length of the humerus in most tyrannosaurids, and only 45% in
|
||
<emphasis box="[867,1071,1162,1181]" italics="true" pageId="12" pageNumber="3">
|
||
<taxonomicName baseAuthorityName="Maleev" baseAuthorityYear="1955" box="[867,1067,1162,1181]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="species" species="bataar">Tyrannosaurus bataar</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
The olecranon process is moderately well developed but short (
|
||
<figureCitation box="[1070,1166,1189,1209]" pageId="12" pageNumber="3" startId="13.[99,167,855,870]" targetBox="[107,710,99,834]" targetPageId="13">fig. 5.16A</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="12.[747,1364,1018,1898]" pageId="12" pageNumber="3">
|
||
The carpus of tyrannosaurids is only poorly preserved. Five elements (three proximal, two distal) have been recovered for
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[747,859,1277,1296]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[747,859,1277,1296]" italics="true" pageId="12" pageNumber="3">Gorgosaurus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<emphasis box="[904,1035,1276,1295]" italics="true" pageId="12" pageNumber="3">
|
||
<taxonomicName baseAuthorityName="Parks" baseAuthorityYear="1933" box="[904,1031,1276,1295]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="genus">Albertosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
four for
|
||
<emphasis box="[1113,1231,1276,1295]" italics="true" pageId="12" pageNumber="3">
|
||
<taxonomicName baseAuthorityName="Maleev" baseAuthorityYear="1974" box="[1113,1227,1276,1295]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="genus">Tarbosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
three for
|
||
<emphasis italics="true" pageId="12" pageNumber="3">
|
||
<taxonomicName baseAuthorityName="Russell" baseAuthorityYear="1970" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="genus">Daspletosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and only two for
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1021,1159,1306,1324]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1021,1159,1306,1324]" italics="true" pageId="12" pageNumber="3">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
(Holtz 2001a; Chure, in press). However, this might well be an artifact of preservation rather than a taxonomic feature. Homologies are hindered by the apomorphically reduced nature of these elements, which lack well-developed facets or trochlea. A single large distal element caps parts of the proximal ends of metacarpals I and II. In fact, in a juvenile tyrannosaurid from the Horseshoe Canyon Formation of Alberta, Canada (Russell 1970a; Holtz 2001a) and in
|
||
<taxonomicName box="[776,882,1535,1553]" class="Reptilia" family="Not" genus="Eotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[776,882,1535,1553]" italics="true" pageId="12" pageNumber="3">Eotyrannus</emphasis>
|
||
</taxonomicName>
|
||
(Naish, pers. comm.) this element does in fact possess transverse trochlea and a semilunate shape. In adult tyrannosaurids this element is poorly formed and lacks these features, perhaps reflecting a shift away from manual prehension during tyrannosauroid evolution and tyrannosaurid ontogeny.
|
||
</paragraph>
|
||
<paragraph blockId="12.[747,1364,1018,1898]" lastBlockId="13.[99,716,990,1927]" lastPageId="13" lastPageNumber="4" pageId="12" pageNumber="3">
|
||
The manus of
|
||
<taxonomicName box="[933,1037,1678,1696]" class="Reptilia" family="Not" genus="Eotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[933,1037,1678,1696]" italics="true" pageId="12" pageNumber="3">Eotyrannus</emphasis>
|
||
</taxonomicName>
|
||
is tridactyl and well developed (Hutt et al. 2001); in contrast, the manus of tyrannosaurids was functionally didactyl, with only metacarpals I and II possessing phalanges (
|
||
<figureCitation box="[857,937,1763,1783]" pageId="12" pageNumber="3" startId="13.[747,815,719,734]" targetBox="[804,1308,100,693]" targetPageId="13">fig. 5.17</figureCitation>
|
||
).The condition in
|
||
<taxonomicName box="[1115,1239,1764,1782]" class="Reptilia" family="Not" genus="Dryptosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1115,1239,1764,1782]" italics="true" pageId="12" pageNumber="3">Dryptosaurus</emphasis>
|
||
</taxonomicName>
|
||
