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<document id="68B926902B842A2D39D84035EB950371" ID-DOI="10.26879/702" ID-ISSN="1094-8074" ID-Zenodo-Dep="11063310" IM.bibliography_approvedBy="julia" IM.illustrations_approvedBy="julia" IM.materialsCitations_approvedBy="julia" IM.metadata_approvedBy="felipe" IM.tables_approvedBy="julia" IM.taxonomicNames_approvedBy="julia" IM.treatments_approvedBy="julia" checkinTime="1714014708037" checkinUser="felipe" docAuthor="Poole, Karen E." docDate="2016" docId="6D2D8786A2775F0C740FFEC626804DBE" docLanguage="en" docName="PalaeontolElectron.25.3.a30.pdf" docOrigin="Palaeontologia Electronica (a 30) 25 (3)" docSource="http://dx.doi.org/10.26879/702" docStyle="DocumentStyle:322FC57CC99045555090E8A84CFB029A.4:PalaeontolElectron.2018-.journal_article" docStyleId="322FC57CC99045555090E8A84CFB029A" docStyleName="PalaeontolElectron.2018-.journal_article" docStyleVersion="4" docTitle="Animalia Sereno 1986" docType="treatment" docVersion="2" lastPageNumber="17" masterDocId="9114FFFEA2675F1C753EFFB622124921" masterDocTitle="Phylogeny of iguanodontian dinosaurs and the evolution of quadrupedality" masterLastPageNumber="65" masterPageNumber="1" pageNumber="17" updateTime="1714997633796" updateUser="julia" zenodo-license-document="CC-BY-NC-SA-4.0">
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<mods:title id="F55879D17F8798CF8C6C0EF46B075958">Phylogeny of iguanodontian dinosaurs and the evolution of quadrupedality</mods:title>
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<subSubSection id="AD9E651BA2775F0C740FFEC620B548AB" box="[305,679,368,394]" pageId="16" pageNumber="17" type="nomenclature">
<paragraph id="E53B3690A2775F0C740FFEC620B548AB" blockId="16.[305,679,368,394]" box="[305,679,368,394]" pageId="16" pageNumber="17">
<taxonomicName id="22844D13A2775F0C740FFEC623ED48AB" authority="Sereno, 1986" authorityName="Sereno" authorityYear="1986" box="[305,511,368,394]" kingdom="Animalia" pageId="16" pageNumber="17" rank="clade">IGUANODONTIA</taxonomicName>
<bibRefCitation id="81154B61A2775F0C7739FEC620B548AB" author="Sereno, P. C." box="[519,679,368,394]" journalOrPublisher="National Geographic Research" pageId="16" pageNumber="17" pagination="234 - 256" part="2" refId="ref22197" refString="Sereno, P. C. 1986. Phylogeny of the bird-hipped dinosaurs (Order Ornithischia). National Geographic Research, 2: 234 - 256." title="Phylogeny of the bird-hipped dinosaurs (Order Ornithischia)" type="journal article" year="1986">Sereno, 1986</bibRefCitation>
</paragraph>
</subSubSection>
<subSubSection id="AD9E651BA2775F0C75FEFE2D26804DBE" pageId="16" pageNumber="17" type="description">
<paragraph id="E53B3690A2775F0C75FEFE2D231E4B15" blockId="16.[192,793,411,1908]" pageId="16" pageNumber="17">
<emphasis id="D7F0EA82A2775F0C75FEFE2D23FE4895" bold="true" box="[192,492,411,436]" pageId="16" pageNumber="17">Phylogenetic definition.</emphasis>
The most inclusive clade containing
<taxonomicName id="22844D13A2775F0C7476FE0D210A48F5" authority="Parks, 1922" authorityName="Parks" authorityYear="1922" box="[328,792,443,468]" class="Reptilia" family="Lambeosauridae" genus="Parasaurolophus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="walkeri">
<emphasis id="D7F0EA82A2775F0C7476FE0D206148F5" box="[328,627,443,468]" italics="true" pageId="16" pageNumber="17">Parasaurolophus walkeri</emphasis>
