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<document id="EE0A1460C772878325A4CE0B8B037DB4" ID-DOI="10.5194/fr-18-17-2015" ID-ISSN="2193-0074" ID-Zenodo-Dep="11045469" IM.bibliography_approvedBy="tatiana" IM.materialsCitations_approvedBy="tatiana" IM.metadata_approvedBy="juliana" IM.taxonomicNames_approvedBy="juliana" IM.treatments_approvedBy="tatiana" checkinTime="1712925722956" checkinUser="felipe" docAuthor="Spindler, F., Scott, D. &amp; Reisz, R. R." docDate="2015" docId="2C5C8787FF89FFC4FF3DFA3F3478CC95" docLanguage="en" docName="FossRec.18.1.17-30.pdf" docOrigin="Fossil Record 18 (1)" docSource="http://dx.doi.org/10.5194/fr-18-17-2015" docStyle="DocumentStyle:1B3764FFAC190C6D25C29F616DE4358D.2:FossRec.2014-2021.journal_article" docStyleId="1B3764FFAC190C6D25C29F616DE4358D" docStyleName="FossRec.2014-2021.journal_article" docStyleVersion="2" docTitle="Ianthodon schultzei Kissel and Reisz 2004" docType="treatment" docVersion="3" lastPageNumber="25" masterDocId="D065FFFFFF88FFCDFF8CFFEE3538CD73" masterDocTitle="New information on the cranial and postcranial anatomy of the early synapsid Ianthodon schultzei (Sphenacomorpha: Sphenacodontia), and its evolutionary significance" masterLastPageNumber="30" masterPageNumber="17" pageNumber="18" updateTime="1731610443596" updateUser="juliana" zenodo-license-document="CC-BY-4.0">
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<mods:title id="BC6377712255969125DE0178FE36D77E">New information on the cranial and postcranial anatomy of the early synapsid Ianthodon schultzei (Sphenacomorpha: Sphenacodontia), and its evolutionary significance</mods:title>
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<mods:namePart id="9BD63D70B87E890B015B892A4C1FEC9A">Spindler, F.</mods:namePart>
<mods:affiliation id="A60A869B14880DFA676CF9A1B45FF815">TU Bergakademie Freiberg, Institut für Geologie, Bernhard-von-Cotta-Strasse 2, 09599 Freiberg, Germany</mods:affiliation>
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<mods:namePart id="5F511AC2737E0BD3174D4BF9E3EF1384">Scott, D.</mods:namePart>
<mods:affiliation id="FB7C3ADA44170CB2692CA0E871533E23">Department of Biology, University of Toronto Mississauga, 3359 Mississauga Road, Mississauga, Ontario, L 5 L 1 C 6, Canada</mods:affiliation>
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<mods:namePart id="BEC6D53E411F71D2EA5FF21C4B4AE6A5">Reisz, R. R.</mods:namePart>
<mods:affiliation id="F1FCA835F4FC732B15A52584AF6F7B9A">Department of Biology, University of Toronto Mississauga, 3359 Mississauga Road, Mississauga, Ontario, L 5 L 1 C 6, Canada</mods:affiliation>
<mods:nameIdentifier id="AA9913EAD773A19325FFBD9991C63517" type="email">robert.reisz@utoronto.ca</mods:nameIdentifier>
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<mods:date id="7E54BE39673B270B02EDEAF7FEB1D172">2015</mods:date>
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<mods:number id="C0B8D38D6A2EF2B282D2CDC76EAD4720">2014-10-06</mods:number>
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<mods:identifier id="4E302681003135BBBA088FFF54231DD7" type="Zenodo-Dep">11045469</mods:identifier>
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<treatment id="2C5C8787FF89FFC4FF3DFA3F3478CC95" LSID="urn:lsid:plazi:treatment:2C5C8787FF89FFC4FF3DFA3F3478CC95" httpUri="http://treatment.plazi.org/id/2C5C8787FF89FFC4FF3DFA3F3478CC95" lastPageId="9" lastPageNumber="25" pageId="1" pageNumber="18">
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<paragraph id="A44A3691FF89FFCCFF3DFA3F37BEC894" blockId="1.[177,646,1488,1512]" box="[177,646,1488,1512]" pageId="1" pageNumber="18">
<heading id="FF0281FDFF89FFCCFF3DFA3F37BEC894" box="[177,646,1488,1512]" fontSize="10" level="2" pageId="1" pageNumber="18" reason="5">
<taxonomicName id="63F54D12FF89FFCCFF3DFA3F37BEC894" authority="Kissel and Reisz (2004)" authorityName="Kissel and Reisz" authorityYear="2004" box="[177,646,1488,1512]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="18" phylum="Chordata" rank="species" species="schultzei" subOrder="Eupelycosauria">
<emphasis id="9681EA83FF89FFCCFF3DFA3F34BAC89B" box="[177,386,1489,1512]" italics="true" pageId="1" pageNumber="18">Ianthodon schultzei</emphasis>
<bibRefCitation id="C0644B60FF89FFCCFE04FA3E37BEC894" author="Kissel, R. A. &amp; Reisz, R. R." box="[392,646,1488,1511]" pageId="1" pageNumber="18" pagination="409 - 428" refId="ref9945" refString="Kissel, R. A. and Reisz, R. R.: Synapsid fauna of the Upper Pennsylvanian Rock Lake Shale near Garnett, Kansas and the diversity pattern of early amniotes, in: Recent Advances in the Origin and Early Radiation of Vertebrates, edited by: Arratia, G., Wilson, M. V. H., and Cloutier, R., Verlag Dr. F. Pfeil, Munchen, 409 - 428, 2004." type="book chapter" year="2004">Kissel and Reisz (2004)</bibRefCitation>
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="ECEF651AFF89FFCCFF1BF9E53072CE76" pageId="1" pageNumber="18" type="materials_examined">
<paragraph id="A44A3691FF89FFCCFF1BF9E537B6CAF2" blockId="1.[124,767,1547,1921]" pageId="1" pageNumber="18">
<materialsCitation id="149D3CCCFF89FFCCFF1BF9E537B6CAF2" collectionCode="KUVP" pageId="1" pageNumber="18" specimenCode="KUVP 133735" specimenCount="2" typeStatus="holotype">
<emphasis id="9681EA83FF89FFCCFF1BF9E535C0CB51" box="[151,248,1547,1570]" italics="true" pageId="1" pageNumber="18">
<typeStatus id="7B4E8833FF89FFCCFF1BF9E535C0CB51" box="[151,248,1547,1570]" pageId="1" pageNumber="18" type="holotype">Holotype</typeStatus>
</emphasis>
.
<specimenCode id="F4539EEAFF89FFCCFE86F9E53495CB51" box="[266,429,1547,1570]" collectionCode="KUVP" pageId="1" pageNumber="18">KUVP 133735</specimenCode>
consists of a nearly complete skull including both mandibles and an anterior postcranial skeleton including vertebrae, ribs, right scapula and coracoid as well as left humerus (
<figureCitation id="3CCE2A14FF89FFCCFE4BF9853731CBF1" box="[455,521,1643,1666]" captionStart="Figure 1" captionStartId="2.[124,191,1699,1720]" captionTargetBox="[310,1278,235,1659]" captionTargetId="figure-9@2.[302,1285,226,1667]" captionTargetPageId="2" captionText="Figure 1. Ianthodon schultzei holotype KUVP 133735, combined slab, with outlines of skull bones (outlines by D. Scott, made prior to Kissel and Reisz, 2004, Fig. 2; now exposed as embedded counter slab) and a disarticulated skeleton of the diapsid reptile Petrolacosaurus kansensis (shaded areas); a – angular; ar – articular; c – vertebra centrum; cau – caudal neural arch; cle – cleithrum; co – anterior coracoid; cp – cultriform process of parasphenoid–basisphenoid complex; cr – cervical rib; d – dentary; dr – dorsal rib; h – humerus; n – nasal; na – presacral neural arch; pal – palatine; pc – posterior coronoid; pm – premaxilla; pra – prearticular; ps – parasphenoid–basisphenoid complex; pt – pterygoid; qj – quadratojugal; sa – surangular; sc – scapula; soc – supraoccipital; sp – splenial; v – vomer." figureDoi="http://doi.org/10.5281/zenodo.11045471" httpUri="https://zenodo.org/record/11045471/files/figure.png" pageId="1" pageNumber="18">Fig. 1</figureCitation>
). The specimen is incompletely ossified, indicating a juvenile individual. On the same slab, a partial skeleton of
<taxonomicName id="63F54D12FF89FFCCFE4BF945374BCBB1" authorityName="Lane" authorityYear="1945" box="[455,627,1707,1730]" family="Petrolacosauridae" genus="Petrolacosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="18" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF89FFCCFE4BF945374BCBB1" box="[455,627,1707,1730]" italics="true" pageId="1" pageNumber="18">Petrolacosaurus</emphasis>
</taxonomicName>
is preserved. Palatal, mandibular, occipital and postcranial elements are newly assigned to the holotype. The skull was removed from the slab during preparation (
<figureCitation id="3CCE2A14FF89FFCCFE3CF8E434CCCA52" box="[432,500,1802,1825]" captionStart="Figure 2" captionStartId="3.[124,191,1147,1168]" captionTargetBox="[133,756,179,1112]" captionTargetId="figure-9@3.[132,759,177,1115]" captionTargetPageId="3" captionText="Figure 2. Ianthodon schultzei holotype KUVP 133735, slab in present condition, combined with photograph of skull area (lower left) before its removal." figureDoi="http://doi.org/10.5281/zenodo.11045473" httpUri="https://zenodo.org/record/11045473/files/figure.png" pageId="1" pageNumber="18">Fig. 2</figureCitation>
). After documenting the originally exposed side (
<bibRefCitation id="C0644B60FF89FFCCFE0EF8C43745CA32" author="Kissel, R. A. &amp; Reisz, R. R." box="[386,637,1834,1857]" pageId="1" pageNumber="18" pagination="409 - 428" refId="ref9945" refString="Kissel, R. A. and Reisz, R. R.: Synapsid fauna of the Upper Pennsylvanian Rock Lake Shale near Garnett, Kansas and the diversity pattern of early amniotes, in: Recent Advances in the Origin and Early Radiation of Vertebrates, edited by: Arratia, G., Wilson, M. V. H., and Cloutier, R., Verlag Dr. F. Pfeil, Munchen, 409 - 428, 2004." type="book chapter" year="2004">Kissel and Reisz, 2004</bibRefCitation>
,
<figureCitation id="3CCE2A14FF89FFCCFD04F8C437F1CA32" box="[648,713,1834,1857]" captionStart="Figure 2" captionStartId="3.[124,191,1147,1168]" captionTargetBox="[133,756,179,1112]" captionTargetId="figure-9@3.[132,759,177,1115]" captionTargetPageId="3" captionText="Figure 2. Ianthodon schultzei holotype KUVP 133735, slab in present condition, combined with photograph of skull area (lower left) before its removal." figureDoi="http://doi.org/10.5281/zenodo.11045473" httpUri="https://zenodo.org/record/11045473/files/figure.png" pageId="1" pageNumber="18">Fig. 2</figureCitation>
), the skull was embedded and prepared from the other side, now exposing the labial surface of the maxilla (
<figureCitation id="3CCE2A14FF89FFCCFDCCF88437B9CAF2" box="[576,641,1898,1921]" captionStart="Figure 3" captionStartId="4.[124,191,1359,1380]" captionTargetBox="[304,1282,177,1325]" captionTargetId="figure-9@4.[302,1285,177,1328]" captionTargetPageId="4" captionText="Figure 3. Ianthodon schultzei holotype KUVP 133735. (a) Redocumented skull; (b) dislocated premaxillae. Fr – frontal; j – jugal; l – lacrimal; m – maxilla; p – parietal; po – postorbital; pof – postfrontal; pp – postparietal; prf – prefrontal; sq – squamosal; st – supratemporal; t – tabular." figureDoi="http://doi.org/10.5281/zenodo.11045475" httpUri="https://zenodo.org/record/11045475/files/figure.png" pageId="1" pageNumber="18">Fig. 3</figureCitation>
).
</materialsCitation>
</paragraph>
<paragraph id="A44A3691FF89FFCCFCC2FF5E36AFCC34" blockId="1.[819,1462,176,773]" pageId="1" pageNumber="18">
<emphasis id="9681EA83FF89FFCCFCC2FF5E3126CDB4" box="[846,1054,176,199]" italics="true" pageId="1" pageNumber="18">Referred specimens</emphasis>
.
