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<document ID-DOI="10.1093/zoolinnean/zlac033" ID-GBIF-Dataset="d18be66e-8788-47d3-bba3-11ad8bb0f126" ID-ISSN="0024-4082" ID-Zenodo-Dep="7922143" IM.bibliography_approvedBy="juliana" IM.illustrations_approvedBy="juliana" IM.materialsCitations_approvedBy="juliana" IM.metadata_approvedBy="juliana" IM.taxonomicNames_approvedBy="juliana" IM.treatments_approvedBy="juliana" checkinTime="1683701348189" checkinUser="plazi" docAuthor="Macungo, Zanildo, Benoit, Julien, Fernandez, Vincent &amp; Araújo, Ricardo M N" docDate="2023" docId="3F6A87ABFFB1FFE771E5FE79FC25FA3E" docLanguage="en" docName="zlac033.pdf" docOrigin="Zoological Journal of the Linnean Society 198 (1)" docSource="https://academic.oup.com/zoolinnean/article/198/1/1/6596559" docStyle="DocumentStyle:36B3BD6A90C22AB4F7F465C853188CC8.7:ZoolJLinnSoc.2017-.journal_article" docStyleId="36B3BD6A90C22AB4F7F465C853188CC8" docStyleName="ZoolJLinnSoc.2017-.journal_article" docStyleVersion="7" docTitle="Kawingasaurus Cox 1972" docType="treatment" docVersion="3" lastPageNumber="25" masterDocId="C353FFD3FFA9FFFF7115FFA9FFFCFFDA" masterDocTitle="X-ray microcomputed and synchrotron tomographic analysis of the basicranial axis of emydopoid dicynodonts: implications for fossoriality and phylogeny" masterLastPageNumber="46" masterPageNumber="1" pageNumber="25" updateTime="1683750160436" updateUser="ExternalLinkService">
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<mods:title>X-ray microcomputed and synchrotron tomographic analysis of the basicranial axis of emydopoid dicynodonts: implications for fossoriality and phylogeny</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Macungo, Zanildo</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Benoit, Julien</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Fernandez, Vincent</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Araújo, Ricardo M N</mods:namePart>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
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<mods:title>Zoological Journal of the Linnean Society</mods:title>
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<mods:part>
<mods:date>2023</mods:date>
<mods:detail type="pubDate">
<mods:number>2023-05-01</mods:number>
</mods:detail>
<mods:detail type="volume">
<mods:number>198</mods:number>
</mods:detail>
<mods:detail type="issue">
<mods:number>1</mods:number>
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<mods:url>https://academic.oup.com/zoolinnean/article/198/1/1/6596559</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">10.1093/zoolinnean/zlac033</mods:identifier>
<mods:identifier type="GBIF-Dataset">d18be66e-8788-47d3-bba3-11ad8bb0f126</mods:identifier>
<mods:identifier type="ISSN">0024-4082</mods:identifier>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.7922177" ID-Zenodo-Dep="7922177" LSID="urn:lsid:plazi:treatment:3F6A87ABFFB1FFE771E5FE79FC25FA3E" httpUri="http://treatment.plazi.org/id/3F6A87ABFFB1FFE771E5FE79FC25FA3E" lastPageNumber="25" pageId="24" pageNumber="25">
<subSubSection pageId="24" pageNumber="25" type="nomenclature">
<paragraph blockId="24.[240,665,463,488]" box="[240,665,463,488]" pageId="24" pageNumber="25">
<heading box="[240,665,463,488]" centered="true" fontSize="9" level="2" pageId="24" pageNumber="25" reason="2">
<emphasis box="[240,541,464,487]" italics="true" pageId="24" pageNumber="25">
<taxonomicName authorityName="Cox" authorityYear="1972" box="[240,434,464,487]" class="Reptilia" family="Cistecephalidae" genus="Kawingasaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Therapsida" pageId="24" pageNumber="25" phylum="Chordata" rank="genus">
<smallCapsWord baselines="481,482" box="[240,434,464,487]" lowerCaseFontSize="8" mainFontSize="10" normCase="title" normString="Kawingasaurus" pageId="24" pageNumber="25">KAWINGASAURUS</smallCapsWord>
</taxonomicName>
<smallCapsWord baselines="482" box="[441,541,467,487]" lowerCaseFontSize="8" mainFontSize="10" normCase="lower" normString="fossilis" pageId="24" pageNumber="25">FOSSILIS</smallCapsWord>
</emphasis>
<bibRefCitation author="Cox CB" box="[547,665,463,488]" pageId="24" pageNumber="25" pagination="73 - 189" refId="ref36081" refString="Cox CB. 1972. A new digging dicynodont from the Upper Permian of Tanzania. In: Josey KA, Kemp TS, eds. Studies in Vertebrate eVolution. Edinburgh: Oliver and Boyd, 73 - 189." type="book chapter" year="1972">
<smallCapsWord baselines="482,482" box="[547,596,463,488]" lowerCaseFontSize="8" mainFontSize="10" normCase="title" normString="Cox" pageId="24" pageNumber="25">COX</smallCapsWord>
, 1972
</bibRefCitation>
</heading>
</paragraph>
<paragraph blockId="24.[145,762,536,558]" box="[145,762,536,558]" pageId="24" pageNumber="25">
<emphasis box="[145,260,536,557]" italics="true" pageId="24" pageNumber="25">Specimen:</emphasis>
GPIT-PV-117032 (formerly GPIT/RE/9272).