<bibRefCitation id="81154B61A2775F0C77BCFE0D210A48F5" author="Parks, W. A." box="[642,792,443,468]" journalOrPublisher="University of Toronto Studies, Geological Series" pageId="16" pageNumber="17" pagination="1 - 32" part="13" refId="ref21238" refString="Parks, W. A. 1922. Parasaurolophus walkeri: a new genus and species of crested trachodont dinosaur. University of Toronto Studies, Geological Series, 13: 1 - 32." title="Parasaurolophus walkeri: a new genus and species of crested trachodont dinosaur" type="journal article" year="1922">Parks, 1922</bibRefCitation>
</taxonomicName>
but not
<taxonomicName id="22844D13A2775F0C7424FE6D20BA48D5" authority="Huxley, 1869" authorityName="Huxley" authorityYear="1869" box="[282,680,475,500]" class="Reptilia" family="Hypsilophodontidae" genus="Hypsilophodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="foxii">
<emphasis id="D7F0EA82A2775F0C7424FE6D201048D5" box="[282,514,475,500]" italics="true" pageId="16" pageNumber="17">Hypsilophodon foxii</emphasis>
<bibRefCitation id="81154B61A2775F0C7734FE6D20BA48D5" author="Huxley, T. H." box="[522,680,475,500]" journalOrPublisher="Abstracts of the Proceedings of the Geological Society (of London)" pageId="16" pageNumber="17" pagination="3 - 4" part="204" refId="ref18591" refString="Huxley, T. H. 1869. On Hypsilophodon, a new genus of Dinosauria. Abstracts of the Proceedings of the Geological Society (of London), 204: 3 - 4." title="On Hypsilophodon, a new genus of Dinosauria" type="journal article" year="1869">Huxley, 1869</bibRefCitation>
</taxonomicName>
, or
<taxonomicName id="22844D13A2775F0C77EAFE6D20884B35" authority="Gilmore, 1913" authorityName="Gilmore" authorityYear="1913" class="Reptilia" family="Hypsilophodontidae" genus="Thescelosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="neglectus">
<emphasis id="D7F0EA82A2775F0C77EAFE6D23D94B35" italics="true" pageId="16" pageNumber="17">Thescelosaurus neglectus</emphasis>
<bibRefCitation id="81154B61A2775F0C74DCFE4D20884B35" author="Gilmore, C. W." box="[482,666,507,533]" journalOrPublisher="Smithsonian Miscellaneous Collections" pageId="16" pageNumber="17" pagination="1 - 5" part="61" refId="ref17689" refString="Gilmore, C. W. 1913. A new dinosaur from the Lance Formation of Wyoming. Smithsonian Miscellaneous Collections, 61 (5): 1 - 5." title="A new dinosaur from the Lance Formation of Wyoming" type="journal article" year="1913">Gilmore, 1913</bibRefCitation>
</taxonomicName>
(
<bibRefCitation id="81154B61A2775F0C7786FE4D23104B15" author="Sereno, P. C." journalOrPublisher="TaxonSearch" pageId="16" pageNumber="17" publicationUrl="http://www.taxonsearch.org/Archive/stem-archosauria-1.0.php" refId="ref22354" refString="Sereno, P. C. 2005. Stem Archosauria - TaxonSearch. http: // www. taxonsearch. org / Archive / stem-archosauria- 1.0. php" title="Stem Archosauria" type="url" year="2005">Sereno, 2005</bibRefCitation>
).
</paragraph>
<paragraph id="E53B3690A2775F0C75FEFD8D230E4A95" blockId="16.[192,793,411,1908]" pageId="16" pageNumber="17">
<emphasis id="D7F0EA82A2775F0C75FEFD8D20734B75" bold="true" box="[192,609,571,596]" pageId="16" pageNumber="17">Unambiguous synapomorphies.</emphasis>
In the parsimony analysis, Iguanodontia is characterized by a maxilla with a broad and triangular dorsal process of the maxilla (21.1), a quadrate extending ventrally such that the quadratojugal is well removed from the mandibular condyle (60.1), a single wear facet on each cheek tooth (131.1), opisthocoelus post-axial cervical vertebrae (157.1), a distinct indentation on the scapula superior to the glenoid, termed here the supraglenoid fossa (199.1), and a manual digit III with three or fewer phalanges (236.1).