is uncertain: at least one digit bore an immense trenchant claw, but which digit this was is uncertain (Carpenter et al. 1997). Metacarpal III in tyrannosaurids is greatly reduced in length and thickness relative to all other theropods and lacks a distal articulation. Metacarpal II is the longest element in the manus: it is about twice as long as metacarpal I in most tyrannosaurids but only about 70% longer in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[306,446,1049,1067]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis box="[306,446,1049,1067]" italics="true" pageId="13" pageNumber="4">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
and only 30%–60% longer in
|
||
<emphasis box="[127,248,1077,1096]" italics="true" pageId="13" pageNumber="4">
|
||
<taxonomicName baseAuthorityName="Maleev" baseAuthorityYear="1974" box="[127,244,1077,1096]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="genus">Tarbosaurus</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Metacarpal III is longer than metacarpal I (the primitive condition in
|
||
<taxonomicName authorityName="Marsh" authorityYear="1881" box="[319,426,1105,1124]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Theropoda" pageId="13" pageNumber="4" phylum="Chordata" rank="order">Theropoda</taxonomicName>
|
||
) in
|
||
<emphasis box="[462,715,1105,1125]" italics="true" pageId="13" pageNumber="4">
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[462,577,1106,1125]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="genus">Gorgosaurus</taxonomicName>
|
||
,
|
||
<taxonomicName baseAuthorityName="Parks" baseAuthorityYear="1933" box="[585,712,1105,1124]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="genus">Albertosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and
|
||
<emphasis box="[141,289,1134,1153]" italics="true" pageId="13" pageNumber="4">
|
||
<taxonomicName baseAuthorityName="Russell" baseAuthorityYear="1970" box="[141,285,1134,1153]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="genus">Daspletosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
but in
|
||
<taxonomicName baseAuthorityName="Maleev" baseAuthorityYear="1974" box="[354,467,1134,1153]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis box="[354,467,1134,1153]" italics="true" pageId="13" pageNumber="4">Tarbosaurus</emphasis>
|
||
</taxonomicName>
|
||
metacarpal III is reduced, so that it is shorter than metacarpal I. Metacarpal III is present in
|
||
<emphasis box="[127,305,1192,1210]" italics="true" pageId="13" pageNumber="4">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[127,301,1192,1210]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
as attested by the notch in the proximal end of metacarpal II for its articulation, but this element has not yet been found (Carpenter and Smith 2001), and it is thus uncertain whether it was as reduced as in
|
||
<emphasis box="[486,607,1277,1296]" italics="true" pageId="13" pageNumber="4">
|
||
<taxonomicName baseAuthorityName="Maleev" baseAuthorityYear="1974" box="[486,603,1277,1296]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="genus">Tarbosaurus</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
These proportions might reflect the general trend in theropod evolution of reduced size of manual elements from digit V toward digit I (Wagner and Gauthier 1999), with metacarpals III and II more reduced in the derived tyrannosaurines relative to the other, more primitive tyrannosauroids. The phalangeal formula of tyrannosaurids is 2-3-0-X-X. Phalanx 1 of digit I is the longest, being longer than metacarpal II in
|
||
<emphasis box="[436,715,1478,1497]" italics="true" pageId="13" pageNumber="4">
|
||
<taxonomicName baseAuthorityName="Parks" baseAuthorityYear="1933" box="[436,566,1478,1497]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="genus">Albertosaurus</taxonomicName>
|
||
,
|
||
<taxonomicName baseAuthorityName="Russell" baseAuthorityYear="1970" box="[574,715,1478,1497]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="genus">Daspletosaurus</taxonomicName>
|
||
</emphasis>
|
||
(incorrectly restored as phalanx 1 of digit II in Russell 1970a), and
|
||
<taxonomicName baseAuthorityName="Maleev" baseAuthorityYear="1974" box="[145,258,1536,1555]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis box="[145,258,1536,1555]" italics="true" pageId="13" pageNumber="4">Tarbosaurus</emphasis>
|
||
</taxonomicName>
|
||
but subequal to metacarpal II in
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[601,716,1536,1555]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis box="[601,716,1536,1555]" italics="true" pageId="13" pageNumber="4">Gorgosaurus</emphasis>
|
||
</taxonomicName>
|
||
and shorter than it in
|
||