<collectionCode id="C2E4AE54FF89FFCCFBA7FF5E314BCDB4" box="[1067,1139,176,199]" pageId="1" pageNumber="18">KUVP</collectionCode>
133736, left maxilla;
<collectionCode id="C2E4AE54FF89FFCCFAD3FF5E30B9CDB4" box="[1375,1409,176,199]" pageId="1" pageNumber="18">FO</collectionCode>
176, right maxilla. These referrals are based not only on the shape of the maxilla but also on tooth shape and the reduced number of precaniniform teeth relative to that in
<taxonomicName id="63F54D12FF89FFCCFA95FEFE36A8CC34" baseAuthorityName="Kissel and Reisz" baseAuthorityYear="2004" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="18" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF89FFCCFA95FEFE36A8CC34" italics="true" pageId="1" pageNumber="18">Haptodus garnettensis</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="A44A3691FF89FFCCFCC2FEBE3072CE76" blockId="1.[819,1462,176,773]" pageId="1" pageNumber="18">
<emphasis id="9681EA83FF89FFCCFCC2FEBE3129CC14" box="[846,1041,336,359]" italics="true" pageId="1" pageNumber="18">Revised diagnosis</emphasis>
: Small sphenacodontian characterized by the presence of three premaxillary tooth positions and with conical marginal teeth that overlap each other at the base. It differs from
<taxonomicName id="63F54D12FF89FFCCFB9FFE41303DCCB5" baseAuthorityName="Kissel and Reisz" baseAuthorityYear="2004" box="[1043,1285,431,454]" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="18" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF89FFCCFB9FFE41303DCCB5" box="[1043,1285,431,454]" italics="true" pageId="1" pageNumber="18">Haptodus garnettensis</emphasis>
</taxonomicName>
in having fewer marginal teeth, with spaces for up to 20 teeth in the maxilla, rather than 23. There are at least 21 tooth positions in the dentary, rather than 24. Distal marginal teeth are slenderer and more distinctly recurved than in
<taxonomicName id="63F54D12FF89FFCCFB37FDC1301ACF35" authorityName="Gaudry" authorityYear="1886" box="[1211,1314,559,582]" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="18" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF89FFCCFB37FDC1301ACF35" box="[1211,1314,559,582]" italics="true" pageId="1" pageNumber="18">Haptodus</emphasis>
</taxonomicName>
, with narrow, rather than bulbous, tips. It differs from
<taxonomicName id="63F54D12FF89FFCCFB60FDA130A8CF15" baseAuthorityName="Kissel and Reisz" baseAuthorityYear="2004" box="[1260,1424,591,614]" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="18" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF89FFCCFB60FDA130A8CF15" box="[1260,1424,591,614]" italics="true" pageId="1" pageNumber="18">H. garnettensis</emphasis>
</taxonomicName>
in having four or fewer maxillary teeth anterior to the enlarged pair of teeth on this element rather than six. The teeth on the transverse flange of the pterygoid are smaller than in
<taxonomicName id="63F54D12FF89FFCCFAE7FD403660CF96" authorityName="Gaudry" authorityYear="1886" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="18" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF89FFCCFAE7FD403660CF96" italics="true" pageId="1" pageNumber="18">Haptodus</emphasis>
</taxonomicName>
. It differs from other sphenacodonts in having the pineal foramen located at the midpoint of parietal length.
</paragraph>
</subSubSection>
<subSubSection id="ECEF651AFF89FFCBFCBFFCC03401CB50" lastPageId="6" lastPageNumber="23" pageId="1" pageNumber="18" type="description">
<paragraph id="A44A3691FF89FFCCFCBFFCC036CACE35" blockId="1.[819,1010,814,838]" box="[819,1010,814,838]" pageId="1" pageNumber="18">
<emphasis id="9681EA83FF89FFCCFCBFFCC036CACE35" bold="true" box="[819,1010,814,838]" pageId="1" pageNumber="18">2.1 Description</emphasis>
</paragraph>
<paragraph id="A44A3691FF89FFCCFCBFFC8036FCC830" blockId="1.[819,1462,877,1538]" pageId="1" pageNumber="18">
<taxonomicName id="63F54D12FF89FFCCFCBFFC80313CCEF6" authorityName="Kissel and Reisz" authorityYear="2004" box="[819,1028,878,901]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="18" phylum="Chordata" rank="species" species="schultzei">
<emphasis id="9681EA83FF89FFCCFCBFFC80313CCEF6" box="[819,1028,878,901]" italics="true" pageId="1" pageNumber="18">Ianthodon schultzei</emphasis>
</taxonomicName>
is known from a single juvenile skeleton with delicate bones, differing from contemporary specimens of
<taxonomicName id="63F54D12FF89FFCCFCDDFC433179CEB7" baseAuthorityName="Kissel and Reisz" baseAuthorityYear="2004" box="[849,1089,941,964]" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="18" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF89FFCCFCDDFC433179CEB7" box="[849,1089,941,964]" italics="true" pageId="1" pageNumber="18">Haptodus garnettensis</emphasis>
</taxonomicName>
even when of similar size. The reconstructed skull length of
<taxonomicName id="63F54D12FF89FFCCFBD9FC2331FACE97" authorityName="Kissel &amp; Reisz" authorityYear="2004" box="[1109,1218,973,996]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="18" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF89FFCCFBD9FC2331FACE97" box="[1109,1218,973,996]" italics="true" pageId="1" pageNumber="18">Ianthodon</emphasis>
</taxonomicName>
is slightly less than
<quantity id="630D9B74FF89FFCCFA17FC23366EC977" metricMagnitude="-1" metricUnit="m" metricValue="1.0" pageId="1" pageNumber="18" unit="cm" value="10.0">10 cm</quantity>
, similar in length to the juvenile and smallest specimens
<collectionCode id="C2E4AE54FF89FFCCFCBFFBE33664C957" box="[819,860,1037,1060]" country="Canada" name="McGill University, Redpath Museum" pageId="1" pageNumber="18" type="Museum">RM</collectionCode>
14,156,
<collectionCode id="C2E4AE54FF89FFCCFC30FBE336DDC957" box="[956,997,1037,1060]" country="Canada" name="McGill University, Redpath Museum" pageId="1" pageNumber="18" type="Museum">RM</collectionCode>
14,157 and
<collectionCode id="C2E4AE54FF89FFCCFBE1FBE33190C957" box="[1133,1192,1037,1060]" country="Canada" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34778" lsid="urn:lsid:biocol.org:col:34778" name="Royal Ontario Museum" pageId="1" pageNumber="18" type="Museum">ROM</collectionCode>
29872 of
<taxonomicName id="63F54D12FF89FFCCFA95FBE336A8C937" baseAuthorityName="Kissel and Reisz" baseAuthorityYear="2004" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="18" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF89FFCCFA95FBE336A8C937" italics="true" pageId="1" pageNumber="18">Haptodus garnettensis</emphasis>
</taxonomicName>
(see
<bibRefCitation id="C0644B60FF89FFCCFC44FBC3316AC937" author="Currie, P. J." box="[968,1106,1069,1092]" pageId="1" pageNumber="18" pagination="927 - 942" refId="ref9556" refString="Currie, P. J.: A new haptodontine sphenacodont (Reptilia: Pelycosauria) from the Upper Pennsylvanian of North America, J. Paleontol., 51, 927 - 942, 1977." type="journal article" year="1977">Currie, 1977</bibRefCitation>
;
<bibRefCitation id="C0644B60FF89FFCCFBD1FBC331D2C937" author="Laurin, M." box="[1117,1258,1069,1092]" pageId="1" pageNumber="18" pagination="200 - 229" refId="ref10031" refString="Laurin, M.: Anatomy and Relationships of Haptodus garnettensis, a Pennsylvanian synapsid from Kansas, J. Vertebr. Paleontol., 13, 200 - 229, 1993." type="journal article" year="1993">Laurin, 1993</bibRefCitation>
). As reconstructed, the skull of
<taxonomicName id="63F54D12FF89FFCCFC3CFBA33125C917" authorityName="Kissel &amp; Reisz" authorityYear="2004" box="[944,1053,1101,1124]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="18" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF89FFCCFC3CFBA33125C917" box="[944,1053,1101,1124]" italics="true" pageId="1" pageNumber="18">Ianthodon</emphasis>
</taxonomicName>
is slenderer than
<taxonomicName id="63F54D12FF89FFCCFB5AFBA33040C917" baseAuthorityName="Kissel and Reisz" baseAuthorityYear="2004" box="[1238,1400,1101,1124]" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="18" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF89FFCCFB5AFBA33040C917" box="[1238,1400,1101,1124]" italics="true" pageId="1" pageNumber="18">H. garnettensis</emphasis>
</taxonomicName>
specimens of the same size in the region of the snout and the anterior mandible. As far as can be discerned, the postcranial proportions of
<taxonomicName id="63F54D12FF89FFCCFB88FB423149C9B0" authorityName="Kissel &amp; Reisz" authorityYear="2004" box="[1028,1137,1196,1219]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="18" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF89FFCCFB88FB423149C9B0" box="[1028,1137,1196,1219]" italics="true" pageId="1" pageNumber="18">Ianthodon</emphasis>
</taxonomicName>
are nearly the same as in
<collectionCode id="C2E4AE54FF89FFCCFA01FB42308EC9B0" box="[1421,1462,1196,1219]" country="Canada" name="McGill University, Redpath Museum" pageId="1" pageNumber="18" type="Museum">RM</collectionCode>
14,156, while the higher number of precaniniform maxillary teeth and the more rectangular shape of the humerus entepicondyle distinguish the
<typeStatus id="7B4E8833FF89FFCCFBA2FAE231B3C850" box="[1070,1163,1292,1315]" pageId="1" pageNumber="18" type="holotype">holotype</typeStatus>
of
<taxonomicName id="63F54D12FF89FFCCFB21FAE23077C850" baseAuthorityName="Kissel and Reisz" baseAuthorityYear="2004" box="[1197,1359,1292,1315]" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="18" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF89FFCCFB21FAE23077C850" box="[1197,1359,1292,1315]" italics="true" pageId="1" pageNumber="18">H. garnettensis</emphasis>
</taxonomicName>
from that of
<taxonomicName id="63F54D12FF89FFCCFCDCFAC23685C830" authorityName="Kissel &amp; Reisz" authorityYear="2004" box="[848,957,1324,1347]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="18" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF89FFCCFCDCFAC23685C830" box="[848,957,1324,1347]" italics="true" pageId="1" pageNumber="18">Ianthodon</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="A44A3691FF89FFCCFCC2FAA2365FCB71" blockId="1.[819,1462,877,1538]" pageId="1" pageNumber="18">
The
<typeStatus id="7B4E8833FF89FFCCFCF1FAA236E2C810" box="[893,986,1356,1379]" pageId="1" pageNumber="18" type="holotype">holotype</typeStatus>
skeleton is preserved on the same block with some skeletal elements of
<taxonomicName id="63F54D12FF89FFCCFBCAFA8231CAC8F0" authorityName="Lane" authorityYear="1945" box="[1094,1266,1388,1411]" family="Petrolacosauridae" genus="Petrolacosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="18" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF89FFCCFBCAFA8231CAC8F0" box="[1094,1266,1388,1411]" italics="true" pageId="1" pageNumber="18">Petrolacosaurus</emphasis>
</taxonomicName>
, from which it can easily be distinguished on anatomical grounds. The humeri, although of similar size, show clear differences such as proportions and the positions of the foramen and supinator process.
</paragraph>
<paragraph id="A44A3691FF89FFCCFCBFF9C53686CB30" blockId="1.[819,958,1579,1603]" box="[819,958,1579,1603]" pageId="1" pageNumber="18">
<emphasis id="9681EA83FF89FFCCFCBFF9C53686CB30" bold="true" box="[819,958,1579,1603]" pageId="1" pageNumber="18">2.1.1 Skull</emphasis>
</paragraph>
<paragraph id="A44A3691FF89FFCEFCBFF9853490CB71" blockId="1.[819,1462,1643,1921]" lastBlockId="3.[124,767,1324,1921]" lastPageId="3" lastPageNumber="20" pageId="1" pageNumber="18">
The dorsal skull roof of
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133735 has already been described in detail by
<bibRefCitation id="C0644B60FF89FFCCFBBBF9643073CBD2" author="Kissel, R. A. &amp; Reisz, R. R." box="[1079,1355,1674,1697]" pageId="1" pageNumber="18" pagination="409 - 428" refId="ref9945" refString="Kissel, R. A. and Reisz, R. R.: Synapsid fauna of the Upper Pennsylvanian Rock Lake Shale near Garnett, Kansas and the diversity pattern of early amniotes, in: Recent Advances in the Origin and Early Radiation of Vertebrates, edited by: Arratia, G., Wilson, M. V. H., and Cloutier, R., Verlag Dr. F. Pfeil, Munchen, 409 - 428, 2004." type="book chapter" year="2004">Kissel and Reisz (2004)</bibRefCitation>
, and this need not be repeated here. The skull elements are spread across the slabs that form the fossil block (
<figureCitation id="3CCE2A14FF89FFCCFB72F9243004CB92" box="[1278,1340,1738,1761]" captionStart="Figure 1" captionStartId="2.[124,191,1699,1720]" captionTargetBox="[310,1278,235,1659]" captionTargetId="figure-9@2.[302,1285,226,1667]" captionTargetPageId="2" captionText="Figure 1. Ianthodon schultzei holotype KUVP 133735, combined slab, with outlines of skull bones (outlines by D. Scott, made prior to Kissel and Reisz, 2004, Fig. 2; now exposed as embedded counter slab) and a disarticulated skeleton of the diapsid reptile Petrolacosaurus kansensis (shaded areas); a – angular; ar – articular; c – vertebra centrum; cau – caudal neural arch; cle – cleithrum; co – anterior coracoid; cp – cultriform process of parasphenoid–basisphenoid complex; cr – cervical rib; d – dentary; dr – dorsal rib; h – humerus; n – nasal; na – presacral neural arch; pal – palatine; pc – posterior coronoid; pm – premaxilla; pra – prearticular; ps – parasphenoid–basisphenoid complex; pt – pterygoid; qj – quadratojugal; sa – surangular; sc – scapula; soc – supraoccipital; sp – splenial; v – vomer." figureDoi="http://doi.org/10.5281/zenodo.11045471" httpUri="https://zenodo.org/record/11045471/files/figure.png" pageId="1" pageNumber="18">Fig. 1</figureCitation>
), with several elements trending off the edges. Most of the skull roof is preserved on the original small block, mostly disarticulated. The nasals, quadratojugal and premaxillae have clearly moved away from the other skull elements. A concentration of palatal, occipital and mandibular elements is found in close proximity to each other on one of the other blocks, together with other skeletal elements. Partly because of the juvenile condition of the specimen, it is somewhat difficult to interpret the exact edges of some of the elements. We are, however, confident that these bones belonged to one individual because of the consistency in anatomy, level of ossification and lack of duplication.