</paragraph>
</subSubSection>
<subSubSection pageId="24" pageNumber="25" type="description">
<paragraph blockId="24.[145,763,598,1908]" pageId="24" pageNumber="25">
<emphasis box="[145,342,598,619]" italics="true" pageId="24" pageNumber="25">Orbitosphenoid:</emphasis>
The orbitosphenoid (
<figureCitation box="[625,696,598,620]" captionStart="Figure 9" captionStartId="25.[163,241,1543,1565]" captionTargetBox="[307,1303,200,1501]" captionTargetId="figure-26@25.[301,1305,216,1504]" captionTargetPageId="25" captionText="Figure 9. The anterior and ventral portions of the braincase of Kaaeingasaurus fossilis (GPIT-PV-117032): orbitosphenoid (purple) in (A) anterior, (B) dorsal and (C) left lateral views; basioccipital (lime green), parabasisphenoid (light blue) and pterygoid (yellow) in (D) dorsal, (E) left lateral and (F) ventral views; parabasisphenoid in (G) ventral view; and basioccipital in (H) posterior view. Horizontal CT-image through the parabasisphenoid and basioccipital (I). aptr, anterior (palatal) ramus of the pterygoid; bo, basioccipital; boc, basioccipital condyle; bocf, basioccipital longitudinal furrow; bolp, basioccipital lateral process; bovr, basioccipital median ridge; bovs, basioccipital vestibular space; bpt, basisphenoidal tubera; bt, basioccipital tubera; bvex, basioccipitalbasisphenoidal ventral excavation; clp, clinoid process; ds, dorsum sellae; dsu, orbitosphenoid dorsal sulcus; ic, internal carotid foramina; jf, jugular foramen; obasu, orbitosphenoid lateral sulci; obgu, orbitosphenoid dorsal gutter; obno, lateral notch; obpl, orbitosphenoid dorsal plate; obvp, orbitosphenoid medial vertical process; obw, orbitosphenoid wings; ofc, olfactory cavity; pbs, parabasisphenoid; psgr, parasphenoidal groove; pt, pterygoid; ptqr, posterior (quadrate) ramus of the pterygoid; stu, sella turcica; tra, trabecular spaces; ve, vestibule. Note that the articular surface of the exoccipital is represented by the horizontal square pattern in (D). The upper scale bar equals 5 mm and 1 mm in the lower. The upper scale serves for (A-C) and the lower for (D-H)." figureDoi="http://doi.org/10.5281/zenodo.7922171" httpUri="https://zenodo.org/record/7922171/files/figure.png" pageId="24" pageNumber="25">Fig. 9</figureCitation>
) is a complex composed of two dorsally directed wings posteriorly and the vertical median process anteriorly often called the mesethmoid in cistecephalids (
<bibRefCitation author="Keyser AW" pageId="24" pageNumber="25" pagination="1 - 71" refId="ref37958" refString="Keyser AW. 1973. A preliminary study of the type area of the Cistecephalus Zone of the Beaufort Series, and a revision of the anomodont family Cistecephalidae. Geological SurVey of South Africa 62: 1 - 71." type="journal article" year="1973">Keyser, 1973</bibRefCitation>
;
<bibRefCitation author="Laass M &amp; Kaestner A" box="[213,477,721,743]" pageId="24" pageNumber="25" pagination="1033 - 1057" refId="ref38292" refString="Laass M, Kaestner A. 2017. Evidence for convergent evolution of a neocortex-like structure in a Late Permian therapsid. Journal of Morphology 278: 1033 - 1057." type="journal article" year="2017">Laass &amp; Kaestner, 2017</bibRefCitation>
;
<bibRefCitation author="Angielczyk KD &amp; Benoit J &amp; Rubidge BS" box="[489,746,721,743]" pageId="24" pageNumber="25" pagination="405 - 446" refId="ref33801" refString="Angielczyk KD, Benoit J, Rubidge BS. 2019. A new tusked cistecephalid dicynodont (Therapsida, Anomodontia) from the Upper Permian Upper Madumabisa Mudstone Formation, Luangwa Basin, Zambia. Papers in Palaeontology 7: 405 - 446." type="journal article" year="2019">
Angielczyk
<emphasis box="[622,678,721,743]" italics="true" pageId="24" pageNumber="25">et al.</emphasis>
, 2019
</bibRefCitation>
). It contacts the frontal dorsally and the parasphenoid ventrally. The orbitosphenoid is nearly rectangular in lateral view and hosts the olfactory bulbs on its dorsal portion.
</paragraph>
<paragraph blockId="24.[145,763,598,1908]" pageId="24" pageNumber="25">
The wings (obw) in GPIT-PV-117032 have the usual Y-shaped cross-section. The lateral wings are oriented almost vertically in anterior view and connect to the body of the bone at a right angle (
<figureCitation box="[535,618,966,988]" captionStart="Figure 9" captionStartId="25.[163,241,1543,1565]" captionTargetBox="[307,1303,200,1501]" captionTargetId="figure-26@25.[301,1305,216,1504]" captionTargetPageId="25" captionText="Figure 9. The anterior and ventral portions of the braincase of Kaaeingasaurus fossilis (GPIT-PV-117032): orbitosphenoid (purple) in (A) anterior, (B) dorsal and (C) left lateral views; basioccipital (lime green), parabasisphenoid (light blue) and pterygoid (yellow) in (D) dorsal, (E) left lateral and (F) ventral views; parabasisphenoid in (G) ventral view; and basioccipital in (H) posterior view. Horizontal CT-image through the parabasisphenoid and basioccipital (I). aptr, anterior (palatal) ramus of the pterygoid; bo, basioccipital; boc, basioccipital condyle; bocf, basioccipital longitudinal furrow; bolp, basioccipital lateral process; bovr, basioccipital median ridge; bovs, basioccipital vestibular space; bpt, basisphenoidal tubera; bt, basioccipital tubera; bvex, basioccipitalbasisphenoidal ventral excavation; clp, clinoid process; ds, dorsum sellae; dsu, orbitosphenoid dorsal sulcus; ic, internal carotid foramina; jf, jugular foramen; obasu, orbitosphenoid lateral sulci; obgu, orbitosphenoid dorsal gutter; obno, lateral notch; obpl, orbitosphenoid dorsal plate; obvp, orbitosphenoid medial vertical process; obw, orbitosphenoid wings; ofc, olfactory cavity; pbs, parabasisphenoid; psgr, parasphenoidal groove; pt, pterygoid; ptqr, posterior (quadrate) ramus of the pterygoid; stu, sella turcica; tra, trabecular spaces; ve, vestibule. Note that the articular surface of the exoccipital is represented by the horizontal square pattern in (D). The upper scale bar equals 5 mm and 1 mm in the lower. The upper scale serves for (A-C) and the lower for (D-H)." figureDoi="http://doi.org/10.5281/zenodo.7922171" httpUri="https://zenodo.org/record/7922171/files/figure.png" pageId="24" pageNumber="25">Fig. 9A</figureCitation>