<emphasis box="[319,464,1565,1583]" italics="true" pageId="13" pageNumber="4">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[319,460,1565,1583]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
The phalanges are robust and bear smooth ginglymoid articular facets. The unguals are proportionately shorter and stockier in tyrannosaurids than in
|
||
<taxonomicName box="[99,223,1651,1669]" class="Reptilia" family="Not" genus="Dryptosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis box="[99,223,1651,1669]" italics="true" pageId="13" pageNumber="4">Dryptosaurus</emphasis>
|
||
</taxonomicName>
|
||
or
|
||
<emphasis box="[257,369,1651,1669]" italics="true" pageId="13" pageNumber="4">
|
||
<taxonomicName box="[257,365,1651,1669]" class="Reptilia" family="Not" genus="Eotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="genus">Eotyrannus</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
The distal end tapers to a point in these two basal forms, as well as in
|
||
<emphasis box="[455,716,1679,1698]" italics="true" pageId="13" pageNumber="4">
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[455,572,1679,1698]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="genus">Gorgosaurus</taxonomicName>
|
||
,
|
||
<taxonomicName baseAuthorityName="Parks" baseAuthorityYear="1933" box="[582,712,1679,1698]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="genus">Albertosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and
|
||
<emphasis box="[144,275,1708,1726]" italics="true" pageId="13" pageNumber="4">
|
||
<taxonomicName box="[144,271,1708,1726]" class="Reptilia" family="Not" genus="Dryptosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="genus">Dryptosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
but is thicker throughout its length, with a blunter tip, in species of
|
||
<emphasis box="[354,502,1737,1755]" italics="true" pageId="13" pageNumber="4">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[354,498,1737,1755]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
The manual unguals are more laterally compressed than those of the pes and bear prominent grooves for the claw sheath. The flexor tubercles are less well developed in tyrannosaurids than in
|
||
<taxonomicName box="[563,687,1823,1841]" class="Reptilia" family="Not" genus="Dryptosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis box="[563,687,1823,1841]" italics="true" pageId="13" pageNumber="4">Dryptosaurus</emphasis>
|
||
</taxonomicName>
|
||
or
|
||
<emphasis box="[99,211,1852,1870]" italics="true" pageId="13" pageNumber="4">
|
||
<taxonomicName box="[99,207,1852,1870]" class="Reptilia" family="Not" genus="Eotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="genus">Eotyrannus</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
</paragraph>
|
||
<caption httpUri="https://zenodo.org/record/3374558/files/figure.png" pageId="13" pageNumber="4" startId="13.[747,815,719,734]" targetBox="[804,1308,100,693]" targetPageId="13">
|
||
<paragraph blockId="13.[747,1330,717,807]" pageId="13" pageNumber="4">
|
||
FIGURE 5.17. Manus of tyrannosaurids: A, right metacarpal III, digit and metacarpal II, and digit and metacarpal I of
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[1158,1324,742,759]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="species" species="libratus">
|
||
<emphasis box="[1158,1324,742,759]" italics="true" pageId="13" pageNumber="4">Gorgosaurus libratus</emphasis>
|
||
</taxonomicName>
|
||
in medial view; B, left manus of
|
||
<taxonomicName baseAuthorityName="Maleev" baseAuthorityYear="1955" box="[1021,1177,766,783]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis box="[1021,1177,766,783]" italics="true" pageId="13" pageNumber="4">Tarbosaurus bataar</emphasis>
|
||
</taxonomicName>
|
||
in cranial view. Scale = 5 cm. (After Lambe 1917a; and Maleev 1974.)
|
||
</paragraph>
|
||
</caption>
|
||
<caption httpUri="https://zenodo.org/record/3374560/files/figure.png" pageId="13" pageNumber="4" startId="13.[99,167,855,870]" targetBox="[107,710,99,834]" targetPageId="13">
|
||
<paragraph blockId="13.[99,683,853,943]" pageId="13" pageNumber="4">
|
||
FIGURE 5.16. Forearm elements of
|
||
<emphasis box="[401,554,854,870]" italics="true" pageId="13" pageNumber="4">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[401,550,854,870]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
|
||
:
|
||
</emphasis>
|
||
A, right ulna in cranial, lateral, caudal, medial, proximal, and distal views; B, right radius in cranial, lateral, caudal, medial, proximal, and distal views. Scale = 10 cm. (After Carpenter and Smith 2001.)