</paragraph>
<paragraph id="E53B3690A2775F0C75C4FC0D20E84DD5" blockId="16.[192,793,411,1908]" pageId="16" pageNumber="17">
Two other synapomorphies recovered for this clade are elongate prezygopophyses on the distal caudal vertebrae (185.1), and chevrons that are strongly and asymmetrically expanded distally (188.1). The former of these is found only in
<taxonomicName id="22844D13A2775F0C77E0FB8D23514D55" authorityName=", Coria and Salgado" authorityYear="1996" class="Reptilia" family="Hypsilophodontidae" genus="Gasparinisaura" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="D7F0EA82A2775F0C77E0FB8D23514D55" italics="true" pageId="16" pageNumber="17">Gasparinisaura</emphasis>
</taxonomicName>
and
<taxonomicName id="22844D13A2775F0C74B7FBED202C4D55" box="[393,574,1115,1140]" family="Hypsilophodontidae" genus="Leaellynasaura" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="D7F0EA82A2775F0C74B7FBED202C4D55" box="[393,574,1115,1140]" italics="true" pageId="16" pageNumber="17">Leaellynasaura</emphasis>
</taxonomicName>
, and the latter in these genera plus
<taxonomicName id="22844D13A2775F0C748CFBCD20474DB5" box="[434,597,1147,1172]" family="Hypsilophodontidae" genus="Parksosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="D7F0EA82A2775F0C748CFBCD20474DB5" box="[434,597,1147,1172]" italics="true" pageId="16" pageNumber="17">Parksosaurus</emphasis>
</taxonomicName>
and
<taxonomicName id="22844D13A2775F0C779EFBCD232E4D95" authorityName="Calvo, Porfiri &amp; Novas" authorityYear="2007" genus="Macrogryphosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="D7F0EA82A2775F0C779EFBCD232E4D95" italics="true" pageId="16" pageNumber="17">Macrogryphosaurus</emphasis>
</taxonomicName>
. While they are present at the base of the clade, these characters are not widespread, and therefore not useful in diagnosing the clade.
</paragraph>
<paragraph id="E53B3690A2775F0C75C4FB4D201E4F35" blockId="16.[192,793,411,1908]" pageId="16" pageNumber="17">
There is only one overlapping character here with the diagnosis of
<bibRefCitation id="81154B61A2775F0C74FEFAAD20624C15" author="Sereno, P. C." box="[448,624,1307,1333]" journalOrPublisher="National Geographic Research" pageId="16" pageNumber="17" pagination="234 - 256" part="2" refId="ref22197" refString="Sereno, P. C. 1986. Phylogeny of the bird-hipped dinosaurs (Order Ornithischia). National Geographic Research, 2: 234 - 256." title="Phylogeny of the bird-hipped dinosaurs (Order Ornithischia)" type="journal article" year="1986">Sereno (1986)</bibRefCitation>
; the reduction of phalanges in digit III. The presence of “leaf-shaped” or mamillated denticles is more restricted within Styracosterna, and while most iguanodontians lack premaxillary teeth, both
<taxonomicName id="22844D13A2775F0C7764FA2D20F14C95" box="[602,739,1435,1460]" class="Reptilia" family="Hadrosauridae" genus="Talenkauen" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="D7F0EA82A2775F0C7764FA2D20F14C95" box="[602,739,1435,1460]" italics="true" pageId="16" pageNumber="17">Talenkauen</emphasis>
</taxonomicName>
and
<taxonomicName id="22844D13A2775F0C75FEFA0D23A54CF5" box="[192,439,1467,1492]" class="Reptilia" family="Hypsilophodontidae" genus="Tenontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="dossi">
<emphasis id="D7F0EA82A2775F0C75FEFA0D23A54CF5" box="[192,439,1467,1492]" italics="true" pageId="16" pageNumber="17">Tenontosaurus dossi</emphasis>
</taxonomicName>
have one premaxillary tooth. Iguanodontia is recovered with jackknife support of 19 and Bremer support of 4.
</paragraph>
<paragraph id="E53B3690A2775F0C75C4F9AD27134A3E" blockId="16.[192,793,411,1908]" lastBlockId="16.[840,1441,166,1183]" pageId="16" pageNumber="17">
Within the Bayesian topology, Iguanodontia (PP=0.43) lacks the basal pectinate region found in the parsimony analysis and is instead composed of the sister groups of rhabdodontoids and Dryomorpha.