). The dorsal vertical portion of the wings is thin, whereas the horizontal portion is thicker. In anterior view, the base of the wing is excavated by a shallow sulcus (obasu,
<figureCitation box="[145,233,1089,1111]" captionStart="Figure 9" captionStartId="25.[163,241,1543,1565]" captionTargetBox="[307,1303,200,1501]" captionTargetId="figure-26@25.[301,1305,216,1504]" captionTargetPageId="25" captionText="Figure 9. The anterior and ventral portions of the braincase of Kaaeingasaurus fossilis (GPIT-PV-117032): orbitosphenoid (purple) in (A) anterior, (B) dorsal and (C) left lateral views; basioccipital (lime green), parabasisphenoid (light blue) and pterygoid (yellow) in (D) dorsal, (E) left lateral and (F) ventral views; parabasisphenoid in (G) ventral view; and basioccipital in (H) posterior view. Horizontal CT-image through the parabasisphenoid and basioccipital (I). aptr, anterior (palatal) ramus of the pterygoid; bo, basioccipital; boc, basioccipital condyle; bocf, basioccipital longitudinal furrow; bolp, basioccipital lateral process; bovr, basioccipital median ridge; bovs, basioccipital vestibular space; bpt, basisphenoidal tubera; bt, basioccipital tubera; bvex, basioccipitalbasisphenoidal ventral excavation; clp, clinoid process; ds, dorsum sellae; dsu, orbitosphenoid dorsal sulcus; ic, internal carotid foramina; jf, jugular foramen; obasu, orbitosphenoid lateral sulci; obgu, orbitosphenoid dorsal gutter; obno, lateral notch; obpl, orbitosphenoid dorsal plate; obvp, orbitosphenoid medial vertical process; obw, orbitosphenoid wings; ofc, olfactory cavity; pbs, parabasisphenoid; psgr, parasphenoidal groove; pt, pterygoid; ptqr, posterior (quadrate) ramus of the pterygoid; stu, sella turcica; tra, trabecular spaces; ve, vestibule. Note that the articular surface of the exoccipital is represented by the horizontal square pattern in (D). The upper scale bar equals 5 mm and 1 mm in the lower. The upper scale serves for (A-C) and the lower for (D-H)." figureDoi="http://doi.org/10.5281/zenodo.7922171" httpUri="https://zenodo.org/record/7922171/files/figure.png" pageId="24" pageNumber="25">Fig. 9A</figureCitation>
) that descends ventromedially for a short distance. The anteriormost half of the orbitosphenoid comprises the median vertical process (obvp), which forms a horizontal dorsal plate (obpl) dorsally and descends to form a thick vertical plate ventrally that articulates with the parasphenoid rostrum (
<figureCitation captionStart="Figure 9" captionStartId="25.[163,241,1543,1565]" captionTargetBox="[307,1303,200,1501]" captionTargetId="figure-26@25.[301,1305,216,1504]" captionTargetPageId="25" captionText="Figure 9. The anterior and ventral portions of the braincase of Kaaeingasaurus fossilis (GPIT-PV-117032): orbitosphenoid (purple) in (A) anterior, (B) dorsal and (C) left lateral views; basioccipital (lime green), parabasisphenoid (light blue) and pterygoid (yellow) in (D) dorsal, (E) left lateral and (F) ventral views; parabasisphenoid in (G) ventral view; and basioccipital in (H) posterior view. Horizontal CT-image through the parabasisphenoid and basioccipital (I). aptr, anterior (palatal) ramus of the pterygoid; bo, basioccipital; boc, basioccipital condyle; bocf, basioccipital longitudinal furrow; bolp, basioccipital lateral process; bovr, basioccipital median ridge; bovs, basioccipital vestibular space; bpt, basisphenoidal tubera; bt, basioccipital tubera; bvex, basioccipitalbasisphenoidal ventral excavation; clp, clinoid process; ds, dorsum sellae; dsu, orbitosphenoid dorsal sulcus; ic, internal carotid foramina; jf, jugular foramen; obasu, orbitosphenoid lateral sulci; obgu, orbitosphenoid dorsal gutter; obno, lateral notch; obpl, orbitosphenoid dorsal plate; obvp, orbitosphenoid medial vertical process; obw, orbitosphenoid wings; ofc, olfactory cavity; pbs, parabasisphenoid; psgr, parasphenoidal groove; pt, pterygoid; ptqr, posterior (quadrate) ramus of the pterygoid; stu, sella turcica; tra, trabecular spaces; ve, vestibule. Note that the articular surface of the exoccipital is represented by the horizontal square pattern in (D). The upper scale bar equals 5 mm and 1 mm in the lower. The upper scale serves for (A-C) and the lower for (D-H)." figureDoi="http://doi.org/10.5281/zenodo.7922171" httpUri="https://zenodo.org/record/7922171/files/figure.png" pageId="24" pageNumber="25">Fig. 9A</figureCitation>
). The dorsal horizontal plate articulates with the frontal dorsally. The median process separates the olfactory cavities (ofc,