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="13.[99,716,990,1927]" lastBlockId="13.[747,1364,874,1926]" pageId="13" pageNumber="4">
|
||
The ilium is long and bladelike in tyrannosauroids, only slightly shorter than the femur in albertosaurines and longer than the femur in tyrannosaurines (
|
||
<figureCitation box="[1089,1166,874,894]" pageId="13" pageNumber="4" startId="14.[99,166,1107,1122]" targetBox="[111,706,101,1081]" targetPageId="14">fig. 5.18</figureCitation>
|
||
; Holtz 2001b; Currie 2003a). The dorsal portion of the cranial edge of the preacetabular blade is concave in
|
||
<taxonomicName authorityName="Madsen" authorityYear="1974" box="[976,1103,932,951]" class="Reptilia" family="Allosauridae" genus="Stokesosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis box="[976,1103,932,951]" italics="true" pageId="13" pageNumber="4">Stokesosaurus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1149,1318,932,951]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
. The preacetabular blade is deep in lateral view due to an apomorphic hooklike ventral projection extending from the preacetabular blade.This ventral projection is supported at its base by a medial shelf contacting the cranial sacral ribs (
|
||
<figureCitation box="[1137,1234,1046,1066]" pageId="13" pageNumber="4" startId="14.[99,166,1107,1122]" targetBox="[111,706,101,1081]" targetPageId="14">fig. 5.18A</figureCitation>
|
||
). The ventral projection and the preacetabular blade meet at a notch on the cranial margin of the ilium. The dorsal margins of the ilia converge more closely along the midline in
|
||
<taxonomicName authorityName="Madsen" authorityYear="1974" box="[1189,1319,1133,1152]" class="Reptilia" family="Allosauridae" genus="Stokesosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1189,1319,1133,1152]" italics="true" pageId="13" pageNumber="4">Stokesosaurus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[747,916,1161,1180]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
than in most other theropods, sometimes contacting the sacral spinous processes and nearly contacting each other. On the lateral surface of the ilium, dorsal to the acetabulum, is a prominent vertical midline crest. The pubic peduncle is more massive than the ischial and longer craniocaudally than wide transversely. In
|
||
<emphasis box="[1109,1364,1305,1324]" italics="true" pageId="13" pageNumber="4">
|
||
<taxonomicName baseAuthorityName="Osmolska" baseAuthorityYear="1996" box="[1109,1218,1306,1324]" class="Reptilia" family="Not" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="genus">Bagaraatan</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Madsen" authorityYear="1974" box="[1230,1360,1305,1324]" class="Reptilia" family="Allosauridae" genus="Stokesosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="genus">Stokesosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and tyrannosaurids the cranial surface of the pubic peduncle is concave (Rauhut 2003), and there is a ventral flange extending from it (Hutchinson 2001a).The supracetabular crest is reduced. The postacetabular blade is longer than the preacetabular blade and has a squared end (more tapered in
|
||
<taxonomicName baseAuthorityName="Maleev" baseAuthorityYear="1974" box="[1135,1248,1448,1467]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1135,1248,1448,1467]" italics="true" pageId="13" pageNumber="4">Tarbosaurus</emphasis>
|
||
</taxonomicName>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="13.[747,1364,874,1926]" pageId="13" pageNumber="4">
|
||
The pubis is long and rodlike and extends subvertically relative to the axis of the sacrum (
|
||
<figureCitation box="[1078,1177,1505,1525]" pageId="13" pageNumber="4" startId="14.[99,166,1107,1122]" targetBox="[111,706,101,1081]" targetPageId="14">fig. 5.18B</figureCitation>
|
||
). The shaft curves cranially in
|
||
<taxonomicName box="[871,995,1535,1553]" class="Reptilia" family="Not" genus="Dryptosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis box="[871,995,1535,1553]" italics="true" pageId="13" pageNumber="4">Dryptosaurus</emphasis>
|
||
</taxonomicName>
|
||
and tyrannosaurids, and a cranially projecting, crestlike pubic tubercle is present near the proximal end of the shaft (Hutchinson 2001a). The pubic boot is large in tyrannosaurids, about half as long craniocaudally as the pubis is long dorsoventrally in albertosaurines and two-thirds as long in tyrannosaurines. The pubic boot is equally long cranially as caudally and narrow transversely, and it tapers caudally to a point (less so cranially). The obturator foramen opens ventrally to form an obturator notch.The pubic shafts are closely appressed throughout the distal two-thirds of the pubis length; there is no cranial pubic foramen.
|
||
</paragraph>
|
||
<paragraph blockId="13.[747,1364,874,1926]" lastBlockId="14.[99,716,1219,1927]" lastPageId="14" lastPageNumber="5" pageId="13" pageNumber="4">
|
||
The ischium is rodlike, slender, points caudoventrally, and tapers to a point (
|
||
<figureCitation box="[918,1013,1877,1897]" pageId="13" pageNumber="4" startId="14.[99,166,1107,1122]" targetBox="[111,706,101,1081]" targetPageId="14">fig. 5.18B</figureCitation>
|
||
). The obturator process is triangular and proximally placed. In tyrannosaurids a ventrally extending sheet of bone continues the obturator process down at least half of the length of the ischial shaft. There is a prominent semicircular scar on the caudolateral surface of the proximal end of the ischium, caudal to the obturator process and ventral to the iliac articulation (Holtz 2001b). Hutchinson (2001a) and Carrano and Hutchinson (2002) reconstruct this as an attachment for M. flexor tibialis internus part 3.