</paragraph>
<caption id="F08A6619FF8AFFCFFFF0F94D31A4CA39" ID-DOI="http://doi.org/10.5281/zenodo.11045471" ID-Zenodo-Dep="11045471" httpUri="https://zenodo.org/record/11045471/files/figure.png" pageId="2" pageNumber="19" startId="2.[124,191,1699,1720]" targetBox="[310,1278,235,1659]" targetPageId="2" targetType="figure">
<paragraph id="A44A3691FF8AFFCFFFF0F94D31A4CA39" blockId="2.[124,1462,1699,1866]" pageId="2" pageNumber="19">
<emphasis id="9681EA83FF8AFFCFFFF0F94D35E2CBCB" bold="true" box="[124,218,1699,1720]" pageId="2" pageNumber="19">Figure 1.</emphasis>
<taxonomicName id="63F54D12FF8AFFCFFF6DF94D34A7CBCB" authorityName="Kissel and Reisz" authorityYear="2004" box="[225,415,1699,1720]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="2" pageNumber="19" phylum="Chordata" rank="species" species="schultzei">
<emphasis id="9681EA83FF8AFFCFFF6DF94D34A7CBCB" box="[225,415,1699,1720]" italics="true" pageId="2" pageNumber="19">Ianthodon schultzei</emphasis>
</taxonomicName>
holotype KUVP 133735, combined slab, with outlines of skull bones (outlines by D. Scott, made prior to
<bibRefCitation id="C0644B60FF8AFFCFFFF0F92E3463CBA6" author="Kissel, R. A. &amp; Reisz, R. R." box="[124,347,1728,1749]" pageId="2" pageNumber="19" pagination="409 - 428" refId="ref9945" refString="Kissel, R. A. and Reisz, R. R.: Synapsid fauna of the Upper Pennsylvanian Rock Lake Shale near Garnett, Kansas and the diversity pattern of early amniotes, in: Recent Advances in the Origin and Early Radiation of Vertebrates, edited by: Arratia, G., Wilson, M. V. H., and Cloutier, R., Verlag Dr. F. Pfeil, Munchen, 409 - 428, 2004." type="book chapter" year="2004">Kissel and Reisz, 2004</bibRefCitation>
, Fig. 2; now exposed as embedded counter slab) and a disarticulated skeleton of the diapsid reptile
<taxonomicName id="63F54D12FF8AFFCFFA97F92F35E0CB80" authorityName="Lane" authorityYear="1945" family="Petrolacosauridae" genus="Petrolacosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="2" pageNumber="19" phylum="Chordata" rank="species" species="kansensis">
<emphasis id="9681EA83FF8AFFCFFA97F92F35E0CB80" italics="true" pageId="2" pageNumber="19">Petrolacosaurus kansensis</emphasis>
</taxonomicName>
(shaded areas); a – angular; ar – articular; c – vertebra centrum; cau – caudal neural arch; cle – cleithrum; co – anterior coracoid; cp – cultriform process of parasphenoid–basisphenoid complex; cr – cervical rib; d – dentary; dr – dorsal rib; h – humerus; n – nasal; na – presacral neural arch; pal – palatine; pc – posterior coronoid; pm – premaxilla; pra – prearticular; ps – parasphenoid–basisphenoid complex; pt – pterygoid; qj – quadratojugal; sa – surangular; sc – scapula; soc – supraoccipital; sp – splenial; v – vomer.
</paragraph>
</caption>
<caption id="F08A6619FF8BFFCEFFF0FB953466C9B8" ID-DOI="http://doi.org/10.5281/zenodo.11045473" ID-Zenodo-Dep="11045473" httpUri="https://zenodo.org/record/11045473/files/figure.png" pageId="3" pageNumber="20" startId="3.[124,191,1147,1168]" targetBox="[133,756,179,1112]" targetPageId="3" targetType="figure">
<paragraph id="A44A3691FF8BFFCEFFF0FB953466C9B8" blockId="3.[124,767,1147,1227]" pageId="3" pageNumber="20">
<emphasis id="9681EA83FF8BFFCEFFF0FB9535E5C9E3" bold="true" box="[124,221,1147,1168]" pageId="3" pageNumber="20">Figure 2.</emphasis>
<taxonomicName id="63F54D12FF8BFFCEFF64FB953491C9E3" authorityName="Kissel and Reisz" authorityYear="2004" box="[232,425,1147,1168]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="20" phylum="Chordata" rank="species" species="schultzei">
<emphasis id="9681EA83FF8BFFCEFF64FB953491C9E3" box="[232,425,1147,1168]" italics="true" pageId="3" pageNumber="20">Ianthodon schultzei</emphasis>
</taxonomicName>
holotype KUVP 133735, slab in present condition, combined with photograph of skull area (lower left) before its removal.
</paragraph>
</caption>
<paragraph id="A44A3691FF8BFFCEFF1BF9E5308FCC74" blockId="3.[124,767,1324,1921]" lastBlockId="3.[819,1463,176,1921]" pageId="3" pageNumber="20">The premaxilla is an unusually slender element, especially in view of the large size of the first tooth. All three rami of the premaxilla are slender. The nasal process is slightly broader transversely than the maxillary process, whereas the vomerine process is the smallest and most slender of the three rami. The slenderness of the maxillary process and of the body of the bone suggests that the teeth were not deeply rooted, a condition similar to that seen in the maxilla, despite the unusually large size of the first tooth. The right premaxilla, the better preserved of the pair, bears three marginal teeth. The first tooth is broad at the base, although slightly exaggerated by compression, and much taller than the second tooth. These teeth are only slightly curved posteriorly, and this curvature is restricted to the tip. The upper crowns are densely striated on the lingual side, and moderately striated on the labial side.</paragraph>
<paragraph id="A44A3691FF8BFFCEFCC2FEFE3657CF96" blockId="3.[819,1463,176,1921]" pageId="3" pageNumber="20">
The maxilla of the
<typeStatus id="7B4E8833FF8BFFCEFB97FEFE3140CC54" box="[1051,1144,272,295]" pageId="3" pageNumber="20" type="holotype">holotype</typeStatus>
<collectionCode id="C2E4AE54FF8BFFCEFBF3FEFE31FFCC54" box="[1151,1223,272,295]" pageId="3" pageNumber="20">KUVP</collectionCode>
133735 is poorly preserved, but two referred specimens provide valuable additional information about the anatomy of this element. These specimens are identified with confidence as belonging to
<taxonomicName id="63F54D12FF8BFFCEFCBFFE613698CCD5" authorityName="Kissel &amp; Reisz" authorityYear="2004" box="[819,928,399,422]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="20" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF8BFFCEFCBFFE613698CCD5" box="[819,928,399,422]" italics="true" pageId="3" pageNumber="20">Ianthodon</emphasis>
</taxonomicName>
because of the unique dentition that they share with the
<typeStatus id="7B4E8833FF8BFFCEFC1CFE4136C8CCB5" box="[912,1008,431,454]" pageId="3" pageNumber="20" type="holotype">holotype</typeStatus>
. Overall, the shape of the teeth is quite unusual, not seen in other coeval amniotes. The teeth have a broad base, but very slender crowns, tapering rapidly crownward without any bulbous thickening. The slender crowns are recurved. This is in strong contrast to the teeth of
<emphasis id="9681EA83FF8BFFCEFAC3FDC13684CF15" italics="true" pageId="3" pageNumber="20">
<taxonomicName id="63F54D12FF8BFFCEFAC3FDC1368FCF15" baseAuthorityName="Kissel and Reisz" baseAuthorityYear="2004" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="20" phylum="Chordata" rank="species" species="garnettensis">Haptodus garnettensis</taxonomicName>
,
</emphasis>
which are characterized by their overall robustness, as well as a slight bulbousness below the crown. These maxillae also differ from those of
<taxonomicName id="63F54D12FF8BFFCEFB12FD61303DCFD5" authorityName="Gaudry" authorityYear="1886" box="[1182,1285,655,678]" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="20" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF8BFFCEFB12FD61303DCFD5" box="[1182,1285,655,678]" italics="true" pageId="3" pageNumber="20">Haptodus</emphasis>
</taxonomicName>
in the outline of the dorsal edge and in the presence of fewer precaniniform teeth.
</paragraph>
<paragraph id="A44A3691FF8BFFCEFCC2FD00308FC9B0" blockId="3.[819,1463,176,1921]" pageId="3" pageNumber="20">
The dorsal blade of the maxilla is low. Best seen in the medially exposed referred specimens, it increases in height gradually along the anterior one-third of the bone and above the alveolar shelf, reaching its maximum above the enlarged pair of teeth. Its dorsal edge extends posteriorly along four or five tooth positions before it starts to slope ventrally, reaching the alveolar shelf by the nineteenth tooth position. Anteriorly, the sutural contact with the premaxilla covers most of the alveolar shelf of the first maxillary tooth. Between the second and sixth tooth positions, the alveolar shelf is smooth, defining the maxillary contribution to the elongate internal naris. There is little or no dorsal expansion of the alveolar shelf in the region of the enlarged teeth. The medioventral part of the dorsal blade is slightly swollen above the alveolar shelf but less so than in
<taxonomicName id="63F54D12FF8BFFCEFBA3FB423027C9B0" baseAuthorityName="Kissel and Reisz" baseAuthorityYear="2004" box="[1071,1311,1196,1219]" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="20" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF8BFFCEFBA3FB423027C9B0" box="[1071,1311,1196,1219]" italics="true" pageId="3" pageNumber="20">Haptodus garnettensis</emphasis>
</taxonomicName>
(
<collectionCode id="C2E4AE54FF8BFFCEFAA0FB42306FC9B0" box="[1324,1367,1196,1219]" country="Canada" name="McGill University, Redpath Museum" pageId="3" pageNumber="20" type="Museum">RM</collectionCode>
14,157).
</paragraph>
<paragraph id="A44A3691FF8BFFCEFCC2FB2236ECCA52" blockId="3.[819,1463,176,1921]" pageId="3" pageNumber="20">
The maxillary tooth count can be calculated for
<taxonomicName id="63F54D12FF8BFFCEFAC5FB22308EC990" authorityName="Kissel &amp; Reisz" authorityYear="2004" box="[1353,1462,1228,1251]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="20" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF8BFFCEFAC5FB22308EC990" box="[1353,1462,1228,1251]" italics="true" pageId="3" pageNumber="20">Ianthodon</emphasis>
</taxonomicName>
because the two referred maxillae are nearly complete. There are at least 18 positions preserved in
<collectionCode id="C2E4AE54FF8BFFCEFB34FAE23038C850" box="[1208,1280,1292,1315]" pageId="3" pageNumber="20">KUVP</collectionCode>
133736, and it is likely that there would have been up to two more tooth positions in the maxilla posteriorly. As seen in the referred specimens (
<figureCitation id="3CCE2A14FF8BFFCEFCF6FA823683C8F0" box="[890,955,1388,1411]" captionStart="Figure 4" captionStartId="5.[124,191,815,836]" captionTargetBox="[377,1208,177,783]" captionTargetId="figure-9@5.[377,1208,177,783]" captionTargetPageId="5" captionText="Figure 4. Ianthodon schultzei. (a) Referred left maxilla KUVP 133736; (b) referred right maxilla FO 176." figureDoi="http://doi.org/10.5281/zenodo.11045477" httpUri="https://zenodo.org/record/11045477/files/figure.png" pageId="3" pageNumber="20">Fig. 4</figureCitation>
), either three (
<collectionCode id="C2E4AE54FF8BFFCEFBD9FA823140C8F0" box="[1109,1144,1388,1411]" pageId="3" pageNumber="20">FO</collectionCode>
176) or four (
<collectionCode id="C2E4AE54FF8BFFCEFA81FA82306FC8F0" box="[1293,1367,1388,1411]" pageId="3" pageNumber="20">KUVP</collectionCode>
133736) anterior teeth increase in size towards the pair of enlarged teeth. As in other sphenacodonts, there are two tooth positions for the largest teeth of the maxilla, and, generally, their location is indicated by the center of growth on the maxilla. Posterior to this region, the teeth gradually decrease in size posteriorly. All teeth are broad at their bases, remaining largely uncurved through most of the crown, with tips curving posteriorly (
<bibRefCitation id="C0644B60FF8BFFCEFB9CF9853199CBF1" author="Laurin, M." box="[1040,1185,1643,1666]" pageId="3" pageNumber="20" pagination="200 - 229" refId="ref10031" refString="Laurin, M.: Anatomy and Relationships of Haptodus garnettensis, a Pennsylvanian synapsid from Kansas, J. Vertebr. Paleontol., 13, 200 - 229, 1993." type="journal article" year="1993">Laurin, 1993</bibRefCitation>
). Striations are present in the presumed crown portions of the maxillary teeth. As seen in the spectrum of juvenile to adult specimens of
<taxonomicName id="63F54D12FF8BFFCEFAC3F945368DCB92" baseAuthorityName="Kissel and Reisz" baseAuthorityYear="2004" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="20" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF8BFFCEFAC3F945368DCB92" italics="true" pageId="3" pageNumber="20">Haptodus garnettensis</emphasis>
</taxonomicName>
, as well as other non-therapsid synapsids, significant ontogenetic changes of the tooth crown
<typeStatus id="7B4E8833FF8BFFCEFABBF904305DCA72" box="[1335,1381,1770,1793]" pageId="3" pageNumber="20">type</typeStatus>
are not to be expected.