<figureCitation box="[505,600,1334,1356]" captionStart="Figure 9" captionStartId="25.[163,241,1543,1565]" captionTargetBox="[307,1303,200,1501]" captionTargetId="figure-26@25.[301,1305,216,1504]" captionTargetPageId="25" captionText="Figure 9. The anterior and ventral portions of the braincase of Kaaeingasaurus fossilis (GPIT-PV-117032): orbitosphenoid (purple) in (A) anterior, (B) dorsal and (C) left lateral views; basioccipital (lime green), parabasisphenoid (light blue) and pterygoid (yellow) in (D) dorsal, (E) left lateral and (F) ventral views; parabasisphenoid in (G) ventral view; and basioccipital in (H) posterior view. Horizontal CT-image through the parabasisphenoid and basioccipital (I). aptr, anterior (palatal) ramus of the pterygoid; bo, basioccipital; boc, basioccipital condyle; bocf, basioccipital longitudinal furrow; bolp, basioccipital lateral process; bovr, basioccipital median ridge; bovs, basioccipital vestibular space; bpt, basisphenoidal tubera; bt, basioccipital tubera; bvex, basioccipitalbasisphenoidal ventral excavation; clp, clinoid process; ds, dorsum sellae; dsu, orbitosphenoid dorsal sulcus; ic, internal carotid foramina; jf, jugular foramen; obasu, orbitosphenoid lateral sulci; obgu, orbitosphenoid dorsal gutter; obno, lateral notch; obpl, orbitosphenoid dorsal plate; obvp, orbitosphenoid medial vertical process; obw, orbitosphenoid wings; ofc, olfactory cavity; pbs, parabasisphenoid; psgr, parasphenoidal groove; pt, pterygoid; ptqr, posterior (quadrate) ramus of the pterygoid; stu, sella turcica; tra, trabecular spaces; ve, vestibule. Note that the articular surface of the exoccipital is represented by the horizontal square pattern in (D). The upper scale bar equals 5 mm and 1 mm in the lower. The upper scale serves for (A-C) and the lower for (D-H)." figureDoi="http://doi.org/10.5281/zenodo.7922171" httpUri="https://zenodo.org/record/7922171/files/figure.png" pageId="24" pageNumber="25">Fig. 9A</figureCitation>
), which are relatively expanded in
<emphasis box="[421,607,1366,1387]" italics="true" pageId="24" pageNumber="25">Kaaeingasaurus</emphasis>
compared to
<taxonomicName box="[145,277,1396,1417]" class="Reptilia" family="Emydopidae" genus="Myosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Therapsida" pageId="24" pageNumber="25" phylum="Chordata" rank="genus">
<emphasis box="[145,277,1396,1417]" italics="true" pageId="24" pageNumber="25">Myosaurus</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Huxley" authorityYear="1868" box="[342,483,1396,1417]" class="Reptilia" genus="Pristerodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Therapsida" pageId="24" pageNumber="25" phylum="Chordata" rank="genus">
<emphasis box="[342,483,1396,1417]" italics="true" pageId="24" pageNumber="25">Pristerodon</emphasis>
</taxonomicName>
(height-to-width ratio 2.0 in
<emphasis box="[218,401,1427,1448]" italics="true" pageId="24" pageNumber="25">Kaaeingasaurus</emphasis>
vs.
<quantity box="[447,513,1426,1448]" metricMagnitude="-2" metricUnit="m" metricValue="4.318" pageId="24" pageNumber="25" unit="in" value="1.7">1.7 in</quantity>
<taxonomicName box="[521,648,1427,1448]" class="Reptilia" family="Emydopidae" genus="Myosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Therapsida" pageId="24" pageNumber="25" phylum="Chordata" rank="genus">
<emphasis box="[521,648,1427,1448]" italics="true" pageId="24" pageNumber="25">Myosaurus</emphasis>
</taxonomicName>
vs.
<quantity box="[695,760,1426,1448]" metricMagnitude="-2" metricUnit="m" metricValue="4.572" pageId="24" pageNumber="25" unit="in" value="1.8">1.8 in</quantity>
<taxonomicName authorityName="Huxley" authorityYear="1868" box="[145,277,1457,1478]" class="Reptilia" genus="Pristerodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Therapsida" pageId="24" pageNumber="25" phylum="Chordata" rank="genus">
<emphasis box="[145,277,1457,1478]" italics="true" pageId="24" pageNumber="25">Pristerodon</emphasis>
</taxonomicName>
). In dorsal view, the horizontal plate of the vertical process is triangular and is excavated by a long anteroposterior sulcus (dsu,
<figureCitation box="[523,604,1518,1540]" captionStart="Figure 9" captionStartId="25.[163,241,1543,1565]" captionTargetBox="[307,1303,200,1501]" captionTargetId="figure-26@25.[301,1305,216,1504]" captionTargetPageId="25" captionText="Figure 9. The anterior and ventral portions of the braincase of Kaaeingasaurus fossilis (GPIT-PV-117032): orbitosphenoid (purple) in (A) anterior, (B) dorsal and (C) left lateral views; basioccipital (lime green), parabasisphenoid (light blue) and pterygoid (yellow) in (D) dorsal, (E) left lateral and (F) ventral views; parabasisphenoid in (G) ventral view; and basioccipital in (H) posterior view. Horizontal CT-image through the parabasisphenoid and basioccipital (I). aptr, anterior (palatal) ramus of the pterygoid; bo, basioccipital; boc, basioccipital condyle; bocf, basioccipital longitudinal furrow; bolp, basioccipital lateral process; bovr, basioccipital median ridge; bovs, basioccipital vestibular space; bpt, basisphenoidal tubera; bt, basioccipital tubera; bvex, basioccipitalbasisphenoidal ventral excavation; clp, clinoid process; ds, dorsum sellae; dsu, orbitosphenoid dorsal sulcus; ic, internal carotid foramina; jf, jugular foramen; obasu, orbitosphenoid lateral sulci; obgu, orbitosphenoid dorsal gutter; obno, lateral notch; obpl, orbitosphenoid dorsal plate; obvp, orbitosphenoid medial vertical process; obw, orbitosphenoid wings; ofc, olfactory cavity; pbs, parabasisphenoid; psgr, parasphenoidal groove; pt, pterygoid; ptqr, posterior (quadrate) ramus of the pterygoid; stu, sella turcica; tra, trabecular spaces; ve, vestibule. Note that the articular surface of the exoccipital is represented by the horizontal square pattern in (D). The upper scale bar equals 5 mm and 1 mm in the lower. The upper scale serves for (A-C) and the lower for (D-H)." figureDoi="http://doi.org/10.5281/zenodo.7922171" httpUri="https://zenodo.org/record/7922171/files/figure.png" pageId="24" pageNumber="25">Fig. 9B</figureCitation>
).