<taxonomicName id="22844D13A2775F0C75C1F92D23A14F95" authorityName=", Coria and Salgado" authorityYear="1996" box="[255,435,1691,1716]" class="Reptilia" family="Hypsilophodontidae" genus="Gasparinisaura" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="D7F0EA82A2775F0C75C1F92D23A14F95" box="[255,435,1691,1716]" italics="true" pageId="16" pageNumber="17">Gasparinisaura</emphasis>
</taxonomicName>
,
<taxonomicName id="22844D13A2775F0C74F8F92D20694F95" box="[454,635,1691,1716]" family="Hypsilophodontidae" genus="Leaellynasaura" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="D7F0EA82A2775F0C74F8F92D20694F95" box="[454,635,1691,1716]" italics="true" pageId="16" pageNumber="17">Leaellynasaura</emphasis>
</taxonomicName>
, and
<taxonomicName id="22844D13A2775F0C77FBF92D23734FF5" authorityName="Calvo, Porfiri &amp; Novas" authorityYear="2007" genus="Macrogryphosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="D7F0EA82A2775F0C77FBF92D23734FF5" italics="true" pageId="16" pageNumber="17">Macrogryphosaurus</emphasis>
</taxonomicName>
are excluded from Iguanodontia, and are recovered instead with the hypsilophodontids. This rearrangement of taxa leads to different synapomorphies for Iguanodontia between the parsimony and Bayesian analyses. Synapomorphies for the Bayesian topology include: premaxilla flaring laterally to form a floor of the narial fossa (3.1), small antorbital fenestra (31.1), predentary with denticulate oral margin (87.1), ventral process of predentary deeply bifurcated (89.1), cheek teeth with crowns tapering toward the root (127.1, 128.1), cheek teeth that are closely packed without spaces between roots (126.1), cheek teeth with one wear facet on each tooth (131.1), cheek teeth lacking a basal ridge (“cingulum”) (148.1), caudal vertebrae with distal facets for chevrons much larger than proximal facets (183.0), humerus with an elongate deltopectoral crest (&gt;43% humeral length) (214.1), manual digit III with three or fewer phalanges (236.1), first phalanx of manual digits II-IV more than twice the length of the second phalanx (239.1), ischium with an untwisted shaft (283.1), ischium with an expanded distal end (288.1), femur with a cranial intercondylar sulcus (300.1), and a caudal intercondylar sulcus partially roofed by the medial condyle (302.1).
</paragraph>
<paragraph id="E53B3690A2775F0C7676FC9026004D3E" blockId="16.[840,1441,166,1183]" pageId="16" pageNumber="17">
<emphasis id="D7F0EA82A2775F0C7676FC9021D14A1E" bold="true" box="[840,963,806,831]" pageId="16" pageNumber="17">Topology.</emphasis>
In the parsimony analysis (
<figureCitation id="7DBF2A15A2775F0C703CFC9027644A1E" box="[1282,1398,806,831]" captionStart="FIGURE 6" captionStartId="10.[194,285,1048,1071]" captionTargetBox="[204,1427,175,1009]" captionTargetId="graphics-257@10.[204,1427,175,1010]" captionTargetPageId="10" captionText="FIGURE 6. Parsimony (left) and Bayesian (right) trees plotted together. Strict consensus of 84 MPTs after pruning Oryctodromeus, Atlascopscosaurus, Planicoxa, Cumnoria, Cedrorestes, and NHMUK R28860. Jackknife values above 20 (with 10% chance of character removal) are shown above and to the left of their respective nodes. Bremer supports above one are shown below and to the left of their respective nodes, and bolded. CI=0.272, RI=0.634. Maximum clade credibility tree produced by Bayesian analysis showing posterior probabilities below and to the right of their respective node." figureDoi="http://doi.org/10.5281/zenodo.11063322" httpUri="https://zenodo.org/record/11063322/files/figure.png" pageId="16" pageNumber="17">Figures 6</figureCitation>
,
<figureCitation id="7DBF2A15A2775F0C70BFFC9027814A1E" box="[1409,1427,806,831]" captionStart="FIGURE 7" captionStartId="11.[200,291,1774,1797]" captionTargetBox="[198,1439,164,1749]" captionTargetId="figure-91@11.[193,1440,164,1750]" captionTargetPageId="11" captionText="FIGURE 7. Time scaled parsimony tree, showing assigned age ranges and broad-scale geographic data. Jackknife values above 20 (with 10% chance of character removal) are shown above and to the left of their respective nodes. Bremer supports above one are shown below and to the left of their respective nodes, and bolded." figureDoi="http://doi.org/10.5281/zenodo.11063324" httpUri="https://zenodo.org/record/11063324/files/figure.png" pageId="16" pageNumber="17">7</figureCitation>