</paragraph>
<paragraph id="A44A3691FF8BFFC9FCC2F8C437DDCB07" blockId="3.[819,1463,176,1921]" lastBlockId="4.[124,767,1502,1907]" lastPageId="4" lastPageNumber="21" pageId="3" pageNumber="20">
When compared to the known maxillae of
<taxonomicName id="63F54D12FF8BFFCEFA96F8C436A8CA12" baseAuthorityName="Kissel and Reisz" baseAuthorityYear="2004" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="20" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF8BFFCEFA96F8C436A8CA12" italics="true" pageId="3" pageNumber="20">Haptodus garnettensis</emphasis>
</taxonomicName>
where a higher tooth count is present both in adult and in juvenile stages (see
<bibRefCitation id="C0644B60FF8BFFCEFB14F884301BCAF2" author="Currie, P. J." box="[1176,1315,1898,1921]" pageId="3" pageNumber="20" pagination="927 - 942" refId="ref9556" refString="Currie, P. J.: A new haptodontine sphenacodont (Reptilia: Pelycosauria) from the Upper Pennsylvanian of North America, J. Paleontol., 51, 927 - 942, 1977." type="journal article" year="1977">Currie, 1977</bibRefCitation>
), the reduced dentition of
<taxonomicName id="63F54D12FF8CFFC9FF72FA303453C886" authorityName="Kissel &amp; Reisz" authorityYear="2004" box="[254,363,1502,1525]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="21" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF8CFFC9FF72FA303453C886" box="[254,363,1502,1525]" italics="true" pageId="4" pageNumber="21">Ianthodon</emphasis>
</taxonomicName>
is of taxonomic significance. This reduced number of maxillary teeth is related to differences in the precaniniform tooth count, a feature that appears to correspond to the reduced premaxillary tooth number in this taxon relative to the condition in
<taxonomicName id="63F54D12FF8CFFC9FE16F9B337D9CB07" authority="(Laurin, 1993)" baseAuthorityName="Laurin" baseAuthorityYear="1993" box="[410,737,1629,1652]" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="21" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF8CFFC9FE16F9B33704CB07" box="[410,572,1629,1652]" italics="true" pageId="4" pageNumber="21">H. garnettensis</emphasis>
(
<bibRefCitation id="C0644B60FF8CFFC9FDC6F9B337E1CB07" author="Laurin, M." box="[586,729,1629,1652]" pageId="4" pageNumber="21" pagination="200 - 229" refId="ref10031" refString="Laurin, M.: Anatomy and Relationships of Haptodus garnettensis, a Pennsylvanian synapsid from Kansas, J. Vertebr. Paleontol., 13, 200 - 229, 1993." type="journal article" year="1993">Laurin, 1993</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
<caption id="F08A6619FF8CFFC9FFF0FAA135DAC8EC" ID-DOI="http://doi.org/10.5281/zenodo.11045475" ID-Zenodo-Dep="11045475" httpUri="https://zenodo.org/record/11045475/files/figure.png" pageId="4" pageNumber="21" startId="4.[124,191,1359,1380]" targetBox="[304,1282,177,1325]" targetPageId="4" targetType="figure">
<paragraph id="A44A3691FF8CFFC9FFF0FAA135DAC8EC" blockId="4.[124,1462,1359,1439]" pageId="4" pageNumber="21">
<emphasis id="9681EA83FF8CFFC9FFF0FAA135E2C817" bold="true" box="[124,218,1359,1380]" pageId="4" pageNumber="21">Figure 3.</emphasis>
<taxonomicName id="63F54D12FF8CFFC9FF6FFABE3499C816" authorityName="Kissel and Reisz" authorityYear="2004" box="[227,417,1360,1381]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="21" phylum="Chordata" rank="species" species="schultzei">
<emphasis id="9681EA83FF8CFFC9FF6FFABE3499C816" box="[227,417,1360,1381]" italics="true" pageId="4" pageNumber="21">Ianthodon schultzei</emphasis>
</taxonomicName>
holotype KUVP 133735.
<emphasis id="9681EA83FF8CFFC9FD24FAA137FCC817" bold="true" box="[680,708,1359,1380]" pageId="4" pageNumber="21">(a)</emphasis>
Redocumented skull;
<emphasis id="9681EA83FF8CFFC9FC28FAA136F9C817" bold="true" box="[932,961,1359,1380]" pageId="4" pageNumber="21">(b)</emphasis>
dislocated premaxillae. Fr – frontal; j – jugal; l – lacrimal; m – maxilla; p – parietal; po – postorbital; pof – postfrontal; pp – postparietal; prf – prefrontal; sq – squamosal; st – supratemporal; t – tabular.
</paragraph>
</caption>
<paragraph id="A44A3691FF8CFFC9FF1BF99331F5CBC7" blockId="4.[124,767,1502,1907]" lastBlockId="4.[819,1462,1502,1907]" pageId="4" pageNumber="21">
The tall lacrimal of
<emphasis id="9681EA83FF8CFFC9FEF8F9933776CBE7" box="[372,590,1661,1684]" italics="true" pageId="4" pageNumber="21">
<taxonomicName id="63F54D12FF8CFFC9FEF8F9933771CBE7" authorityName="Kissel and Reisz" authorityYear="2004" box="[372,585,1661,1684]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="21" phylum="Chordata" rank="species" species="schultzei">Ianthodon schultzei</taxonomicName>
,
</emphasis>
despite its relatively poor preservation, is sufficiently complete for determining its outline, showing it to be as tall as the maxilla or even slightly taller. This is consistent with the lacrimal height of other basal sphenacodontians (
<bibRefCitation id="C0644B60FF8CFFC9FDAEF9133797CA67" author="Laurin, M." box="[546,687,1789,1812]" pageId="4" pageNumber="21" pagination="200 - 229" refId="ref10031" refString="Laurin, M.: Anatomy and Relationships of Haptodus garnettensis, a Pennsylvanian synapsid from Kansas, J. Vertebr. Paleontol., 13, 200 - 229, 1993." type="journal article" year="1993">Laurin, 1993</bibRefCitation>
,
<figureCitation id="3CCE2A14FF8CFFC9FD35F91337CDCA67" box="[697,757,1789,1812]" captionStart="Figure 3" captionStartId="4.[124,191,1359,1380]" captionTargetBox="[304,1282,177,1325]" captionTargetId="figure-9@4.[302,1285,177,1328]" captionTargetPageId="4" captionText="Figure 3. Ianthodon schultzei holotype KUVP 133735. (a) Redocumented skull; (b) dislocated premaxillae. Fr – frontal; j – jugal; l – lacrimal; m – maxilla; p – parietal; po – postorbital; pof – postfrontal; pp – postparietal; prf – prefrontal; sq – squamosal; st – supratemporal; t – tabular." figureDoi="http://doi.org/10.5281/zenodo.11045475" httpUri="https://zenodo.org/record/11045475/files/figure.png" pageId="4" pageNumber="21">Fig. 3</figureCitation>
) but contrasts with the still taller lacrimal of
<taxonomicName id="63F54D12FF8CFFC9FDEDF8F337C6CA47" authorityName="Credner" authorityYear="1888" box="[609,766,1821,1844]" genus="Palaeohatteria" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="21" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF8CFFC9FDEDF8F337C6CA47" box="[609,766,1821,1844]" italics="true" pageId="4" pageNumber="21">Palaeohatteria</emphasis>
</taxonomicName>
(
<bibRefCitation id="C0644B60FF8CFFC9FF0FF8D2341ACA20" author="Credner, H." box="[131,290,1852,1875]" pageId="4" pageNumber="21" pagination="490 - 558" refId="ref9593" refString="Credner, H.: Die Stegocephalen und Saurier aus dem Rothliegenden des Plauen'schen Grundes bei Dresden, vii. Theil: Palaeohatteria longicaudata CRED, Z. Deut. Geol. Ges., 40, 490 - 558, 1888." type="journal article" year="1888">Credner, 1888</bibRefCitation>
, pl. 25,
<figureCitation id="3CCE2A14FF8CFFC9FEF2F8D234F8CA20" box="[382,448,1852,1875]" captionStart="Figure 4" captionStartId="5.[124,191,815,836]" captionTargetBox="[377,1208,177,783]" captionTargetId="figure-9@5.[377,1208,177,783]" captionTargetPageId="5" captionText="Figure 4. Ianthodon schultzei. (a) Referred left maxilla KUVP 133736; (b) referred right maxilla FO 176." figureDoi="http://doi.org/10.5281/zenodo.11045477" httpUri="https://zenodo.org/record/11045477/files/figure.png" pageId="4" pageNumber="21">Fig. 4</figureCitation>
) and the slender elements in ophiacodontids and eothyridids. The latter basal synapsids have distinct, elongate maxilla–lacrimal sutures, and their lacrimals tend to be slender anteriorly. As in edaphosaurids and basal sphenacodontians,
<taxonomicName id="63F54D12FF8CFFC9FBE3F9F031DCCB46" authorityName="Kissel and Reisz" authorityYear="2004" box="[1135,1252,1566,1589]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="21" phylum="Chordata" rank="species" species="schultzei">
<emphasis id="9681EA83FF8CFFC9FBE3F9F031DCCB46" box="[1135,1252,1566,1589]" italics="true" pageId="4" pageNumber="21">I. schultzei</emphasis>
</taxonomicName>
has a tall lacrimal, indicating that the snout was proportionately taller than in eothyridids or ophiacodontids. Nevertheless, details of the lacrimal foramina and its precise sutural contacts are not available for description or evaluation.
</paragraph>
<paragraph id="A44A3691FF8CFFC8FCC2F95337C7CE8D" blockId="4.[819,1462,1502,1907]" lastBlockId="5.[124,767,904,1915]" lastPageId="5" lastPageNumber="22" pageId="4" pageNumber="21">
In the original description (
<bibRefCitation id="C0644B60FF8CFFC9FBF1F95330B8CBA7" author="Kissel, R. A. &amp; Reisz, R. R." box="[1149,1408,1725,1748]" pageId="4" pageNumber="21" pagination="409 - 428" refId="ref9945" refString="Kissel, R. A. and Reisz, R. R.: Synapsid fauna of the Upper Pennsylvanian Rock Lake Shale near Garnett, Kansas and the diversity pattern of early amniotes, in: Recent Advances in the Origin and Early Radiation of Vertebrates, edited by: Arratia, G., Wilson, M. V. H., and Cloutier, R., Verlag Dr. F. Pfeil, Munchen, 409 - 428, 2004." type="book chapter" year="2004">Kissel and Reisz, 2004</bibRefCitation>
), the left prefrontal was identified as an element covering the maxillary tooth row, and the right was located between the lacrimal and frontal. The supraorbital bar of the “left” currently shows a ridge that may indicate the dorsal surface of the skull, and therefore this element more likely represents the right prefrontal. The anterior tips of the two prefrontals point toward one other and are exposed in medial view. A prefrontal pocket is now clearly observed as the depression in the innermost prefrontal in
<bibRefCitation id="C0644B60FF8DFFC8FE38FC093791CE8D" author="Kissel, R. A. &amp; Reisz, R. R." box="[436,681,999,1022]" pageId="5" pageNumber="22" pagination="409 - 428" refId="ref9945" refString="Kissel, R. A. and Reisz, R. R.: Synapsid fauna of the Upper Pennsylvanian Rock Lake Shale near Garnett, Kansas and the diversity pattern of early amniotes, in: Recent Advances in the Origin and Early Radiation of Vertebrates, edited by: Arratia, G., Wilson, M. V. H., and Cloutier, R., Verlag Dr. F. Pfeil, Munchen, 409 - 428, 2004." type="book chapter" year="2004">Kissel and Reisz (2004</bibRefCitation>
,
<figureCitation id="3CCE2A14FF8DFFC8FD3EFC0937CACE8D" box="[690,754,999,1022]" captionStart="Figure 2" captionStartId="3.[124,191,1147,1168]" captionTargetBox="[133,756,179,1112]" captionTargetId="figure-9@3.[132,759,177,1115]" captionTargetPageId="3" captionText="Figure 2. Ianthodon schultzei holotype KUVP 133735, slab in present condition, combined with photograph of skull area (lower left) before its removal." figureDoi="http://doi.org/10.5281/zenodo.11045473" httpUri="https://zenodo.org/record/11045473/files/figure.png" pageId="5" pageNumber="22">Fig. 2</figureCitation>
).
</paragraph>
<caption id="F08A6619FF8DFFC8FFF0FCC1314CCE37" ID-DOI="http://doi.org/10.5281/zenodo.11045477" ID-Zenodo-Dep="11045477" box="[124,1140,815,836]" httpUri="https://zenodo.org/record/11045477/files/figure.png" pageId="5" pageNumber="22" startId="5.[124,191,815,836]" targetBox="[377,1208,177,783]" targetPageId="5" targetType="figure">
<paragraph id="A44A3691FF8DFFC8FFF0FCC1314CCE37" blockId="5.[124,1140,815,836]" box="[124,1140,815,836]" pageId="5" pageNumber="22">
<emphasis id="9681EA83FF8DFFC8FFF0FCC135EFCE37" bold="true" box="[124,215,815,836]" pageId="5" pageNumber="22">Figure 4.</emphasis>
<taxonomicName id="63F54D12FF8DFFC8FF51FCC134A0CE37" authorityName="Kissel and Reisz" authorityYear="2004" box="[221,408,815,836]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="22" phylum="Chordata" rank="species" species="schultzei">
<emphasis id="9681EA83FF8DFFC8FF51FCC134A0CE37" box="[221,408,815,836]" italics="true" pageId="5" pageNumber="22">Ianthodon schultzei</emphasis>
</taxonomicName>
.