</paragraph>
<paragraph blockId="24.[145,763,598,1908]" lastBlockId="24.[809,1425,197,342]" pageId="24" pageNumber="25">
In lateral view, the orbitosphenoid in
<emphasis box="[581,760,1550,1571]" italics="true" pageId="24" pageNumber="25">Kaaeingasaurus</emphasis>
is relatively longer anteroposteriorly than in
<taxonomicName box="[145,270,1611,1632]" class="Reptilia" family="Emydopidae" genus="Myosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Therapsida" pageId="24" pageNumber="25" phylum="Chordata" rank="genus">
<emphasis box="[145,270,1611,1632]" italics="true" pageId="24" pageNumber="25">Myosaurus</emphasis>
</taxonomicName>
(
<figureCitation box="[282,360,1610,1632]" captionStart="Figure 9" captionStartId="25.[163,241,1543,1565]" captionTargetBox="[307,1303,200,1501]" captionTargetId="figure-26@25.[301,1305,216,1504]" captionTargetPageId="25" captionText="Figure 9. The anterior and ventral portions of the braincase of Kaaeingasaurus fossilis (GPIT-PV-117032): orbitosphenoid (purple) in (A) anterior, (B) dorsal and (C) left lateral views; basioccipital (lime green), parabasisphenoid (light blue) and pterygoid (yellow) in (D) dorsal, (E) left lateral and (F) ventral views; parabasisphenoid in (G) ventral view; and basioccipital in (H) posterior view. Horizontal CT-image through the parabasisphenoid and basioccipital (I). aptr, anterior (palatal) ramus of the pterygoid; bo, basioccipital; boc, basioccipital condyle; bocf, basioccipital longitudinal furrow; bolp, basioccipital lateral process; bovr, basioccipital median ridge; bovs, basioccipital vestibular space; bpt, basisphenoidal tubera; bt, basioccipital tubera; bvex, basioccipitalbasisphenoidal ventral excavation; clp, clinoid process; ds, dorsum sellae; dsu, orbitosphenoid dorsal sulcus; ic, internal carotid foramina; jf, jugular foramen; obasu, orbitosphenoid lateral sulci; obgu, orbitosphenoid dorsal gutter; obno, lateral notch; obpl, orbitosphenoid dorsal plate; obvp, orbitosphenoid medial vertical process; obw, orbitosphenoid wings; ofc, olfactory cavity; pbs, parabasisphenoid; psgr, parasphenoidal groove; pt, pterygoid; ptqr, posterior (quadrate) ramus of the pterygoid; stu, sella turcica; tra, trabecular spaces; ve, vestibule. Note that the articular surface of the exoccipital is represented by the horizontal square pattern in (D). The upper scale bar equals 5 mm and 1 mm in the lower. The upper scale serves for (A-C) and the lower for (D-H)." figureDoi="http://doi.org/10.5281/zenodo.7922171" httpUri="https://zenodo.org/record/7922171/files/figure.png" pageId="24" pageNumber="25">Fig. 9C</figureCitation>
). The wings are well separated from the mesethmoid and their lateral surfaces are slightly convex. Their dorsal margins are horizontal and flat to articulate with the frontal, as in
<taxonomicName authorityName="Huxley" authorityYear="1868" box="[566,702,1702,1723]" class="Reptilia" genus="Pristerodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Therapsida" pageId="24" pageNumber="25" phylum="Chordata" rank="genus">
<emphasis box="[566,702,1702,1723]" italics="true" pageId="24" pageNumber="25">Pristerodon</emphasis>
</taxonomicName>
. The ventral margin of the wing expands posteriorly and horizontally, giving it a C-shaped posterior margin. The anterior margin of the wings appears S-shaped in lateral view. In dorsal view, the gutter (obgu) formed by the two wings is anteroposteriorly short and inclined posteriorly. There is a notch between the median vertical process and the orbitosphenoid wings in lateral view (obno). In lateral view, the vertical plate of the median vertical process has a horizontally oval outline topped by the rectangular shape of the dorsal plate dorsally.