|
||
</paragraph>
|
||
<caption httpUri="https://zenodo.org/record/3374562/files/figure.png" pageId="14" pageNumber="5" startId="14.[99,166,1107,1122]" targetBox="[111,706,101,1081]" targetPageId="14">
|
||
<paragraph blockId="14.[99,689,1105,1171]" pageId="14" pageNumber="5">
|
||
FIGURE 5.18. Pelvis of
|
||
<emphasis box="[296,451,1106,1122]" italics="true" pageId="14" pageNumber="5">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[296,447,1106,1122]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="5" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
A, cranial view of ilium and sacrum (arrow indicates tyrannosaurid medial shelf); B, right lateral view of pelvis. Scale = 10 cm. (After Osborn 1917.)
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="14.[99,716,1219,1927]" lastBlockId="14.[747,1364,101,1926]" pageId="14" pageNumber="5">
|
||
The hindlimb of tyrannosauroids is long relative to their body size (Holtz 1995b; Currie 2003a). As in all nonavian theropods, the tibia becomes proportionately shorter relative to the femur as the body size increases (Holtz 1995b); consequently, using the proportions of the hindlimb elements without reference to overall body size is problematic for taxonomy, particularly for forms (e.g., tyrannosaurids) that ranged in size from about 1 m for juveniles to 14 m in adults. In adult tyrannosauroids of small body size and in juveniles of the larger taxa the tibia is longer than the femur; in larger individuals the tibia is often as short as the femur or even shorter (Holtz 1995b; Currie 2000). However, in tyrannosauroids the tibia is proportionately longer than in any other nonavian theropod (other than ornithomimosaurs and
|
||
<taxonomicName authorityName="Janensch" authorityYear="1920" box="[313,445,1793,1812]" class="Reptilia" family="Ornithomimidae" genus="Elaphrosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis box="[313,445,1793,1812]" italics="true" pageId="14" pageNumber="5">Elaphrosaurus</emphasis>
|
||
</taxonomicName>
|
||
) of the same femur length, and the point at which tibia length is less than femur length occurs in much larger individuals than in carnosaurs, ceratosaurs, and other taxa (Holtz 1995b). Similarly, the metatarsi of tyrannosaurids are much longer and more gracile than those of other theropods of the same femur length, again with the exception of ornithomimosaurs and
|
||
<taxonomicName authorityName="Janensch" authorityYear="1920" box="[976,1105,130,149]" class="Reptilia" family="Ornithomimidae" genus="Elaphrosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis box="[976,1105,130,149]" italics="true" pageId="14" pageNumber="5">Elaphrosaurus</emphasis>
|
||
</taxonomicName>
|
||
[Holtz 1995b; Paul 2000]).
|
||
</paragraph>
|
||
<paragraph blockId="14.[747,1364,101,1926]" pageId="14" pageNumber="5">
|
||
The femur is a robust element, straight in cranial view (
|
||
<figureCitation pageId="14" pageNumber="5" startId="15.[267,335,652,667]" targetBox="[267,1195,99,626]" targetPageId="15">fig. 5.19A, B</figureCitation>
|
||
). As in most theropods, it is curved, so that it is concave caudally; this curvature is greater in even the largest tyrannosaurids than in carnosaurs. The well-defined, hemispherical head projects medially and dorsally. The greater trochanter is cleft from and lower than the femoral head. The alariform cranial trochanter is as high as or higher than the greater trochanter and can equal or exceed the height of the dorsalmost point of the femoral head. The fourth trochanter is powerfully developed. In the distal condyles the flexor sulcus is well developed and the extensor groove is deep and prominent. The scar for M. tibialis cranialis is well defined (a triangular shape) just above the tibial condyle on the cranial face of the shaft.