<emphasis id="9681EA83FF8DFFC8FE29FCC134F8CE37" bold="true" box="[421,448,815,836]" pageId="5" pageNumber="22">(a)</emphasis>
Referred left maxilla KUVP 133736;
<emphasis id="9681EA83FF8DFFC8FCA3FCC13674CE37" bold="true" box="[815,844,815,836]" pageId="5" pageNumber="22">(b)</emphasis>
referred right maxilla FO 176.
</paragraph>
</caption>
<paragraph id="A44A3691FF8DFFC8FF1BFBE93585C9ED" blockId="5.[124,767,904,1915]" pageId="5" pageNumber="22">
As with many other elements, the squamosal is already well-described. However, the element identified as the left squamosal by
<bibRefCitation id="C0644B60FF8DFFC8FEADFBA9370CC92D" author="Kissel, R. A. &amp; Reisz, R. R." box="[289,564,1095,1118]" pageId="5" pageNumber="22" pagination="409 - 428" refId="ref9945" refString="Kissel, R. A. and Reisz, R. R.: Synapsid fauna of the Upper Pennsylvanian Rock Lake Shale near Garnett, Kansas and the diversity pattern of early amniotes, in: Recent Advances in the Origin and Early Radiation of Vertebrates, edited by: Arratia, G., Wilson, M. V. H., and Cloutier, R., Verlag Dr. F. Pfeil, Munchen, 409 - 428, 2004." type="book chapter" year="2004">Kissel and Reisz (2004)</bibRefCitation>
bears an ascending rim that might mark the inflection towards the occipital plane.
</paragraph>
<paragraph id="A44A3691FF8DFFC8FF1BFB48372BC8CF" blockId="5.[124,767,904,1915]" pageId="5" pageNumber="22">
The quadratojugal is nearly complete and quite similar to those in edaphosaurids and basal sphenacodonts. The narrow ventral portion that would have touched the base of the quadrate is covered by a vertebra, but the dorsal process of the quadratojugal is exposed next to the coracoid. It is sheetlike above the ventral quadrate buttress and is slightly bifurcate at its thin dorsal end. It lacks an anterior process. Overall, its preserved portion is similar to that seen in
<taxonomicName id="63F54D12FF8DFFC8FD1BFA6835C6C8CE" baseAuthorityName="Laurin" baseAuthorityYear="1993" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="22" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF8DFFC8FD1BFA6835C6C8CE" italics="true" pageId="5" pageNumber="22">Haptodus garnettensis</emphasis>
</taxonomicName>
and other sphenacodonts.
</paragraph>
<paragraph id="A44A3691FF8DFFC8FF1BFA2B342BCBEF" blockId="5.[124,767,904,1915]" pageId="5" pageNumber="22">Both vomers can be identified, partially covered by other elements and exposed in partial dorsal view. Anteriorly, its narrow tip has a short indentation for contact with the premaxilla. This element broadens posteriorly, but is covered by the mandible. Very small teeth are observed below the lateral margin, where the vomer would have formed the medial edge of the choana.</paragraph>
<paragraph id="A44A3691FF8DFFC8FF1BF94B3772CA68" blockId="5.[124,767,904,1915]" pageId="5" pageNumber="22">
The rough dimensions of the palatine are discernible on the central block (
<figureCitation id="3CCE2A14FF8DFFC8FEB1F92A3446CBA8" box="[317,382,1732,1755]" captionStart="Figure 5" captionStartId="6.[124,191,1303,1324]" captionTargetBox="[339,1246,177,1272]" captionTargetId="figure-9@6.[339,1246,177,1272]" captionTargetPageId="6" captionText="Figure 5. Ianthodon schultzei holotype KUVP 133735. (a) Close-up of central block; (b) detail of right posterior coronoid with eroded denticles; (c) detail of right pterygoid transverse flange dentition in dorsolateral aspect. Ic – intercentrum; pt-a – pterygoid anterior ramus; pt-q – quadrate ramus of pterygoid." figureDoi="http://doi.org/10.5281/zenodo.11045479" httpUri="https://zenodo.org/record/11045479/files/figure.png" pageId="5" pageNumber="22">Fig. 5</figureCitation>
), near a series of neural arches. The choanal notch of the palatine is also exposed, although the outline of the anterior region is incomplete.
</paragraph>
<paragraph id="A44A3691FF8DFFC8FF1BF8CA307BC8CF" blockId="5.[124,767,904,1915]" lastBlockId="5.[819,1463,904,1883]" pageId="5" pageNumber="22">
Both pterygoids are preserved. Measured from the level of the thickened transverse flange, the anterior process is about twice the length of the quadrate ramus. The dorsal blade emerging from the medial rim of the palatine ramus is as low as in
<taxonomicName id="63F54D12FF8DFFC8FC28FC4631AFCECC" baseAuthorityName="Laurin" baseAuthorityYear="1993" box="[932,1175,936,959]" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="22" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF8DFFC8FC28FC4631AFCECC" box="[932,1175,936,959]" italics="true" pageId="5" pageNumber="22">Haptodus garnettensis</emphasis>
</taxonomicName>
(
<collectionCode id="C2E4AE54FF8DFFC8FB24FC4931DDCECD" box="[1192,1253,935,958]" country="Canada" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34778" lsid="urn:lsid:biocol.org:col:34778" name="Royal Ontario Museum" pageId="5" pageNumber="22" type="Museum">ROM</collectionCode>
43606) but with a steeper posterior edge. The transverse flange bears at least three teeth. Regarding the structural similarity with the pterygoid of
<taxonomicName id="63F54D12FF8DFFC8FC01FBE931B9C96D" baseAuthorityName="Laurin" baseAuthorityYear="1993" box="[909,1153,1031,1054]" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="22" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF8DFFC8FC01FBE931B9C96D" box="[909,1153,1031,1054]" italics="true" pageId="5" pageNumber="22">Haptodus garnettensis</emphasis>
</taxonomicName>
, there are most likely more tooth positions, particularly since the teeth of this row are proportionately smaller in
<taxonomicName id="63F54D12FF8DFFC8FBDDFBA93186C92D" authorityName="Kissel &amp; Reisz" authorityYear="2004" box="[1105,1214,1095,1118]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="22" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF8DFFC8FBDDFBA93186C92D" box="[1105,1214,1095,1118]" italics="true" pageId="5" pageNumber="22">Ianthodon</emphasis>
</taxonomicName>
. The high quadrate ramus equals the proportions seen in other basal synapsids. As is typical for basal sphenacodontians, the basipterygoid articulation lies more ventrally on the body of the pterygoid than in
<taxonomicName id="63F54D12FF8DFFC8FC0EFB293139C9AD" authorityName="Cope" authorityYear="1878" box="[898,1025,1223,1246]" family="Sphenacodontidae" genus="Dimetrodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="22" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF8DFFC8FC0EFB293139C9AD" box="[898,1025,1223,1246]" italics="true" pageId="5" pageNumber="22">Dimetrodon</emphasis>
</taxonomicName>
and other sphenacodontids but more dorsally than in varanopids and ophiacodontid synapsids (compare to
<bibRefCitation id="C0644B60FF8DFFC8FC0EFAE8314FC86E" author="Romer, A. S. &amp; Price, L. I." box="[898,1143,1286,1309]" pageId="5" pageNumber="22" pagination="1 - 538" refId="ref10605" refString="Romer, A. S. and Price, L. I.: Review of the Pelycosauria, Geol. S. Am. S., 28, 1 - 538, 1940." type="journal article" year="1940">Romer and Price, 1940</bibRefCitation>
; Reisz, 1986). A peg-like process marks the area where the epipterygoid articulates with a subvertical notch in the anterior edge of the quadrate ramus, as seen also in most stem sphenacodonts (not in
<taxonomicName id="63F54D12FF8DFFC8FAA3FA88366AC8EE" authorityName="Credner" authorityYear="1888" genus="Palaeohatteria" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="22" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF8DFFC8FAA3FA88366AC8EE" italics="true" pageId="5" pageNumber="22">Palaeohatteria</emphasis>
</taxonomicName>
),
<taxonomicName id="63F54D12FF8DFFC8FCE5FA6836C0C8EE" baseAuthorityName="Brink and Reisz" baseAuthorityYear="2014" box="[873,1016,1414,1437]" family="Sphenacodontidae" genus="Sphenacodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="22" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF8DFFC8FCE5FA6836C0C8EE" box="[873,1016,1414,1437]" italics="true" pageId="5" pageNumber="22">Sphenacodon</emphasis>
</taxonomicName>
(
<bibRefCitation id="C0644B60FF8DFFC8FB8BFA6831ADC8EE" author="Eberth, D. A." box="[1031,1173,1414,1437]" pageId="5" pageNumber="22" pagination="1 - 39" refId="ref9636" refString="Eberth, D. A.: The skull of Sphenacodon ferocior, and comparisons with other sphenacodontines (Reptilia: Pelycosauria), New Mexico Bureau of Mines and Mineral Resources, Circular, 190, 1 - 39, 1985." type="journal article" year="1985">Eberth, 1985</bibRefCitation>
, Fig. 21) and
<taxonomicName id="63F54D12FF8DFFC8FAA6FA68308EC8EE" authorityName="Reisz &amp; Berman" authorityYear="1986" box="[1322,1462,1414,1437]" family="Edaphosauridae" genus="Ianthasaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="22" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF8DFFC8FAA6FA68308EC8EE" box="[1322,1462,1414,1437]" italics="true" pageId="5" pageNumber="22">Ianthasaurus</emphasis>
</taxonomicName>
(
<collectionCode id="C2E4AE54FF8DFFC8FCB6FA4B364FC8CF" box="[826,887,1445,1468]" country="Canada" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34778" lsid="urn:lsid:biocol.org:col:34778" name="Royal Ontario Museum" pageId="5" pageNumber="22" type="Museum">ROM</collectionCode>
59933, contra
<bibRefCitation id="C0644B60FF8DFFC8FB9BFA4B300FC8CF" author="Mazierski, D. M. &amp; Reisz, R. R." box="[1047,1335,1445,1468]" pageId="5" pageNumber="22" pagination="901 - 912" refId="ref10183" refString="Mazierski, D. M. and Reisz, R. R.: Description of a new specimen of Ianthasaurus hardestiorum (Eupelycosauria: Edaphosauridae) and a re-evaluation of edaphosaurid phylogeny, Can. J. Earth Sci., 47, 901 - 912, 2010." type="journal article" year="2010">Mazierski and Reisz, 2010</bibRefCitation>
).
</paragraph>
<paragraph id="A44A3691FF8DFFC8FCC2FA2B3645CBEF" blockId="5.[819,1463,904,1883]" pageId="5" pageNumber="22">
As with the palatine, no significant difference could be found in the parasphenoid–basisphenoid complex of
<taxonomicName id="63F54D12FF8DFFC8FCD6F9EB310DCB6F" authorityName="Kissel and Reisz" authorityYear="2004" box="[858,1077,1541,1564]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="22" phylum="Chordata" rank="species" species="schultzei">
<emphasis id="9681EA83FF8DFFC8FCD6F9EB310DCB6F" box="[858,1077,1541,1564]" italics="true" pageId="5" pageNumber="22">Ianthodon schultzei</emphasis>
</taxonomicName>
and
<taxonomicName id="63F54D12FF8DFFC8FBF0F9EB304ECB6F" baseAuthorityName="Laurin" baseAuthorityYear="1993" box="[1148,1398,1541,1564]" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="22" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF8DFFC8FBF0F9EB304ECB6F" box="[1148,1398,1541,1564]" italics="true" pageId="5" pageNumber="22">Haptodus garnettensis</emphasis>
</taxonomicName>
. The basipterygoid processes point anterolaterally and have flat articular facets. Compared to
<taxonomicName id="63F54D12FF8DFFC8FBFBF9AB3051CB2F" baseAuthorityName="Laurin" baseAuthorityYear="1993" box="[1143,1385,1605,1628]" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="22" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF8DFFC8FBFBF9AB3051CB2F" box="[1143,1385,1605,1628]" italics="true" pageId="5" pageNumber="22">Haptodus garnettensis</emphasis>
</taxonomicName>
(
<collectionCode id="C2E4AE54FF8DFFC8FAF5F9AB308ECB2F" box="[1401,1462,1605,1628]" country="Canada" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34778" lsid="urn:lsid:biocol.org:col:34778" name="Royal Ontario Museum" pageId="5" pageNumber="22" type="Museum">ROM</collectionCode>
43602, 43604), the cultriform process is proportionally longer.