</paragraph>
<paragraph blockId="24.[809,1426,382,1508]" pageId="24" pageNumber="25">
<emphasis box="[809,926,382,403]" italics="true" pageId="24" pageNumber="25">Pterygoid:</emphasis>
The pterygoid (pt,
<figureCitation box="[1160,1223,382,404]" captionStart="Figure 9" captionStartId="25.[163,241,1543,1565]" captionTargetBox="[307,1303,200,1501]" captionTargetId="figure-26@25.[301,1305,216,1504]" captionTargetPageId="25" captionText="Figure 9. The anterior and ventral portions of the braincase of Kaaeingasaurus fossilis (GPIT-PV-117032): orbitosphenoid (purple) in (A) anterior, (B) dorsal and (C) left lateral views; basioccipital (lime green), parabasisphenoid (light blue) and pterygoid (yellow) in (D) dorsal, (E) left lateral and (F) ventral views; parabasisphenoid in (G) ventral view; and basioccipital in (H) posterior view. Horizontal CT-image through the parabasisphenoid and basioccipital (I). aptr, anterior (palatal) ramus of the pterygoid; bo, basioccipital; boc, basioccipital condyle; bocf, basioccipital longitudinal furrow; bolp, basioccipital lateral process; bovr, basioccipital median ridge; bovs, basioccipital vestibular space; bpt, basisphenoidal tubera; bt, basioccipital tubera; bvex, basioccipitalbasisphenoidal ventral excavation; clp, clinoid process; ds, dorsum sellae; dsu, orbitosphenoid dorsal sulcus; ic, internal carotid foramina; jf, jugular foramen; obasu, orbitosphenoid lateral sulci; obgu, orbitosphenoid dorsal gutter; obno, lateral notch; obpl, orbitosphenoid dorsal plate; obvp, orbitosphenoid medial vertical process; obw, orbitosphenoid wings; ofc, olfactory cavity; pbs, parabasisphenoid; psgr, parasphenoidal groove; pt, pterygoid; ptqr, posterior (quadrate) ramus of the pterygoid; stu, sella turcica; tra, trabecular spaces; ve, vestibule. Note that the articular surface of the exoccipital is represented by the horizontal square pattern in (D). The upper scale bar equals 5 mm and 1 mm in the lower. The upper scale serves for (A-C) and the lower for (D-H)." figureDoi="http://doi.org/10.5281/zenodo.7922171" httpUri="https://zenodo.org/record/7922171/files/figure.png" pageId="24" pageNumber="25">Fig. 9</figureCitation>
D-F) displays the typical X-shape dicynodont morphology in ventral view. In GPIT-PV-117032, the pterygoid is almost completely preserved except for the right quadrate ramus. The quadrate ramus in
<emphasis box="[1033,1215,506,527]" italics="true" pageId="24" pageNumber="25">Kaaeingasaurus</emphasis>
diverges from the median sagittal plane at an angle of about 80°. The anterior palatal ramus of the pterygoid has a concave lateral surface observed in dorsal view, and it forms an angle of 30° with the median sagittal axis of the skull. The angle between the palatal and quadrate rami of the pterygoid is 83°. As observed in lateral view, the palatal ramus is taller than the quadrate ramus (
<figureCitation box="[818,904,750,772]" captionStart="Figure 9" captionStartId="25.[163,241,1543,1565]" captionTargetBox="[307,1303,200,1501]" captionTargetId="figure-26@25.[301,1305,216,1504]" captionTargetPageId="25" captionText="Figure 9. The anterior and ventral portions of the braincase of Kaaeingasaurus fossilis (GPIT-PV-117032): orbitosphenoid (purple) in (A) anterior, (B) dorsal and (C) left lateral views; basioccipital (lime green), parabasisphenoid (light blue) and pterygoid (yellow) in (D) dorsal, (E) left lateral and (F) ventral views; parabasisphenoid in (G) ventral view; and basioccipital in (H) posterior view. Horizontal CT-image through the parabasisphenoid and basioccipital (I). aptr, anterior (palatal) ramus of the pterygoid; bo, basioccipital; boc, basioccipital condyle; bocf, basioccipital longitudinal furrow; bolp, basioccipital lateral process; bovr, basioccipital median ridge; bovs, basioccipital vestibular space; bpt, basisphenoidal tubera; bt, basioccipital tubera; bvex, basioccipitalbasisphenoidal ventral excavation; clp, clinoid process; ds, dorsum sellae; dsu, orbitosphenoid dorsal sulcus; ic, internal carotid foramina; jf, jugular foramen; obasu, orbitosphenoid lateral sulci; obgu, orbitosphenoid dorsal gutter; obno, lateral notch; obpl, orbitosphenoid dorsal plate; obvp, orbitosphenoid medial vertical process; obw, orbitosphenoid wings; ofc, olfactory cavity; pbs, parabasisphenoid; psgr, parasphenoidal groove; pt, pterygoid; ptqr, posterior (quadrate) ramus of the pterygoid; stu, sella turcica; tra, trabecular spaces; ve, vestibule. Note that the articular surface of the exoccipital is represented by the horizontal square pattern in (D). The upper scale bar equals 5 mm and 1 mm in the lower. The upper scale serves for (A-C) and the lower for (D-H)." figureDoi="http://doi.org/10.5281/zenodo.7922171" httpUri="https://zenodo.org/record/7922171/files/figure.png" pageId="24" pageNumber="25">Fig. 9E</figureCitation>
). The posterior end of the quadrate ramus deepens ventromedially in lateral view. Medially, the posteriormost region of the quadrate ramus has a convex surface that articulates with the quadrate.
</paragraph>
<paragraph blockId="24.[809,1426,382,1508]" pageId="24" pageNumber="25">
The palatal ramus of the pterygoid is thicker ventrally than the quadrate ramus and terminates anteriorly in an irregular suture with the ectopterygoid (
<figureCitation box="[1301,1380,934,956]" captionStart="Figure 9" captionStartId="25.[163,241,1543,1565]" captionTargetBox="[307,1303,200,1501]" captionTargetId="figure-26@25.[301,1305,216,1504]" captionTargetPageId="25" captionText="Figure 9. The anterior and ventral portions of the braincase of Kaaeingasaurus fossilis (GPIT-PV-117032): orbitosphenoid (purple) in (A) anterior, (B) dorsal and (C) left lateral views; basioccipital (lime green), parabasisphenoid (light blue) and pterygoid (yellow) in (D) dorsal, (E) left lateral and (F) ventral views; parabasisphenoid in (G) ventral view; and basioccipital in (H) posterior view. Horizontal CT-image through the parabasisphenoid and basioccipital (I). aptr, anterior (palatal) ramus of the pterygoid; bo, basioccipital; boc, basioccipital condyle; bocf, basioccipital longitudinal furrow; bolp, basioccipital lateral process; bovr, basioccipital median ridge; bovs, basioccipital vestibular space; bpt, basisphenoidal tubera; bt, basioccipital tubera; bvex, basioccipitalbasisphenoidal ventral excavation; clp, clinoid process; ds, dorsum sellae; dsu, orbitosphenoid dorsal sulcus; ic, internal carotid foramina; jf, jugular foramen; obasu, orbitosphenoid lateral sulci; obgu, orbitosphenoid dorsal gutter; obno, lateral notch; obpl, orbitosphenoid dorsal plate; obvp, orbitosphenoid medial vertical process; obw, orbitosphenoid wings; ofc, olfactory cavity; pbs, parabasisphenoid; psgr, parasphenoidal groove; pt, pterygoid; ptqr, posterior (quadrate) ramus of the pterygoid; stu, sella turcica; tra, trabecular spaces; ve, vestibule. Note that the articular surface of the exoccipital is represented by the horizontal square pattern in (D). The upper scale bar equals 5 mm and 1 mm in the lower. The upper scale serves for (A-C) and the lower for (D-H)." figureDoi="http://doi.org/10.5281/zenodo.7922171" httpUri="https://zenodo.org/record/7922171/files/figure.png" pageId="24" pageNumber="25">Fig. 9F</figureCitation>
). In ventral view, the two pterygoids meet at the midline of the median horizontal plate in
<emphasis box="[1202,1385,996,1017]" italics="true" pageId="24" pageNumber="25">Kaaeingasaurus</emphasis>
, as in most cistecephalids. The ventral exposure of the pterygoid median plate in
<emphasis box="[1134,1323,1057,1078]" italics="true" pageId="24" pageNumber="25">Kaaeingasaurus</emphasis>
is short compared to the parabasisphenoid at the median sagittal axis (
<figureCitation box="[975,1059,1118,1140]" captionStart="Figure 9" captionStartId="25.[163,241,1543,1565]" captionTargetBox="[307,1303,200,1501]" captionTargetId="figure-26@25.[301,1305,216,1504]" captionTargetPageId="25" captionText="Figure 9. The anterior and ventral portions of the braincase of Kaaeingasaurus fossilis (GPIT-PV-117032): orbitosphenoid (purple) in (A) anterior, (B) dorsal and (C) left lateral views; basioccipital (lime green), parabasisphenoid (light blue) and pterygoid (yellow) in (D) dorsal, (E) left lateral and (F) ventral views; parabasisphenoid in (G) ventral view; and basioccipital in (H) posterior view. Horizontal CT-image through the parabasisphenoid and basioccipital (I). aptr, anterior (palatal) ramus of the pterygoid; bo, basioccipital; boc, basioccipital condyle; bocf, basioccipital longitudinal furrow; bolp, basioccipital lateral process; bovr, basioccipital median ridge; bovs, basioccipital vestibular space; bpt, basisphenoidal tubera; bt, basioccipital tubera; bvex, basioccipitalbasisphenoidal ventral excavation; clp, clinoid process; ds, dorsum sellae; dsu, orbitosphenoid dorsal sulcus; ic, internal carotid foramina; jf, jugular foramen; obasu, orbitosphenoid lateral sulci; obgu, orbitosphenoid dorsal gutter; obno, lateral notch; obpl, orbitosphenoid dorsal plate; obvp, orbitosphenoid medial vertical process; obw, orbitosphenoid wings; ofc, olfactory cavity; pbs, parabasisphenoid; psgr, parasphenoidal groove; pt, pterygoid; ptqr, posterior (quadrate) ramus of the pterygoid; stu, sella turcica; tra, trabecular spaces; ve, vestibule. Note that the articular surface of the exoccipital is represented by the horizontal square pattern in (D). The upper scale bar equals 5 mm and 1 mm in the lower. The upper scale serves for (A-C) and the lower for (D-H)." figureDoi="http://doi.org/10.5281/zenodo.7922171" httpUri="https://zenodo.org/record/7922171/files/figure.png" pageId="24" pageNumber="25">Fig. 9F</figureCitation>
), as noted in
<bibRefCitation author="Angielczyk KD &amp; Benoit J &amp; Rubidge BS" pageId="24" pageNumber="25" pagination="405 - 446" refId="ref33801" refString="Angielczyk KD, Benoit J, Rubidge BS. 2019. A new tusked cistecephalid dicynodont (Therapsida, Anomodontia) from the Upper Permian Upper Madumabisa Mudstone Formation, Luangwa Basin, Zambia. Papers in Palaeontology 7: 405 - 446." type="journal article" year="2019">
Angielczyk
<emphasis box="[1365,1425,1118,1140]" italics="true" pageId="24" pageNumber="25">et al.</emphasis>
(2019)
</bibRefCitation>
.
<emphasis box="[898,1083,1149,1170]" italics="true" pageId="24" pageNumber="25">Kaaeingasaurus</emphasis>
lacks the crista oesophagea, with the median pterygoid plate being flat (
<figureCitation box="[1327,1410,1179,1201]" captionStart="Figure 9" captionStartId="25.[163,241,1543,1565]" captionTargetBox="[307,1303,200,1501]" captionTargetId="figure-26@25.[301,1305,216,1504]" captionTargetPageId="25" captionText="Figure 9. The anterior and ventral portions of the braincase of Kaaeingasaurus fossilis (GPIT-PV-117032): orbitosphenoid (purple) in (A) anterior, (B) dorsal and (C) left lateral views; basioccipital (lime green), parabasisphenoid (light blue) and pterygoid (yellow) in (D) dorsal, (E) left lateral and (F) ventral views; parabasisphenoid in (G) ventral view; and basioccipital in (H) posterior view. Horizontal CT-image through the parabasisphenoid and basioccipital (I). aptr, anterior (palatal) ramus of the pterygoid; bo, basioccipital; boc, basioccipital condyle; bocf, basioccipital longitudinal furrow; bolp, basioccipital lateral process; bovr, basioccipital median ridge; bovs, basioccipital vestibular space; bpt, basisphenoidal tubera; bt, basioccipital tubera; bvex, basioccipitalbasisphenoidal ventral excavation; clp, clinoid process; ds, dorsum sellae; dsu, orbitosphenoid dorsal sulcus; ic, internal carotid foramina; jf, jugular foramen; obasu, orbitosphenoid lateral sulci; obgu, orbitosphenoid dorsal gutter; obno, lateral notch; obpl, orbitosphenoid dorsal plate; obvp, orbitosphenoid medial vertical process; obw, orbitosphenoid wings; ofc, olfactory cavity; pbs, parabasisphenoid; psgr, parasphenoidal groove; pt, pterygoid; ptqr, posterior (quadrate) ramus of the pterygoid; stu, sella turcica; tra, trabecular spaces; ve, vestibule. Note that the articular surface of the exoccipital is represented by the horizontal square pattern in (D). The upper scale bar equals 5 mm and 1 mm in the lower. The upper scale serves for (A-C) and the lower for (D-H)." figureDoi="http://doi.org/10.5281/zenodo.7922171" httpUri="https://zenodo.org/record/7922171/files/figure.png" pageId="24" pageNumber="25">Fig. 9F</figureCitation>
), and also lacks an interpterygoid vacuity, as noted by
<bibRefCitation author="Cox CB" box="[809,928,1241,1262]" pageId="24" pageNumber="25" pagination="73 - 189" refId="ref36081" refString="Cox CB. 1972. A new digging dicynodont from the Upper Permian of Tanzania. In: Josey KA, Kemp TS, eds. Studies in Vertebrate eVolution. Edinburgh: Oliver and Boyd, 73 - 189." type="book chapter" year="1972">Cox (1972)</bibRefCitation>
. An interpterygoid vacuity is also essentially absent in
<taxonomicName box="[933,1100,1271,1292]" class="Reptilia" family="Cistecephalidae" genus="Cistecephalus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Therapsida" pageId="24" pageNumber="25" phylum="Chordata" rank="genus">
<emphasis box="[933,1100,1271,1292]" italics="true" pageId="24" pageNumber="25">Cistecephalus</emphasis>
</taxonomicName>
(
<bibRefCitation author="Keyser AW" box="[1119,1276,1272,1293]" pageId="24" pageNumber="25" pagination="1 - 71" refId="ref37958" refString="Keyser AW. 1973. A preliminary study of the type area of the Cistecephalus Zone of the Beaufort Series, and a revision of the anomodont family Cistecephalidae. Geological SurVey of South Africa 62: 1 - 71." type="journal article" year="1973">Keyser, 1973</bibRefCitation>
;
<bibRefCitation author="Angielczyk KD &amp; Benoit J &amp; Rubidge BS" pageId="24" pageNumber="25" pagination="405 - 446" refId="ref33801" refString="Angielczyk KD, Benoit J, Rubidge BS. 2019. A new tusked cistecephalid dicynodont (Therapsida, Anomodontia) from the Upper Permian Upper Madumabisa Mudstone Formation, Luangwa Basin, Zambia. Papers in Palaeontology 7: 405 - 446." type="journal article" year="2019">
Angielczyk
<emphasis box="[809,875,1302,1324]" italics="true" pageId="24" pageNumber="25">et al.</emphasis>
, 2019
</bibRefCitation>
), contrasting with the condition in cistecephalids like
<emphasis box="[1024,1181,1333,1354]" italics="true" pageId="24" pageNumber="25">Kembaaeacela</emphasis>
and
<taxonomicName baseAuthorityName="Cluver" baseAuthorityYear="1974" box="[1237,1425,1333,1354]" class="Reptilia" family="Cistecephalidae" genus="Cistecephaloides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Therapsida" pageId="24" pageNumber="25" phylum="Chordata" rank="genus">
<emphasis box="[1237,1425,1333,1354]" italics="true" pageId="24" pageNumber="25">Cistecephaloides</emphasis>
</taxonomicName>
(
<bibRefCitation author="Cluver MA" box="[818,985,1363,1385]" pageId="24" pageNumber="25" pagination="137 - 155" refId="ref35849" refString="Cluver MA. 1974 a. The skull and mandible of a new cistecephalid dicyndont. Annals of the South African Museum 64: 137 - 155." type="journal article" year="1974">Cluver, 1974a</bibRefCitation>
;
<bibRefCitation author="Kammerer CF &amp; Bandyopadhyay S &amp; Ray S" box="[1002,1276,1363,1385]" pageId="24" pageNumber="25" pagination="569 - 584" refId="ref37567" refString="Kammerer CF, Bandyopadhyay S, Ray S. 2016. A new taxon of cistecephalid dicynodont from the Upper Permian Kundaram Formation of India. Papers in Palaeontology 2: 569 - 584." type="journal article" year="2016">
Kammerer
<emphasis box="[1141,1203,1363,1385]" italics="true" pageId="24" pageNumber="25">et al.</emphasis>
, 2016
</bibRefCitation>
;
<bibRefCitation author="Angielczyk KD &amp; Benoit J &amp; Rubidge BS" pageId="24" pageNumber="25" pagination="405 - 446" refId="ref33801" refString="Angielczyk KD, Benoit J, Rubidge BS. 2019. A new tusked cistecephalid dicynodont (Therapsida, Anomodontia) from the Upper Permian Upper Madumabisa Mudstone Formation, Luangwa Basin, Zambia. Papers in Palaeontology 7: 405 - 446." type="journal article" year="2019">
Angielczyk
<emphasis box="[809,873,1394,1416]" italics="true" pageId="24" pageNumber="25">et al.</emphasis>
, 2019
</bibRefCitation>
), which have small openings for this structure. The quadrate ramus is broad posteriorly but mediolaterally compressed towards the median plate in ventral view.
</paragraph>
</subSubSection>
</treatment>
</document>
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