|
||
</paragraph>
|
||
<paragraph blockId="14.[747,1364,101,1926]" pageId="14" pageNumber="5">
|
||
The tibia, although powerfully built, is never as robust and stocky as in carnosaurs and other basal tetanurans (
|
||
<figureCitation box="[1233,1352,559,579]" pageId="14" pageNumber="5" startId="15.[267,335,652,667]" targetBox="[267,1195,99,626]" targetPageId="15">fig. 5.19C, D</figureCitation>
|
||
). The cnemial process is well developed and curves out of the lateral surface of the tibia. The incisura tibialis (excavation on the cranial portion) occupies more than two-thirds of the lateral surface of the proximal tibia. The proximolateral condyle is pinched cranially and caudally, forming a waist between the condyle and the main body of the tibia. The crista fibularis is well developed and proximally placed. On the craniodistal surface there is a prominent facet for the large ascending process of the astragalus. The fibula is slender. The proximal end is expanded to 75% or more of the craniocaudal length of the tibia. The medial surface of the proximal half of the fibula is deeply excavated. There is a distinct cranial tubercle distal to the cranial expansion, formed by two parallel longitudinal ridges, the lateral one being larger. The distal end is tapered but contacts the tarsus.
|
||
</paragraph>
|
||
<paragraph blockId="14.[747,1364,101,1926]" pageId="14" pageNumber="5">The astragalus of tyrannosauroids bears a tall, triangular ascending process that extends for one-quarter to one-third of the vertical distance of the femur. Some individuals bear a deep fossa at the base of the ascending fossa (Mader and Bradley 1989). The calcaneum is small but supports the distal end of the fibula and contributes to the lateral condyle.Two distal tarsals, disklike elements nearly equal in size, are present.</paragraph>
|
||
<paragraph blockId="14.[747,1364,101,1926]" lastBlockId="15.[99,716,1678,1927]" lastPageId="15" lastPageNumber="6" pageId="14" pageNumber="5">
|
||
The metatarsus of tyrannosauroids is long and slender (
|
||
<figureCitation pageId="14" pageNumber="5" startId="15.[283,351,1547,1562]" targetBox="[283,1174,779,1520]" targetPageId="15">figs. 5.20</figureCitation>
|
||
,
|
||
<figureCitation box="[802,847,1247,1267]" pageId="14" pageNumber="5" startId="16.[99,166,958,973]" targetBox="[106,708,104,934]" targetPageId="16">5.21</figureCitation>
|
||
). Metatarsals II and IV are subcylindrical and nearly equal in length, although IV is longer; metatarsal III is longer than the others. Metatarsal III is pinched proximally to form a slender, solid bony splint and is wedge-shaped distally. Buttressing surfaces from metatarsals II and IV contact the sides of metatarsal III. Metatarsal III has not been recovered for nontyrannosaurid tyrannosauroids, so it is not certain whether it was fully pinched in these taxa. Metatarsal III contributes only slightly to the proximal surface of the metatarsus, forming a crescentic shape (never an oval,
|
||
<emphasis box="[1083,1142,1506,1524]" italics="true" pageId="14" pageNumber="5">contra</emphasis>
|
||
Holtz 1995b) limited to the caudal half of that surface. This compact, slender metatarsal structure has been termed the
|
||
<emphasis box="[1095,1242,1564,1582]" italics="true" pageId="14" pageNumber="5">arctometatarsus</emphasis>
|
||
(Holtz 1994, 1995b). The arctometatarsi of tyrannosaurids are never proximally co-ossified. There is a contact on the dorsal part of the proximal ends of metatarsals II and IV, and metatarsals II and IV contact at midshaft on the plantar surface. Juvenile or small adult tyrannosaurid metatarsi might be confused with those of large ornithomimids (Currie 2000), but in the latter metatarsal III has an oval rather than a crescentic proximal surface. The distal articular surfaces of metatarsals II and IV are elongate and condylar, while that of metatarsal III has a nearly rectangular ginglymoid form. Metatarsal II can be distinguished from IV in that the distal articulation of the former is nearly square in outline, while that of the latter is narrower and more triangular; in dorsal view metatarsal II generally has a greater area and is wider mediolaterally than metatarsal IV.Metatarsal I is greatly reduced in size and positioned slightly distal to midshaft and slightly shifted plantarly on metatarsal II (but not fully retroverted as in avialans). Metatarsal V has a cranially facing, rugose tuberosity (Brochu 2002).