</paragraph>
<paragraph id="A44A3691FF8DFFCBFCC2F94B35FDC892" blockId="5.[819,1463,904,1883]" lastBlockId="6.[124,766,1451,1505]" lastPageId="6" lastPageNumber="23" pageId="5" pageNumber="22">The supraoccipital is subrectangular in outline, with a shallow embayment for the foramen magnum in its central margin. There is a modest vertical median ridge that widens toward the embayment, where the supraoccipital articulated with the exoccipitals on either side of the foramen magnum. Other elements of the braincase may be preserved, but are difficult to identify with confidence because of poor preservation.</paragraph>
<caption id="F08A6619FF8EFFCBFFF0FAF934E8C814" ID-DOI="http://doi.org/10.5281/zenodo.11045479" ID-Zenodo-Dep="11045479" httpUri="https://zenodo.org/record/11045479/files/figure.png" pageId="6" pageNumber="23" startId="6.[124,191,1303,1324]" targetBox="[339,1246,177,1272]" targetPageId="6" targetType="figure">
<paragraph id="A44A3691FF8EFFCBFFF0FAF934E8C814" blockId="6.[124,1462,1303,1383]" pageId="6" pageNumber="23">
<emphasis id="9681EA83FF8EFFCBFFF0FAF935E2C85F" bold="true" box="[124,218,1303,1324]" pageId="6" pageNumber="23">Figure 5.</emphasis>
<taxonomicName id="63F54D12FF8EFFCBFF6EFAF63498C85E" authorityName="Kissel and Reisz" authorityYear="2004" box="[226,416,1304,1325]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="6" pageNumber="23" phylum="Chordata" rank="species" species="schultzei">
<emphasis id="9681EA83FF8EFFCBFF6EFAF63498C85E" box="[226,416,1304,1325]" italics="true" pageId="6" pageNumber="23">Ianthodon schultzei</emphasis>
</taxonomicName>
holotype KUVP 133735.
<emphasis id="9681EA83FF8EFFCBFD28FAF937F8C85F" bold="true" box="[676,704,1303,1324]" pageId="6" pageNumber="23">(a)</emphasis>
Close-up of central block;
<emphasis id="9681EA83FF8EFFCBFC5EFAF936D7C85F" bold="true" box="[978,1007,1303,1324]" pageId="6" pageNumber="23">(b)</emphasis>
detail of right posterior coronoid with eroded denticles;
<emphasis id="9681EA83FF8EFFCBFF53FADB35C1C839" bold="true" box="[223,249,1333,1354]" pageId="6" pageNumber="23">(c)</emphasis>
detail of right pterygoid transverse flange dentition in dorsolateral aspect. Ic – intercentrum; pt-a – pterygoid anterior ramus; pt-q – quadrate ramus of pterygoid.
</paragraph>
</caption>
<paragraph id="A44A3691FF8EFFCBFFF0F9E53401CB50" blockId="6.[124,313,1547,1571]" box="[124,313,1547,1571]" pageId="6" pageNumber="23">
<emphasis id="9681EA83FF8EFFCBFFF0F9E53401CB50" bold="true" box="[124,313,1547,1571]" pageId="6" pageNumber="23">2.1.2 Mandible</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="ECEF651AFF8EFFC4FFF0F9A53478CC95" lastPageId="9" lastPageNumber="26" pageId="6" pageNumber="23" type="materials_examined">
<paragraph id="A44A3691FF8EFFCBFFF0F9A53645CB52" blockId="6.[124,767,1611,1921]" lastBlockId="6.[819,1463,1451,1920]" pageId="6" pageNumber="23">
Both mandibles are preserved, with the better-articulated left ramus measuring approximately
<quantity id="630D9B74FF8EFFCBFD81F985376DCBF1" box="[525,597,1643,1666]" metricMagnitude="-2" metricUnit="m" metricValue="8.4" pageId="6" pageNumber="23" unit="mm" value="84.0">84 mm</quantity>
in total length. In lateral view, the mandible is only slightly bent in its tooth-bearing anterior half. The posterior end of the dentary, together with the dislocated surangular, forms a moderate bulge of the coronoid region, resembling that of the
<taxonomicName id="63F54D12FF8EFFCBFD44F9043478CA52" baseAuthorityName="Laurin" baseAuthorityYear="1993" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="6" pageNumber="23" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF8EFFCBFD44F9043478CA52" italics="true" pageId="6" pageNumber="23">Haptodus garnettensis</emphasis>
</taxonomicName>
specimen
<specimenCode id="F4539EEAFF8EFFCBFE38F8E43706CA52" box="[436,574,1802,1825]" collectionCode="ROM" country="Canada" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34778" lsid="urn:lsid:biocol.org:col:34778" name="Royal Ontario Museum" pageId="6" pageNumber="23" type="Museum">ROM 30099</specimenCode>
. The anterior part of the mandible is dorsoventrally slenderer than in all known specimens of
<taxonomicName id="63F54D12FF8EFFCBFE82F8A434C7CA12" baseAuthorityName="Laurin" baseAuthorityYear="1993" box="[270,511,1866,1889]" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="6" pageNumber="23" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF8EFFCBFE82F8A434C7CA12" box="[270,511,1866,1889]" italics="true" pageId="6" pageNumber="23">Haptodus garnettensis</emphasis>
</taxonomicName>
, including the youngest known mandible in
<specimenCode id="F4539EEAFF8EFFCBFEDFF88434E5CAF2" box="[339,477,1898,1921]" collectionCode="ROM" country="Canada" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34778" lsid="urn:lsid:biocol.org:col:34778" name="Royal Ontario Museum" pageId="6" pageNumber="23" type="Museum">ROM 29872</specimenCode>
. This juvenile is of similar size to the
<taxonomicName id="63F54D12FF8EFFCBFC25FA45312EC8B1" authorityName="Kissel &amp; Reisz" authorityYear="2004" box="[937,1046,1451,1474]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="6" pageNumber="23" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF8EFFCBFC25FA45312EC8B1" box="[937,1046,1451,1474]" italics="true" pageId="6" pageNumber="23">Ianthodon</emphasis>
</taxonomicName>
<typeStatus id="7B4E8833FF8EFFCBFB92FA453143C8B1" box="[1054,1147,1451,1474]" pageId="6" pageNumber="23" type="holotype">holotype</typeStatus>
but is much more robust and shows an initial dorsoventral thickening of the symphyseal area, a region that is strong in the largest specimen
<specimenCode id="F4539EEAFF8EFFCBFAF6FA043641CB52" collectionCode="ROM" country="Canada" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34778" lsid="urn:lsid:biocol.org:col:34778" name="Royal Ontario Museum" pageId="6" pageNumber="23" type="Museum">ROM 43604</specimenCode>
.
</paragraph>
<paragraph id="A44A3691FF8EFFCAFCC2F9C43585CCB5" blockId="6.[819,1463,1451,1920]" lastBlockId="7.[124,767,176,1921]" lastPageId="7" lastPageNumber="24" pageId="6" pageNumber="23">
The dentary is very slender in dorsoventral view, with a thickened alveolar shelf that occupies almost half the height on the lingual side. The slight concavity of the tooth margin corresponds to the moderate convexity of the maxilla. There are 21 teeth preserved, with one certain gap for an unpreserved and maybe enlarged third tooth near the tip of the left dentary. Applying the pattern of the tooth density seen in the middle region, there might be even more positions, for a possible maximum of 25. The tooth bases are broad, but often appear to overlap each other, partly due to shape and partly due to compaction. The tooth morphology is readily distinguishable from that of
<taxonomicName id="63F54D12FF8FFFCAFE3DFF5E379BCDB4" baseAuthorityName="Laurin" baseAuthorityYear="1993" box="[433,675,176,199]" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="24" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF8FFFCAFE3DFF5E379BCDB4" box="[433,675,176,199]" italics="true" pageId="7" pageNumber="24">Haptodus garnettensis</emphasis>
</taxonomicName>
both by size of the base and the narrowness of the tip (see
<bibRefCitation id="C0644B60FF8FFFCAFD3DFF3E358CCC74" author="Laurin, M." pageId="7" pageNumber="24" pagination="200 - 229" refId="ref10031" refString="Laurin, M.: Anatomy and Relationships of Haptodus garnettensis, a Pennsylvanian synapsid from Kansas, J. Vertebr. Paleontol., 13, 200 - 229, 1993." type="journal article" year="1993">Laurin, 1993</bibRefCitation>
,
<figureCitation id="3CCE2A14FF8FFFCAFF4EFF1E342ACC74" box="[194,274,240,263]" captionStart="Figure 4" captionStartId="5.[124,191,815,836]" captionTargetBox="[377,1208,177,783]" captionTargetId="figure-9@5.[377,1208,177,783]" captionTargetPageId="5" captionText="Figure 4. Ianthodon schultzei. (a) Referred left maxilla KUVP 133736; (b) referred right maxilla FO 176." figureDoi="http://doi.org/10.5281/zenodo.11045477" httpUri="https://zenodo.org/record/11045477/files/figure.png" pageId="7" pageNumber="24">Fig. 4b</figureCitation>
) as well as by the overall delicate morphology of the crown. The level of compaction of the teeth and the overall reduced thickness of the dentine suggest that the teeth in
<taxonomicName id="63F54D12FF8FFFCAFF5DFEBE3406CC14" authorityName="Kissel &amp; Reisz" authorityYear="2004" box="[209,318,336,359]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="24" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF8FFFCAFF5DFEBE3406CC14" box="[209,318,336,359]" italics="true" pageId="7" pageNumber="24">Ianthodon</emphasis>
</taxonomicName>
reflect trophic specializations that are different from
<emphasis id="9681EA83FF8FFFCAFF75FE9E3450CCF4" box="[249,360,368,391]" italics="true" pageId="7" pageNumber="24">
<taxonomicName id="63F54D12FF8FFFCAFF75FE9E345CCCF4" authorityName="Gaudry" authorityYear="1886" box="[249,356,368,391]" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="24" phylum="Chordata" rank="genus">Haptodus</taxonomicName>
.
</emphasis>
The teeth in the posterior region of the dentary increase in broadness as they become smaller posteriorly.
</paragraph>
<paragraph id="A44A3691FF8FFFCAFF1BFE2137C9CFF6" blockId="7.[124,767,176,1921]" pageId="7" pageNumber="24">Dislocated from the articulated mandibles, there is a single splenial next to the premaxillae. Both the anterior and posterior tips are delicate and poorly preserved. The dorsoventral extension is high, suggesting that the splenial had a thin exposure in lateral aspect. Its posterior end has a slight dorsal expansion, possibly contributing to the coronoid eminence.</paragraph>
<paragraph id="A44A3691FF8FFFCAFF1BFD603714CEB7" blockId="7.[124,767,176,1921]" pageId="7" pageNumber="24">
The angular of
<taxonomicName id="63F54D12FF8FFFCAFEC9FD613721CFD5" authorityName="Kissel and Reisz" authorityYear="2004" box="[325,537,655,678]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="24" phylum="Chordata" rank="species" species="schultzei">
<emphasis id="9681EA83FF8FFFCAFEC9FD613721CFD5" box="[325,537,655,678]" italics="true" pageId="7" pageNumber="24">Ianthodon schultzei</emphasis>
</taxonomicName>
is slender and obviously not as tall or as massive as in
<taxonomicName id="63F54D12FF8FFFCAFD8AFD4037C0CFB6" baseAuthorityName="Laurin" baseAuthorityYear="1993" box="[518,760,686,709]" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="24" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF8FFFCAFD8AFD4037C0CFB6" box="[518,760,686,709]" italics="true" pageId="7" pageNumber="24">Haptodus garnettensis</emphasis>
</taxonomicName>
, especially relative to the surangular height; it has a flat ventral lamina that does not extend as far below the prearticular level as in
<taxonomicName id="63F54D12FF8FFFCAFF63FCE034D8CE56" baseAuthorityName="Laurin" baseAuthorityYear="1993" box="[239,480,782,805]" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="24" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF8FFFCAFF63FCE034D8CE56" box="[239,480,782,805]" italics="true" pageId="7" pageNumber="24">Haptodus garnettensis</emphasis>
</taxonomicName>
(compare to
<bibRefCitation id="C0644B60FF8FFFCAFDE2FCE037C3CE56" author="Laurin, M." box="[622,763,782,805]" pageId="7" pageNumber="24" pagination="200 - 229" refId="ref10031" refString="Laurin, M.: Anatomy and Relationships of Haptodus garnettensis, a Pennsylvanian synapsid from Kansas, J. Vertebr. Paleontol., 13, 200 - 229, 1993." type="journal article" year="1993">Laurin, 1993</bibRefCitation>
, Fig. 10). Both genera share the gently convex ventral edge of the angular, which is thin in cross section but not developed as a reflected lamina. Its ventral edge is marked by slight rugosities. On the medial side, the angular has a strong longitudinal bar for the prearticular contact.
</paragraph>
<paragraph id="A44A3691FF8FFFCAFF1BFC2335EFC870" blockId="7.[124,767,176,1921]" pageId="7" pageNumber="24">
The right surangular is exposed medially, showing the posterior articulation for the articular and the anterodorsal groove to host the dentary. A gently developed ridge runs along its dorsal edge where an aponeurosis would have attached for the adductor muscles. The posterior border of the adductor fossa is marked by a small triangular process for the anterodorsal process of the articular. There are no clear differences from the surangular of
<taxonomicName id="63F54D12FF8FFFCAFE41FB423785C9B0" baseAuthorityName="Laurin" baseAuthorityYear="1993" box="[461,701,1196,1219]" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="24" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF8FFFCAFE41FB423785C9B0" box="[461,701,1196,1219]" italics="true" pageId="7" pageNumber="24">Haptodus garnettensis</emphasis>
</taxonomicName>
, apart from being slightly narrower dorsoventrally (
<bibRefCitation id="C0644B60FF8FFFCAFDE6FB2237C3C990" author="Laurin, M." box="[618,763,1228,1251]" pageId="7" pageNumber="24" pagination="200 - 229" refId="ref10031" refString="Laurin, M.: Anatomy and Relationships of Haptodus garnettensis, a Pennsylvanian synapsid from Kansas, J. Vertebr. Paleontol., 13, 200 - 229, 1993." type="journal article" year="1993">Laurin, 1993</bibRefCitation>
, Fig. 10).