), the basally branching portion of Iguanodontia forms a pectinate topology outside of Dryomorpha, which includes
<emphasis id="D7F0EA82A2775F0C712BFC30264B4AFE" italics="true" pageId="16" pageNumber="17">
<taxonomicName id="22844D13A2775F0C712BFC3026DE4ABE" authorityName=", Coria and Salgado" authorityYear="1996" box="[1045,1228,902,927]" class="Reptilia" family="Hypsilophodontidae" genus="Gasparinisaura" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">Gasparinisaura</taxonomicName>
,
<taxonomicName id="22844D13A2775F0C71DAFC30278E4ABE" box="[1252,1436,902,927]" family="Hypsilophodontidae" genus="Leaellynasaura" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">Leaellynasaura</taxonomicName>
,
<taxonomicName id="22844D13A2775F0C7676FC1026244A9E" authorityName="Calvo, Porfiri &amp; Novas" authorityYear="2007" box="[840,1078,934,959]" genus="Macrogryphosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">Macrogryphosaurus</taxonomicName>
,
<taxonomicName id="22844D13A2775F0C7166FC1026F64A9E" box="[1112,1252,934,959]" class="Reptilia" family="Hadrosauridae" genus="Talenkauen" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">Talenkauen</taxonomicName>
,
<taxonomicName id="22844D13A2775F0C703BFC1027894A9E" box="[1285,1435,934,959]" family="Dryosauridae" genus="Valdosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">Valdosaurus</taxonomicName>
,
<taxonomicName id="22844D13A2775F0C7676FC7021D54AFE" authorityName="Coria &amp; Calvo" authorityYear="2002" box="[840,967,966,991]" class="Reptilia" genus="Anabisetia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">Anabisetia</taxonomicName>
,
<taxonomicName id="22844D13A2775F0C76E3FC7026464AFE" box="[989,1108,966,991]" genus="Trinisaura" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">Trinisaura</taxonomicName>
,
</emphasis>
and
<taxonomicName id="22844D13A2775F0C7196FC7027464AFE" authorityName="SAM" authorityYear="2732" box="[1192,1364,966,991]" family="Dryosauridae" genus="Kangnasaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis id="D7F0EA82A2775F0C7196FC7027464AFE" box="[1192,1364,966,991]" italics="true" pageId="16" pageNumber="17">Kangnasaurus</emphasis>
</taxonomicName>
. It is supported by a jackknife value of 19 and a Bremmer support of 4.
</paragraph>
<paragraph id="E53B3690A2775F0C76BCFB9026804DBE" blockId="16.[840,1441,166,1183]" pageId="16" pageNumber="17">
In the Bayesian analysis (
<figureCitation id="7DBF2A15A2775F0C7190FB9027314D1E" box="[1198,1315,1062,1087]" captionStart="FIGURE 6" captionStartId="10.[194,285,1048,1071]" captionTargetBox="[204,1427,175,1009]" captionTargetId="graphics-257@10.[204,1427,175,1010]" captionTargetPageId="10" captionText="FIGURE 6. Parsimony (left) and Bayesian (right) trees plotted together. Strict consensus of 84 MPTs after pruning Oryctodromeus, Atlascopscosaurus, Planicoxa, Cumnoria, Cedrorestes, and NHMUK R28860. Jackknife values above 20 (with 10% chance of character removal) are shown above and to the left of their respective nodes. Bremer supports above one are shown below and to the left of their respective nodes, and bolded. CI=0.272, RI=0.634. Maximum clade credibility tree produced by Bayesian analysis showing posterior probabilities below and to the right of their respective node." figureDoi="http://doi.org/10.5281/zenodo.11063322" httpUri="https://zenodo.org/record/11063322/files/figure.png" pageId="16" pageNumber="17">Figures 6</figureCitation>
,
<figureCitation id="7DBF2A15A2775F0C7010FB9027524D1E" box="[1326,1344,1062,1087]" captionStart="FIGURE 8" captionStartId="12.[196,286,1624,1647]" captionTargetBox="[195,1436,163,1599]" captionTargetId="graphics-57@12.[196,1437,163,1599]" captionTargetPageId="12" captionText="FIGURE 8. Maximum Clade Credibility tree produced by Bayesian analysis. Posterior probabilities are shown to the left of their nodes. The geologic timescale is shown across the top. Tips represent the average age found for each OTU across all sampled trees." figureDoi="http://doi.org/10.5281/zenodo.11063326" httpUri="https://zenodo.org/record/11063326/files/figure.png" pageId="16" pageNumber="17">8</figureCitation>
), Iguanodontia bifurcates into rhabdodontoids and dryomorphans. Iguanodontia is supported by a posterior probability of 0.43.
</paragraph>
</subSubSection>
</treatment>
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