|
||
</paragraph>
|
||
<caption httpUri="https://zenodo.org/record/3374564/files/figure.png" pageId="15" pageNumber="6" startId="15.[267,335,652,667]" targetBox="[267,1195,99,626]" targetPageId="15">
|
||
<paragraph blockId="15.[267,1171,650,716]" pageId="15" pageNumber="6">
|
||
FIGURE 5.19. Femora and tibiae:A, right femur of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[699,848,652,668]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="6" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[699,848,652,668]" italics="true" pageId="15" pageNumber="6">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
in medial and cranial views; B, left femur of
|
||
<taxonomicName baseAuthorityName="Maleev" baseAuthorityYear="1955" box="[345,501,675,692]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="6" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis box="[345,501,675,692]" italics="true" pageId="15" pageNumber="6">Tarbosaurus bataar</emphasis>
|
||
</taxonomicName>
|
||
in caudal view; C, left tibia of
|
||
<taxonomicName baseAuthorityName="Maleev" baseAuthorityYear="1955" box="[758,914,675,692]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="6" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis box="[758,914,675,692]" italics="true" pageId="15" pageNumber="6">Tarbosaurus bataar</emphasis>
|
||
</taxonomicName>
|
||
in cranial view; D, left tibia of
|
||
<taxonomicName authorityName="Gilmore" authorityYear="1933" box="[267,427,699,716]" class="Reptilia" family="Tyrannosauridae" genus="Alectrosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="6" phylum="Chordata" rank="species" species="olseni">
|
||
<emphasis box="[267,427,699,716]" italics="true" pageId="15" pageNumber="6">Alectrosaurus olseni</emphasis>
|
||
</taxonomicName>
|
||
in cranial view. Scale = 20 cm. (After Osborn 1917; Maleev 1974; and Gilmore 1933a.)
|
||
</paragraph>
|
||
</caption>
|
||
<caption httpUri="https://zenodo.org/record/3374566/files/figure.png" pageId="15" pageNumber="6" startId="15.[283,351,1547,1562]" targetBox="[283,1174,779,1520]" targetPageId="15">
|
||
<paragraph blockId="15.[283,1166,1545,1611]" pageId="15" pageNumber="6">
|
||
FIGURE 5.20. Pedes: A, right metatarsus of
|
||
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[652,818,1546,1563]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="6" phylum="Chordata" rank="species" species="libratus">
|
||
<emphasis box="[652,818,1546,1563]" italics="true" pageId="15" pageNumber="6">Gorgosaurus libratus</emphasis>
|
||
</taxonomicName>
|
||
in proximal (with distal tarsals in articu- lation), cranial, and distal views; B, left metatarsus of
|
||
<taxonomicName authorityName="Gilmore" authorityYear="1933" box="[738,897,1570,1587]" class="Reptilia" family="Tyrannosauridae" genus="Alectrosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="6" phylum="Chordata" rank="species" species="olseni">
|
||
<emphasis box="[738,897,1570,1587]" italics="true" pageId="15" pageNumber="6">Alectrosaurus olseni</emphasis>
|
||
</taxonomicName>
|
||
in cranial view; C, left pes of
|
||
<taxonomicName baseAuthorityName="Maleev" baseAuthorityYear="1955" box="[283,440,1594,1611]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="6" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis box="[283,440,1594,1611]" italics="true" pageId="15" pageNumber="6">Tarbosaurus bataar</emphasis>
|
||
</taxonomicName>
|
||
in cranial view.Scale = 10 cm.(After Lambe 1917a; Gilmore 1933a; and Maleev 1974.)
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="15.[99,716,1678,1927]" lastBlockId="15.[747,1364,1677,1927]" pageId="15" pageNumber="6">
|
||
The arctometatarsalian condition is as well developed in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[99,274,1880,1898]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="6" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[99,274,1880,1898]" italics="true" pageId="15" pageNumber="6">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
as in other tyrannosaurids: metatarsal III is reduced and compressed proximally, and the metatarsus is long, slender, and compact (Holtz 1995b; Brochu 2002). However, outdated restorations of incorrectly mounted hindlimbs of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[747,798,1736,1754]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="6" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[747,798,1736,1754]" italics="true" pageId="15" pageNumber="6">T. rex</emphasis>
|
||
</taxonomicName>
|
||
(with a thicker metatarsal III visible for its entire length throughout the metatarsus and subequal in length to the other weight-bearing metatarsals) continue to be published in modern vertebrate paleontological literature (e.g., Carroll 1988, 1997; Benton 1997).