</paragraph>
<paragraph id="A44A3691FF8FFFCAFF1BFAE237E4C8B1" blockId="7.[124,767,176,1921]" pageId="7" pageNumber="24">The posterior coronoid is a slender, elongate element with a narrow anterior process and a bifurcate posterior region. The body of the left posterior coronoid, exposed in medial view, is covered by more than 20 small teeth. Another denticulate coronoid element is preserved, but it is largely covered by other elements, making precise identification difficult.</paragraph>
<paragraph id="A44A3691FF8FFFCAFF1BFA2536C0CCF5" blockId="7.[124,767,176,1921]" lastBlockId="7.[819,1462,176,390]" pageId="7" pageNumber="24">
Both articulars are preserved, one in articulation with the rest of the left mandible, the other isolated and displaced in front of the right dentary. No detailed observations on the presence of a pterygoideus or a retroarticular process can be made. There are extremely elongate, slender prearticulars preserved in place on the left mandible and separately from the right mandible, the latter displaced posteriorly to lie under the humerus of
<taxonomicName id="63F54D12FF8FFFCAFED9F9453739CBB1" authorityName="Lane" authorityYear="1945" box="[341,513,1707,1730]" family="Petrolacosauridae" genus="Petrolacosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="24" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF8FFFCAFED9F9453739CBB1" box="[341,513,1707,1730]" italics="true" pageId="7" pageNumber="24">Petrolacosaurus</emphasis>
</taxonomicName>
and a rib. Its posterior process is slightly bifurcate. Both seem untwisted, contrasting with all edaphosaurids and basal sphenacodonts. Based on our experience with the compaction properties of bones in the Garnett fossil beds, we can determine that this is not a diagenetic artifact. Unfortunately, the juvenile
<taxonomicName id="63F54D12FF8FFFCAFD1BF8A435C6CAF2" baseAuthorityName="Laurin" baseAuthorityYear="1993" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="24" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF8FFFCAFD1BF8A435C6CAF2" italics="true" pageId="7" pageNumber="24">Haptodus garnettensis</emphasis>
</taxonomicName>
specimen
<collectionCode id="C2E4AE54FF8FFFCAFEFCF8843493CAF2" box="[368,427,1898,1921]" country="Canada" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34778" lsid="urn:lsid:biocol.org:col:34778" name="Royal Ontario Museum" pageId="7" pageNumber="24" type="Museum">ROM</collectionCode>
29872 is not exposed for direct comparison, but the rather adult
<collectionCode id="C2E4AE54FF8FFFCAFB14FF5E31EBCDB4" box="[1176,1235,176,199]" country="Canada" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34778" lsid="urn:lsid:biocol.org:col:34778" name="Royal Ontario Museum" pageId="7" pageNumber="24" type="Museum">ROM</collectionCode>
30099 has a twisted prearticular. Thus, the
<taxonomicName id="63F54D12FF8FFFCAFBAFFF3E31A8CD94" authorityName="Kissel &amp; Reisz" authorityYear="2004" box="[1059,1168,208,231]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="24" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF8FFFCAFBAFFF3E31A8CD94" box="[1059,1168,208,231]" italics="true" pageId="7" pageNumber="24">Ianthodon</emphasis>
</taxonomicName>
prearticular appears to preserve the plesiomorphic condition. Along with the articular, the main body of the prearticular is strongly built, and, despite its juvenile stage, the articular is well ossified and suggests that it had large quadrate condyles, although not preserved on the slab.
</paragraph>
<paragraph id="A44A3691FF8FFFCAFCBFFE41315FCCB4" blockId="7.[819,1127,431,455]" box="[819,1127,431,455]" pageId="7" pageNumber="24">
<emphasis id="9681EA83FF8FFFCAFCBFFE41315FCCB4" bold="true" box="[819,1127,431,455]" pageId="7" pageNumber="24">2.1.3 Postcranial skeleton</emphasis>
</paragraph>
<paragraph id="A44A3691FF8FFFCAFCBFFE0130AACE97" blockId="7.[819,1463,495,1921]" pageId="7" pageNumber="24">
Several elements of the postcranial axial skeleton, mostly from the dorsal region, have been recovered (
<figureCitation id="3CCE2A14FF8FFFCAFA9DFDE1306ACF55" box="[1297,1362,527,550]" captionStart="Figure 6" captionStartId="8.[124,191,1123,1144]" captionTargetBox="[384,1208,182,1091]" captionTargetId="figure-9@8.[377,1208,177,1092]" captionTargetPageId="8" captionText="Figure 6. Ianthodon schultzei cranial and skeletal reconstruction. Three-dimensional arrangement and projections based on a wax maquette. Skull in dorsal, ventral and lateral view; mandible in lateral and medial view." figureDoi="http://doi.org/10.5281/zenodo.11045481" httpUri="https://zenodo.org/record/11045481/files/figure.png" pageId="7" pageNumber="24">Fig. 6</figureCitation>
). The average centrum length is about
<quantity id="630D9B74FF8FFFCAFBFEFDC13186CF35" box="[1138,1214,559,582]" metricMagnitude="-2" metricUnit="m" metricValue="1.0" pageId="7" pageNumber="24" unit="mm" value="10.0">10 mm</quantity>
. A blunt ridge seems to be present on the ventral side of each centrum. Dorsal neural arches measure approximately
<quantity id="630D9B74FF8FFFCAFBF1FD8031FDCFF6" box="[1149,1221,622,645]" metricMagnitude="-2" metricUnit="m" metricValue="2.4" pageId="7" pageNumber="24" unit="mm" value="24.0">24 mm</quantity>
from articulation with the centrum to the dorsal edge of the spine. All arches are delicate, but do not differ significantly from those of
<taxonomicName id="63F54D12FF8FFFCAFAE7FD4036DBCF96" baseAuthorityName="Laurin" baseAuthorityYear="1993" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="24" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF8FFFCAFAE7FD4036DBCF96" italics="true" pageId="7" pageNumber="24">Haptodus garnettensis</emphasis>
</taxonomicName>
. Laterally, they are not, or only shallowly, excavated; this also depends on the level of compression. The zygapophyses are short. Unlike in
<taxonomicName id="63F54D12FF8FFFCAFB54FCE03677CE36" baseAuthorityName="Laurin" baseAuthorityYear="1993" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="24" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF8FFFCAFB54FCE03677CE36" italics="true" pageId="7" pageNumber="24">Haptodus garnettensis</emphasis>
</taxonomicName>
, the postzygapohyses are widely spaced and single, not showing the broad double-lobed plate as in
<collectionCode id="C2E4AE54FF8FFFCAFA9DFCA03074CE16" box="[1297,1356,846,869]" country="Canada" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34778" lsid="urn:lsid:biocol.org:col:34778" name="Royal Ontario Museum" pageId="7" pageNumber="24" type="Museum">ROM</collectionCode>
43604 or
<collectionCode id="C2E4AE54FF8FFFCAFCBFFC833656CEF7" box="[819,878,877,900]" country="Canada" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34778" lsid="urn:lsid:biocol.org:col:34778" name="Royal Ontario Museum" pageId="7" pageNumber="24" type="Museum">ROM</collectionCode>
29872. All diapophyses are moderate lateral extensions and have sub-rounded cross sections. The diapophysial laminae are unreduced, connecting the central edge of the neural arch to the very tip of the diapophysis by a convex blade.
</paragraph>
<paragraph id="A44A3691FF8FFFCAFCC2FC033054C9F0" blockId="7.[819,1463,495,1921]" pageId="7" pageNumber="24">
Relatively large intercentra appear to be present between the supraoccipital and one of the posterior coronoids, indicating that
<taxonomicName id="63F54D12FF8FFFCAFC06FBC336CFC937" authorityName="Kissel &amp; Reisz" authorityYear="2004" box="[906,1015,1069,1092]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="24" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF8FFFCAFC06FBC336CFC937" box="[906,1015,1069,1092]" italics="true" pageId="7" pageNumber="24">Ianthodon</emphasis>
</taxonomicName>
has the plesiomorphic tetrapod condition, in contrast to the tendency of reduction of these vertebral elements in advanced pelycosaur-grade sphenacodonts.
</paragraph>
<paragraph id="A44A3691FF8FFFCAFCC2FB623079C850" blockId="7.[819,1463,495,1921]" pageId="7" pageNumber="24">No elements posterior to the middle trunk are known, except for a single neural arch with a shortened spine that might belong to the proximal caudal series. The anterior zygapophyses are shortened compared to average dorsal vertebrae, and the spine is half as tall as in the dorsals.</paragraph>
<paragraph id="A44A3691FF8FFFCAFCC2FAC23007CB31" blockId="7.[819,1463,495,1921]" pageId="7" pageNumber="24">All dorsal ribs are long and weakly curved, with the strongest flexion in the proximal portion. At least one cervical rib is identified with confidence, bearing a straight, stout shaft that flares distally. An anterior process is present on the cervical rib, presumably a plesiomorphic character. In dorsal ribs, the tuberculum is prominent, but not separated from the capitulum by a notch. Like in other basal synapsids, a lamina is present in the dichocephalous rib head, allowing no aperture between the diapophysial lamina and the rib.</paragraph>
<paragraph id="A44A3691FF8FFFC5FCC2F9A534BDC87D" blockId="7.[819,1463,495,1921]" lastBlockId="8.[124,767,1239,1900]" lastPageId="8" lastPageNumber="25" pageId="7" pageNumber="24">
Between the cranial elements described by
<bibRefCitation id="C0644B60FF8FFFCAFACCF9A5368ECBF1" author="Kissel, R. A. &amp; Reisz, R. R." pageId="7" pageNumber="24" pagination="409 - 428" refId="ref9945" refString="Kissel, R. A. and Reisz, R. R.: Synapsid fauna of the Upper Pennsylvanian Rock Lake Shale near Garnett, Kansas and the diversity pattern of early amniotes, in: Recent Advances in the Origin and Early Radiation of Vertebrates, edited by: Arratia, G., Wilson, M. V. H., and Cloutier, R., Verlag Dr. F. Pfeil, Munchen, 409 - 428, 2004." type="book chapter" year="2004">Kissel and Reisz (2004</bibRefCitation>
,
<figureCitation id="3CCE2A14FF8FFFCAFC4CF9853130CBF1" box="[960,1032,1643,1666]" captionStart="Figure 2" captionStartId="3.[124,191,1147,1168]" captionTargetBox="[133,756,179,1112]" captionTargetId="figure-9@3.[132,759,177,1115]" captionTargetPageId="3" captionText="Figure 2. Ianthodon schultzei holotype KUVP 133735, slab in present condition, combined with photograph of skull area (lower left) before its removal." figureDoi="http://doi.org/10.5281/zenodo.11045473" httpUri="https://zenodo.org/record/11045473/files/figure.png" pageId="7" pageNumber="24">Figs. 2</figureCitation>
and
<figureCitation id="3CCE2A14FF8FFFCAFBB0F9853172CBF1" box="[1084,1098,1643,1666]" captionStart="Figure 3" captionStartId="4.[124,191,1359,1380]" captionTargetBox="[304,1282,177,1325]" captionTargetId="figure-9@4.[302,1285,177,1328]" captionTargetPageId="4" captionText="Figure 3. Ianthodon schultzei holotype KUVP 133735. (a) Redocumented skull; (b) dislocated premaxillae. Fr – frontal; j – jugal; l – lacrimal; m – maxilla; p – parietal; po – postorbital; pof – postfrontal; pp – postparietal; prf – prefrontal; sq – squamosal; st – supratemporal; t – tabular." figureDoi="http://doi.org/10.5281/zenodo.11045475" httpUri="https://zenodo.org/record/11045475/files/figure.png" pageId="7" pageNumber="24">3</figureCitation>
), there is one labeled with a question mark. Given the postcranial proportions, it matches the shape and size one would expect for the cleithrum. A flat and broadened dorsal tip instantly turns into a rounder cross section, which is again gradually flattened towards its presumed contact with the clavicle. Similar cleithra are present in
<taxonomicName id="63F54D12FF8FFFCAFCDDF8C436D3CA32" authorityName="Cope" authorityYear="1882" box="[849,1003,1834,1857]" family="Edaphosauridae" genus="Edaphosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="24" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF8FFFCAFCDDF8C436D3CA32" box="[849,1003,1834,1857]" italics="true" pageId="7" pageNumber="24">Edaphosaurus</emphasis>
</taxonomicName>
and
<taxonomicName id="63F54D12FF8FFFCAFBAFF8C4319ACA32" authorityName="Cope" authorityYear="1878" box="[1059,1186,1834,1857]" family="Sphenacodontidae" genus="Dimetrodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="24" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF8FFFCAFBAFF8C4319ACA32" box="[1059,1186,1834,1857]" italics="true" pageId="7" pageNumber="24">Dimetrodon</emphasis>
</taxonomicName>
(
<bibRefCitation id="C0644B60FF8FFFCAFB3EF8C4308ACA32" author="Romer, A. S. &amp; Price, L. I." box="[1202,1458,1834,1857]" pageId="7" pageNumber="24" pagination="1 - 538" refId="ref10605" refString="Romer, A. S. and Price, L. I.: Review of the Pelycosauria, Geol. S. Am. S., 28, 1 - 538, 1940." type="journal article" year="1940">Romer and Price, 1940</bibRefCitation>
, pl. 28 B and 38 C) as well as in
<taxonomicName id="63F54D12FF8FFFCAFBF5F8A43028CA12" authorityName="Huene" authorityYear="1925" box="[1145,1296,1866,1889]" genus="Pantelosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="24" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF8FFFCAFBF5F8A43028CA12" box="[1145,1296,1866,1889]" italics="true" pageId="7" pageNumber="24">Pantelosaurus</emphasis>
</taxonomicName>
. It most closely resembles the cleithrum of an undescribed sphenacodont from Garnett, as well as that of the
<typeStatus id="7B4E8833FF80FFC5FD9CFB393755C99D" box="[528,621,1239,1262]" pageId="8" pageNumber="25" type="holotype">holotype</typeStatus>
of
<taxonomicName id="63F54D12FF80FFC5FD1BFB363415C87D" authority="RM" authorityName="RM" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="25" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF80FFC5FD1BFB3635C6C87C" italics="true" pageId="8" pageNumber="25">Haptodus garnettensis</emphasis>
RM
</taxonomicName>
14,156.
</paragraph>
<caption id="F08A6619FF80FFC5FFF0FB8D3664C9E5" ID-DOI="http://doi.org/10.5281/zenodo.11045481" ID-Zenodo-Dep="11045481" httpUri="https://zenodo.org/record/11045481/files/figure.png" pageId="8" pageNumber="25" startId="8.[124,191,1123,1144]" targetBox="[384,1208,182,1091]" targetPageId="8" targetType="figure">
<paragraph id="A44A3691FF80FFC5FFF0FB8D3664C9E5" blockId="8.[124,1463,1123,1174]" pageId="8" pageNumber="25">
<emphasis id="9681EA83FF80FFC5FFF0FB8D35EFC90B" bold="true" box="[124,215,1123,1144]" pageId="8" pageNumber="25">Figure 6.</emphasis>
<taxonomicName id="63F54D12FF80FFC5FF51FB8A34A0C90A" authorityName="Kissel and Reisz" authorityYear="2004" box="[221,408,1124,1145]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="25" phylum="Chordata" rank="species" species="schultzei">
<emphasis id="9681EA83FF80FFC5FF51FB8A34A0C90A" box="[221,408,1124,1145]" italics="true" pageId="8" pageNumber="25">Ianthodon schultzei</emphasis>
</taxonomicName>
cranial and skeletal reconstruction. Three-dimensional arrangement and projections based on a wax maquette. Skull in dorsal, ventral and lateral view; mandible in lateral and medial view.
</paragraph>
</caption>
<paragraph id="A44A3691FF80FFC5FF1BFAF93058C8BD" blockId="8.[124,767,1239,1900]" lastBlockId="8.[819,1463,1239,1900]" pageId="8" pageNumber="25">
The replacement bones of the pectoral girdle are delicately constructed and remain unfused in this juvenile specimen. The scapula is
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tall and
<quantity id="630D9B74FF80FFC5FD90FAB9375CC81D" box="[540,612,1367,1390]" metricMagnitude="-2" metricUnit="m" metricValue="1.9" pageId="8" pageNumber="25" unit="mm" value="19.0">19 mm</quantity>
wide in midlongitudinal extension. The bone is well-ossified, as is typical for even very young individuals in basal amniotes. As seen in
<taxonomicName id="63F54D12FF80FFC5FF17FA593400C8BD" authorityName="Credner" authorityYear="1888" box="[155,312,1463,1486]" genus="Palaeohatteria" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="25" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF80FFC5FF17FA593400C8BD" box="[155,312,1463,1486]" italics="true" pageId="8" pageNumber="25">Palaeohatteria</emphasis>
</taxonomicName>
, the scapula and anterior coracoid ossify earlier in ontogeny than the posterior coracoid, possibly to support the glenoid fossa. The ontogenetic stage of the
<typeStatus id="7B4E8833FF80FFC5FD4AFA183592CB5E" pageId="8" pageNumber="25" type="holotype">holotype</typeStatus>
of
<taxonomicName id="63F54D12FF80FFC5FF41F9F834A6CB5E" authorityName="Kissel and Reisz" authorityYear="2004" box="[205,414,1558,1581]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="25" phylum="Chordata" rank="species" species="schultzei">
<emphasis id="9681EA83FF80FFC5FF41F9F834A6CB5E" box="[205,414,1558,1581]" italics="true" pageId="8" pageNumber="25">Ianthodon schultzei</emphasis>
</taxonomicName>
is between the average condition found in
<taxonomicName id="63F54D12FF80FFC5FF53F9D83729CB3E" baseAuthorityName="Credner" baseAuthorityYear="1888" box="[223,529,1590,1613]" genus="Palaeohatteria" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="25" phylum="Chordata" rank="species" species="longicaudata">
<emphasis id="9681EA83FF80FFC5FF53F9D83729CB3E" box="[223,529,1590,1613]" italics="true" pageId="8" pageNumber="25">Palaeohatteria longicaudata</emphasis>
</taxonomicName>
and that of
<taxonomicName id="63F54D12FF80FFC5FD1BF9D835C6CB1E" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="25" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF80FFC5FD1BF9D835C6CB1E" italics="true" pageId="8" pageNumber="25">Haptodus garnettensis</emphasis>
</taxonomicName>
, as the scapula and anterior coracoid are still unfused, but the ventral margin of the scapula is fully differentiated and is not as simple as the dorsal end like in
<taxonomicName id="63F54D12FF80FFC5FD07F9783596CBBE" authorityName="Credner" authorityYear="1888" genus="Palaeohatteria" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="25" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF80FFC5FD07F9783596CBBE" italics="true" pageId="8" pageNumber="25">Palaeohatteria</emphasis>
</taxonomicName>
juveniles. The lateral foramen (supraglenoid foramen) is located above the glenoid fossa, at about one quarter of the total scapular height, and slightly posterior to the edge of the supraglenoid ridge. There is no indication of a notch interrupting the proximal anterior margin. In
<taxonomicName id="63F54D12FF80FFC5FDA7F8DB37A7CA3F" authorityName="Kissel and Reisz" authorityYear="2004" box="[555,671,1845,1868]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="25" phylum="Chordata" rank="species" species="schultzei">
<emphasis id="9681EA83FF80FFC5FDA7F8DB37A7CA3F" box="[555,671,1845,1868]" italics="true" pageId="8" pageNumber="25">I. schultzei</emphasis>
</taxonomicName>
, the dorsal end of the scapula flares anteroposteriorly, in strong contrast to the condition in
<taxonomicName id="63F54D12FF80FFC5FBB6FB363014C99C" box="[1082,1324,1240,1263]" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="25" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF80FFC5FBB6FB363014C99C" box="[1082,1324,1240,1263]" italics="true" pageId="8" pageNumber="25">Haptodus garnettensis</emphasis>
</taxonomicName>
(compare to
<bibRefCitation id="C0644B60FF80FFC5FCBFFB193686C87D" author="Laurin, M." box="[819,958,1271,1294]" pageId="8" pageNumber="25" pagination="200 - 229" refId="ref10031" refString="Laurin, M.: Anatomy and Relationships of Haptodus garnettensis, a Pennsylvanian synapsid from Kansas, J. Vertebr. Paleontol., 13, 200 - 229, 1993." type="journal article" year="1993">Laurin, 1993</bibRefCitation>
, Fig. 19). Unfortunately, the distal portion of the juvenile
<typeStatus id="7B4E8833FF80FFC5FC02FAF936D3C85D" box="[910,1003,1303,1326]" pageId="8" pageNumber="25" type="holotype">holotype</typeStatus>
of
<taxonomicName id="63F54D12FF80FFC5FB80FAF9318AC85D" box="[1036,1202,1303,1327]" family="Haptodontidae" genus="Haptodus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="25" phylum="Chordata" rank="species" species="garnettensis">
<emphasis id="9681EA83FF80FFC5FB80FAF93119C85C" box="[1036,1057,1303,1327]" italics="true" pageId="8" pageNumber="25">H</emphasis>
.
<emphasis id="9681EA83FF80FFC5FBBCFAF9318AC85D" box="[1072,1202,1303,1326]" italics="true" pageId="8" pageNumber="25">garnettensis</emphasis>
</taxonomicName>
is not preserved (
<bibRefCitation id="C0644B60FF80FFC5FAE6FAF93653C83D" author="Currie, P. J." pageId="8" pageNumber="25" pagination="927 - 942" refId="ref9556" refString="Currie, P. J.: A new haptodontine sphenacodont (Reptilia: Pelycosauria) from the Upper Pennsylvanian of North America, J. Paleontol., 51, 927 - 942, 1977." type="journal article" year="1977">Currie, 1977</bibRefCitation>
, and pers. obs. by F. Spindler). No basal sphenacodontian shows a distally flaring scapula in early ontogeny that is compensated for by further ossification to form a more rectangular outline. Therefore, we assume that the flared scapular blade is an age-independent character of
<taxonomicName id="63F54D12FF80FFC5FB6BFA593062C8BD" authorityName="Kissel and Reisz" authorityYear="2004" box="[1255,1370,1463,1486]" genus="Ianthodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="25" phylum="Chordata" rank="species" species="schultzei">
<emphasis id="9681EA83FF80FFC5FB6BFA593062C8BD" box="[1255,1370,1463,1486]" italics="true" pageId="8" pageNumber="25">I. schultzei</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="A44A3691FF80FFC5FCC2FA3836A6CB5E" blockId="8.[819,1463,1239,1900]" pageId="8" pageNumber="25">
The anterior coracoid is a simple oval disc with delicate radiating texture, measuring 32 × 21.
<quantity id="630D9B74FF80FFC5FB32FA1831C0CB7E" box="[1214,1272,1526,1549]" metricMagnitude="-3" metricUnit="m" metricValue="5.0" pageId="8" pageNumber="25" unit="mm" value="5.0">5 mm</quantity>
along the orthogonal axes.
</paragraph>
<paragraph id="A44A3691FF80FFC4FCC2F9D83478CC95" blockId="8.[819,1463,1239,1900]" lastBlockId="9.[124,767,176,486]" lastPageId="9" lastPageNumber="26" pageId="8" pageNumber="25">
A left humerus is the only known appendicular element. It measures
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in length,
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in mid-diaphysial width and
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in width at the distal epiphysis. As the specimen is a juvenile, it resembles juvenile specimens of
<taxonomicName id="63F54D12FF80FFC5FACEF978365DCBBE" authorityName="Credner" authorityYear="1888" genus="Palaeohatteria" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="25" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF80FFC5FACEF978365DCBBE" italics="true" pageId="8" pageNumber="25">Palaeohatteria</emphasis>
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. Its proximal head is strongly distorted by crushing, revealing the broken edge of the tubercle for the M. latissimus dorsi and the beginning of the articulation area. The distal end is relatively well-preserved. In typical sphenacodontian fashion, the distal dorsal surface of the bone has an elongate groove on the entepicondyle for the entepicondylar foramen. The ectepidondylar ridge on the dorsal surface of the bone is modestly developed. The supinator process, as in
<taxonomicName id="63F54D12FF81FFC4FD07FF3E3596CC74" authorityName="Credner" authorityYear="1888" genus="Palaeohatteria" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="9" pageNumber="26" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF81FFC4FD07FF3E3596CC74" italics="true" pageId="9" pageNumber="26">Palaeohatteria</emphasis>
</taxonomicName>
and other basal sphenacodontians, is a blade-like structure that extends distally without flaring significantly anteriorly. In effect, its anterior edge is nearly parallel to that of the ectepicondylar ridge. In overall shape, it resembles
<taxonomicName id="63F54D12FF81FFC4FFF0FE9E3460CCF4" authorityName="Romer" authorityYear="1937" box="[124,344,368,391]" family="Sphenacodontidae" genus="Dimetrodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="kempae">
<emphasis id="9681EA83FF81FFC4FFF0FE9E3460CCF4" box="[124,344,368,391]" italics="true" pageId="9" pageNumber="26">Dimetrodon kempae</emphasis>
</taxonomicName>
,
<taxonomicName id="63F54D12FF81FFC4FEE6FE9E34CBCCF4" authorityName="Romer" authorityYear="1937" box="[362,499,368,391]" genus="Lupeosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="9" pageNumber="26" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF81FFC4FEE6FE9E34CBCCF4" box="[362,499,368,391]" italics="true" pageId="9" pageNumber="26">Lupeosaurus</emphasis>
</taxonomicName>
and
<taxonomicName id="63F54D12FF81FFC4FDBDFE9E374BCCF4" box="[561,627,368,391]" family="Caseidae" genus="Casea" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="9" pageNumber="26" phylum="Chordata" rank="genus">
<emphasis id="9681EA83FF81FFC4FDBDFE9E374BCCF4" box="[561,627,368,391]" italics="true" pageId="9" pageNumber="26">Casea</emphasis>
</taxonomicName>
(see
<bibRefCitation id="C0644B60FF81FFC4FD3AFE81341FCCD5" author="Romer, A. S. &amp; Price, L. I." pageId="9" pageNumber="26" pagination="1 - 538" refId="ref10605" refString="Romer, A. S. and Price, L. I.: Review of the Pelycosauria, Geol. S. Am. S., 28, 1 - 538, 1940." type="journal article" year="1940">Romer and Price, 1940</bibRefCitation>
, Figs. 31 and 32). There is no evidence that the distal end of the supinator process enclosed an ectepicondylar foramen.
</paragraph>
</subSubSection>
</treatment>
</document>