|
||
</paragraph>
|
||
<paragraph blockId="15.[747,1364,1677,1927]" lastBlockId="16.[99,716,1132,1295]" lastPageId="16" lastPageNumber="7" pageId="15" pageNumber="6">
|
||
The pedal phalangeal formula is 2-3-4-5-0. Digit III is the longest, and digits II and IV are subequal in length. Digit I is reduced but does bear an ungual. The subcylindrical phalanges are wider than tall. The pedal unguals are stouter and less laterally compressed than the manual unguals and considerably larger. The grooves for the claw sheaths are well developed. The unguals are blunter and less tapered than those of carnosaurs such as
|
||
<taxonomicName authorityName="Currie & Zhao" authorityYear="1993" box="[174,258,1276,1295]" class="Reptilia" family="Neovenatoridae" genus="Sinraptor" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="genus">
|
||
<emphasis box="[174,258,1276,1295]" italics="true" pageId="16" pageNumber="7">Sinraptor</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<emphasis box="[305,410,1276,1295]" italics="true" pageId="16" pageNumber="7">
|
||
<taxonomicName box="[305,406,1276,1295]" class="Reptilia" family="Allosauridae" genus="Allosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="genus">Allosaurus</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
</paragraph>
|
||
<caption httpUri="https://zenodo.org/record/3374568/files/figure.png" pageId="16" pageNumber="7" startId="16.[99,166,958,973]" targetBox="[106,708,104,934]" targetPageId="16">
|
||
<paragraph blockId="16.[99,691,956,1022]" pageId="16" pageNumber="7">
|
||
FIGURE 5.21. Plots comparing metatarsus length to femur length
|
||
<emphasis box="[99,156,981,997]" italics="true" pageId="16" pageNumber="7">(upper)</emphasis>
|
||
and tibia length
|
||
<emphasis box="[301,356,981,998]" italics="true" pageId="16" pageNumber="7">(lower)</emphasis>
|
||
in tyrannosaurids (solid triangles), ornithomimosaurs (open squares), and other nonavian theropods (x’s).
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="16.[99,411,1364,1388]" box="[99,411,1364,1388]" pageId="16" pageNumber="7">
|
||
<heading bold="true" box="[99,411,1364,1388]" fontSize="10" level="1" pageId="16" pageNumber="7" reason="0">
|
||
<emphasis bold="true" box="[99,411,1364,1388]" pageId="16" pageNumber="7">Systematics and Evolution</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="16.[99,716,1420,1926]" pageId="16" pageNumber="7">
|
||
Tyrannosaurids were considered members of Carnosauria for many decades (Huene 1932; Colbert 1955; Romer 1956; Walker 1964; Steel 1970; Gauthier 1986; Bonaparte et al. 1990; Molnar et al. 1990). However, as early as the 1920s Huene (1923, 1926c) and Matthew and Brown (1922) suggested that these giant theropods shared a closer phylogenetic position with small coelurosaurian dinosaurs, in particular
|
||
<taxonomicName authorityName="Osborn" authorityYear="1903" box="[411,532,1592,1611]" class="Reptilia" family="Compsognathidae" genus="Ornitholestes" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="genus">
|
||
<emphasis box="[411,532,1592,1611]" italics="true" pageId="16" pageNumber="7">Ornitholestes</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="Marsh" authorityYear="1890" class="Reptilia" family="Ornithomimidae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="family">Ornithomimidae</taxonomicName>
|
||
, than with other large-bodied theropods, such as
|
||
<taxonomicName box="[617,715,1621,1640]" class="Reptilia" family="Allosauridae" genus="Allosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="genus">
|
||
<emphasis box="[617,715,1621,1640]" italics="true" pageId="16" pageNumber="7">Allosaurus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<emphasis box="[145,282,1649,1668]" italics="true" pageId="16" pageNumber="7">
|
||
<taxonomicName authorityName="Buckland" authorityYear="1822" box="[145,278,1649,1668]" class="Reptilia" family="Megalosauridae" genus="Megalosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="genus">Megalosaurus</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
This hypothesis has since been supported in subsequent numerical phylogenetic analyses (Novas 1992b; Pérez-Moreno et al. 1993, 1994; Holtz 1994, 1998a, 2001a; Sereno et al. 1994, 1996; Forster et al. 1998; Makovicky and Sues 1998; Sereno 1999a; Currie and Carpenter 2000; Norell et al. 2000; Rauhut 2003). Subsequently,
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[493,662,1793,1812]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
and Tyrannosauroidea are now included within the taxon
|
||
<taxonomicName authorityName="Huene" authorityYear="1914" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Avetheropoda" pageId="16" pageNumber="7" phylum="Chordata" rank="subOrder" subOrder="Coelurosauria">Coelurosauria</taxonomicName>
|
||
rather than in Carnosauria.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |