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<document id="CB0F7E2E61CEB9FB56AEF8746EA018BA" ID-DOI="10.1093/zoolinnean/zlad035" ID-ISSN="0024-4082" ID-Zenodo-Dep="8426147" IM.bibliography_approvedBy="valdenar" IM.illustrations_approvedBy="felipe" IM.materialsCitations_approvedBy="valdenar" IM.metadata_approvedBy="felipe" IM.tables_approvedBy="valdenar" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="valdenar" checkinTime="1696940290583" checkinUser="plazi" docAuthor="Spiekman, Stephan N. F." docDate="2023" docId="03A2222F31119F0B42B4FD833975FBE1" docLanguage="en" docName="zlad035.pdf" docOrigin="Zoological Journal of the Linnean Society 199 (2)" docSource="http://dx.doi.org/10.1093/zoolinnean/zlad035" docStyle="DocumentStyle:4F230B9370E98E256D973D6DFB57F36C.1:ZoolJLinnSoc.2023-.journal_article" docStyleId="4F230B9370E98E256D973D6DFB57F36C" docStyleName="ZoolJLinnSoc.2023-.journal_article" docStyleVersion="1" docTitle="Saltoposuchus connectens Huene 1921" docType="treatment" docVersion="3" lastPageNumber="385" masterDocId="FF9B5A57311D9F144118FF80380CFFD5" masterDocTitle="A revision and histological investigation of Saltoposuchus connectens (Archosauria: Crocodylomorpha) from the Norian (Late Triassic) of south-western Germany" masterLastPageNumber="391" masterPageNumber="354" pageNumber="366" updateTime="1697046361159" updateUser="valdenar">
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<mods:title id="CA9EFB17ACAD0470ECECEE14C35DEFD4">A revision and histological investigation of Saltoposuchus connectens (Archosauria: Crocodylomorpha) from the Norian (Late Triassic) of south-western Germany</mods:title>
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<mods:name id="D7E6F6A0CE5753B10C6FB053256559E4" type="personal">
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<mods:namePart id="10D1E7C650A6FD6EE9945CAA79923B59">Spiekman, Stephan N. F.</mods:namePart>
<mods:affiliation id="AC427FD5EEC62D3DA1C2A4CC61830BB5">Staatliches Museum für Naturkunde Stuưgart, Rosenstein 1, 70191 Stuưgart, Germany</mods:affiliation>
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<mods:title id="576D677A2A63AA4317A9A71FCB92965C">Zoological Journal of the Linnean Society</mods:title>
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<mods:date id="AB3678684114C1156E19AC055A0BB18E">2023</mods:date>
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<mods:number id="801D6169BB466C897CDCE84954576061">2023-06-20</mods:number>
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<mods:number id="2C3D858DBE6C8F28251EA2160033BE38">199</mods:number>
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<mods:identifier id="259ED05BFF7F4E6ABD5A57980DFF15F4" type="DOI">10.1093/zoolinnean/zlad035</mods:identifier>
<mods:identifier id="A68D3DA2AEB6DD3D7D255476AB7D4520" type="ISSN">0024-4082</mods:identifier>
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<treatment id="03A2222F31119F0B42B4FD833975FBE1" LSID="urn:lsid:plazi:treatment:03A2222F31119F0B42B4FD833975FBE1" httpUri="http://treatment.plazi.org/id/03A2222F31119F0B42B4FD833975FBE1" lastPageId="31" lastPageNumber="385" pageId="12" pageNumber="366">
<subSubSection id="C311C0B231119F1842B4FD833D3DFDC8" box="[940,1329,515,542]" pageId="12" pageNumber="366" type="nomenclature">
<paragraph id="8BB4933931119F1842B4FD833D3DFDC8" blockId="12.[940,1329,515,542]" box="[940,1329,515,542]" pageId="12" pageNumber="366">
<heading id="D0FC245531119F1842B4FD833D3DFDC8" bold="true" box="[940,1329,515,542]" centered="true" fontSize="11" level="2" pageId="12" pageNumber="366" reason="2">
<taxonomicName id="4C0BE8BA31119F1842B4FD833D3DFDC8" authority="Huene 1921" authorityName="Huene" authorityYear="1921" box="[940,1329,515,542]" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="12" pageNumber="366" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B31119F1842B4FD833D3DFDC8" bold="true" box="[940,1329,515,542]" pageId="12" pageNumber="366">
<emphasis id="B97F4F2B31119F1842B4FD833CA5FDCB" bold="true" box="[940,1193,515,542]" italics="true" pageId="12" pageNumber="366">Saltoposuchus connectens</emphasis>
<bibRefCitation id="EF9AEEC831119F1845B6FD833D3DFDC8" author="Huene FR" box="[1198,1329,515,541]" pageId="12" pageNumber="366" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene 1921</bibRefCitation>
</emphasis>
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C311C0B231119F194232FDC4392DFCFA" lastPageId="13" lastPageNumber="367" pageId="12" pageNumber="366" type="diagnosis">
<paragraph id="8BB4933931119F184232FDC43B9AFAD8" blockId="12.[810,1459,580,1293]" pageId="12" pageNumber="366">
<emphasis id="B97F4F2B31119F184232FDC43B9CFD8E" box="[810,912,580,603]" italics="true" pageId="12" pageNumber="366">Diagnosis:</emphasis>
A medium-sized non-crocodyliform crocodylomorph (femoral length SMNS 12596:
<quantity id="4CF33EDC31119F18456BFDE33CC2FDAE" box="[1139,1230,611,635]" metricMagnitude="-1" metricUnit="m" metricValue="1.06" pageId="12" pageNumber="366" unit="mm" value="106.0">106 mm</quantity>
) distinguished by the following combination of character states (autapomorphies among non-crocodyliform crocodylomorphs are indicated with an asterix): enlarged anterior maxillary teeth (also present in
<taxonomicName id="4C0BE8BA31119F184232FD603BFBFD2D" box="[810,1015,736,760]" class="Reptilia" genus="Junggarsuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="12" pageNumber="366" phylum="Chordata" rank="species" species="sloani">
<emphasis id="B97F4F2B31119F184232FD603BFBFD2D" box="[810,1015,736,760]" italics="true" pageId="12" pageNumber="366">Junggarsuchus sloani</emphasis>
</taxonomicName>
); antorbital fossa well defined, forming a complete circumference around the antorbital fenestra (also present in CM 29894,
<taxonomicName id="4C0BE8BA31119F184515FC9F3CE6FCE2" authorityName="Clark &amp; Sues" authorityYear="2002" box="[1037,1258,799,823]" class="Reptilia" family="Sphenosuchidae" genus="Kayentasuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="12" pageNumber="366" phylum="Chordata" rank="species" species="walkeri">
<emphasis id="B97F4F2B31119F184515FC9F3CE6FCE2" box="[1037,1258,799,823]" italics="true" pageId="12" pageNumber="366">Kayentasuchus walkeri</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C0BE8BA31119F184407FC9F3B72FC83" class="Reptilia" family="Sphenosuchidae" genus="Dromicosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="12" pageNumber="366" phylum="Chordata" rank="species" species="grallator">
<emphasis id="B97F4F2B31119F184407FC9F3B72FC83" italics="true" pageId="12" pageNumber="366">Dromicosuchus grallator</emphasis>
</taxonomicName>
); antorbital fenestra that is approximately half the size of the orbit*; an oval fossa on the ventral surface of the palatine outlined by a radially striated surface medially*; a dorsally expanded anterior end of the dentary (also present in
<taxonomicName id="4C0BE8BA31119F184440FC1C3B66FC06" authorityName="Reig" authorityYear="1963" class="Polyplacophora" genus="Trialestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodilomorphi" pageId="12" pageNumber="366" phylum="Mollusca" rank="species" species="romeri">
<emphasis id="B97F4F2B31119F184440FC1C3B66FC06" italics="true" pageId="12" pageNumber="366">Trialestes romeri</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0BE8BA31119F1842B9FC3C3D02FC01" authority=", AMNH FARB" authorityName="AMNH FARB" box="[929,1294,956,980]" class="Reptilia" family="Sphenosuchidae" genus="Hesperosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="12" pageNumber="366" phylum="Chordata" rank="species" species="agilis">
<emphasis id="B97F4F2B31119F1842B9FC3C3C65FC01" box="[929,1129,956,980]" italics="true" pageId="12" pageNumber="366">Hesperosuchus agilis</emphasis>
, AMNH FARB
</taxonomicName>
6758); splenial slightly involved in the symphysis (also present in
<taxonomicName id="4C0BE8BA31119F18443AFC5B3B7EFBC7" authorityName="Clark &amp; Sues" authorityYear="2002" class="Reptilia" family="Sphenosuchidae" genus="Kayentasuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="12" pageNumber="366" phylum="Chordata" rank="species" species="walkeri">
<emphasis id="B97F4F2B31119F18443AFC5B3B7EFBC7" italics="true" pageId="12" pageNumber="366">Kayentasuchus walkeri</emphasis>
</taxonomicName>
); and a notch on the lateral side of the proximal head of the tibia*.
<taxonomicName id="4C0BE8BA31119F18428EFB993C87FBE4" authorityName="Huene" authorityYear="1921" box="[918,1163,1049,1073]" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="12" pageNumber="366" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B31119F18428EFB993C87FBE4" box="[918,1163,1049,1073]" italics="true" pageId="12" pageNumber="366">Saltoposuchus connectens</emphasis>
</taxonomicName>
is additionally distinguished from
<taxonomicName id="4C0BE8BA31119F18427EFBB93C49FB84" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[870,1093,1081,1105]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="12" pageNumber="366" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B31119F18427EFBB93C49FB84" box="[870,1093,1081,1105]" italics="true" pageId="12" pageNumber="366">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0BE8BA31119F184567FBB93D99FB84" authorityName=", Clark and Sues" authorityYear="2002" box="[1151,1429,1081,1105]" class="Reptilia" family="Sphenosuchidae" genus="Litargosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="12" pageNumber="366" phylum="Chordata" rank="species" species="leptorhynchus">
<emphasis id="B97F4F2B31119F184567FBB93D99FB84" box="[1151,1429,1081,1105]" italics="true" pageId="12" pageNumber="366">Litargosuchus leptorhynchus</emphasis>
</taxonomicName>
in possessing a longitudinal crest along the dorsolateral surface of the squamosal; a distinct depression for the supratemporal fossa on the posterior end of the dorsal surface of the frontal; strongly sculpted and transversely wide osteoderms; a wide and short, robust ischial shass; and a robust femur that is longer than the tibia and fibula.
</paragraph>
<paragraph id="8BB4933931119F184232FAAE3B94F9B5" blockId="12.[810,1459,1326,1632]" pageId="12" pageNumber="366">
<materialsCitation id="3B63996431119F184232FAAE3B94F9B5" collectionCode="SMNS" pageId="12" pageNumber="366" specimenCode="SMNS 12597" specimenCount="3" typeStatus="holotype">
<emphasis id="B97F4F2B31119F184232FAAE3B86FA93" box="[810,906,1326,1350]" italics="true" pageId="12" pageNumber="366">
<typeStatus id="54B02D9B31119F184232FAAE3B89FA93" box="[810,901,1326,1350]" pageId="12" pageNumber="366">Holotype</typeStatus>
:
</emphasis>
<specimenCode id="DBAD3B4231119F184289FAAE3C11FA93" box="[913,1053,1326,1350]" collectionCode="SMNS" country="Germany" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34840" lsid="urn:lsid:biocol.org:col:34840" name="Staatliches Museum fuer Naturkund Stuttgart" pageId="12" pageNumber="366">SMNS 12597</specimenCode>
a–d (
<figureCitation id="13308FBC31119F18454DFAAE3CB6FA93" box="[1109,1210,1326,1350]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="12" pageNumber="366">Fig. 2A–E</figureCitation>
), an associated skeleton comprising a partial skull and much of the postcranium. An isolated less angular
<specimenCode id="DBAD3B4231119F1842C1FAED3C69FA50" box="[985,1125,1389,1413]" collectionCode="SMNS" country="Germany" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34840" lsid="urn:lsid:biocol.org:col:34840" name="Staatliches Museum fuer Naturkund Stuttgart" pageId="12" pageNumber="366">SMNS 12597</specimenCode>
e (
<figureCitation id="13308FBC31119F184599FAED3CC4FA50" box="[1153,1224,1389,1413]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="12" pageNumber="366">Fig. 2F</figureCitation>
) and isolated right calcaneum
<specimenCode id="DBAD3B4231119F184298FA0C3C05FA71" box="[896,1033,1420,1444]" collectionCode="SMNS" country="Germany" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34840" lsid="urn:lsid:biocol.org:col:34840" name="Staatliches Museum fuer Naturkund Stuttgart" pageId="12" pageNumber="366">SMNS 12597</specimenCode>
f (
<figureCitation id="13308FBC31119F18453BFA0C3C81FA71" box="[1059,1165,1420,1444]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="12" pageNumber="366">Fig. 2G, H</figureCitation>
) were previously considered to belong to the holotype specimen (
<bibRefCitation id="EF9AEEC831119F1845B0FA2C3D2BFA11" author="Huene FR" box="[1192,1319,1452,1476]" pageId="12" pageNumber="366" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene 1921</bibRefCitation>
) but are here shown to not belong to the same individual as the holotype (see descriptions of respective bones below). Their assignment to
<taxonomicName id="4C0BE8BA31119F184251F9893C4CF9F4" authorityName="Huene" authorityYear="1921" box="[841,1088,1545,1569]" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="12" pageNumber="366" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B31119F184251F9893C4CF9F4" box="[841,1088,1545,1569]" italics="true" pageId="12" pageNumber="366">Saltoposuchus connectens</emphasis>
</taxonomicName>
is tentatively maintained based on their corresponding morphology to other elements in
<specimenCode id="DBAD3B4231119F184475F9A93B61F9B5" collectionCode="SMNS" country="Germany" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34840" lsid="urn:lsid:biocol.org:col:34840" name="Staatliches Museum fuer Naturkund Stuttgart" pageId="12" pageNumber="366">SMNS 12597</specimenCode>
a–d.
</materialsCitation>
</paragraph>
<paragraph id="8BB4933931119F194232F90138C6FF12" blockId="12.[810,1459,1665,1971]" lastBlockId="13.[129,773,144,199]" lastPageId="13" lastPageNumber="367" pageId="12" pageNumber="366">
<emphasis id="B97F4F2B31119F184232F9013BD6F94C" box="[810,986,1665,1689]" italics="true" pageId="12" pageNumber="366">Referred material:</emphasis>
SMNS 12597e–f (
<figureCitation id="13308FBC31119F184583F9013D0FF94F" box="[1179,1283,1665,1690]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="12" pageNumber="366">Fig. 2F–H</figureCitation>
), isolated less angular and isolated right calcaneum; SMNS 12596 (
<figureCitation id="13308FBC31119F184420F9213D9CF96C" box="[1336,1424,1697,1721]" captionStart-0="Figure 3" captionStart-1="Figure 4" captionStart-2="Figure 5" captionStartId-0="4.[113,178,1499,1523]" captionStartId-1="5.[129,194,1154,1178]" captionStartId-2="6.[113,178,1777,1801]" captionTargetBox-0="[147,1426,144,1471]" captionTargetBox-1="[164,1441,148,1126]" captionTargetBox-2="[221,1353,149,1749]" captionTargetId-0="figure-217@4.[146,1426,144,1471]" captionTargetId-1="figure-363@5.[161,1441,144,1126]" captionTargetId-2="figure-5@6.[220,1353,149,1749]" captionTargetPageId-0="4" captionTargetPageId-1="5" captionTargetPageId-2="6" captionText-0="Figure 3. Part of Saltoposuchus connectens referred specimen SMNS 12596. A, SMNS 12596a, cranial block; B, SMNS 12596b, two dentaries exposed in ventrolateral view; C, SMNS 12596c exposing a less femur in anterolateral view; D, exposing the opposite side of SMNS 12596c, Abbreviations:de, dentary; dr, dorsal rib; dv, dorsal vertebra; fe, femur; ga, gastralia; l., less; mx, maxilla; na, nasal; os, osteoderm; q, quadrate; r., right; sq, squamosal; sr, sacral rib; sv, sacral vertebra; tp, transverse process." captionText-1="Figure 4. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, SMNS 12596d, cervical vertebra exposed in less lateral view; B, interpretative drawing of (A); C, SMNS 12596e, posterior cervical or anterior dorsal vertebra exposed in an angled right anterolateral view; D, interpretative drawing of (C); E, SMNS 12596c, close-up of second sacral vertebra in right lateral view and osteoderm in dorsal view; F, interpretative drawing of (E); G, SMNS 12596f, proximal caudal vertebra block exposed in less lateral view; H, interpretative drawing of (G). Abbreviations: accs, accessory anterior spine; dp, diapophysis; izl, interzygapophyseal lamina; ncs, neurocentral suture; ns, neural spine; os, osteoderm; pap, parapophysis; pcdl, posterior centrodiapophyseal lamina; pofl, posterior flange; poz, postzygaphysis; prdl, prezygodiapophyseal lamina; prz, prezygapophysis; sr, sacral rib; tp, transverse process." captionText-2="Figure 5. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, B, SMNS 12596g, right pubis in (A) anterior and (B) angled medial views; C, SMNS 12596h, right ischium exposed in dorsomedial view and two proximal caudal vertebrae exposed in dorsal view; D, SMNS 12596i, tentative less ulna in posteromedial view; E, F, G, H, SMNS 12596j, right femur in (E) posteromedial, (F) anterolateral, (G) proximal, and (H) distal views; I, proximal half of the right femur in posterolateral view; J, K, SMNS 12596k, right tibia and fibula, (J) with the tibia in posteromedial and the fibula in lateral(?) view and (K) with the tibia in anterolateral and the fibula in medial(?) view. Abbreviations: alcr, anterolateral crest; amt, anteromedial tuber; car, caudal rib; cav, caudal vertebra; cr, crest; crtf, crista tibiofibularis; fi, fibula; gr, groove; ilf, ilium facet; iss, ischial shass; lc, lateral condyle; ltr, lesser trochanter; mc, medial condyle; no, notch; ns, neural spine; obf, obturator foramen; olp, olecranon process; os, osteoderm; poz, postzygapophysis; pr., proximal; prz, prezygapophysis; pus, pubic shass; put, pubic tubercle; r., right; ti, tibia; tr, trochanter." figureDoi-0="http://doi.org/10.5281/zenodo.8426153" figureDoi-1="http://doi.org/10.5281/zenodo.8426157" figureDoi-2="http://doi.org/10.5281/zenodo.8426159" httpUri-0="https://zenodo.org/record/8426153/files/figure.png" httpUri-1="https://zenodo.org/record/8426157/files/figure.png" httpUri-2="https://zenodo.org/record/8426159/files/figure.png" pageId="12" pageNumber="366">Figs 3–5</figureCitation>
), a disarticulated skeleton including several skull elements; SMNS 55009 (
<figureCitation id="13308FBC31119F184264F95F3BB5F922" box="[892,953,1759,1783]" captionStart="Figure 6" captionStartId="7.[129,194,1605,1629]" captionTargetBox="[161,1441,657,1577]" captionTargetId="figure-310@7.[161,1441,657,1577]" captionTargetPageId="7" captionText="Figure 6. Part of Saltoposuchus connectens referred specimen SMNS 55009. A, SMNS 55009a, dorsal vertebral centrum in less lateral view; B, SMNS 55009d, proximal caudal vertebral centrum in angled lateral view; C, SMNS 55009b, first sacral vertebra in less lateral view; D, interpretative drawing of (C); E, SMNS 55009c, proximal caudal vertebra in posterior view; F, interpretative drawing of (E). Abbreviations: car, caudal rib; ns, neural spine; pas, posterior articular surface; poz, postzygapophysis; prz, prezygapophysis; sr, sacral rib; sut, suture." figureDoi="http://doi.org/10.5281/zenodo.8426161" httpUri="https://zenodo.org/record/8426161/files/figure.png" pageId="12" pageNumber="366">Figs 6</figureCitation>
,
<figureCitation id="13308FBC31119F1842DDF9603BDEF922" box="[965,978,1760,1783]" captionStart="Figure 7" captionStartId="8.[113,178,1656,1680]" captionTargetBox="[146,1426,144,1628]" captionTargetId="figure-102@8.[146,1426,144,1628]" captionTargetPageId="8" captionText="Figure 7. Saltoposuchus connectens referred specimen SMNS 55009, continued. A, B, C, D, E, F, SMNS 55009e, right tibia in (A) posterior, (B) anterior, (C) medial, (D) lateral, (E) proximal, and (F) distal views; G, H, SMNS 55009h, metatarsal block in (G) dorsal(?) and (H) opposing views; I, J, K, L, SMNS 55009g, right calcaneum in (I) medial, (J) lateral, (K) proximal, and (L) distal views. Abbreviations: als, anterolateral side; asf, astragalus facet; calt, calcaneal tuber; gr, groove; latc, lateral condyle; lg, longitudinal groove; mt, metatarsal; no, notch; pme, posterior extension; pms, posteromedial side; tr, trough." figureDoi="http://doi.org/10.5281/zenodo.8426163" httpUri="https://zenodo.org/record/8426163/files/figure.png" pageId="12" pageNumber="366">7</figureCitation>
), several postcervical vertebrae and part of the right hindlimb, including an incomplete pes; SMNS 12591a (
<figureCitation id="13308FBC31119F18422DF89E3B7AF8E3" box="[821,886,1822,1846]" captionStart="Figure 8" captionStartId="9.[129,194,1772,1796]" captionTargetBox="[178,1429,144,1744]" captionTargetId="figure-6@9.[174,1429,144,1744]" captionTargetPageId="9" captionText="Figure 8. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in right lateral view; B, interpretative drawing of A; C, less lateral view; D, interpretative drawing of (C). Abbreviations:an, angular; cc, crista cranii; de, dentary; fr, frontal; j, jugal; la, lacrimal; ls, laterosphenoid; mx, maxilla; na, nasal; pa, parietal; pl, palatine; po, postorbital; prf, prefrontal; pt, pterygoid; q, quadrate; qj, quadratojugal; sa, surangular; sq, squamosal." figureDoi="http://doi.org/10.5281/zenodo.8426167" httpUri="https://zenodo.org/record/8426167/files/figure.png" pageId="12" pageNumber="366">Figs 8</figureCitation>
,
<figureCitation id="13308FBC31119F18429FF89E3B98F8E3" box="[903,916,1822,1846]" captionStart="Figure 9" captionStartId="10.[113,178,1772,1796]" captionTargetBox="[222,1352,144,1744]" captionTargetId="figure-6@10.[220,1352,144,1744]" captionTargetPageId="10" captionText="Figure 9. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in dorsal view; B, interpretative drawing of (A); C, ventral view; D, interpretative drawing of (C). Abbreviations:an, angular; antfo, antorbital fossa; bpp, basipterygoid process; crb, ceratobranchial I; de, dentary; ect, ectopterygoid; fr, frontal; j, jugal; la, lacrimal; mx, maxilla; na, nasal; pa, parietal; pbs, parabasisphenoid; pl, palatine; plf, palatine fossa; po, postorbital; pra, pre-articular; prf, prefrontal; pt, pterygoid; ptmr, pterygoid medial rim; sa, surangular; spl, splenial; sq, squamosal; vo, vomer." figureDoi="http://doi.org/10.5281/zenodo.8426169" httpUri="https://zenodo.org/record/8426169/files/figure.png" pageId="12" pageNumber="366">9</figureCitation>
), a largely complete, strongly dorsoventrally compressed skull and mandible. This specimen has undergone acidbased preparation since the description of this specimen by
<bibRefCitation id="EF9AEEC831119F184232F8FC3C28F841" author="Sereno PC &amp; Wild R" box="[810,1060,1916,1940]" pageId="12" pageNumber="366" pagination="435 - 58" refId="ref32722" refString="Sereno PC, Wild R. Procompsognathus: theropod, ' thecodont' or both? Journal of Vertebrate Paleontology 1992; 12: 435 - 58." type="journal article" year="1992">Sereno and Wild (1992)</bibRefCitation>
, revealing additional morphological information, particularly on the palate (
<figureCitation id="13308FBC31119F1845B0F81B3D1CF866" box="[1192,1296,1947,1971]" captionStart="Figure 9" captionStartId="10.[113,178,1772,1796]" captionTargetBox="[222,1352,144,1744]" captionTargetId="figure-6@10.[220,1352,144,1744]" captionTargetPageId="10" captionText="Figure 9. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in dorsal view; B, interpretative drawing of (A); C, ventral view; D, interpretative drawing of (C). Abbreviations:an, angular; antfo, antorbital fossa; bpp, basipterygoid process; crb, ceratobranchial I; de, dentary; ect, ectopterygoid; fr, frontal; j, jugal; la, lacrimal; mx, maxilla; na, nasal; pa, parietal; pbs, parabasisphenoid; pl, palatine; plf, palatine fossa; po, postorbital; pra, pre-articular; prf, prefrontal; pt, pterygoid; ptmr, pterygoid medial rim; sa, surangular; spl, splenial; sq, squamosal; vo, vomer." figureDoi="http://doi.org/10.5281/zenodo.8426169" httpUri="https://zenodo.org/record/8426169/files/figure.png" pageId="12" pageNumber="366">Fig. 9C, D</figureCitation>
); SMNS 12352 (
<figureCitation id="13308FBC31109F194193FF1038D8FF7D" box="[139,212,144,168]" captionStart="Figure 10" captionStartId="14.[113,178,1772,1796]" captionTargetBox="[197,1376,144,1744]" captionTargetId="figure-5@14.[197,1376,144,1744]" captionTargetPageId="14" captionText="Figure 10. Saltoposuchus connectens referred specimen SMNS 12352.A, SMNS 12352a with the snout region exposed in angled right lateral view; B, interpretative drawing of (A); C, opposite side of SMNS 12352a exposing the posterior portion of the less maxilla and lacrimal in angled lateral view; D, interpretative drawing of (C); E, SMNS 12352b, carpus and manus in dorsal view; F, interpretative drawing of (E); G, SMNS 12352b in medial view; H, interpretative drawing of (G). Abbreviations; antfe, antorbital fenestra; antfo, antorbital fossa; dc, distal carpal; de, dentary; exp, extensor pit; fr, frontal; j, jugal; la, lacrimal; lad, lacrimal duct; mc, metacarpal; mx, maxilla; na, nasal; ph, phalanx; pm, premaxilla; prf, prefrontal; ra, radiale; ul, ulnare." figureDoi="http://doi.org/10.5281/zenodo.8426171" httpUri="https://zenodo.org/record/8426171/files/figure.png" pageId="13" pageNumber="367">Fig. 10</figureCitation>
), a largely complete, well preserved snout, and a less manus.
</paragraph>
<paragraph id="8BB4933931109F194199FF693A0FFE8A" blockId="13.[129,777,233,351]" pageId="13" pageNumber="367">
<emphasis id="B97F4F2B31109F194199FF69396CFED4" box="[129,352,233,257]" italics="true" pageId="13" pageNumber="367">Locality and horizon:</emphasis>
The discontinued ‘Weisser Steinbruch’ sandstone quarries near Pfaffenhofen. Middle Stubensandstein Member of the Löwenstein Formation, middle Keuper (Norian, Late Triassic;
<bibRefCitation id="EF9AEEC831109F194015FEC739FEFE8A" author="Sues H-D &amp; Schoch RR" box="[269,498,327,351]" pageId="13" pageNumber="367" pagination="859 - 72" refId="ref32785" refString="Sues H-D, Schoch RR. Reassessment of cf. Halticosaurus orbitoangulatus from the Upper Triassic (Norian) of Germany - a pseudosuchian, not a dinosaur. Zoological Journal of the Linnean Society 2013; 168: 859 - 72." type="journal article" year="2013">Sues and Schoch 2013</bibRefCitation>
).
</paragraph>
<paragraph id="8BB4933931109F194199FE00392DFCFA" blockId="13.[129,778,384,815]" pageId="13" pageNumber="367">
<emphasis id="B97F4F2B31109F194199FE003961FE4D" box="[129,365,384,408]" italics="true" pageId="13" pageNumber="367">Ontogenetic assessment:</emphasis>
The relative sizes of the
<taxonomicName id="4C0BE8BA31109F194399FE0038EBFE62" authorityName="Huene" authorityYear="1921" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="13" pageNumber="367" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B31109F194399FE0038EBFE62" italics="true" pageId="13" pageNumber="367">Saltoposuchus connectens</emphasis>
</taxonomicName>
specimens are compared in
<tableCitation id="C689A68231109F19433AFE1F3A7EFE62" box="[546,626,415,439]" captionStart="Table 2" captionStartId="7.[131,186,143,167]" captionText="Table 2. Comparison of measurements among the Saltoposuchus connectensspecimens. A full list of measurements is presented in the Supporting Information, Table S1" pageId="13" pageNumber="367">Table 2</tableCitation>
. Size discrepancy between the smallest (SMNS 12591a) and largest specimen (SMNS 55009) is inferred at 58.1%. The right femur of SMNS 12596, the second-largest specimen (90.2%), has been histologically sectioned. The interpretation of its relative age is somewhat ambiguous but the individual probably represents a sub-adult, since it is neither an early juvenile, based on the relative size and robusticity of its skeleton, nor skeletally mature, based on the presence of open neurocentral sutures in some of the presacral vertebrae and the absence of an outer circumferential layer (OCL; also known as an external fundamental system, EFS) and secondary osteons in the histological section (see discussion below).
</paragraph>
</subSubSection>
<subSubSection id="C311C0B231109F0B409DFCD33975FBE1" lastPageId="31" lastPageNumber="385" pageId="13" pageNumber="367" type="description">
<paragraph id="8BB4933931109F19409DFCD33A09FCB8" blockId="13.[389,517,851,877]" box="[389,517,851,877]" pageId="13" pageNumber="367">
<emphasis id="B97F4F2B31109F19409DFCD33A09FCB8" bold="true" box="[389,517,851,877]" pageId="13" pageNumber="367">Description</emphasis>
</paragraph>
<paragraph id="8BB4933931109F1940B5FCFD39D1FC42" blockId="13.[429,477,893,919]" box="[429,477,893,919]" pageId="13" pageNumber="367">
<emphasis id="B97F4F2B31109F1940B5FCFD39D1FC42" box="[429,477,893,919]" italics="true" pageId="13" pageNumber="367">Skull</emphasis>
</paragraph>
<paragraph id="8BB4933931109F194199FC233904FAA4" blockId="13.[128,778,931,1394]" pageId="13" pageNumber="367">
<emphasis id="B97F4F2B31109F194199FC2338FFFC6E" box="[129,243,931,955]" italics="true" pageId="13" pageNumber="367">Premaxilla:</emphasis>
SMNS 12352a is the only specimen to preserve remains of the premaxilla and even in this specimen only a badly broken right premaxilla and a small fragment of the posterodorsal process of the less premaxilla are present (
<figureCitation id="13308FBC31109F1943FBFB8138C7FBED" captionStart="Figure 10" captionStartId="14.[113,178,1772,1796]" captionTargetBox="[197,1376,144,1744]" captionTargetId="figure-5@14.[197,1376,144,1744]" captionTargetPageId="14" captionText="Figure 10. Saltoposuchus connectens referred specimen SMNS 12352.A, SMNS 12352a with the snout region exposed in angled right lateral view; B, interpretative drawing of (A); C, opposite side of SMNS 12352a exposing the posterior portion of the less maxilla and lacrimal in angled lateral view; D, interpretative drawing of (C); E, SMNS 12352b, carpus and manus in dorsal view; F, interpretative drawing of (E); G, SMNS 12352b in medial view; H, interpretative drawing of (G). Abbreviations; antfe, antorbital fenestra; antfo, antorbital fossa; dc, distal carpal; de, dentary; exp, extensor pit; fr, frontal; j, jugal; la, lacrimal; lad, lacrimal duct; mc, metacarpal; mx, maxilla; na, nasal; ph, phalanx; pm, premaxilla; prf, prefrontal; ra, radiale; ul, ulnare." figureDoi="http://doi.org/10.5281/zenodo.8426171" httpUri="https://zenodo.org/record/8426171/files/figure.png" pageId="13" pageNumber="367">Fig. 10A, B</figureCitation>
). The premaxilla connects to the maxilla posteriorly and to the nasal posterodorsally. The right premaxilla preserves two teeth and is incomplete anteriorly. It bears a prominent posterodorsal process, which wedges between, and overlaps with, both the maxilla and nasal, as in other non-crocodyliform crocodylomorphs like
<taxonomicName id="4C0BE8BA31109F19407DFB3D3B08FB00" authority="(Sues et al. 2003)" baseAuthorityName="Sues" baseAuthorityYear="2003" box="[357,772,1213,1237]" class="Reptilia" family="Sphenosuchidae" genus="Dromicosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="13" pageNumber="367" phylum="Chordata" rank="species" species="grallator">
<emphasis id="B97F4F2B31109F19407DFB3D3A5DFB00" box="[357,593,1213,1237]" italics="true" pageId="13" pageNumber="367">Dromicosuchus grallator</emphasis>
(
<bibRefCitation id="EF9AEEC831109F194347FB3D3AF5FB00" author="Sues H-D &amp; Olsen PE &amp; Carter JG" box="[607,761,1213,1237]" pageId="13" pageNumber="367" pagination="329 - 43" refId="ref32828" refString="Sues H-D, Olsen PE, Carter JG et al. A new crocodylomorph archosaur from the Upper Triassic of North Carolina. Journal of Vertebrate Paleontology 2003; 23: 329 - 43." type="journal article" year="2003">
Sues
<emphasis id="B97F4F2B31109F19438BFB3E3ACDFB00" box="[659,705,1213,1237]" italics="true" pageId="13" pageNumber="367">et al.</emphasis>
2003
</bibRefCitation>
)
</taxonomicName>
, differing from the tight suture between the premaxilla and maxilla seen in crocodyliforms (e.g.
<bibRefCitation id="EF9AEEC831109F1940DBFB7C3AB7FAC1" author="Colbert EH &amp; Mook CC" box="[451,699,1276,1300]" pageId="13" pageNumber="367" pagination="143 - 82" refId="ref31130" refString="Colbert EH, Mook CC. The ancestral crocodilian Protosuchus. Bulletin of the American Museum of Natural History 1951; 97: 143 - 82." type="journal article" year="1951">Colbert and Mook 1951</bibRefCitation>
). Thus, the maxilla did not contribute to the margin of the external naris in
<taxonomicName id="4C0BE8BA31109F194183FABB399CFA87" authorityName="Huene" authorityYear="1921" box="[155,400,1339,1363]" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="13" pageNumber="367" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B31109F194183FABB399CFA87" box="[155,400,1339,1363]" italics="true" pageId="13" pageNumber="367">Saltoposuchus connectens</emphasis>
</taxonomicName>
, as was also indicated by
<bibRefCitation id="EF9AEEC831109F19438DFABB390EFAA7" author="Sereno PC &amp; Wild R" pageId="13" pageNumber="367" pagination="435 - 58" refId="ref32722" refString="Sereno PC, Wild R. Procompsognathus: theropod, ' thecodont' or both? Journal of Vertebrate Paleontology 1992; 12: 435 - 58." type="journal article" year="1992">Sereno and Wild (1992)</bibRefCitation>
.
</paragraph>
<paragraph id="8BB4933931109F194198FA133CE2FBC0" blockId="13.[128,778,1427,1984]" lastBlockId="13.[825,1475,144,1984]" pageId="13" pageNumber="367">
<emphasis id="B97F4F2B31109F194198FA1338DAFA7E" box="[128,214,1427,1451]" italics="true" pageId="13" pageNumber="367">Maxilla:</emphasis>
The maxilla is preserved in SMNS 12596a, SMNS 12352a, and SMNS 12591a, and it thus allows for comparison betweenaspecimenunambiguouslyaưributableto
<taxonomicName id="4C0BE8BA31109F194399FA5238EAF9DC" authorityName="Huene" authorityYear="1921" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="13" pageNumber="367" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B31109F194399FA5238EAF9DC" italics="true" pageId="13" pageNumber="367">Saltoposuchus connectens</emphasis>
</taxonomicName>
(SMNS 12596) and specimens with a controversial taxonomic identification (SMNS 12352, SMNS 12591a;
<tableCitation id="C689A68231109F1943C9F9913881F99D" captionStart="Table 1" captionStartId="2.[114,169,143,167]" captionText="Table 1. Summary of the taxonomic assignments in relevant studies of the specimens that now comprise Saltoposuchus connectens" pageId="13" pageNumber="367">Table 1</tableCitation>
). In fact, most of the features put forward to distinguish these specimens taxonomically have revolved around the maxilla (
<bibRefCitation id="EF9AEEC831109F194193F9EF3919F952" author="Ostrom JH" box="[139,277,1647,1671]" pageId="13" pageNumber="367" pagination="179 - 95" refId="ref32208" refString="Ostrom JH. Procompsognathus - Theropod or Thecodont? Palaeontographica A 1981; 175: 179 - 95." type="journal article" year="1981">Ostrom 1981</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831109F194039F9EF3A27F952" author="Knoll F &amp; Rohrberg K" box="[289,555,1647,1671]" pageId="13" pageNumber="367" pagination="109 - 15" refId="ref31840" refString="Knoll F, Rohrberg K. CT scanning, rapid prototyping and re-examination of a partial skull of a basal crocodylomorph from the Late Triassic of Germany. Swiss Journal of Geosciences 2012; 105: 109 - 15." type="journal article" year="2012">Knoll and Rohrberg 2012</bibRefCitation>
). SMNS 12596a preserves a disarticulated, virtually complete, less maxilla (
<figureCitation id="13308FBC31109F1943A8F90E3AF5F973" box="[688,761,1678,1702]" captionStart="Figure 3" captionStartId="4.[113,178,1499,1523]" captionTargetBox="[147,1426,144,1471]" captionTargetId="figure-217@4.[146,1426,144,1471]" captionTargetPageId="4" captionText="Figure 3. Part of Saltoposuchus connectens referred specimen SMNS 12596. A, SMNS 12596a, cranial block; B, SMNS 12596b, two dentaries exposed in ventrolateral view; C, SMNS 12596c exposing a less femur in anterolateral view; D, exposing the opposite side of SMNS 12596c, Abbreviations:de, dentary; dr, dorsal rib; dv, dorsal vertebra; fe, femur; ga, gastralia; l., less; mx, maxilla; na, nasal; os, osteoderm; q, quadrate; r., right; sq, squamosal; sr, sacral rib; sv, sacral vertebra; tp, transverse process." figureDoi="http://doi.org/10.5281/zenodo.8426153" httpUri="https://zenodo.org/record/8426153/files/figure.png" pageId="13" pageNumber="367">Fig. 3A</figureCitation>
). Based on the length of this element, this specimen is considerably larger than SMNS 12352 and SMNS 12591a (
<tableCitation id="C689A68231109F19438DF94C3AEDF930" box="[661,737,1740,1765]" captionStart="Table 2" captionStartId="7.[131,186,143,167]" captionText="Table 2. Comparison of measurements among the Saltoposuchus connectensspecimens. A full list of measurements is presented in the Supporting Information, Table S1" pageId="13" pageNumber="367">Table 2</tableCitation>
). It preserves four teeth, with the largest, the second counted from anterior, being partially shissed out of its alveolus, exposing much of the root. At least eight nutrient foramina are present dorsal to the alveolar margin in lateral view. SMNS 12352a has a virtually complete right maxilla, which preserves 14 teeth (
<figureCitation id="13308FBC31109F19436EF8E93AEFF854" box="[630,739,1897,1921]" captionStart="Figure 10" captionStartId="14.[113,178,1772,1796]" captionTargetBox="[197,1376,144,1744]" captionTargetId="figure-5@14.[197,1376,144,1744]" captionTargetPageId="14" captionText="Figure 10. Saltoposuchus connectens referred specimen SMNS 12352.A, SMNS 12352a with the snout region exposed in angled right lateral view; B, interpretative drawing of (A); C, opposite side of SMNS 12352a exposing the posterior portion of the less maxilla and lacrimal in angled lateral view; D, interpretative drawing of (C); E, SMNS 12352b, carpus and manus in dorsal view; F, interpretative drawing of (E); G, SMNS 12352b in medial view; H, interpretative drawing of (G). Abbreviations; antfe, antorbital fenestra; antfo, antorbital fossa; dc, distal carpal; de, dentary; exp, extensor pit; fr, frontal; j, jugal; la, lacrimal; lad, lacrimal duct; mc, metacarpal; mx, maxilla; na, nasal; ph, phalanx; pm, premaxilla; prf, prefrontal; ra, radiale; ul, ulnare." figureDoi="http://doi.org/10.5281/zenodo.8426171" httpUri="https://zenodo.org/record/8426171/files/figure.png" pageId="13" pageNumber="367">Fig. 10A, B</figureCitation>
). A single tooth is clearly missing and therefore the maxillary tooth count is tentatively established at 15. Many nutrient foramina can be observed dorsal to the alveolar margin. Both maxillae of SMNS 12591a preserve nine teeth, an incomplete count (
<figureCitation id="13308FBC31109F194484FF2F3B4AFF33" captionStart="Figure 8" captionStartId="9.[129,194,1772,1796]" captionTargetBox="[178,1429,144,1744]" captionTargetId="figure-6@9.[174,1429,144,1744]" captionTargetPageId="9" captionText="Figure 8. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in right lateral view; B, interpretative drawing of A; C, less lateral view; D, interpretative drawing of (C). Abbreviations:an, angular; cc, crista cranii; de, dentary; fr, frontal; j, jugal; la, lacrimal; ls, laterosphenoid; mx, maxilla; na, nasal; pa, parietal; pl, palatine; po, postorbital; prf, prefrontal; pt, pterygoid; q, quadrate; qj, quadratojugal; sa, surangular; sq, squamosal." figureDoi="http://doi.org/10.5281/zenodo.8426167" httpUri="https://zenodo.org/record/8426167/files/figure.png" pageId="13" pageNumber="367">Fig. 8</figureCitation>
). On both sides, the posteriormost teeth are only represented by small protuberances. The posterior extent of the maxillary teeth is similar to that in SMNS 12352a, namely roughly at the level of the ventral foot of the lacrimal. This thus corroborates the interpretation of
<bibRefCitation id="EF9AEEC831109F194539FECC3D3CFEB1" author="Sereno PC &amp; Wild R" box="[1057,1328,332,356]" pageId="13" pageNumber="367" pagination="435 - 58" refId="ref32722" refString="Sereno PC, Wild R. Procompsognathus: theropod, ' thecodont' or both? Journal of Vertebrate Paleontology 1992; 12: 435 - 58." type="journal article" year="1992">Sereno and Wild (1992)</bibRefCitation>
and opposes
<bibRefCitation id="EF9AEEC831109F194221FEEB3BD7FE56" author="Ostrom JH" box="[825,987,363,387]" pageId="13" pageNumber="367" pagination="179 - 95" refId="ref32208" refString="Ostrom JH. Procompsognathus - Theropod or Thecodont? Palaeontographica A 1981; 175: 179 - 95." type="journal article" year="1981">Ostrom (1981)</bibRefCitation>
, who considered both specimens to be distinguishable taxonomically in part based on a difference in posterior extent of the maxillary teeth. The angle formed between the main body of the maxilla and its ascending process is 24° in SMNS 12596a, 29° in SMNS 12352a, and 9° in SMNS 12591a. The very sharp angle of SMNS 12591a can clearly be aưributable to the considerable dorsoventral compression of this specimen, as has been pointed out by previous authors (e.g.
<bibRefCitation id="EF9AEEC831109F194454FDC63BA9FDAB" author="Sereno PC &amp; Wild R" pageId="13" pageNumber="367" pagination="435 - 58" refId="ref32722" refString="Sereno PC, Wild R. Procompsognathus: theropod, ' thecodont' or both? Journal of Vertebrate Paleontology 1992; 12: 435 - 58." type="journal article" year="1992">Sereno and Wild 1992</bibRefCitation>
). The alveolar margin is slightly convex in lateral view in its anterior half and very slightly concave in its posterior half in both SMNS 12596a and SMNS 12352a. In SMNS 12591a this curvature is much less distinct, but this too is clearly aưributable to its dorsoventral compression. This compression also led to the misinterpretation of
<bibRefCitation id="EF9AEEC831109F194541FC823CF7FCCF" author="Ostrom JH" box="[1113,1275,770,794]" pageId="13" pageNumber="367" pagination="179 - 95" refId="ref32208" refString="Ostrom JH. Procompsognathus - Theropod or Thecodont? Palaeontographica A 1981; 175: 179 - 95." type="journal article" year="1981">Ostrom (1981)</bibRefCitation>
that the antorbital fenestra of SMNS 12591a is restricted to the anterior portion of the antorbital fossa, and that the lacrimal had a considerable extent anteriorly. The element considered to be the anterior part of the lacrimal in the antorbital fossa in fact represents the palatine, which has been displaced dorsally on its medial side due to dorsoventral compression, as was already indicated by
<bibRefCitation id="EF9AEEC831109F194465FC3F3B74FC23" author="Huene FR" pageId="13" pageNumber="367" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene (1921</bibRefCitation>
: fig. 22), and which has been corroborated by other authors (
<bibRefCitation id="EF9AEEC831109F194266FC7D3C6EFBC0" author="Sereno PC &amp; Wild R" box="[894,1122,1021,1045]" pageId="13" pageNumber="367" pagination="435 - 58" refId="ref32722" refString="Sereno PC, Wild R. Procompsognathus: theropod, ' thecodont' or both? Journal of Vertebrate Paleontology 1992; 12: 435 - 58." type="journal article" year="1992">Sereno and Wild 1992</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831109F194575FC7D3CD1FBC0" author="Allen RC" box="[1133,1245,1021,1045]" pageId="13" pageNumber="367" refId="ref30591" refString="Allen RC. The anatomy and systematics of Terrestrisuchus gracilis (Archosauria, Crocodylomorpha). Unpublished Ph. D. Thesis, Northern Illinois University, 2010." type="book" year="2010">Allen 2010</bibRefCitation>
).
</paragraph>
<paragraph id="8BB4933931109F1B424DFB9C3B05FED3" blockId="13.[825,1475,144,1984]" lastBlockId="15.[129,778,144,262]" lastPageId="15" lastPageNumber="369" pageId="13" pageNumber="367">
The outline of the antorbital cavity and the relative contribution of the antorbital fenestra and fossa are virtually identical between SMNS 12596a and SMNS 12352a (
<figureCitation id="13308FBC31109F194416FBDB3D51FBA6" box="[1294,1373,1115,1139]" captionStart="Figure 3" captionStartId="4.[113,178,1499,1523]" captionTargetBox="[147,1426,144,1471]" captionTargetId="figure-217@4.[146,1426,144,1471]" captionTargetPageId="4" captionText="Figure 3. Part of Saltoposuchus connectens referred specimen SMNS 12596. A, SMNS 12596a, cranial block; B, SMNS 12596b, two dentaries exposed in ventrolateral view; C, SMNS 12596c exposing a less femur in anterolateral view; D, exposing the opposite side of SMNS 12596c, Abbreviations:de, dentary; dr, dorsal rib; dv, dorsal vertebra; fe, femur; ga, gastralia; l., less; mx, maxilla; na, nasal; os, osteoderm; q, quadrate; r., right; sq, squamosal; sr, sacral rib; sv, sacral vertebra; tp, transverse process." figureDoi="http://doi.org/10.5281/zenodo.8426153" httpUri="https://zenodo.org/record/8426153/files/figure.png" pageId="13" pageNumber="367">Figs 3A</figureCitation>
,
<figureCitation id="13308FBC31109F194473FBDB3DBDFBA7" box="[1387,1457,1115,1139]" captionStart="Figure 10" captionStartId="14.[113,178,1772,1796]" captionTargetBox="[197,1376,144,1744]" captionTargetId="figure-5@14.[197,1376,144,1744]" captionTargetPageId="14" captionText="Figure 10. Saltoposuchus connectens referred specimen SMNS 12352.A, SMNS 12352a with the snout region exposed in angled right lateral view; B, interpretative drawing of (A); C, opposite side of SMNS 12352a exposing the posterior portion of the less maxilla and lacrimal in angled lateral view; D, interpretative drawing of (C); E, SMNS 12352b, carpus and manus in dorsal view; F, interpretative drawing of (E); G, SMNS 12352b in medial view; H, interpretative drawing of (G). Abbreviations; antfe, antorbital fenestra; antfo, antorbital fossa; dc, distal carpal; de, dentary; exp, extensor pit; fr, frontal; j, jugal; la, lacrimal; lad, lacrimal duct; mc, metacarpal; mx, maxilla; na, nasal; ph, phalanx; pm, premaxilla; prf, prefrontal; ra, radiale; ul, ulnare." figureDoi="http://doi.org/10.5281/zenodo.8426171" httpUri="https://zenodo.org/record/8426171/files/figure.png" pageId="13" pageNumber="367">10A, B</figureCitation>
). Both have a considerable anterior contribution of the antorbital fossa, but neither form a pit in the anteroventral corner of the fossa, in contrast to
<taxonomicName id="4C0BE8BA31109F194517FB393CE0FB04" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[1039,1260,1209,1233]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="13" pageNumber="367" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B31109F194517FB393CE0FB04" box="[1039,1260,1209,1233]" italics="true" pageId="13" pageNumber="367">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9AEEC831109F1945E6FB393D76FB04" author="Crush PJ" box="[1278,1402,1209,1233]" pageId="13" pageNumber="367" refId="ref31157" refString="Crush PJ. An early, terrestrial, crocodile from South Wales. Unpublished Ph. D. Thesis, University College London, 1980." type="book" year="1980">Crush 1980</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831109F194490FB393DB0FB04" author="Crush PJ" box="[1416,1468,1209,1233]" pageId="13" pageNumber="367" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">1984</bibRefCitation>
,
<emphasis id="B97F4F2B31109F194221FB593B7AFB25" box="[825,886,1241,1264]" italics="true" pageId="13" pageNumber="367">contra</emphasis>
<bibRefCitation id="EF9AEEC831109F194266FB583BFEFB25" author="Clark JM" box="[894,1010,1240,1264]" pageId="13" pageNumber="367" refId="ref30947" refString="Clark JM. Phylogenetic relationships of the crocodylomorphs archosaurs. Unpublished Ph. D. Thesis, University of Chicago, 1986." type="book" year="1986">Clark 1986</bibRefCitation>
). The interpretation by
<bibRefCitation id="EF9AEEC831109F1945E9FB583B88FAC5" author="Knoll F &amp; Rohrberg K" pageId="13" pageNumber="367" pagination="109 - 15" refId="ref31840" refString="Knoll F, Rohrberg K. CT scanning, rapid prototyping and re-examination of a partial skull of a basal crocodylomorph from the Late Triassic of Germany. Swiss Journal of Geosciences 2012; 105: 109 - 15." type="journal article" year="2012">Knoll and Rohrberg (2012)</bibRefCitation>
that the anterior corner of the antorbital fenestra is more acute in SMNS 12596a, and that in SMNS 12352a the anterior end of the antorbital fenestra diverges from the ventral margin of the antorbital fossa, are both rejected. Both specimens clearly possess a rounded anterior end of the antorbital fenestra and the anterior outline of the antorbital fossa is virtually indistinguishable between both specimens. The relative distribution of the antorbital fossa within the antorbital cavity is less extensive in SMNS 12591a, but this is aưributable to the incomplete preservation of the thin lamina that comprises the antorbital fossa (
<figureCitation id="13308FBC31109F194484F9913B4AF99C" captionStart="Figure 8" captionStartId="9.[129,194,1772,1796]" captionTargetBox="[178,1429,144,1744]" captionTargetId="figure-6@9.[174,1429,144,1744]" captionTargetPageId="9" captionText="Figure 8. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in right lateral view; B, interpretative drawing of A; C, less lateral view; D, interpretative drawing of (C). Abbreviations:an, angular; cc, crista cranii; de, dentary; fr, frontal; j, jugal; la, lacrimal; ls, laterosphenoid; mx, maxilla; na, nasal; pa, parietal; pl, palatine; po, postorbital; prf, prefrontal; pt, pterygoid; q, quadrate; qj, quadratojugal; sa, surangular; sq, squamosal." figureDoi="http://doi.org/10.5281/zenodo.8426167" httpUri="https://zenodo.org/record/8426167/files/figure.png" pageId="13" pageNumber="367">Fig. 8</figureCitation>
). The considerable extent of the thin lamina of the antorbital fossa within the antorbital cavity, and the corresponding relatively small opening of the antorbital fenestra, seen particularly in SMNS 12596a and SMNS 12352a (
<figureCitation id="13308FBC31109F1945DCF90F3D1DF972" box="[1220,1297,1679,1703]" captionStart="Figure 3" captionStartId="4.[113,178,1499,1523]" captionTargetBox="[147,1426,144,1471]" captionTargetId="figure-217@4.[146,1426,144,1471]" captionTargetPageId="4" captionText="Figure 3. Part of Saltoposuchus connectens referred specimen SMNS 12596. A, SMNS 12596a, cranial block; B, SMNS 12596b, two dentaries exposed in ventrolateral view; C, SMNS 12596c exposing a less femur in anterolateral view; D, exposing the opposite side of SMNS 12596c, Abbreviations:de, dentary; dr, dorsal rib; dv, dorsal vertebra; fe, femur; ga, gastralia; l., less; mx, maxilla; na, nasal; os, osteoderm; q, quadrate; r., right; sq, squamosal; sr, sacral rib; sv, sacral vertebra; tp, transverse process." figureDoi="http://doi.org/10.5281/zenodo.8426153" httpUri="https://zenodo.org/record/8426153/files/figure.png" pageId="13" pageNumber="367">Figs 3A</figureCitation>
,
<figureCitation id="13308FBC31109F194405F90F3D6EF973" box="[1309,1378,1679,1703]" captionStart="Figure 10" captionStartId="14.[113,178,1772,1796]" captionTargetBox="[197,1376,144,1744]" captionTargetId="figure-5@14.[197,1376,144,1744]" captionTargetPageId="14" captionText="Figure 10. Saltoposuchus connectens referred specimen SMNS 12352.A, SMNS 12352a with the snout region exposed in angled right lateral view; B, interpretative drawing of (A); C, opposite side of SMNS 12352a exposing the posterior portion of the less maxilla and lacrimal in angled lateral view; D, interpretative drawing of (C); E, SMNS 12352b, carpus and manus in dorsal view; F, interpretative drawing of (E); G, SMNS 12352b in medial view; H, interpretative drawing of (G). Abbreviations; antfe, antorbital fenestra; antfo, antorbital fossa; dc, distal carpal; de, dentary; exp, extensor pit; fr, frontal; j, jugal; la, lacrimal; lad, lacrimal duct; mc, metacarpal; mx, maxilla; na, nasal; ph, phalanx; pm, premaxilla; prf, prefrontal; ra, radiale; ul, ulnare." figureDoi="http://doi.org/10.5281/zenodo.8426171" httpUri="https://zenodo.org/record/8426171/files/figure.png" pageId="13" pageNumber="367">10A, B</figureCitation>
), reflects the anatomy seen in most non-crocodyliform crocodylomorphs except
<taxonomicName id="4C0BE8BA31109F194290F94D3C53F930" box="[904,1119,1741,1765]" genus="Carnufex" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="13" pageNumber="367" phylum="Chordata" rank="species" species="carolinensis">
<emphasis id="B97F4F2B31109F194290F94D3C53F930" box="[904,1119,1741,1765]" italics="true" pageId="13" pageNumber="367">Carnufex carolinensis</emphasis>
</taxonomicName>
(e.g.
<bibRefCitation id="EF9AEEC831109F1945BAF94D3D2EF930" author="Crush PJ" box="[1186,1314,1741,1765]" pageId="13" pageNumber="367" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831109F19442CF94D3DB0F930" author="Walker AD" box="[1332,1468,1741,1765]" pageId="13" pageNumber="367" pagination="1 - 120" refId="ref33031" refString="Walker AD. A revision of Sphenosuchus acutus Haughton, a crocodylomorph reptile from the Elliot Formation (Late Triassic or Early Jurassic) of South Africa. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1990; 330: 1 - 120." type="journal article" year="1990">Walker 1990</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831109F194221F96D3C39F8D0" author="Ruebenstahl AA &amp; Klein MD &amp; Yi H" box="[825,1077,1773,1797]" pageId="13" pageNumber="367" pagination="2463 - 556" refId="ref32564" refString="Ruebenstahl AA, Klein MD, Yi H et al. Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros. The Anatomical Record 2022; 305: 2463 - 556." type="journal article" year="2022">
Ruebenstahl
<emphasis id="B97F4F2B31109F1942DCF96D3BF4F8D0" box="[964,1016,1773,1797]" italics="true" pageId="13" pageNumber="367">et al.</emphasis>
2022
</bibRefCitation>
), but conflicts with the morphology seen in early theropod dinosaurs, including coelophysoids (
<bibRefCitation id="EF9AEEC831109F19425CF8AB3BB7F896" author="Raath MA" box="[836,955,1835,1859]" pageId="13" pageNumber="367" refId="ref32399" refString="Raath MA. The anatomy of the Triassic theropod Syntarsus rhodesiensis (Saurischia: Podokesauridae) and a consideration of its biology. Unpublished Ph. D. Thesis, Rhodes University, 1977." type="book" year="1977">Raath 1977</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831109F1942DFF8AB3C5DF896" author="Colbert EH" box="[967,1105,1835,1859]" pageId="13" pageNumber="367" pagination="1 - 160" refId="ref31108" refString="Colbert EH. The Triassic dinosaur Coelophysis. Museum of Northern Arizona, Bulletin 1989; 57: 1 - 160." type="journal article" year="1989">Colbert 1989</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831109F194545F8AB3D1BF896" author="Nesbiu SJ &amp; Smith ND &amp; Irmis RB" box="[1117,1303,1835,1859]" pageId="13" pageNumber="367" pagination="1530 - 3" refId="ref32154" refString="Nesbiu SJ, Smith ND, Irmis RB et al. A complete skeleton of a Late Triassic saurischian and the early evolution of dinosaurs. Science 2009; 326: 1530 - 3." type="journal article" year="2009">
Nesbiư
<emphasis id="B97F4F2B31109F1945B6F8AC3CD2F896" box="[1198,1246,1835,1859]" italics="true" pageId="13" pageNumber="367">et al.</emphasis>
2009
</bibRefCitation>
). The acid preparation of SMNS 12591a has also revealed much of its palatal region, including what are very likely the posterior portions of the palatal processes of the maxillae (
<figureCitation id="13308FBC31109F1945A2F8093D28F874" box="[1210,1316,1929,1953]" captionStart="Figure 9" captionStartId="10.[113,178,1772,1796]" captionTargetBox="[222,1352,144,1744]" captionTargetId="figure-6@10.[220,1352,144,1744]" captionTargetPageId="10" captionText="Figure 9. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in dorsal view; B, interpretative drawing of (A); C, ventral view; D, interpretative drawing of (C). Abbreviations:an, angular; antfo, antorbital fossa; bpp, basipterygoid process; crb, ceratobranchial I; de, dentary; ect, ectopterygoid; fr, frontal; j, jugal; la, lacrimal; mx, maxilla; na, nasal; pa, parietal; pbs, parabasisphenoid; pl, palatine; plf, palatine fossa; po, postorbital; pra, pre-articular; prf, prefrontal; pt, pterygoid; ptmr, pterygoid medial rim; sa, surangular; spl, splenial; sq, squamosal; vo, vomer." figureDoi="http://doi.org/10.5281/zenodo.8426169" httpUri="https://zenodo.org/record/8426169/files/figure.png" pageId="13" pageNumber="367">Fig. 9C, D</figureCitation>
). They meet in the midline and are located quite far posteriorly in the snout. This provides a clear indication of the crocodylomorph affinities of this specimen, as it represents a typical character of this group that foreshadows the maxillary ‘secondary palate’ of later crocodylomorphs (
<bibRefCitation id="EF9AEEC831129F1B405DFF6E39C4FED3" author="Nesbiu SJ" box="[325,456,238,262]" pageId="15" pageNumber="369" pagination="1 - 292" refId="ref32089" refString="Nesbiu SJ. The early evolution of archosaurs: relationships and the origin of major clades. Bulletin of the American Museum of Natural History 2011; 352: 1 - 292." type="journal article" year="2011">Nesbiư 2011</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831129F1B40C9FF6E3AF5FED3" author="Dollman KN &amp; Choiniere JN" box="[465,761,238,262]" pageId="15" pageNumber="369" pagination="2766 - 90" refId="ref31204" refString="Dollman KN, Choiniere JN. Palate evolution in early-branching crocodylomorphs: Implications for homology, systematics, and ecomorphology. The Anatomical Record 2022; 305: 2766 - 90." type="journal article" year="2022">Dollman and Choiniere 2022</bibRefCitation>
).
</paragraph>
<caption id="DF74C3B131139F1A4169F96C3A3AF845" ID-DOI="http://doi.org/10.5281/zenodo.8426171" ID-Zenodo-Dep="8426171" httpUri="https://zenodo.org/record/8426171/files/figure.png" pageId="14" pageNumber="368" startId="14.[113,178,1772,1796]" targetBox="[197,1376,144,1744]" targetPageId="14" targetType="figure">
<paragraph id="8BB4933931139F1A4169F96C3A3AF845" blockId="14.[113,1454,1772,1936]" pageId="14" pageNumber="368">
<emphasis id="B97F4F2B31139F1A4169F96C38DAF8D1" bold="true" box="[113,214,1772,1796]" pageId="14" pageNumber="368">Figure 10.</emphasis>
<taxonomicName id="4C0BE8BA31139F1A41C2F96C39B1F8D1" authorityName="Huene" authorityYear="1921" box="[218,445,1772,1796]" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="14" pageNumber="368" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B31139F1A41C2F96C39B1F8D1" box="[218,445,1772,1796]" italics="true" pageId="14" pageNumber="368">Saltoposuchus connectens</emphasis>
</taxonomicName>
referred specimen SMNS 12352. A, SMNS 12352a with the snout region exposed in angled right lateral view; B, interpretative drawing of (A); C, opposite side of SMNS 12352a exposing the posterior portion of the less maxilla and lacrimal in angled lateral view; D, interpretative drawing of (C); E, SMNS 12352b, carpus and manus in dorsal view; F, interpretative drawing of (E); G, SMNS 12352b in medial view; H, interpretative drawing of (G). Abbreviations; antfe, antorbital fenestra; antfo, antorbital fossa; dc, distal carpal; de, dentary; exp, extensor pit; fr, frontal; j, jugal; la, lacrimal; lad, lacrimal duct; mc, metacarpal; mx, maxilla; na, nasal; ph, phalanx; pm, premaxilla; prf, prefrontal; ra, radiale; ul, ulnare.
</paragraph>
</caption>
<paragraph id="8BB4933931129F1B4199FEA7393DFBC5" blockId="15.[128,778,295,1040]" pageId="15" pageNumber="369">
<emphasis id="B97F4F2B31129F1B4199FEA738CDFEEA" box="[129,193,295,319]" italics="true" pageId="15" pageNumber="369">Nasal:</emphasis>
Clearly identifiable nasals are preserved in SMNS 12352a and SMNS 12591a. In addition,
<bibRefCitation id="EF9AEEC831129F1B40CDFEC73A6AFE8A" author="Crush PJ" box="[469,614,327,351]" pageId="15" pageNumber="369" refId="ref31157" refString="Crush PJ. An early, terrestrial, crocodile from South Wales. Unpublished Ph. D. Thesis, University College London, 1980." type="book" year="1980">Crush (1980)</bibRefCitation>
identified a flat bone fragment on SMNS 12597b as a poorly preserved nasal. However, due to the incompleteness of this element it is not possible to corroborate this identification with confidence, although its preserved morphology does correspond with that of the nasal (
<figureCitation id="13308FBC31129F1B4193FE6338DBFE2E" box="[139,215,483,507]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="15" pageNumber="369">Fig. 2C</figureCitation>
). In SMNS 12352a and SMNS 12591a, the nasals are mostly complete, although their anterior ends are missing (
<figureCitation id="13308FBC31129F1B43F8FD8338B5FDEC" captionStart="Figure 9" captionStartId="10.[113,178,1772,1796]" captionTargetBox="[222,1352,144,1744]" captionTargetId="figure-6@10.[220,1352,144,1744]" captionTargetPageId="10" captionText="Figure 9. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in dorsal view; B, interpretative drawing of (A); C, ventral view; D, interpretative drawing of (C). Abbreviations:an, angular; antfo, antorbital fossa; bpp, basipterygoid process; crb, ceratobranchial I; de, dentary; ect, ectopterygoid; fr, frontal; j, jugal; la, lacrimal; mx, maxilla; na, nasal; pa, parietal; pbs, parabasisphenoid; pl, palatine; plf, palatine fossa; po, postorbital; pra, pre-articular; prf, prefrontal; pt, pterygoid; ptmr, pterygoid medial rim; sa, surangular; spl, splenial; sq, squamosal; vo, vomer." figureDoi="http://doi.org/10.5281/zenodo.8426169" httpUri="https://zenodo.org/record/8426169/files/figure.png" pageId="15" pageNumber="369">Figs 9A, B</figureCitation>
,
<figureCitation id="13308FBC31129F1B41DCFDA23905FDEC" box="[196,265,546,570]" captionStart="Figure 10" captionStartId="14.[113,178,1772,1796]" captionTargetBox="[197,1376,144,1744]" captionTargetId="figure-5@14.[197,1376,144,1744]" captionTargetPageId="14" captionText="Figure 10. Saltoposuchus connectens referred specimen SMNS 12352.A, SMNS 12352a with the snout region exposed in angled right lateral view; B, interpretative drawing of (A); C, opposite side of SMNS 12352a exposing the posterior portion of the less maxilla and lacrimal in angled lateral view; D, interpretative drawing of (C); E, SMNS 12352b, carpus and manus in dorsal view; F, interpretative drawing of (E); G, SMNS 12352b in medial view; H, interpretative drawing of (G). Abbreviations; antfe, antorbital fenestra; antfo, antorbital fossa; dc, distal carpal; de, dentary; exp, extensor pit; fr, frontal; j, jugal; la, lacrimal; lad, lacrimal duct; mc, metacarpal; mx, maxilla; na, nasal; ph, phalanx; pm, premaxilla; prf, prefrontal; ra, radiale; ul, ulnare." figureDoi="http://doi.org/10.5281/zenodo.8426171" httpUri="https://zenodo.org/record/8426171/files/figure.png" pageId="15" pageNumber="369">10A, B</figureCitation>
). The nasals are longitudinally elongate and transversely narrow elements that articulate with the premaxilla anteriorly, the maxilla and lacrimal laterally, the prefrontal posterolaterally, and the frontal posteriorly. The nasal gradually widens posteriorly and has straight lateral and medial margins. Posteriorly, the nasals form a V-shaped articulation with the frontals, which wedge between the nasals anteromedially, as can best be observed in SMNS 12352a (
<figureCitation id="13308FBC31129F1B40EBFD7D3A68FCC0" box="[499,612,765,789]" captionStart="Figure 10" captionStartId="14.[113,178,1772,1796]" captionTargetBox="[197,1376,144,1744]" captionTargetId="figure-5@14.[197,1376,144,1744]" captionTargetPageId="14" captionText="Figure 10. Saltoposuchus connectens referred specimen SMNS 12352.A, SMNS 12352a with the snout region exposed in angled right lateral view; B, interpretative drawing of (A); C, opposite side of SMNS 12352a exposing the posterior portion of the less maxilla and lacrimal in angled lateral view; D, interpretative drawing of (C); E, SMNS 12352b, carpus and manus in dorsal view; F, interpretative drawing of (E); G, SMNS 12352b in medial view; H, interpretative drawing of (G). Abbreviations; antfe, antorbital fenestra; antfo, antorbital fossa; dc, distal carpal; de, dentary; exp, extensor pit; fr, frontal; j, jugal; la, lacrimal; lad, lacrimal duct; mc, metacarpal; mx, maxilla; na, nasal; ph, phalanx; pm, premaxilla; prf, prefrontal; ra, radiale; ul, ulnare." figureDoi="http://doi.org/10.5281/zenodo.8426171" httpUri="https://zenodo.org/record/8426171/files/figure.png" pageId="15" pageNumber="369">Fig. 10A, B</figureCitation>
). This configuration corresponds to that seen in many early crocodylomorphs (
<bibRefCitation id="EF9AEEC831129F1B4193FCBC3905FC81" author="Crush PJ" box="[139,265,828,852]" pageId="15" pageNumber="369" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831129F1B4000FCBC39AAFC81" author="Colbert EH" box="[280,422,828,852]" pageId="15" pageNumber="369" pagination="1 - 160" refId="ref31108" refString="Colbert EH. The Triassic dinosaur Coelophysis. Museum of Northern Arizona, Bulletin 1989; 57: 1 - 160." type="journal article" year="1989">Colbert 1989</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831129F1B40AEFCBC3A31FC81" author="Walker AD" box="[438,573,828,852]" pageId="15" pageNumber="369" pagination="1 - 120" refId="ref33031" refString="Walker AD. A revision of Sphenosuchus acutus Haughton, a crocodylomorph reptile from the Elliot Formation (Late Triassic or Early Jurassic) of South Africa. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1990; 330: 1 - 120." type="journal article" year="1990">Walker 1990</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831129F1B4354FCBC38B9FCA6" author="Ruebenstahl AA &amp; Klein MD &amp; Yi H" pageId="15" pageNumber="369" pagination="2463 - 556" refId="ref32564" refString="Ruebenstahl AA, Klein MD, Yi H et al. Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros. The Anatomical Record 2022; 305: 2463 - 556." type="journal article" year="2022">
Ruebenstahl
<emphasis id="B97F4F2B31129F1B43CFFCBD3B06FC81" box="[727,778,828,852]" italics="true" pageId="15" pageNumber="369">et al.</emphasis>
2022
</bibRefCitation>
) and several other Triassic pseudosuchians (
<bibRefCitation id="EF9AEEC831129F1B4365FCDB3B0EFCA6" author="Nesbiu SJ" box="[637,770,859,883]" pageId="15" pageNumber="369" pagination="1 - 292" refId="ref32089" refString="Nesbiu SJ. The early evolution of archosaurs: relationships and the origin of major clades. Bulletin of the American Museum of Natural History 2011; 352: 1 - 292." type="journal article" year="2011">Nesbiư 2011</bibRefCitation>
: ch. 43). The dorsal surface of the nasals is distinctly piưed in SMNS 12352a (
<figureCitation id="13308FBC31129F1B403EFC1A3977FC67" box="[294,379,922,946]" captionStart="Figure 10" captionStartId="14.[113,178,1772,1796]" captionTargetBox="[197,1376,144,1744]" captionTargetId="figure-5@14.[197,1376,144,1744]" captionTargetPageId="14" captionText="Figure 10. Saltoposuchus connectens referred specimen SMNS 12352.A, SMNS 12352a with the snout region exposed in angled right lateral view; B, interpretative drawing of (A); C, opposite side of SMNS 12352a exposing the posterior portion of the less maxilla and lacrimal in angled lateral view; D, interpretative drawing of (C); E, SMNS 12352b, carpus and manus in dorsal view; F, interpretative drawing of (E); G, SMNS 12352b in medial view; H, interpretative drawing of (G). Abbreviations; antfe, antorbital fenestra; antfo, antorbital fossa; dc, distal carpal; de, dentary; exp, extensor pit; fr, frontal; j, jugal; la, lacrimal; lad, lacrimal duct; mc, metacarpal; mx, maxilla; na, nasal; ph, phalanx; pm, premaxilla; prf, prefrontal; ra, radiale; ul, ulnare." figureDoi="http://doi.org/10.5281/zenodo.8426171" httpUri="https://zenodo.org/record/8426171/files/figure.png" pageId="15" pageNumber="369">Fig. 10A</figureCitation>
). In this specimen, the dorsal surface of the nasal is transversely convex, particularly anteriorly, whereas in SMNS 12591a this surface is flat, resulting from its dorsoventral compression.
</paragraph>
<paragraph id="8BB4933931129F1B4199FBB43A5DF86F" blockId="15.[129,778,1076,1978]" pageId="15" pageNumber="369">
<emphasis id="B97F4F2B31129F1B4199FBB438EDFB99" box="[129,225,1076,1100]" italics="true" pageId="15" pageNumber="369">Lacrimal:</emphasis>
Both lacrimals are preserved in both SMNS 12352a and SMNS 12591a, although only a portion of the shass is exposed for the less lacrimal of SMNS 12352a (
<figureCitation id="13308FBC31129F1B4356FBF33A86FB5E" box="[590,650,1139,1163]" captionStart="Figure 8" captionStartId="9.[129,194,1772,1796]" captionTargetBox="[178,1429,144,1744]" captionTargetId="figure-6@9.[174,1429,144,1744]" captionTargetPageId="9" captionText="Figure 8. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in right lateral view; B, interpretative drawing of A; C, less lateral view; D, interpretative drawing of (C). Abbreviations:an, angular; cc, crista cranii; de, dentary; fr, frontal; j, jugal; la, lacrimal; ls, laterosphenoid; mx, maxilla; na, nasal; pa, parietal; pl, palatine; po, postorbital; prf, prefrontal; pt, pterygoid; q, quadrate; qj, quadratojugal; sa, surangular; sq, squamosal." figureDoi="http://doi.org/10.5281/zenodo.8426167" httpUri="https://zenodo.org/record/8426167/files/figure.png" pageId="15" pageNumber="369">Figs 8</figureCitation>
,
<figureCitation id="13308FBC31129F1B438EFBF33AAFFB5E" box="[662,675,1139,1163]" captionStart="Figure 9" captionStartId="10.[113,178,1772,1796]" captionTargetBox="[222,1352,144,1744]" captionTargetId="figure-6@10.[220,1352,144,1744]" captionTargetPageId="10" captionText="Figure 9. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in dorsal view; B, interpretative drawing of (A); C, ventral view; D, interpretative drawing of (C). Abbreviations:an, angular; antfo, antorbital fossa; bpp, basipterygoid process; crb, ceratobranchial I; de, dentary; ect, ectopterygoid; fr, frontal; j, jugal; la, lacrimal; mx, maxilla; na, nasal; pa, parietal; pbs, parabasisphenoid; pl, palatine; plf, palatine fossa; po, postorbital; pra, pre-articular; prf, prefrontal; pt, pterygoid; ptmr, pterygoid medial rim; sa, surangular; spl, splenial; sq, squamosal; vo, vomer." figureDoi="http://doi.org/10.5281/zenodo.8426169" httpUri="https://zenodo.org/record/8426169/files/figure.png" pageId="15" pageNumber="369">9</figureCitation>
,
<figureCitation id="13308FBC31129F1B43B6FBF33AF4FB5E" box="[686,760,1139,1163]" captionStart="Figure 10" captionStartId="14.[113,178,1772,1796]" captionTargetBox="[197,1376,144,1744]" captionTargetId="figure-5@14.[197,1376,144,1744]" captionTargetPageId="14" captionText="Figure 10. Saltoposuchus connectens referred specimen SMNS 12352.A, SMNS 12352a with the snout region exposed in angled right lateral view; B, interpretative drawing of (A); C, opposite side of SMNS 12352a exposing the posterior portion of the less maxilla and lacrimal in angled lateral view; D, interpretative drawing of (C); E, SMNS 12352b, carpus and manus in dorsal view; F, interpretative drawing of (E); G, SMNS 12352b in medial view; H, interpretative drawing of (G). Abbreviations; antfe, antorbital fenestra; antfo, antorbital fossa; dc, distal carpal; de, dentary; exp, extensor pit; fr, frontal; j, jugal; la, lacrimal; lad, lacrimal duct; mc, metacarpal; mx, maxilla; na, nasal; ph, phalanx; pm, premaxilla; prf, prefrontal; ra, radiale; ul, ulnare." figureDoi="http://doi.org/10.5281/zenodo.8426171" httpUri="https://zenodo.org/record/8426171/files/figure.png" pageId="15" pageNumber="369">10A–D</figureCitation>
). Although both lacrimals are virtually complete in SMNS 12591a, they are deformed and poorly preserved, and the right lacrimal of SMNS 12352a is the best representative of this element. It has an inverted L-shape, with a dorsoventrally oriented shass and an anterodorsal process (
<figureCitation id="13308FBC31129F1B40A2FA903A3DFAF2" box="[442,561,1296,1320]" captionStart="Figure 10" captionStartId="14.[113,178,1772,1796]" captionTargetBox="[197,1376,144,1744]" captionTargetId="figure-5@14.[197,1376,144,1744]" captionTargetPageId="14" captionText="Figure 10. Saltoposuchus connectens referred specimen SMNS 12352.A, SMNS 12352a with the snout region exposed in angled right lateral view; B, interpretative drawing of (A); C, opposite side of SMNS 12352a exposing the posterior portion of the less maxilla and lacrimal in angled lateral view; D, interpretative drawing of (C); E, SMNS 12352b, carpus and manus in dorsal view; F, interpretative drawing of (E); G, SMNS 12352b in medial view; H, interpretative drawing of (G). Abbreviations; antfe, antorbital fenestra; antfo, antorbital fossa; dc, distal carpal; de, dentary; exp, extensor pit; fr, frontal; j, jugal; la, lacrimal; lad, lacrimal duct; mc, metacarpal; mx, maxilla; na, nasal; ph, phalanx; pm, premaxilla; prf, prefrontal; ra, radiale; ul, ulnare." figureDoi="http://doi.org/10.5281/zenodo.8426171" httpUri="https://zenodo.org/record/8426171/files/figure.png" pageId="15" pageNumber="369">Fig. 10A, B</figureCitation>
). At its ventral end, the lacrimal connects to the jugal posteriorly and the maxilla anteriorly. Posterodorsally, the lacrimal connects to the prefrontal, dorsomedially to the nasal, and anteriorly again to the maxilla. The lacrimal also forms the anteroventral portion of the orbital margin and the posterodorsal and posterior margins of the antorbital fenestra. The shass has a prominent anterolateral crest along its ventral half, and the portion of the antorbital fossa directly anterior to it is thus deeply excavated (
<figureCitation id="13308FBC31129F1B4326F98A3AA2F9F7" box="[574,686,1546,1570]" captionStart="Figure 10" captionStartId="14.[113,178,1772,1796]" captionTargetBox="[197,1376,144,1744]" captionTargetId="figure-5@14.[197,1376,144,1744]" captionTargetPageId="14" captionText="Figure 10. Saltoposuchus connectens referred specimen SMNS 12352.A, SMNS 12352a with the snout region exposed in angled right lateral view; B, interpretative drawing of (A); C, opposite side of SMNS 12352a exposing the posterior portion of the less maxilla and lacrimal in angled lateral view; D, interpretative drawing of (C); E, SMNS 12352b, carpus and manus in dorsal view; F, interpretative drawing of (E); G, SMNS 12352b in medial view; H, interpretative drawing of (G). Abbreviations; antfe, antorbital fenestra; antfo, antorbital fossa; dc, distal carpal; de, dentary; exp, extensor pit; fr, frontal; j, jugal; la, lacrimal; lad, lacrimal duct; mc, metacarpal; mx, maxilla; na, nasal; ph, phalanx; pm, premaxilla; prf, prefrontal; ra, radiale; ul, ulnare." figureDoi="http://doi.org/10.5281/zenodo.8426171" httpUri="https://zenodo.org/record/8426171/files/figure.png" pageId="15" pageNumber="369">Fig. 10A, B</figureCitation>
), similar to the condition seen in crocodylomorphs like
<taxonomicName id="4C0BE8BA31129F1B439BF9AA390BF9B4" authorityName=", Clark and Sues" authorityYear="2002" class="Reptilia" family="Sphenosuchidae" genus="Litargosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="369" phylum="Chordata" rank="species" species="leptorhynchus">
<emphasis id="B97F4F2B31129F1B439BF9AA390BF9B4" italics="true" pageId="15" pageNumber="369">Litargosuchus leptorhynchus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0BE8BA31129F1B4004F9C93A18F9B4" box="[284,532,1609,1633]" class="Reptilia" family="Sphenosuchidae" genus="Dromicosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="15" pageNumber="369" phylum="Chordata" rank="species" species="grallator">
<emphasis id="B97F4F2B31129F1B4004F9C93A18F9B4" box="[284,532,1609,1633]" italics="true" pageId="15" pageNumber="369">Dromicosuchus grallator</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0BE8BA31129F1B4331F9C93B08F9B4" box="[553,772,1609,1633]" genus="Carnufex" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="369" phylum="Chordata" rank="species" species="carolinensis">
<emphasis id="B97F4F2B31129F1B4331F9C93B08F9B4" box="[553,772,1609,1633]" italics="true" pageId="15" pageNumber="369">Carnufex carolinensis</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C0BE8BA31129F1B41A8F9E839ADF955" authorityName="Simmons" authorityYear="1965" box="[176,417,1640,1664]" class="Reptilia" family="Sphenosuchidae" genus="Dibothrosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="15" pageNumber="369" phylum="Chordata" rank="species" species="elaphros">
<emphasis id="B97F4F2B31129F1B41A8F9E839ADF955" box="[176,417,1640,1664]" italics="true" pageId="15" pageNumber="369">Dibothrosuchus elaphros</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9AEEC831129F1B40ACF9E83A9DF955" author="Clark JM &amp; Sues H-D" box="[436,657,1640,1664]" pageId="15" pageNumber="369" pagination="77 - 95" refId="ref30970" refString="Clark JM, Sues H-D. Two new basal crocodylomorph archosaurs from the Lower Jurassic and the monophyly of the Sphenosuchia. Zoological Journal of the Linnean Society 2002; 136: 77 - 95." type="journal article" year="2002">Clark and Sues 2002</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831129F1B43B8F9E838B9F975" author="Sues H-D &amp; Olsen PE &amp; Carter JG" pageId="15" pageNumber="369" pagination="329 - 43" refId="ref32828" refString="Sues H-D, Olsen PE, Carter JG et al. A new crocodylomorph archosaur from the Upper Triassic of North Carolina. Journal of Vertebrate Paleontology 2003; 23: 329 - 43." type="journal article" year="2003">
Sues
<emphasis id="B97F4F2B31129F1B43C0F9E93B06F955" box="[728,778,1640,1664]" italics="true" pageId="15" pageNumber="369">et al.</emphasis>
2003
</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831129F1B41DAF90839DDF975" author="Drymala SM &amp; Zanno LE" box="[194,465,1672,1696]" pageId="15" pageNumber="369" pagination="0157528" refId="ref31236" refString="Drymala SM, Zanno LE. Osteology of Carnufex carolinensis (Archosauria: Psuedosuchia) from the Pekin Formation of North Carolina and its implications for early crocodylomorph evolution. PLoS One 2016; 11: e 0157528." type="journal article" year="2016">Drymala and Zanno 2016</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831129F1B40C7F9083ADEF975" author="Ruebenstahl AA &amp; Klein MD &amp; Yi H" box="[479,722,1672,1696]" pageId="15" pageNumber="369" pagination="2463 - 556" refId="ref32564" refString="Ruebenstahl AA, Klein MD, Yi H et al. Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros. The Anatomical Record 2022; 305: 2463 - 556." type="journal article" year="2022">
Ruebenstahl
<emphasis id="B97F4F2B31129F1B437EF9083A94F975" box="[614,664,1672,1696]" italics="true" pageId="15" pageNumber="369">et al.</emphasis>
2022
</bibRefCitation>
). An anteriorly directed opening within a concavity on the posterior margin of the lacrimal shass near the tapering ventral end of the prefrontal probably represents the opening for the lacrimal duct, as was also indicated by
<bibRefCitation id="EF9AEEC831129F1B4062F8853A7BF8C8" author="Sereno PC &amp; Wild R" box="[378,631,1797,1821]" pageId="15" pageNumber="369" pagination="435 - 58" refId="ref32722" refString="Sereno PC, Wild R. Procompsognathus: theropod, ' thecodont' or both? Journal of Vertebrate Paleontology 1992; 12: 435 - 58." type="journal article" year="1992">Sereno and Wild (1992)</bibRefCitation>
, although this cannot be determined with certainty. The ventral foot of the shass extends posteroventrally to form a confluent connection with the jugal, similar to that of all other non-crocodyliform crocodylomorphs except
<taxonomicName id="4C0BE8BA31129F1B4088F8023A6AF84F" box="[400,614,1922,1946]" genus="Carnufex" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="369" phylum="Chordata" rank="species" species="carolinensis">
<emphasis id="B97F4F2B31129F1B4088F8023A6AF84F" box="[400,614,1922,1946]" italics="true" pageId="15" pageNumber="369">Carnufex carolinensis</emphasis>
</taxonomicName>
, which lacks a subcircular orbit (
<bibRefCitation id="EF9AEEC831129F1B4021F8213A4DF86C" author="Drymala SM &amp; Zanno LE" box="[313,577,1953,1977]" pageId="15" pageNumber="369" pagination="0157528" refId="ref31236" refString="Drymala SM, Zanno LE. Osteology of Carnufex carolinensis (Archosauria: Psuedosuchia) from the Pekin Formation of North Carolina and its implications for early crocodylomorph evolution. PLoS One 2016; 11: e 0157528." type="journal article" year="2016">Drymala and Zanno 2016</bibRefCitation>
).
</paragraph>
<paragraph id="8BB4933931129F1B424DFF0F3C08FADA" blockId="15.[825,1475,143,1295]" pageId="15" pageNumber="369">
The lacrimal forms a large contribution to the antorbital fossa both directly anterior to the lacrimal shass and along the anterodorsal process (
<figureCitation id="13308FBC31129F1B4501FF4E3C8BFF33" box="[1049,1159,206,230]" captionStart="Figure 10" captionStartId="14.[113,178,1772,1796]" captionTargetBox="[197,1376,144,1744]" captionTargetId="figure-5@14.[197,1376,144,1744]" captionTargetPageId="14" captionText="Figure 10. Saltoposuchus connectens referred specimen SMNS 12352.A, SMNS 12352a with the snout region exposed in angled right lateral view; B, interpretative drawing of (A); C, opposite side of SMNS 12352a exposing the posterior portion of the less maxilla and lacrimal in angled lateral view; D, interpretative drawing of (C); E, SMNS 12352b, carpus and manus in dorsal view; F, interpretative drawing of (E); G, SMNS 12352b in medial view; H, interpretative drawing of (G). Abbreviations; antfe, antorbital fenestra; antfo, antorbital fossa; dc, distal carpal; de, dentary; exp, extensor pit; fr, frontal; j, jugal; la, lacrimal; lad, lacrimal duct; mc, metacarpal; mx, maxilla; na, nasal; ph, phalanx; pm, premaxilla; prf, prefrontal; ra, radiale; ul, ulnare." figureDoi="http://doi.org/10.5281/zenodo.8426171" httpUri="https://zenodo.org/record/8426171/files/figure.png" pageId="15" pageNumber="369">Fig. 10A, B</figureCitation>
). The former contribution represents an anteroposteriorly narrow lamina, whereas the laưer contribution is considerable, entirely filling the dorsal half of the antorbital cavity at its greatest extent, near its posterior end. The antorbital fossa contribution of the anterodorsal process gradually reduces in height anteriorly, providing the process with a triangular outline in lateral view. This condition is similar to that seen in most non-crocodyliform crocodylomorphs, including
<taxonomicName id="4C0BE8BA31129F1B4221FE493C19FE34" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[825,1045,457,481]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="15" pageNumber="369" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B31129F1B4221FE493C19FE34" box="[825,1045,457,481]" italics="true" pageId="15" pageNumber="369">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0BE8BA31129F1B453AFE493D4BFE34" authority=", CM" authorityName="CM" box="[1058,1351,457,481]" class="Reptilia" family="Sphenosuchidae" genus="Dromicosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="15" pageNumber="369" phylum="Chordata" rank="species" species="grallator">
<emphasis id="B97F4F2B31129F1B453AFE493D1DFE34" box="[1058,1297,457,481]" italics="true" pageId="15" pageNumber="369">Dromicosuchus grallator</emphasis>
, CM
</taxonomicName>
29894, and
<taxonomicName id="4C0BE8BA31129F1B4221FE683C04FDD5" box="[825,1032,488,512]" genus="Carnufex" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="369" phylum="Chordata" rank="species" species="carolinensis">
<emphasis id="B97F4F2B31129F1B4221FE683C04FDD5" box="[825,1032,488,512]" italics="true" pageId="15" pageNumber="369">Carnufex carolinensis</emphasis>
</taxonomicName>
(e.g.
<bibRefCitation id="EF9AEEC831129F1B4523FE683CBFFDD5" author="Crush PJ" box="[1083,1203,488,512]" pageId="15" pageNumber="369" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831129F1B45A4FE683D86FDD5" author="Clark JM &amp; Sues H-D" box="[1212,1418,488,512]" pageId="15" pageNumber="369" pagination="77 - 95" refId="ref30970" refString="Clark JM, Sues H-D. Two new basal crocodylomorph archosaurs from the Lower Jurassic and the monophyly of the Sphenosuchia. Zoological Journal of the Linnean Society 2002; 136: 77 - 95." type="journal article" year="2002">Clark and Sues 2002</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831129F1B448BFE683BAEFDF5" author="Sues H-D &amp; Olsen PE &amp; Carter JG" pageId="15" pageNumber="369" pagination="329 - 43" refId="ref32828" refString="Sues H-D, Olsen PE, Carter JG et al. A new crocodylomorph archosaur from the Upper Triassic of North Carolina. Journal of Vertebrate Paleontology 2003; 23: 329 - 43." type="journal article" year="2003">
Sues
<emphasis id="B97F4F2B31129F1B4221FD883B65FDCA" box="[825,873,519,543]" italics="true" pageId="15" pageNumber="369">et al.</emphasis>
2003
</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831129F1B42B6FD873CB5FDCA" author="Drymala SM &amp; Zanno LE" box="[942,1209,519,543]" pageId="15" pageNumber="369" pagination="0157528" refId="ref31236" refString="Drymala SM, Zanno LE. Osteology of Carnufex carolinensis (Archosauria: Psuedosuchia) from the Pekin Formation of North Carolina and its implications for early crocodylomorph evolution. PLoS One 2016; 11: e 0157528." type="journal article" year="2016">Drymala and Zanno 2016</bibRefCitation>
), except for
<taxonomicName id="4C0BE8BA31129F1B4424FD873B7BFDEB" authorityName="Haughton" authorityYear="1915" class="Reptilia" genus="Sphenosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="369" phylum="Chordata" rank="species" species="acutus">
<emphasis id="B97F4F2B31129F1B4424FD873B7BFDEB" italics="true" pageId="15" pageNumber="369">Sphenosuchus acutus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0BE8BA31129F1B42AAFDA73C8CFDEA" box="[946,1152,551,575]" class="Reptilia" genus="Junggarsuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="369" phylum="Chordata" rank="species" species="sloani">
<emphasis id="B97F4F2B31129F1B42AAFDA73C8CFDEA" box="[946,1152,551,575]" italics="true" pageId="15" pageNumber="369">Junggarsuchus sloani</emphasis>
</taxonomicName>
, in which the antorbital fossa contribution of the anterodorsal process of the lacrimal is much more restricted (
<bibRefCitation id="EF9AEEC831129F1B42F4FDE53C7DFDA8" author="Walker AD" box="[1004,1137,613,637]" pageId="15" pageNumber="369" pagination="1 - 120" refId="ref33031" refString="Walker AD. A revision of Sphenosuchus acutus Haughton, a crocodylomorph reptile from the Elliot Formation (Late Triassic or Early Jurassic) of South Africa. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1990; 330: 1 - 120." type="journal article" year="1990">Walker 1990</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831129F1B4567FDE53D79FDA8" author="Ruebenstahl AA &amp; Klein MD &amp; Yi H" box="[1151,1397,613,637]" pageId="15" pageNumber="369" pagination="2463 - 556" refId="ref32564" refString="Ruebenstahl AA, Klein MD, Yi H et al. Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros. The Anatomical Record 2022; 305: 2463 - 556." type="journal article" year="2022">
Ruebenstahl
<emphasis id="B97F4F2B31129F1B441FFDE63D35FDA8" box="[1287,1337,613,637]" italics="true" pageId="15" pageNumber="369">et al.</emphasis>
2022
</bibRefCitation>
). Note that CM 29894 was originally referred to
<taxonomicName id="4C0BE8BA31129F1B45E1FD053BF1FD69" authority="(Clark et al. 2000)" baseAuthorityName="Clark" baseAuthorityYear="2000" class="Reptilia" family="Sphenosuchidae" genus="Hesperosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="15" pageNumber="369" phylum="Chordata" rank="species" species="agilis">
<emphasis id="B97F4F2B31129F1B45E1FD053DCEFD48" box="[1273,1474,645,669]" italics="true" pageId="15" pageNumber="369">Hesperosuchus agilis</emphasis>
(
<bibRefCitation id="EF9AEEC831129F1B425BFD243BFDFD69" author="Clark JM &amp; Sues H-D &amp; Berman DS" box="[835,1009,676,700]" pageId="15" pageNumber="369" pagination="683 - 704" refId="ref31006" refString="Clark JM, Sues H-D, Berman DS. A new specimen of Hesperosuchus agilis from the Upper Triassic of New Mexico and the interrelationships of basal crocodylomorph archosaurs. Journal of Vertebrate Paleontology 2000; 20: 683 - 704." type="journal article" year="2000">
Clark
<emphasis id="B97F4F2B31129F1B429CFD253BBAFD69" box="[900,950,676,700]" italics="true" pageId="15" pageNumber="369">et al.</emphasis>
2000
</bibRefCitation>
)
</taxonomicName>
. Since then, it has been suggested that this specimen, as well as an additional crocodylomorph specimen from the same locality, YPM 41198, might be referable to a separate taxon, based in part on an approximately 20 million year gap between these
<specimenCount id="9D0D58B031129F1B452EFCA23CDFFCEC" box="[1078,1235,801,825]" pageId="15" pageNumber="369" type="generic">two specimens</specimenCount>
, which come from the
<taxonomicName id="4C0BE8BA31129F1B4221FCC13BA6FC8C" baseAuthorityName="Fischer" baseAuthorityYear="1883" box="[825,938,833,857]" class="Reptilia" family="Coelophysidae" genus="Coelophysis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="15" pageNumber="369" phylum="Chordata" rank="genus">
<emphasis id="B97F4F2B31129F1B4221FCC13BA6FC8C" box="[825,938,833,857]" italics="true" pageId="15" pageNumber="369">Coelophysis</emphasis>
</taxonomicName>
ºuarry in the upper part of the Chinle Formation, and the
<typeStatus id="54B02D9B31129F1B4289FCE03BE1FCAD" box="[913,1005,864,888]" pageId="15" pageNumber="369">holotype</typeStatus>
of
<taxonomicName id="4C0BE8BA31129F1B450BFCE03D8CFCAD" authority=", AMNH FARB" authorityName="AMNH FARB" box="[1043,1408,864,888]" class="Reptilia" family="Sphenosuchidae" genus="Hesperosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="15" pageNumber="369" phylum="Chordata" rank="species" species="agilis">
<emphasis id="B97F4F2B31129F1B450BFCE03CD6FCAD" box="[1043,1242,864,888]" italics="true" pageId="15" pageNumber="369">Hesperosuchus agilis</emphasis>
, AMNH FARB
</taxonomicName>
6758, which derives from the Blue Mesa Member in the lower part of the Chinle Formation (
<bibRefCitation id="EF9AEEC831129F1B4545FC1F3CE4FC62" author="Nesbiu SJ" box="[1117,1256,927,951]" pageId="15" pageNumber="369" pagination="1 - 292" refId="ref32089" refString="Nesbiu SJ. The early evolution of archosaurs: relationships and the origin of major clades. Bulletin of the American Museum of Natural History 2011; 352: 1 - 292." type="journal article" year="2011">Nesbiư 2011</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831129F1B45E2FC1F3DBDFC62" author="Irmis RB &amp; Nesbiu SJ &amp; Sues H-D" box="[1274,1457,927,951]" pageId="15" pageNumber="369" pagination="275 - 302" refId="ref31747" refString="Irmis RB, Nesbiu SJ, Sues H-D. Early Crocodylomorpha. Geological Society, London, Special Publications 2013; 379: 275 - 302." type="journal article" year="2013">
Irmis
<emphasis id="B97F4F2B31129F1B4424FC1F3D7EFC62" box="[1340,1394,927,951]" italics="true" pageId="15" pageNumber="369">et al.</emphasis>
2013
</bibRefCitation>
). Although others have suggested that there are no distinct morphological distinctions between AMNH FARB 6758 and CM 29894 (
<bibRefCitation id="EF9AEEC831129F1B4296FC7D3C46FBC0" author="Leardi JM &amp; Pol D &amp; Clark JM" box="[910,1098,1021,1045]" pageId="15" pageNumber="369" pagination="2801" refId="ref31922" refString="Leardi JM, Pol D, Clark JM. Detailed anatomy of the braincase of Macelognathus vagans Marsh, 1884 (Archosauria, Crocodylomorpha) using high resolution tomography and new insights on basal crocodylomorph phylogeny. PeerJ 2017; 5: e 2801." type="journal article" year="2017">
Leardi
<emphasis id="B97F4F2B31129F1B42C2FC7D3C01FBC0" box="[986,1037,1021,1045]" italics="true" pageId="15" pageNumber="369">et al.</emphasis>
2017
</bibRefCitation>
), to avoid confusion separate morphological comparisons are made here to CM 29894 and YPM 41198, and to
<taxonomicName id="4C0BE8BA31129F1B42D7FBBB3C99FB86" box="[975,1173,1083,1107]" class="Reptilia" family="Sphenosuchidae" genus="Hesperosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="15" pageNumber="369" phylum="Chordata" rank="species" species="agilis">
<emphasis id="B97F4F2B31129F1B42D7FBBB3C99FB86" box="[975,1173,1083,1107]" italics="true" pageId="15" pageNumber="369">Hesperosuchus agilis</emphasis>
</taxonomicName>
as represented exclusively by the
<typeStatus id="54B02D9B31129F1B4279FBDA3BB1FBA7" box="[865,957,1114,1138]" pageId="15" pageNumber="369">holotype</typeStatus>
AMNH FARB 6758. The exact anterior extent of the lacrimal and its contribution to the antorbital fossa is hard to establish in SMNS 12352a, but it appears to reach to close to the anterior end of the antorbital fenestra. In both lacrimals of SMNS 12591a, the contribution to the antorbital fossa is almost completely missing.
</paragraph>
<paragraph id="8BB4933931129F044221FAB03AEBFE56" blockId="15.[825,1475,1328,1979]" lastBlockId="16.[113,763,144,387]" lastPageId="16" lastPageNumber="370" pageId="15" pageNumber="369">
<emphasis id="B97F4F2B31129F1B4221FAB03BAEFA9D" box="[825,930,1328,1352]" italics="true" pageId="15" pageNumber="369">Preffontal:</emphasis>
The prefrontal is also preserved in SMNS 12352a and SMNS 12591a. The right prefrontal of SMNS 12352a is well preserved but incomplete, only preserving its anteriormost and ventral portions (
<figureCitation id="13308FBC31129F1B4538FA0E3C99FA73" box="[1056,1173,1422,1447]" captionStart="Figure 10" captionStartId="14.[113,178,1772,1796]" captionTargetBox="[197,1376,144,1744]" captionTargetId="figure-5@14.[197,1376,144,1744]" captionTargetPageId="14" captionText="Figure 10. Saltoposuchus connectens referred specimen SMNS 12352.A, SMNS 12352a with the snout region exposed in angled right lateral view; B, interpretative drawing of (A); C, opposite side of SMNS 12352a exposing the posterior portion of the less maxilla and lacrimal in angled lateral view; D, interpretative drawing of (C); E, SMNS 12352b, carpus and manus in dorsal view; F, interpretative drawing of (E); G, SMNS 12352b in medial view; H, interpretative drawing of (G). Abbreviations; antfe, antorbital fenestra; antfo, antorbital fossa; dc, distal carpal; de, dentary; exp, extensor pit; fr, frontal; j, jugal; la, lacrimal; lad, lacrimal duct; mc, metacarpal; mx, maxilla; na, nasal; ph, phalanx; pm, premaxilla; prf, prefrontal; ra, radiale; ul, ulnare." figureDoi="http://doi.org/10.5281/zenodo.8426171" httpUri="https://zenodo.org/record/8426171/files/figure.png" pageId="15" pageNumber="369">Fig. 10A, B</figureCitation>
). Both prefrontals of SMNS 12591a are virtually complete, but their anterior and ventral portions are poorly preserved due to compression (
<figureCitation id="13308FBC31129F1B4434FA4D3D66FA30" box="[1324,1386,1485,1509]" captionStart="Figure 8" captionStartId="9.[129,194,1772,1796]" captionTargetBox="[178,1429,144,1744]" captionTargetId="figure-6@9.[174,1429,144,1744]" captionTargetPageId="9" captionText="Figure 8. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in right lateral view; B, interpretative drawing of A; C, less lateral view; D, interpretative drawing of (C). Abbreviations:an, angular; cc, crista cranii; de, dentary; fr, frontal; j, jugal; la, lacrimal; ls, laterosphenoid; mx, maxilla; na, nasal; pa, parietal; pl, palatine; po, postorbital; prf, prefrontal; pt, pterygoid; q, quadrate; qj, quadratojugal; sa, surangular; sq, squamosal." figureDoi="http://doi.org/10.5281/zenodo.8426167" httpUri="https://zenodo.org/record/8426167/files/figure.png" pageId="15" pageNumber="369">Figs 8</figureCitation>
,
<figureCitation id="13308FBC31129F1B446EFA4D3D8FFA30" box="[1398,1411,1485,1509]" captionStart="Figure 9" captionStartId="10.[113,178,1772,1796]" captionTargetBox="[222,1352,144,1744]" captionTargetId="figure-6@10.[220,1352,144,1744]" captionTargetPageId="10" captionText="Figure 9. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in dorsal view; B, interpretative drawing of (A); C, ventral view; D, interpretative drawing of (C). Abbreviations:an, angular; antfo, antorbital fossa; bpp, basipterygoid process; crb, ceratobranchial I; de, dentary; ect, ectopterygoid; fr, frontal; j, jugal; la, lacrimal; mx, maxilla; na, nasal; pa, parietal; pbs, parabasisphenoid; pl, palatine; plf, palatine fossa; po, postorbital; pra, pre-articular; prf, prefrontal; pt, pterygoid; ptmr, pterygoid medial rim; sa, surangular; spl, splenial; sq, squamosal; vo, vomer." figureDoi="http://doi.org/10.5281/zenodo.8426169" httpUri="https://zenodo.org/record/8426169/files/figure.png" pageId="15" pageNumber="369">9</figureCitation>
). The prefrontal forms most of the anterior margin of the orbit and articulates with the lacrimal anteriorly and the nasal and frontal dorsomedially. In SMNS 12591a, the prefrontal also connects extensively with the palate, and although this is also the case in several other non-crocodyliform crocodylomorphs (e.g.
<bibRefCitation id="EF9AEEC831129F1B446FF9EA3B61F974" author="Nesbiu SJ" pageId="15" pageNumber="369" pagination="1 - 292" refId="ref32089" refString="Nesbiu SJ. The early evolution of archosaurs: relationships and the origin of major clades. Bulletin of the American Museum of Natural History 2011; 352: 1 - 292." type="journal article" year="2011">Nesbiư 2011</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831129F1B4262F9093C61F974" author="Ruebenstahl AA &amp; Klein MD &amp; Yi H" box="[890,1133,1673,1697]" pageId="15" pageNumber="369" pagination="2463 - 556" refId="ref32564" refString="Ruebenstahl AA, Klein MD, Yi H et al. Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros. The Anatomical Record 2022; 305: 2463 - 556." type="journal article" year="2022">
Ruebenstahl
<emphasis id="B97F4F2B31129F1B451AF90A3C3FF974" box="[1026,1075,1673,1697]" italics="true" pageId="15" pageNumber="369">et al.</emphasis>
2022
</bibRefCitation>
), it cannot be excluded that this is the result of the dorsoventral compression in this specimen. The ventral process of the prefrontal forms a thin strip of bone that tapers ventrally and reaches until a level just ventral to the dorsoventral midpoint of the orbit in lateral view. In dorsal view, the prefrontal has a markedly convex lateral margin, which is best observed on the right side of SMNS 12591a. A similar convexity is also present in
<taxonomicName id="4C0BE8BA31129F1B42F3F8E43CB9F8A9" box="[1003,1205,1892,1916]" class="Reptilia" genus="Junggarsuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="369" phylum="Chordata" rank="species" species="sloani">
<emphasis id="B97F4F2B31129F1B42F3F8E43CB9F8A9" box="[1003,1205,1892,1916]" italics="true" pageId="15" pageNumber="369">Junggarsuchus sloani</emphasis>
</taxonomicName>
, which was described as a prefrontal overhang of the orbit in that taxon (
<bibRefCitation id="EF9AEEC831129F1B4417F8043B61F86E" author="Ruebenstahl AA &amp; Klein MD &amp; Yi H" pageId="15" pageNumber="369" pagination="2463 - 556" refId="ref32564" refString="Ruebenstahl AA, Klein MD, Yi H et al. Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros. The Anatomical Record 2022; 305: 2463 - 556." type="journal article" year="2022">
Ruebenstahl
<emphasis id="B97F4F2B31129F1B448CF8043DCFF849" box="[1428,1475,1924,1948]" italics="true" pageId="15" pageNumber="369">et al.</emphasis>
2022
</bibRefCitation>
). The medial margin is more difficult to discern due to poor preservation, but it is roughly straight in dorsal view. Posteriorly, the prefrontal reaches until just anterior to the dorsal apex of the orbit. As was also indicated by
<bibRefCitation id="EF9AEEC8310D9F0440B0FF4F3AADFF32" author="Sereno PC &amp; Wild R" box="[424,673,207,231]" pageId="16" pageNumber="370" pagination="435 - 58" refId="ref32722" refString="Sereno PC, Wild R. Procompsognathus: theropod, ' thecodont' or both? Journal of Vertebrate Paleontology 1992; 12: 435 - 58." type="journal article" year="1992">Sereno and Wild (1992)</bibRefCitation>
, the prefrontal extends similarly far posteriorly on the ventral side of the skull roof. The presence of a considerable medial extension of the prefrontal on the anterior side of the orbit is widely present among crocodylomorphs (
<bibRefCitation id="EF9AEEC8310D9F04409AFECC3A0AFEB1" author="Nesbiu SJ" box="[386,518,332,356]" pageId="16" pageNumber="370" pagination="1 - 292" refId="ref32089" refString="Nesbiu SJ. The early evolution of archosaurs: relationships and the origin of major clades. Bulletin of the American Museum of Natural History 2011; 352: 1 - 292." type="journal article" year="2011">Nesbiư 2011</bibRefCitation>
: ch. 40) and this feature can also be observed on the right prefrontal of SMNS 12591a.
</paragraph>
<paragraph id="8BB49339310D9F044169FE253924FB58" blockId="16.[113,763,421,1980]" pageId="16" pageNumber="370">
<emphasis id="B97F4F2B310D9F044169FE2538CCFE68" box="[113,192,421,445]" italics="true" pageId="16" pageNumber="370">Frontal:</emphasis>
Disarticulated but virtually complete frontals are preserved and exposed in ventral view in SMNS 12597a (
<figureCitation id="13308FBC310D9F0443C8FE443883FE2E" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="16" pageNumber="370">Figs 2A</figureCitation>
,
<figureCitation id="13308FBC310D9F044187FE6338C6FE2E" box="[159,202,483,507]" captionStart="Figure 11" captionStartId="17.[127,192,1672,1696]" captionTargetBox="[131,1404,147,1640]" captionTargetId="figure-6@17.[127,1407,144,1644]" captionTargetPageId="17" captionText="Figure 11. Details of the cranial anatomy of Saltoposuchus connectens. A, frontals of SMNS 12597a in ventral view; B, less squamosal of SMNS 12596a in dorsal view; C, right squamosal of SMNS 12597b in ventral view; D, less quadrate of SMNS 12597b in posterolateral view; E, less quadrate of SMNS 12596a in anterolateral view; F, enlarged anterior maxillary tooth of SMNS 12352a in angled lateral view.Abbreviations: adp, anterodorsal process; ap, anterior process; dh, dorsal head; fr, frontal; iqf, internal quadrate fenestra; l., less; lcr, longitudinal crest; lr, longitudinal ridge; olb, olfactory bulb; otfl, otoccipital flange; paf, parietal facet; pmp, posteromedial process; pof, postorbital facet; ppeg, posterior peg; prff, prefrontal facet; ptfl, post-temporal flange; ptp, pterygoid process; ptra, pterygoid ramus; qf, quadrate facet; r., right; stfo, supratemporal fossa; vdp, ventrally directed process; vee, ventral end." figureDoi="http://doi.org/10.5281/zenodo.8426173" httpUri="https://zenodo.org/record/8426173/files/figure.png" pageId="16" pageNumber="370">11A</figureCitation>
). Both frontals are also virtually complete in SMNS 12591a and the anterior ends of the frontals are preserved in SMNS 12352a (
<figureCitation id="13308FBC310D9F044004FDA2398BFDEC" box="[284,391,546,570]" captionStart="Figure 9" captionStartId="10.[113,178,1772,1796]" captionTargetBox="[222,1352,144,1744]" captionTargetId="figure-6@10.[220,1352,144,1744]" captionTargetPageId="10" captionText="Figure 9. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in dorsal view; B, interpretative drawing of (A); C, ventral view; D, interpretative drawing of (C). Abbreviations:an, angular; antfo, antorbital fossa; bpp, basipterygoid process; crb, ceratobranchial I; de, dentary; ect, ectopterygoid; fr, frontal; j, jugal; la, lacrimal; mx, maxilla; na, nasal; pa, parietal; pbs, parabasisphenoid; pl, palatine; plf, palatine fossa; po, postorbital; pra, pre-articular; prf, prefrontal; pt, pterygoid; ptmr, pterygoid medial rim; sa, surangular; spl, splenial; sq, squamosal; vo, vomer." figureDoi="http://doi.org/10.5281/zenodo.8426169" httpUri="https://zenodo.org/record/8426169/files/figure.png" pageId="16" pageNumber="370">Figs 9A, B</figureCitation>
,
<figureCitation id="13308FBC310D9F04408CFDA239D7FDEC" box="[404,475,546,570]" captionStart="Figure 10" captionStartId="14.[113,178,1772,1796]" captionTargetBox="[197,1376,144,1744]" captionTargetId="figure-5@14.[197,1376,144,1744]" captionTargetPageId="14" captionText="Figure 10. Saltoposuchus connectens referred specimen SMNS 12352.A, SMNS 12352a with the snout region exposed in angled right lateral view; B, interpretative drawing of (A); C, opposite side of SMNS 12352a exposing the posterior portion of the less maxilla and lacrimal in angled lateral view; D, interpretative drawing of (C); E, SMNS 12352b, carpus and manus in dorsal view; F, interpretative drawing of (E); G, SMNS 12352b in medial view; H, interpretative drawing of (G). Abbreviations; antfe, antorbital fenestra; antfo, antorbital fossa; dc, distal carpal; de, dentary; exp, extensor pit; fr, frontal; j, jugal; la, lacrimal; lad, lacrimal duct; mc, metacarpal; mx, maxilla; na, nasal; ph, phalanx; pm, premaxilla; prf, prefrontal; ra, radiale; ul, ulnare." figureDoi="http://doi.org/10.5281/zenodo.8426171" httpUri="https://zenodo.org/record/8426171/files/figure.png" pageId="16" pageNumber="370">10A, B</figureCitation>
). The frontal is an elongate element with a straight medial margin, which articulates with its antimere, and a more complex lateral margin that for most of its length is distinctly concave in dorsal and ventral view. This concavity forms the dorsal margin of the orbit. At its anterior end, the frontal mostly articulates with the nasal, but also with the prefrontal on its lateral side. As outlined above, the articulation between frontals and nasals is V-shaped, with each frontal possessing a distinctly medially tapering anterior end. The frontalnasal region is badly broken in SMNS 12591a, but the V-shaped suture between the elements can still be discerned. The frontal articulates with the postorbital on its posterolateral end, and with the parietal along its posterior margin. This last articulation is W-shaped in dorsal view in SMNS 12597a and SMNS 12591a, similar to that seen in
<taxonomicName id="4C0BE8BA310D9F04406CFC583A59FC25" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[372,597,984,1008]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="16" pageNumber="370" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B310D9F04406CFC583A59FC25" box="[372,597,984,1008]" italics="true" pageId="16" pageNumber="370">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0BE8BA310D9F04437FFC5838C9FBC5" class="Reptilia" family="Sphenosuchidae" genus="Dromicosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="16" pageNumber="370" phylum="Chordata" rank="species" species="grallator">
<emphasis id="B97F4F2B310D9F04437FFC5838C9FBC5" italics="true" pageId="16" pageNumber="370">Dromicosuchus grallator</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0BE8BA310D9F0441C4FC7839D9FBC5" authorityName="Simmons" authorityYear="1965" box="[220,469,1016,1040]" class="Reptilia" family="Sphenosuchidae" genus="Dibothrosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="16" pageNumber="370" phylum="Chordata" rank="species" species="elaphros">
<emphasis id="B97F4F2B310D9F0441C4FC7839D9FBC5" box="[220,469,1016,1040]" italics="true" pageId="16" pageNumber="370">Dibothrosuchus elaphros</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C0BE8BA310D9F04433BFC783AF6FBC5" box="[547,762,1016,1040]" genus="Almadasuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="16" pageNumber="370" phylum="Chordata" rank="species" species="figarii">
<emphasis id="B97F4F2B310D9F04433BFC783AF6FBC5" box="[547,762,1016,1040]" italics="true" pageId="16" pageNumber="370">Almadasuchus figarii</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9AEEC8310D9F044164FB9738FBFBFA" author="Crush PJ" box="[124,247,1047,1071]" pageId="16" pageNumber="370" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310D9F04401DFB9739A4FBFA" author="Sues H-D &amp; Olsen PE &amp; Carter JG" box="[261,424,1047,1071]" pageId="16" pageNumber="370" pagination="329 - 43" refId="ref32828" refString="Sues H-D, Olsen PE, Carter JG et al. A new crocodylomorph archosaur from the Upper Triassic of North Carolina. Journal of Vertebrate Paleontology 2003; 23: 329 - 43." type="journal article" year="2003">
Sues
<emphasis id="B97F4F2B310D9F044024FB983961FBFA" box="[316,365,1047,1071]" italics="true" pageId="16" pageNumber="370">et al.</emphasis>
2003
</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310D9F0440AEFB973A60FBFA" author="Leardi JM &amp; Pol D &amp; Clark JM" box="[438,620,1047,1071]" pageId="16" pageNumber="370" pagination="48 - 73" refId="ref31967" refString="Leardi JM, Pol D, Clark JM. Braincase anatomy of Almadasuchus figarii (Archosauria, Crocodylomorpha) and a review of the cranial pneumaticity in the origins of Crocodylomorpha. Journal of Anatomy 2020; 237: 48 - 73." type="journal article" year="2020">
Leardi
<emphasis id="B97F4F2B310D9F044318FB983A3DFBFA" box="[512,561,1047,1071]" italics="true" pageId="16" pageNumber="370">et al.</emphasis>
2020
</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310D9F044362FB9738D7FB9A" author="Ruebenstahl AA &amp; Klein MD &amp; Yi H" pageId="16" pageNumber="370" pagination="2463 - 556" refId="ref32564" refString="Ruebenstahl AA, Klein MD, Yi H et al. Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros. The Anatomical Record 2022; 305: 2463 - 556." type="journal article" year="2022">
Ruebenstahl
<emphasis id="B97F4F2B310D9F044169FBB738ADFB9B" box="[113,161,1078,1102]" italics="true" pageId="16" pageNumber="370">et al.</emphasis>
2022
</bibRefCitation>
). The frontal is transversely widest near its posterior end, where it articulates laterally with the dorsomedial process of the postorbital.
</paragraph>
<paragraph id="8BB49339310D9F044195FB143A60F935" blockId="16.[113,763,421,1980]" pageId="16" pageNumber="370">
The dorsal surface of the frontals is mostly flat, but a shallow, straight, longitudinal gulley is present directly medial to the orbital margin and, consequently, the surface directly medial to this gulley is convex. A similar morphology is also seen in
<taxonomicName id="4C0BE8BA310D9F044181FA92398EFAFF" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[153,386,1298,1322]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="16" pageNumber="370" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B310D9F044181FA92398EFAFF" box="[153,386,1298,1322]" italics="true" pageId="16" pageNumber="370">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0BE8BA310D9F044083FA923AB7FAFF" authorityName=", Clark and Sues" authorityYear="2002" box="[411,699,1298,1322]" class="Reptilia" family="Sphenosuchidae" genus="Litargosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="16" pageNumber="370" phylum="Chordata" rank="species" species="leptorhynchus">
<emphasis id="B97F4F2B310D9F044083FA923AB7FAFF" box="[411,699,1298,1322]" italics="true" pageId="16" pageNumber="370">Litargosuchus leptorhynchus</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C0BE8BA310D9F044169FAB13952FA9C" box="[113,350,1329,1353]" class="Reptilia" family="Sphenosuchidae" genus="Dromicosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="16" pageNumber="370" phylum="Chordata" rank="species" species="grallator">
<emphasis id="B97F4F2B310D9F044169FAB13952FA9C" box="[113,350,1329,1353]" italics="true" pageId="16" pageNumber="370">Dromicosuchus grallator</emphasis>
</taxonomicName>
(NHMUK PV R 37731,
<bibRefCitation id="EF9AEEC8310D9F044379FAB138A9FABD" author="Clark JM &amp; Sues H-D" pageId="16" pageNumber="370" pagination="77 - 95" refId="ref30970" refString="Clark JM, Sues H-D. Two new basal crocodylomorph archosaurs from the Lower Jurassic and the monophyly of the Sphenosuchia. Zoological Journal of the Linnean Society 2002; 136: 77 - 95." type="journal article" year="2002">Clark and Sues 2002</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310D9F0441A8FAD03940FABD" author="Sues H-D &amp; Olsen PE &amp; Carter JG" box="[176,332,1360,1384]" pageId="16" pageNumber="370" pagination="329 - 43" refId="ref32828" refString="Sues H-D, Olsen PE, Carter JG et al. A new crocodylomorph archosaur from the Upper Triassic of North Carolina. Journal of Vertebrate Paleontology 2003; 23: 329 - 43." type="journal article" year="2003">
Sues
<emphasis id="B97F4F2B310D9F0441FDFAD13918FABD" box="[229,276,1360,1384]" italics="true" pageId="16" pageNumber="370">et al.</emphasis>
2003
</bibRefCitation>
), whereas a more distinct, medial longitudinal bulge or ridge is present in
<taxonomicName id="4C0BE8BA310D9F0440D7FAF03A9BFA5D" box="[463,663,1392,1416]" class="Reptilia" family="Sphenosuchidae" genus="Hesperosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="16" pageNumber="370" phylum="Chordata" rank="species" species="agilis">
<emphasis id="B97F4F2B310D9F0440D7FAF03A9BFA5D" box="[463,663,1392,1416]" italics="true" pageId="16" pageNumber="370">Hesperosuchus agilis</emphasis>
</taxonomicName>
(AMNH FARB 6758) and non-crocodyliform crocodylomorphs that are generally found to be more closely related to crocodyliforms (
<bibRefCitation id="EF9AEEC8310D9F044164FA4E3905FA30" author="Colbert EH" box="[124,265,1485,1510]" pageId="16" pageNumber="370" pagination="561 - 92" refId="ref31083" refString="Colbert EH. A pseudosuchian reptile from Arizona. Bulletin of the American Museum of Natural History 1952; 99: 561 - 92." type="journal article" year="1952">Colbert 1952</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310D9F044000FA4E39ADFA33" author="Nesbiu SJ" box="[280,417,1486,1510]" pageId="16" pageNumber="370" pagination="1 - 292" refId="ref32089" refString="Nesbiu SJ. The early evolution of archosaurs: relationships and the origin of major clades. Bulletin of the American Museum of Natural History 2011; 352: 1 - 292." type="journal article" year="2011">Nesbiư 2011</bibRefCitation>
: ch. 42). The posterolateral end of the less frontal in SMNS 12591a bears a clear concavity on its dorsal surface, which is anteriorly rounded. This concavity was also indicated by
<bibRefCitation id="EF9AEEC8310D9F04404EF9AC3A5BF991" author="Sereno PC &amp; Wild R" box="[342,599,1580,1604]" pageId="16" pageNumber="370" pagination="435 - 58" refId="ref32722" refString="Sereno PC, Wild R. Procompsognathus: theropod, ' thecodont' or both? Journal of Vertebrate Paleontology 1992; 12: 435 - 58." type="journal article" year="1992">Sereno and Wild (1992)</bibRefCitation>
, and represents the contribution of the frontal to the supratemporal fossa. This frontal contribution is also present in other non-crocodyliform crocodylomorphs (e.g.
<bibRefCitation id="EF9AEEC8310D9F044044F90A39E9F974" author="Colbert EH" box="[348,485,1673,1698]" pageId="16" pageNumber="370" pagination="561 - 92" refId="ref31083" refString="Colbert EH. A pseudosuchian reptile from Arizona. Bulletin of the American Museum of Natural History 1952; 99: 561 - 92." type="journal article" year="1952">Colbert 1952</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310D9F0440F7F90A3A7CF977" author="Walker AD" box="[495,624,1674,1698]" pageId="16" pageNumber="370" pagination="1 - 120" refId="ref33031" refString="Walker AD. A revision of Sphenosuchus acutus Haughton, a crocodylomorph reptile from the Elliot Formation (Late Triassic or Early Jurassic) of South Africa. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1990; 330: 1 - 120." type="journal article" year="1990">Walker 1990</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310D9F044362F90A38D1F914" author="Ruebenstahl AA &amp; Klein MD &amp; Yi H" pageId="16" pageNumber="370" pagination="2463 - 556" refId="ref32564" refString="Ruebenstahl AA, Klein MD, Yi H et al. Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros. The Anatomical Record 2022; 305: 2463 - 556." type="journal article" year="2022">
Ruebenstahl
<emphasis id="B97F4F2B310D9F044169F92A38AEF914" box="[113,162,1705,1729]" italics="true" pageId="16" pageNumber="370">et al.</emphasis>
2022
</bibRefCitation>
), except for
<taxonomicName id="4C0BE8BA310D9F04407BF9293A33F914" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[355,575,1705,1729]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="16" pageNumber="370" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B310D9F04407BF9293A33F914" box="[355,575,1705,1729]" italics="true" pageId="16" pageNumber="370">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0BE8BA310D9F04436CF92938FBF935" authorityName=", Clark and Sues" authorityYear="2002" class="Reptilia" family="Sphenosuchidae" genus="Litargosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="16" pageNumber="370" phylum="Chordata" rank="species" species="leptorhynchus">
<emphasis id="B97F4F2B310D9F04436CF92938FBF935" italics="true" pageId="16" pageNumber="370">Litargosuchus leptorhynchus</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9AEEC8310D9F04401FF948398CF935" author="Crush PJ" box="[263,384,1736,1760]" pageId="16" pageNumber="370" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310D9F044092F9483A50F935" author="Clark JM &amp; Sues H-D" box="[394,604,1736,1760]" pageId="16" pageNumber="370" pagination="77 - 95" refId="ref30970" refString="Clark JM, Sues H-D. Two new basal crocodylomorph archosaurs from the Lower Jurassic and the monophyly of the Sphenosuchia. Zoological Journal of the Linnean Society 2002; 136: 77 - 95." type="journal article" year="2002">Clark and Sues 2002</bibRefCitation>
).
</paragraph>
<paragraph id="8BB49339310D9F044195F9673BD3FD0E" blockId="16.[113,763,421,1980]" lastBlockId="16.[810,1460,143,731]" pageId="16" pageNumber="370">
The ventral surface of the frontals of SMNS 12597a is characterized by a distinct longitudinal ridge, which runs near and parallel to the orbital rim of the bone. The medial edge of the bone is also raised ventrally, and between both ridges the excavated olfactory tract is formed (
<figureCitation id="13308FBC310D9F0440D7F8E53A2DF8A8" box="[463,545,1893,1917]" captionStart="Figure 11" captionStartId="17.[127,192,1672,1696]" captionTargetBox="[131,1404,147,1640]" captionTargetId="figure-6@17.[127,1407,144,1644]" captionTargetPageId="17" captionText="Figure 11. Details of the cranial anatomy of Saltoposuchus connectens. A, frontals of SMNS 12597a in ventral view; B, less squamosal of SMNS 12596a in dorsal view; C, right squamosal of SMNS 12597b in ventral view; D, less quadrate of SMNS 12597b in posterolateral view; E, less quadrate of SMNS 12596a in anterolateral view; F, enlarged anterior maxillary tooth of SMNS 12352a in angled lateral view.Abbreviations: adp, anterodorsal process; ap, anterior process; dh, dorsal head; fr, frontal; iqf, internal quadrate fenestra; l., less; lcr, longitudinal crest; lr, longitudinal ridge; olb, olfactory bulb; otfl, otoccipital flange; paf, parietal facet; pmp, posteromedial process; pof, postorbital facet; ppeg, posterior peg; prff, prefrontal facet; ptfl, post-temporal flange; ptp, pterygoid process; ptra, pterygoid ramus; qf, quadrate facet; r., right; stfo, supratemporal fossa; vdp, ventrally directed process; vee, ventral end." figureDoi="http://doi.org/10.5281/zenodo.8426173" httpUri="https://zenodo.org/record/8426173/files/figure.png" pageId="16" pageNumber="370">Fig 11A</figureCitation>
). Within this tract, a subcircular concavity just anterior to the transversely narrowest point of the frontal forms the olfactory bulb (
<bibRefCitation id="EF9AEEC8310D9F04437BF8233AE6F869" author="Walker AD" box="[611,746,1955,1980]" pageId="16" pageNumber="370" pagination="1 - 120" refId="ref33031" refString="Walker AD. A revision of Sphenosuchus acutus Haughton, a crocodylomorph reptile from the Elliot Formation (Late Triassic or Early Jurassic) of South Africa. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1990; 330: 1 - 120." type="journal article" year="1990">Walker 1990</bibRefCitation>
). A distinct excavation on the anterolateral margin of the ventral surface of the frontal forms the facet for the prefrontal. The anterolateral facet for the prefrontal and the orbital rim are separated by a triangular process that is directed somewhat ventrally. A similarly shaped process is also present in
<taxonomicName id="4C0BE8BA310D9F0445ECFE8D3B42FE91" baseAuthorityName="Crush" baseAuthorityYear="1984" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="16" pageNumber="370" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B310D9F0445ECFE8D3B42FE91" italics="true" pageId="16" pageNumber="370">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0BE8BA310D9F04429CFEAC3C5CFE91" authorityName="Haughton" authorityYear="1915" box="[900,1104,300,324]" class="Reptilia" genus="Sphenosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="16" pageNumber="370" phylum="Chordata" rank="species" species="acutus">
<emphasis id="B97F4F2B310D9F04429CFEAC3C5CFE91" box="[900,1104,300,324]" italics="true" pageId="16" pageNumber="370">Sphenosuchus acutus</emphasis>
</taxonomicName>
(NHMUK PV R 38290b,
<bibRefCitation id="EF9AEEC8310D9F044473FEAC3B52FEB1" author="Walker AD" pageId="16" pageNumber="370" pagination="1 - 120" refId="ref33031" refString="Walker AD. A revision of Sphenosuchus acutus Haughton, a crocodylomorph reptile from the Elliot Formation (Late Triassic or Early Jurassic) of South Africa. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1990; 330: 1 - 120." type="journal article" year="1990">Walker 1990</bibRefCitation>
). On the posterolateral margin of this process, a small but distinct groove is present, which forms an additional articulation facet for the prefrontal. The ventral surface of the posterior end on the right frontal forms a clear facet for the parietal, which thus slightly overlapped the frontal ventrally here. As such, the articulation between the frontal and parietal was complex, with the frontal overlying the parietal dorsally, except for a medial peg on the frontal that articulated on the ventral side of the parietal, as was also described by
<bibRefCitation id="EF9AEEC8310D9F04453FFDC63CBBFD8B" author="Crush PJ" box="[1063,1207,582,606]" pageId="16" pageNumber="370" refId="ref31157" refString="Crush PJ. An early, terrestrial, crocodile from South Wales. Unpublished Ph. D. Thesis, University College London, 1980." type="book" year="1980">Crush (1980)</bibRefCitation>
. The wide posterolateral end of the frontal forms a thin articulation facet for the postorbital, which thus articulated on the ventral side of this frontal process. The posterolateral end of the frontal tapers and terminates in a sharp tip.
</paragraph>
<paragraph id="8BB49339310D9F044232FC803D27F9F2" blockId="16.[810,1460,768,1575]" pageId="16" pageNumber="370">
<emphasis id="B97F4F2B310D9F044232FC803B72FCCD" box="[810,894,768,792]" italics="true" pageId="16" pageNumber="370">Parietal:</emphasis>
A partial parietal is preserved in SMNS 12597b. This element was originally identified as a quadratojugal by
<bibRefCitation id="EF9AEEC8310D9F044475FCA03B78FC82" author="Huene FR" pageId="16" pageNumber="370" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene (1921)</bibRefCitation>
, but later correctly reinterpreted as the posterior half of a less parietal exposed in dorsal view by
<bibRefCitation id="EF9AEEC8310D9F0445A1FCDE3D44FCA3" author="Crush PJ" box="[1209,1352,862,886]" pageId="16" pageNumber="370" refId="ref31157" refString="Crush PJ. An early, terrestrial, crocodile from South Wales. Unpublished Ph. D. Thesis, University College London, 1980." type="book" year="1980">Crush (1980)</bibRefCitation>
, including the posterolateral process (
<figureCitation id="13308FBC310D9F044527FCFD3C85FC40" box="[1087,1161,893,917]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="16" pageNumber="370">Fig. 2C</figureCitation>
). The anterior halves of both parietals are also present in SMNS 12591a (
<figureCitation id="13308FBC310D9F04441EFC1D3D7DFC61" box="[1286,1393,925,949]" captionStart="Figure 9" captionStartId="10.[113,178,1772,1796]" captionTargetBox="[222,1352,144,1744]" captionTargetId="figure-6@10.[220,1352,144,1744]" captionTargetPageId="10" captionText="Figure 9. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in dorsal view; B, interpretative drawing of (A); C, ventral view; D, interpretative drawing of (C). Abbreviations:an, angular; antfo, antorbital fossa; bpp, basipterygoid process; crb, ceratobranchial I; de, dentary; ect, ectopterygoid; fr, frontal; j, jugal; la, lacrimal; mx, maxilla; na, nasal; pa, parietal; pbs, parabasisphenoid; pl, palatine; plf, palatine fossa; po, postorbital; pra, pre-articular; prf, prefrontal; pt, pterygoid; ptmr, pterygoid medial rim; sa, surangular; spl, splenial; sq, squamosal; vo, vomer." figureDoi="http://doi.org/10.5281/zenodo.8426169" httpUri="https://zenodo.org/record/8426169/files/figure.png" pageId="16" pageNumber="370">Fig. 9A, B</figureCitation>
). The parietals are clearly unfused in both specimens. Anteriorly, the parietal only connects to the frontal. As outlined above, the frontal–parietal suture is W-shaped in dorsal view, with a distinct anteromedial process of the parietal and a V-shaped notch directly lateral to this process. The dorsal surface of the parietal is smooth, but possesses a marked depression on its lateral half, and correspondingly, a raised longitudinally-directed platform on its medial half. A distinct sagiưal crest, which is present in several non-crocodyliform crocodylmorphs, including
<taxonomicName id="4C0BE8BA310D9F044232FB563C4EFB3B" authorityName="Bonaparte" authorityYear="1969" box="[810,1090,1238,1262]" class="Polyplacophora" genus="Pseudhesperosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="16" pageNumber="370" phylum="Mollusca" rank="species" species="jachaleri">
<emphasis id="B97F4F2B310D9F044232FB563C4EFB3B" box="[810,1090,1238,1262]" italics="true" pageId="16" pageNumber="370">Pseudhesperosuchus jachaleri</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0BE8BA310D9F044569FB563D35FB3B" authorityName="Haughton" authorityYear="1915" box="[1137,1337,1238,1262]" class="Reptilia" genus="Sphenosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="16" pageNumber="370" phylum="Chordata" rank="species" species="acutus">
<emphasis id="B97F4F2B310D9F044569FB563D35FB3B" box="[1137,1337,1238,1262]" italics="true" pageId="16" pageNumber="370">Sphenosuchus acutus</emphasis>
</taxonomicName>
, is absent as in
<taxonomicName id="4C0BE8BA310D9F04425AFB753C17FAD8" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[834,1051,1269,1293]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="16" pageNumber="370" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B310D9F04425AFB753C17FAD8" box="[834,1051,1269,1293]" italics="true" pageId="16" pageNumber="370">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0BE8BA310D9F04453DFB753D65FAD8" authority=", CM" authorityName="CM" box="[1061,1385,1269,1293]" class="Reptilia" family="Sphenosuchidae" genus="Litargosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="16" pageNumber="370" phylum="Chordata" rank="species" species="leptorhynchus">
<emphasis id="B97F4F2B310D9F04453DFB753D39FAD8" box="[1061,1333,1269,1293]" italics="true" pageId="16" pageNumber="370">Litargosuchus leptorhynchus</emphasis>
, CM
</taxonomicName>
29894, and
<taxonomicName id="4C0BE8BA310D9F04424CFA943C4CFAF9" box="[852,1088,1300,1324]" class="Reptilia" family="Sphenosuchidae" genus="Dromicosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="16" pageNumber="370" phylum="Chordata" rank="species" species="grallator">
<emphasis id="B97F4F2B310D9F04424CFA943C4CFAF9" box="[852,1088,1300,1324]" italics="true" pageId="16" pageNumber="370">Dromicosuchus grallator</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9AEEC8310D9F044556FA943CCAFAF8" author="Crush PJ" box="[1102,1222,1300,1325]" pageId="16" pageNumber="370" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310D9F0445D7FA943D7DFAF8" author="Clark JM &amp; Sues H-D &amp; Berman DS" box="[1231,1393,1300,1325]" pageId="16" pageNumber="370" pagination="683 - 704" refId="ref31006" refString="Clark JM, Sues H-D, Berman DS. A new specimen of Hesperosuchus agilis from the Upper Triassic of New Mexico and the interrelationships of basal crocodylomorph archosaurs. Journal of Vertebrate Paleontology 2000; 20: 683 - 704." type="journal article" year="2000">
Clark
<emphasis id="B97F4F2B310D9F044414FA953D36FAF9" box="[1292,1338,1300,1324]" italics="true" pageId="16" pageNumber="370">et al.</emphasis>
2000
</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310D9F044462FA943BCFFA99" author="Clark JM &amp; Sues H-D" pageId="16" pageNumber="370" pagination="77 - 95" refId="ref30970" refString="Clark JM, Sues H-D. Two new basal crocodylomorph archosaurs from the Lower Jurassic and the monophyly of the Sphenosuchia. Zoological Journal of the Linnean Society 2002; 136: 77 - 95." type="journal article" year="2002">Clark and Sues 2002</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310D9F0442C8FAB43C7EFA99" author="Sues H-D &amp; Olsen PE &amp; Carter JG" box="[976,1138,1332,1356]" pageId="16" pageNumber="370" pagination="329 - 43" refId="ref32828" refString="Sues H-D, Olsen PE, Carter JG et al. A new crocodylomorph archosaur from the Upper Triassic of North Carolina. Journal of Vertebrate Paleontology 2003; 23: 329 - 43." type="journal article" year="2003">
Sues
<emphasis id="B97F4F2B310D9F04451FFAB53C34FA99" box="[1031,1080,1332,1356]" italics="true" pageId="16" pageNumber="370">et al.</emphasis>
2003
</bibRefCitation>
). The lateral depression of the parietal is part of the supratemporal fossa, and the lateral margin of the bone, which is concave in dorsal view, forms the medial margin of the supratemporal fenestra. The medial margin, which forms the suture between the parietals, is straight. The posterior margin of the less parietal of SMNS 12597b is straight and slightly raised. It is directed posterolaterally to anteromedially. The occipital surface of the parietal is not exposed.
</paragraph>
<paragraph id="8BB49339310D9F064232F9C83A72FD8A" blockId="16.[810,1459,1608,1977]" lastBlockId="18.[113,763,144,1171]" lastPageId="18" lastPageNumber="372" pageId="16" pageNumber="370">
<emphasis id="B97F4F2B310D9F044232F9C83B96F9B5" box="[810,922,1608,1632]" italics="true" pageId="16" pageNumber="370">Postorbital:</emphasis>
The postorbital is only preserved in SMNS 12591a (
<figureCitation id="13308FBC310D9F04422DF9E83B7FF955" box="[821,883,1640,1664]" captionStart="Figure 8" captionStartId="9.[129,194,1772,1796]" captionTargetBox="[178,1429,144,1744]" captionTargetId="figure-6@9.[174,1429,144,1744]" captionTargetPageId="9" captionText="Figure 8. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in right lateral view; B, interpretative drawing of A; C, less lateral view; D, interpretative drawing of (C). Abbreviations:an, angular; cc, crista cranii; de, dentary; fr, frontal; j, jugal; la, lacrimal; ls, laterosphenoid; mx, maxilla; na, nasal; pa, parietal; pl, palatine; po, postorbital; prf, prefrontal; pt, pterygoid; q, quadrate; qj, quadratojugal; sa, surangular; sq, squamosal." figureDoi="http://doi.org/10.5281/zenodo.8426167" httpUri="https://zenodo.org/record/8426167/files/figure.png" pageId="16" pageNumber="370">Figs 8</figureCitation>
,
<figureCitation id="13308FBC310D9F044298F9E83BB7F9AA" box="[896,955,1640,1664]" captionStart="Figure 9" captionStartId="10.[113,178,1772,1796]" captionTargetBox="[222,1352,144,1744]" captionTargetId="figure-6@10.[220,1352,144,1744]" captionTargetPageId="10" captionText="Figure 9. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in dorsal view; B, interpretative drawing of (A); C, ventral view; D, interpretative drawing of (C). Abbreviations:an, angular; antfo, antorbital fossa; bpp, basipterygoid process; crb, ceratobranchial I; de, dentary; ect, ectopterygoid; fr, frontal; j, jugal; la, lacrimal; mx, maxilla; na, nasal; pa, parietal; pbs, parabasisphenoid; pl, palatine; plf, palatine fossa; po, postorbital; pra, pre-articular; prf, prefrontal; pt, pterygoid; ptmr, pterygoid medial rim; sa, surangular; spl, splenial; sq, squamosal; vo, vomer." figureDoi="http://doi.org/10.5281/zenodo.8426169" httpUri="https://zenodo.org/record/8426169/files/figure.png" pageId="16" pageNumber="370">9A, B</figureCitation>
). It connects to the frontal dorsally, to the jugal ventrally, and it forms an extensive connection with the squamosal along its posterior side. It is a triradiate bone composed of a dorsomedial process, ventral process, and posterior process. The first two processes are elongate, with the ventral process being the longest, whereas the posterior process is considerably shorter and triangular in outline. The dorsomedial and posterior processes are primarily exposed in dorsal view, whereas the ventral process is deflected relative to the rest of the element, and exposed in lateral view. Although this deflection is at least to some extent the result of the dorsoventral compression of SMNS 12591a, the configuration is similar to that seen in
<taxonomicName id="4C0BE8BA310F9F064169FF2F395EFF12" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[113,338,175,199]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="18" pageNumber="372" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B310F9F064169FF2F395EFF12" box="[113,338,175,199]" italics="true" pageId="18" pageNumber="372">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0BE8BA310F9F064096FF2F3AA9FF12" box="[398,677,175,199]" class="Reptilia" family="Sphenosuchidae" genus="Litargosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="18" pageNumber="372" phylum="Chordata" rank="species" species="leptorhynchus">
<emphasis id="B97F4F2B310F9F064096FF2F3AA9FF12" box="[398,677,175,199]" italics="true" pageId="18" pageNumber="372">Litargosuchus leptorhynchus</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9AEEC8310F9F0643A2FF2F38A9FF32" author="Crush PJ" pageId="18" pageNumber="372" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310F9F0641ABFF4F3982FF32" author="Clark JM &amp; Sues H-D" box="[179,398,207,231]" pageId="18" pageNumber="372" pagination="77 - 95" refId="ref30970" refString="Clark JM, Sues H-D. Two new basal crocodylomorph archosaurs from the Lower Jurassic and the monophyly of the Sphenosuchia. Zoological Journal of the Linnean Society 2002; 136: 77 - 95." type="journal article" year="2002">Clark and Sues 2002</bibRefCitation>
). The anterior margin of the postorbital is distinctly concave in lateral and dorsal view, and forms much of the posterior margin of the orbit. The connection with the frontal is best preserved on the right side of SMNS 12591a, which reveals that the medial margin of the dorsomedial process articulated tightly with the lateral surface of the posterolateral side of the frontal, thus corroborating the interpretation of the postorbital facet on the disarticulated frontals of SMNS 12597a described above. This corresponds to the morphology seen in early crocodylomorphs, whereas in later non-crocodyliform crocodylomorphs, such as
<taxonomicName id="4C0BE8BA310F9F0640A3FD883AB0FDF5" authorityName="Simmons" authorityYear="1965" box="[443,700,520,544]" class="Reptilia" family="Sphenosuchidae" genus="Dibothrosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="18" pageNumber="372" phylum="Chordata" rank="species" species="elaphros">
<emphasis id="B97F4F2B310F9F0640A3FD883AB0FDF5" box="[443,700,520,544]" italics="true" pageId="18" pageNumber="372">Dibothrosuchus elaphros</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0BE8BA310F9F064169FDA73933FDEA" box="[113,319,551,575]" class="Reptilia" genus="Junggarsuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="18" pageNumber="372" phylum="Chordata" rank="species" species="sloani">
<emphasis id="B97F4F2B310F9F064169FDA73933FDEA" box="[113,319,551,575]" italics="true" pageId="18" pageNumber="372">Junggarsuchus sloani</emphasis>
</taxonomicName>
, the dorsomedial process additionally articulates with the parietal (
<bibRefCitation id="EF9AEEC8310F9F064099FDC63A61FD8A" author="Ruebenstahl AA &amp; Klein MD &amp; Yi H" box="[385,621,582,607]" pageId="18" pageNumber="372" pagination="2463 - 556" refId="ref32564" refString="Ruebenstahl AA, Klein MD, Yi H et al. Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros. The Anatomical Record 2022; 305: 2463 - 556." type="journal article" year="2022">
Ruebenstahl
<emphasis id="B97F4F2B310F9F06431EFDC73A39FD8B" box="[518,565,582,606]" italics="true" pageId="18" pageNumber="372">et al.</emphasis>
2022
</bibRefCitation>
).
</paragraph>
<caption id="DF74C3B1310C9F054167F9083B04F89D" ID-DOI="http://doi.org/10.5281/zenodo.8426173" ID-Zenodo-Dep="8426173" httpUri="https://zenodo.org/record/8426173/files/figure.png" pageId="17" pageNumber="371" startId="17.[127,192,1672,1696]" targetBox="[131,1404,147,1640]" targetPageId="17" targetType="figure">
<paragraph id="8BB49339310C9F054167F9083B04F89D" blockId="17.[127,1450,1672,1864]" pageId="17" pageNumber="371">
<emphasis id="B97F4F2B310C9F054167F90838EFF975" bold="true" box="[127,227,1672,1696]" pageId="17" pageNumber="371">Figure 11.</emphasis>
Details of the cranial anatomy of
<taxonomicName id="4C0BE8BA310C9F054305F9083B0CF975" authorityName="Huene" authorityYear="1921" box="[541,768,1672,1696]" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="17" pageNumber="371" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B310C9F054305F9083B0CF975" box="[541,768,1672,1696]" italics="true" pageId="17" pageNumber="371">Saltoposuchus connectens</emphasis>
</taxonomicName>
. A, frontals of SMNS 12597a in ventral view; B, less squamosal of SMNS 12596a in dorsal view; C, right squamosal of SMNS 12597b in ventral view; D, less quadrate of SMNS 12597b in posterolateral view; E, less quadrate of SMNS 12596a in anterolateral view; F, enlarged anterior maxillary tooth of SMNS 12352a in angled lateral view. Abbreviations: adp, anterodorsal process; ap, anterior process; dh, dorsal head; fr, frontal; iqf, internal quadrate fenestra; l., less; lcr, longitudinal crest; lr, longitudinal ridge; olb, olfactory bulb; otfl, otoccipital flange; paf, parietal facet; pmp, posteromedial process; pof, postorbital facet; ppeg, posterior peg; prff, prefrontal facet; ptfl, post-temporal flange; ptp, pterygoid process; ptra, pterygoid ramus; qf, quadrate facet; r., right; stfo, supratemporal fossa; vdp, ventrally directed process; vee, ventral end.
</paragraph>
</caption>
<paragraph id="8BB49339310F9F064195FDE638BAFB46" blockId="18.[113,763,144,1171]" pageId="18" pageNumber="372">
The dorsomedial process is distinctly curved anterodorsally. The posterior process, as well as the proximal bases of the dorsomedial and ventral processes, all overlapped the squamosal dorsally. This wide overlapping articulation between postorbital and squamosal does not occur in theropods, and is typical of non-crocodyliform crocodylomorphs (
<bibRefCitation id="EF9AEEC8310F9F06431FFC833AF9FCCE" author="Sereno PC &amp; Wild R" box="[519,757,770,795]" pageId="18" pageNumber="372" pagination="435 - 58" refId="ref32722" refString="Sereno PC, Wild R. Procompsognathus: theropod, ' thecodont' or both? Journal of Vertebrate Paleontology 1992; 12: 435 - 58." type="journal article" year="1992">Sereno and Wild 1992</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310F9F064169FCA238F6FCEF" author="Nesbiu SJ" box="[113,250,802,826]" pageId="18" pageNumber="372" pagination="1 - 292" refId="ref32089" refString="Nesbiu SJ. The early evolution of archosaurs: relationships and the origin of major clades. Bulletin of the American Museum of Natural History 2011; 352: 1 - 292." type="journal article" year="2011">Nesbiư 2011</bibRefCitation>
: ch. 66). The configuration of SMNS 12591a is particularly reminiscent of that seen in
<taxonomicName id="4C0BE8BA310F9F0640EFFCC13ADCFC8C" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[503,720,833,857]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="18" pageNumber="372" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B310F9F0640EFFCC13ADCFC8C" box="[503,720,833,857]" italics="true" pageId="18" pageNumber="372">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0BE8BA310F9F064169FCE03988FCAD" box="[113,388,864,888]" class="Reptilia" family="Sphenosuchidae" genus="Litargosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="18" pageNumber="372" phylum="Chordata" rank="species" species="leptorhynchus">
<emphasis id="B97F4F2B310F9F064169FCE03988FCAD" box="[113,388,864,888]" italics="true" pageId="18" pageNumber="372">Litargosuchus leptorhynchus</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9AEEC8310F9F06408CFCE03A02FCAC" author="Crush PJ" box="[404,526,864,889]" pageId="18" pageNumber="372" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310F9F064302FCE03AE3FCAC" author="Clark JM &amp; Sues H-D" box="[538,751,864,889]" pageId="18" pageNumber="372" pagination="77 - 95" refId="ref30970" refString="Clark JM, Sues H-D. Two new basal crocodylomorph archosaurs from the Lower Jurassic and the monophyly of the Sphenosuchia. Zoological Journal of the Linnean Society 2002; 136: 77 - 95." type="journal article" year="2002">Clark and Sues 2002</bibRefCitation>
) and thus represents a clear indication of the crocodylomorph identity of this specimen, as was also pointed out by
<bibRefCitation id="EF9AEEC8310F9F0643ABFC1F3927FC02" author="Sereno PC &amp; Wild R" pageId="18" pageNumber="372" pagination="435 - 58" refId="ref32722" refString="Sereno PC, Wild R. Procompsognathus: theropod, ' thecodont' or both? Journal of Vertebrate Paleontology 1992; 12: 435 - 58." type="journal article" year="1992">Sereno and Wild (1992)</bibRefCitation>
. The ventral process of the postorbital articulates on the anterior side of the ascending process of the jugal and tapers to a point ventrally, contrasting with the condition in
<taxonomicName id="4C0BE8BA310F9F0641F9FB9C39C9FBE1" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[225,453,1052,1076]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="18" pageNumber="372" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B310F9F0641F9FB9C39C9FBE1" box="[225,453,1052,1076]" italics="true" pageId="18" pageNumber="372">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0BE8BA310F9F0640C1FB9C3AF9FBE1" box="[473,757,1052,1076]" class="Reptilia" family="Sphenosuchidae" genus="Litargosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="18" pageNumber="372" phylum="Chordata" rank="species" species="leptorhynchus">
<emphasis id="B97F4F2B310F9F0640C1FB9C3AF9FBE1" box="[473,757,1052,1076]" italics="true" pageId="18" pageNumber="372">Litargosuchus leptorhynchus</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C0BE8BA310F9F064186FBBC3965FB86" authorityName="Haughton" authorityYear="1915" box="[158,361,1084,1108]" class="Reptilia" genus="Sphenosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="18" pageNumber="372" phylum="Chordata" rank="species" species="acutus">
<emphasis id="B97F4F2B310F9F064186FBBC3965FB86" box="[158,361,1084,1108]" italics="true" pageId="18" pageNumber="372">Sphenosuchus acutus</emphasis>
</taxonomicName>
, in which the tip of the ventral process is gently rounded (
<bibRefCitation id="EF9AEEC8310F9F064024FBDB39B5FBA6" author="Crush PJ" box="[316,441,1115,1139]" pageId="18" pageNumber="372" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310F9F0640DFFBDB3A40FBA6" author="Walker AD" box="[455,588,1115,1139]" pageId="18" pageNumber="372" pagination="1 - 120" refId="ref33031" refString="Walker AD. A revision of Sphenosuchus acutus Haughton, a crocodylomorph reptile from the Elliot Formation (Late Triassic or Early Jurassic) of South Africa. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1990; 330: 1 - 120." type="journal article" year="1990">Walker 1990</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310F9F064343FBDB38A9FB47" author="Clark JM &amp; Sues H-D" pageId="18" pageNumber="372" pagination="77 - 95" refId="ref30970" refString="Clark JM, Sues H-D. Two new basal crocodylomorph archosaurs from the Lower Jurassic and the monophyly of the Sphenosuchia. Zoological Journal of the Linnean Society 2002; 136: 77 - 95." type="journal article" year="2002">Clark and Sues 2002</bibRefCitation>
).
</paragraph>
<paragraph id="8BB49339310F9F064169FB343CD3FED0" blockId="18.[113,763,1204,1979]" lastBlockId="18.[810,1460,143,1044]" pageId="18" pageNumber="372">
<emphasis id="B97F4F2B310F9F064169FB3438A6FB19" box="[113,170,1204,1228]" italics="true" pageId="18" pageNumber="372">Jugal:</emphasis>
A largely complete less jugal missing the anterior end of the anterior process is preserved in SMNS 12597b (
<figureCitation id="13308FBC310F9F064384FB533AE6FB3E" box="[668,746,1235,1259]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="18" pageNumber="372">Fig. 2C</figureCitation>
). The lateral surface of this element is generally well preserved, but broken in the region where the three processes of the bone meet. This element was originally interpreted as a lacrimal (
<bibRefCitation id="EF9AEEC8310F9F0643ADFAB138A9FABD" author="Huene FR" pageId="18" pageNumber="372" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene 1921</bibRefCitation>
), but reinterpreted as a jugal by
<bibRefCitation id="EF9AEEC8310F9F0640FDFAD03A78FABD" author="Crush PJ" box="[485,628,1360,1384]" pageId="18" pageNumber="372" refId="ref31157" refString="Crush PJ. An early, terrestrial, crocodile from South Wales. Unpublished Ph. D. Thesis, University College London, 1980." type="book" year="1980">Crush (1980)</bibRefCitation>
. The anterior tip of the right jugal is present in SMNS 12352a, whereas both jugals are largely complete in SMNS 12591a (
<figureCitation id="13308FBC310F9F064346FA0F3A92FA72" box="[606,670,1423,1447]" captionStart="Figure 8" captionStartId="9.[129,194,1772,1796]" captionTargetBox="[178,1429,144,1744]" captionTargetId="figure-6@9.[174,1429,144,1744]" captionTargetPageId="9" captionText="Figure 8. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in right lateral view; B, interpretative drawing of A; C, less lateral view; D, interpretative drawing of (C). Abbreviations:an, angular; cc, crista cranii; de, dentary; fr, frontal; j, jugal; la, lacrimal; ls, laterosphenoid; mx, maxilla; na, nasal; pa, parietal; pl, palatine; po, postorbital; prf, prefrontal; pt, pterygoid; q, quadrate; qj, quadratojugal; sa, surangular; sq, squamosal." figureDoi="http://doi.org/10.5281/zenodo.8426167" httpUri="https://zenodo.org/record/8426167/files/figure.png" pageId="18" pageNumber="372">Figs 8</figureCitation>
,
<figureCitation id="13308FBC310F9F0643B5FA0F3AB6FA72" box="[685,698,1423,1447]" captionStart="Figure 9" captionStartId="10.[113,178,1772,1796]" captionTargetBox="[222,1352,144,1744]" captionTargetId="figure-6@10.[220,1352,144,1744]" captionTargetPageId="10" captionText="Figure 9. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in dorsal view; B, interpretative drawing of (A); C, ventral view; D, interpretative drawing of (C). Abbreviations:an, angular; antfo, antorbital fossa; bpp, basipterygoid process; crb, ceratobranchial I; de, dentary; ect, ectopterygoid; fr, frontal; j, jugal; la, lacrimal; mx, maxilla; na, nasal; pa, parietal; pbs, parabasisphenoid; pl, palatine; plf, palatine fossa; po, postorbital; pra, pre-articular; prf, prefrontal; pt, pterygoid; ptmr, pterygoid medial rim; sa, surangular; spl, splenial; sq, squamosal; vo, vomer." figureDoi="http://doi.org/10.5281/zenodo.8426169" httpUri="https://zenodo.org/record/8426169/files/figure.png" pageId="18" pageNumber="372">9</figureCitation>
,
<figureCitation id="13308FBC310F9F0643D1FA0F3873FA13" captionStart="Figure 10" captionStartId="14.[113,178,1772,1796]" captionTargetBox="[197,1376,144,1744]" captionTargetId="figure-5@14.[197,1376,144,1744]" captionTargetPageId="14" captionText="Figure 10. Saltoposuchus connectens referred specimen SMNS 12352.A, SMNS 12352a with the snout region exposed in angled right lateral view; B, interpretative drawing of (A); C, opposite side of SMNS 12352a exposing the posterior portion of the less maxilla and lacrimal in angled lateral view; D, interpretative drawing of (C); E, SMNS 12352b, carpus and manus in dorsal view; F, interpretative drawing of (E); G, SMNS 12352b in medial view; H, interpretative drawing of (G). Abbreviations; antfe, antorbital fenestra; antfo, antorbital fossa; dc, distal carpal; de, dentary; exp, extensor pit; fr, frontal; j, jugal; la, lacrimal; lad, lacrimal duct; mc, metacarpal; mx, maxilla; na, nasal; ph, phalanx; pm, premaxilla; prf, prefrontal; ra, radiale; ul, ulnare." figureDoi="http://doi.org/10.5281/zenodo.8426171" httpUri="https://zenodo.org/record/8426171/files/figure.png" pageId="18" pageNumber="372">10A, B</figureCitation>
). In the laưer specimen, the edges of the anterior process are broken in the less element, and the distal end of the posterior process is missing in the right element. The jugal is a triradiate element, consisting of an anterior, posterior, and dorsal or ascending process. The anterior and posterior processes are both oriented in the horizontal plane relative to the skull, whereas the ascending process is oriented virtually completely perpendicular to the other two processes in both SMNS 12597b and SMNS 12591a, being directed dorsally and very slightly posteriorly.This contrasts with
<taxonomicName id="4C0BE8BA310F9F064015F94839E6F935" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[269,490,1736,1760]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="18" pageNumber="372" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B310F9F064015F94839E6F935" box="[269,490,1736,1760]" italics="true" pageId="18" pageNumber="372">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0BE8BA310F9F064339F9483AF9F935" box="[545,757,1736,1760]" genus="Carnufex" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="18" pageNumber="372" phylum="Chordata" rank="species" species="carolinensis">
<emphasis id="B97F4F2B310F9F064339F9483AF9F935" box="[545,757,1736,1760]" italics="true" pageId="18" pageNumber="372">Carnufex carolinensis</emphasis>
</taxonomicName>
, in which the ascending process is more posterodorsally oriented (NHMUK PV R 7591,
<bibRefCitation id="EF9AEEC8310F9F064047F88739D4F8CA" author="Crush PJ" box="[351,472,1799,1823]" pageId="18" pageNumber="372" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310F9F0640FAF8873AE5F8CA" author="Drymala SM &amp; Zanno LE" box="[482,745,1799,1823]" pageId="18" pageNumber="372" pagination="0157528" refId="ref31236" refString="Drymala SM, Zanno LE. Osteology of Carnufex carolinensis (Archosauria: Psuedosuchia) from the Pekin Formation of North Carolina and its implications for early crocodylomorph evolution. PLoS One 2016; 11: e 0157528." type="journal article" year="2016">Drymala and Zanno 2016</bibRefCitation>
), but corresponds with the morphology seen in
<taxonomicName id="4C0BE8BA310F9F06436CF8A638FBF888" class="Reptilia" family="Sphenosuchidae" genus="Litargosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="18" pageNumber="372" phylum="Chordata" rank="species" species="leptorhynchus">
<emphasis id="B97F4F2B310F9F06436CF8A638FBF888" italics="true" pageId="18" pageNumber="372">Litargosuchus leptorhynchus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0BE8BA310F9F064032F8C539F9F888" authorityName="Haughton" authorityYear="1915" box="[298,501,1861,1885]" class="Reptilia" genus="Sphenosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="18" pageNumber="372" phylum="Chordata" rank="species" species="acutus">
<emphasis id="B97F4F2B310F9F064032F8C539F9F888" box="[298,501,1861,1885]" italics="true" pageId="18" pageNumber="372">Sphenosuchus acutus</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9AEEC8310F9F06431FF8C53A85F88B" author="Walker AD" box="[519,649,1861,1886]" pageId="18" pageNumber="372" pagination="1 - 120" refId="ref33031" refString="Walker AD. A revision of Sphenosuchus acutus Haughton, a crocodylomorph reptile from the Elliot Formation (Late Triassic or Early Jurassic) of South Africa. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1990; 330: 1 - 120." type="journal article" year="1990">Walker 1990</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310F9F06438DF8C538D7F8A8" author="Clark JM &amp; Sues H-D" pageId="18" pageNumber="372" pagination="77 - 95" refId="ref30970" refString="Clark JM, Sues H-D. Two new basal crocodylomorph archosaurs from the Lower Jurassic and the monophyly of the Sphenosuchia. Zoological Journal of the Linnean Society 2002; 136: 77 - 95." type="journal article" year="2002">Clark and Sues 2002</bibRefCitation>
). The anterior process forms the longest of the three processes, followed by the posterior process. The anterior process forms the ventral margin of the orbit, the posterior process forms the ventral margin of the infratemporal fenestra, and the ascending process forms the posteroventral margin of the orbit on its anterior side and the anteroventral margin of the infratemporal fenestra on its posterior side.
</paragraph>
<paragraph id="8BB49339310F9F06425DFE8D3CC2FBC1" blockId="18.[810,1460,143,1044]" pageId="18" pageNumber="372">
The anterior process does not contribute to the antorbital fossa in both SMNS 12352a and SMNS 12591a, similar to other non-crocodyliform crocodylomorphs (
<bibRefCitation id="EF9AEEC8310F9F0645A7FECC3DA1FEB1" author="Sereno PC &amp; Wild R" box="[1215,1453,331,356]" pageId="18" pageNumber="372" pagination="435 - 58" refId="ref32722" refString="Sereno PC, Wild R. Procompsognathus: theropod, ' thecodont' or both? Journal of Vertebrate Paleontology 1992; 12: 435 - 58." type="journal article" year="1992">Sereno and Wild 1992</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310F9F064232FEEB3BA3FE56" author="Nesbiu SJ" box="[810,943,363,387]" pageId="18" pageNumber="372" pagination="1 - 292" refId="ref32089" refString="Nesbiu SJ. The early evolution of archosaurs: relationships and the origin of major clades. Bulletin of the American Museum of Natural History 2011; 352: 1 - 292." type="journal article" year="2011">Nesbiư 2011</bibRefCitation>
: ch. 69). The anterior articulation of the jugal with the maxilla and lacrimal is best observed in SMNS 12352a. The jugal sits in a slot on the dorsolateral surface of the maxilla, whereas the jugal connects to the lacrimal on the dorsomedial surface of the former and the ventral surface of the laưer. The ascending process has a straight to slightly convex posterior margin in lateral view in SMNS 12597b and both jugals of SMNS 12591a, a character unique to certain early crocodylomorphs among archosaurs (
<bibRefCitation id="EF9AEEC8310F9F0642A9FDE53C4CFDA8" author="Nesbiu SJ" box="[945,1088,613,637]" pageId="18" pageNumber="372" pagination="1 - 292" refId="ref32089" refString="Nesbiu SJ. The early evolution of archosaurs: relationships and the origin of major clades. Bulletin of the American Museum of Natural History 2011; 352: 1 - 292." type="journal article" year="2011">Nesbiư 2011</bibRefCitation>
: ch. 73), including
<taxonomicName id="4C0BE8BA310F9F064435FDE53B64FD49" authorityName="Haughton" authorityYear="1915" class="Reptilia" genus="Sphenosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="18" pageNumber="372" phylum="Chordata" rank="species" species="acutus">
<emphasis id="B97F4F2B310F9F064435FDE53B64FD49" italics="true" pageId="18" pageNumber="372">Sphenosuchus acutus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0BE8BA310F9F064265FD053C94FD48" box="[893,1176,645,669]" class="Reptilia" family="Sphenosuchidae" genus="Litargosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="18" pageNumber="372" phylum="Chordata" rank="species" species="leptorhynchus">
<emphasis id="B97F4F2B310F9F064265FD053C94FD48" box="[893,1176,645,669]" italics="true" pageId="18" pageNumber="372">Litargosuchus leptorhynchus</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C0BE8BA310F9F0645F9FD053DBFFD48" box="[1249,1459,645,669]" class="Reptilia" genus="Junggarsuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="18" pageNumber="372" phylum="Chordata" rank="species" species="sloani">
<emphasis id="B97F4F2B310F9F0645F9FD053DBFFD48" box="[1249,1459,645,669]" italics="true" pageId="18" pageNumber="372">Junggarsuchus sloani</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9AEEC8310F9F06422DFD243BB5FD69" author="Walker AD" box="[821,953,676,700]" pageId="18" pageNumber="372" pagination="1 - 120" refId="ref33031" refString="Walker AD. A revision of Sphenosuchus acutus Haughton, a crocodylomorph reptile from the Elliot Formation (Late Triassic or Early Jurassic) of South Africa. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1990; 330: 1 - 120." type="journal article" year="1990">Walker 1990</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310F9F0642DFFD243CACFD69" author="Clark JM &amp; Sues H-D" box="[967,1184,676,700]" pageId="18" pageNumber="372" pagination="77 - 95" refId="ref30970" refString="Clark JM, Sues H-D. Two new basal crocodylomorph archosaurs from the Lower Jurassic and the monophyly of the Sphenosuchia. Zoological Journal of the Linnean Society 2002; 136: 77 - 95." type="journal article" year="2002">Clark and Sues 2002</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310F9F0645B6FD243DAEFD69" author="Ruebenstahl AA &amp; Klein MD &amp; Yi H" box="[1198,1442,676,700]" pageId="18" pageNumber="372" pagination="2463 - 556" refId="ref32564" refString="Ruebenstahl AA, Klein MD, Yi H et al. Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros. The Anatomical Record 2022; 305: 2463 - 556." type="journal article" year="2022">
Ruebenstahl
<emphasis id="B97F4F2B310F9F06442EFD253D6BFD69" box="[1334,1383,676,700]" italics="true" pageId="18" pageNumber="372">et al.</emphasis>
2022
</bibRefCitation>
). Dorsally, the postorbital articulates on the anterior surface of the ascending process, and a faint ridge on the lateral surface of the process demarcates this articular facet on the jugal of SMNS 12597b. At its distal end, the posterior process would have articulated with the quadratojugal on the medial surface of the former, but this articulation is not clearly preserved in any of the specimens. The posterior process gradually tapers posteriorly in lateral view, as can be observed in SMNS 12597b and the less jugal of SMNS 12591a, and terminates in a slightly rounded tip in SMNS 12597b. The lateral surface of the jugal is smooth and transversely convex for most of its length.
</paragraph>
<paragraph id="8BB49339310F9F064232FBB63C1EF974" blockId="18.[810,1459,1078,1979]" pageId="18" pageNumber="372">
<emphasis id="B97F4F2B310F9F064232FBB63B90FB9B" box="[810,924,1078,1102]" italics="true" pageId="18" pageNumber="372">Squamosal:</emphasis>
Both squamosals are preserved in SMNS 12597b (
<figureCitation id="13308FBC310F9F06422DFBD53B8FFBB8" box="[821,899,1109,1133]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="18" pageNumber="372">Fig. 2C</figureCitation>
).
<bibRefCitation id="EF9AEEC8310F9F064285FBD53C3CFBB8" author="Crush PJ" box="[925,1072,1109,1133]" pageId="18" pageNumber="372" refId="ref31157" refString="Crush PJ. An early, terrestrial, crocodile from South Wales. Unpublished Ph. D. Thesis, University College London, 1980." type="book" year="1980">Crush (1980)</bibRefCitation>
considered the less squamosal to be beưer preserved but this element has since been damaged, particularly near its posterior end. Thus, the right element is now the more complete and beưer preserved of the two (
<figureCitation id="13308FBC310F9F064452FB333DAEFB1E" box="[1354,1442,1203,1227]" captionStart="Figure 11" captionStartId="17.[127,192,1672,1696]" captionTargetBox="[131,1404,147,1640]" captionTargetId="figure-6@17.[127,1407,144,1644]" captionTargetPageId="17" captionText="Figure 11. Details of the cranial anatomy of Saltoposuchus connectens. A, frontals of SMNS 12597a in ventral view; B, less squamosal of SMNS 12596a in dorsal view; C, right squamosal of SMNS 12597b in ventral view; D, less quadrate of SMNS 12597b in posterolateral view; E, less quadrate of SMNS 12596a in anterolateral view; F, enlarged anterior maxillary tooth of SMNS 12352a in angled lateral view.Abbreviations: adp, anterodorsal process; ap, anterior process; dh, dorsal head; fr, frontal; iqf, internal quadrate fenestra; l., less; lcr, longitudinal crest; lr, longitudinal ridge; olb, olfactory bulb; otfl, otoccipital flange; paf, parietal facet; pmp, posteromedial process; pof, postorbital facet; ppeg, posterior peg; prff, prefrontal facet; ptfl, post-temporal flange; ptp, pterygoid process; ptra, pterygoid ramus; qf, quadrate facet; r., right; stfo, supratemporal fossa; vdp, ventrally directed process; vee, ventral end." figureDoi="http://doi.org/10.5281/zenodo.8426173" httpUri="https://zenodo.org/record/8426173/files/figure.png" pageId="18" pageNumber="372">Fig. 11C</figureCitation>
). SMNS 12596a possesses a largely complete less squamosal and the anterior portion of a right squamosal is preserved in SMNS 12591a (
<figureCitation id="13308FBC310F9F064290FA913BDBFAFC" box="[904,983,1297,1321]" captionStart="Figure 3" captionStartId="4.[113,178,1499,1523]" captionTargetBox="[147,1426,144,1471]" captionTargetId="figure-217@4.[146,1426,144,1471]" captionTargetPageId="4" captionText="Figure 3. Part of Saltoposuchus connectens referred specimen SMNS 12596. A, SMNS 12596a, cranial block; B, SMNS 12596b, two dentaries exposed in ventrolateral view; C, SMNS 12596c exposing a less femur in anterolateral view; D, exposing the opposite side of SMNS 12596c, Abbreviations:de, dentary; dr, dorsal rib; dv, dorsal vertebra; fe, femur; ga, gastralia; l., less; mx, maxilla; na, nasal; os, osteoderm; q, quadrate; r., right; sq, squamosal; sr, sacral rib; sv, sacral vertebra; tp, transverse process." figureDoi="http://doi.org/10.5281/zenodo.8426153" httpUri="https://zenodo.org/record/8426153/files/figure.png" pageId="18" pageNumber="372">Figs 3A</figureCitation>
,
<figureCitation id="13308FBC310F9F0642FDFA913C13FAFC" box="[997,1055,1297,1321]" captionStart="Figure 8" captionStartId="9.[129,194,1772,1796]" captionTargetBox="[178,1429,144,1744]" captionTargetId="figure-6@9.[174,1429,144,1744]" captionTargetPageId="9" captionText="Figure 8. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in right lateral view; B, interpretative drawing of A; C, less lateral view; D, interpretative drawing of (C). Abbreviations:an, angular; cc, crista cranii; de, dentary; fr, frontal; j, jugal; la, lacrimal; ls, laterosphenoid; mx, maxilla; na, nasal; pa, parietal; pl, palatine; po, postorbital; prf, prefrontal; pt, pterygoid; q, quadrate; qj, quadratojugal; sa, surangular; sq, squamosal." figureDoi="http://doi.org/10.5281/zenodo.8426167" httpUri="https://zenodo.org/record/8426167/files/figure.png" pageId="18" pageNumber="372">8A, B</figureCitation>
,
<figureCitation id="13308FBC310F9F064534FA913C6BFAFC" box="[1068,1127,1297,1321]" captionStart="Figure 9" captionStartId="10.[113,178,1772,1796]" captionTargetBox="[222,1352,144,1744]" captionTargetId="figure-6@10.[220,1352,144,1744]" captionTargetPageId="10" captionText="Figure 9. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in dorsal view; B, interpretative drawing of (A); C, ventral view; D, interpretative drawing of (C). Abbreviations:an, angular; antfo, antorbital fossa; bpp, basipterygoid process; crb, ceratobranchial I; de, dentary; ect, ectopterygoid; fr, frontal; j, jugal; la, lacrimal; mx, maxilla; na, nasal; pa, parietal; pbs, parabasisphenoid; pl, palatine; plf, palatine fossa; po, postorbital; pra, pre-articular; prf, prefrontal; pt, pterygoid; ptmr, pterygoid medial rim; sa, surangular; spl, splenial; sq, squamosal; vo, vomer." figureDoi="http://doi.org/10.5281/zenodo.8426169" httpUri="https://zenodo.org/record/8426169/files/figure.png" pageId="18" pageNumber="372">9A, B</figureCitation>
,
<figureCitation id="13308FBC310F9F06456CFA913C90FAFC" box="[1140,1180,1297,1321]" captionStart="Figure 11" captionStartId="17.[127,192,1672,1696]" captionTargetBox="[131,1404,147,1640]" captionTargetId="figure-6@17.[127,1407,144,1644]" captionTargetPageId="17" captionText="Figure 11. Details of the cranial anatomy of Saltoposuchus connectens. A, frontals of SMNS 12597a in ventral view; B, less squamosal of SMNS 12596a in dorsal view; C, right squamosal of SMNS 12597b in ventral view; D, less quadrate of SMNS 12597b in posterolateral view; E, less quadrate of SMNS 12596a in anterolateral view; F, enlarged anterior maxillary tooth of SMNS 12352a in angled lateral view.Abbreviations: adp, anterodorsal process; ap, anterior process; dh, dorsal head; fr, frontal; iqf, internal quadrate fenestra; l., less; lcr, longitudinal crest; lr, longitudinal ridge; olb, olfactory bulb; otfl, otoccipital flange; paf, parietal facet; pmp, posteromedial process; pof, postorbital facet; ppeg, posterior peg; prff, prefrontal facet; ptfl, post-temporal flange; ptp, pterygoid process; ptra, pterygoid ramus; qf, quadrate facet; r., right; stfo, supratemporal fossa; vdp, ventrally directed process; vee, ventral end." figureDoi="http://doi.org/10.5281/zenodo.8426173" httpUri="https://zenodo.org/record/8426173/files/figure.png" pageId="18" pageNumber="372">11B</figureCitation>
).
<bibRefCitation id="EF9AEEC8310F9F0645ACFA923D46FAFC" author="Huene FR" box="[1204,1354,1297,1321]" pageId="18" pageNumber="372" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene (1921)</bibRefCitation>
originally interpreted the squamosals in both SMNS 12597b and SMNS 12596a as pterygoids, but their identification as squamosals was later established by
<bibRefCitation id="EF9AEEC8310F9F064539FAEF3CBDFA52" author="Crush PJ" box="[1057,1201,1391,1415]" pageId="18" pageNumber="372" refId="ref31157" refString="Crush PJ. An early, terrestrial, crocodile from South Wales. Unpublished Ph. D. Thesis, University College London, 1980." type="book" year="1980">Crush (1980)</bibRefCitation>
.
<bibRefCitation id="EF9AEEC8310F9F0645A5FAEF3D49FA52" author="Allen RC" box="[1213,1349,1391,1415]" pageId="18" pageNumber="372" refId="ref30591" refString="Allen RC. The anatomy and systematics of Terrestrisuchus gracilis (Archosauria, Crocodylomorpha). Unpublished Ph. D. Thesis, Northern Illinois University, 2010." type="book" year="2010">Allen (2010)</bibRefCitation>
misidentified the right squamosal of SMNS 12591a as a displaced quadrate, but its unambiguous identity as a squamosal has been established by
<bibRefCitation id="EF9AEEC8310F9F0642D8FA4E3C59FA30" author="Huene FR" box="[960,1109,1485,1509]" pageId="18" pageNumber="372" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene (1921)</bibRefCitation>
and
<bibRefCitation id="EF9AEEC8310F9F06459EFA4D3D8DFA30" author="Sereno PC &amp; Wild R" box="[1158,1409,1485,1509]" pageId="18" pageNumber="372" pagination="435 - 58" refId="ref32722" refString="Sereno PC, Wild R. Procompsognathus: theropod, ' thecodont' or both? Journal of Vertebrate Paleontology 1992; 12: 435 - 58." type="journal article" year="1992">Sereno and Wild (1992)</bibRefCitation>
. The squamosal connects to the postorbital anteriorly, the quadrate and quadratojugal ventrally, the parietal posteromedially, and the otoccipital posteroventrally. Its medial edge forms the lateral and posterolateral margin of the supratemporal fenestra, whereas its lateroventral edge forms the dorsal margin of the infratemporal fenestra.
</paragraph>
<paragraph id="8BB49339310F9F07425EF9283A25FC62" blockId="18.[810,1459,1078,1979]" lastBlockId="19.[128,778,144,1985]" lastPageId="19" lastPageNumber="373" pageId="18" pageNumber="372">
The squamosal is predominantly made up of a large plate-like surface that is curved strongly medially near its posterior end in dorsal view. This plate is transversely wide in all specimens, which represents a characteristic trait of several non-crocodyliform crocodylomorphs, including
<taxonomicName id="4C0BE8BA310F9F064438F8A63B63F888" baseAuthorityName="Crush" baseAuthorityYear="1984" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="18" pageNumber="372" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B310F9F064438F8A63B63F888" italics="true" pageId="18" pageNumber="372">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0BE8BA310F9F064298F8C53C97F888" box="[896,1179,1861,1885]" class="Reptilia" family="Sphenosuchidae" genus="Litargosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="18" pageNumber="372" phylum="Chordata" rank="species" species="leptorhynchus">
<emphasis id="B97F4F2B310F9F064298F8C53C97F888" box="[896,1179,1861,1885]" italics="true" pageId="18" pageNumber="372">Litargosuchus leptorhynchus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0BE8BA310F9F0645B3F8C53D77F888" box="[1195,1403,1861,1885]" class="Reptilia" genus="Junggarsuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="18" pageNumber="372" phylum="Chordata" rank="species" species="sloani">
<emphasis id="B97F4F2B310F9F0645B3F8C53D77F888" box="[1195,1403,1861,1885]" italics="true" pageId="18" pageNumber="372">Junggarsuchus sloani</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C0BE8BA310F9F064232F8E43BF0F8A9" box="[810,1020,1892,1916]" genus="Almadasuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="18" pageNumber="372" phylum="Chordata" rank="species" species="figarii">
<emphasis id="B97F4F2B310F9F064232F8E43BF0F8A9" box="[810,1020,1892,1916]" italics="true" pageId="18" pageNumber="372">Almadasuchus figarii</emphasis>
</taxonomicName>
(e.g.
<bibRefCitation id="EF9AEEC8310F9F064525F8E43CB7F8A9" author="Crush PJ" box="[1085,1211,1892,1916]" pageId="18" pageNumber="372" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310F9F0645D4F8E43DA1F8A9" author="Clark JM &amp; Sues H-D" box="[1228,1453,1892,1916]" pageId="18" pageNumber="372" pagination="77 - 95" refId="ref30970" refString="Clark JM, Sues H-D. Two new basal crocodylomorph archosaurs from the Lower Jurassic and the monophyly of the Sphenosuchia. Zoological Journal of the Linnean Society 2002; 136: 77 - 95." type="journal article" year="2002">Clark and Sues 2002</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310F9F064232F8043C2EF849" author="Ruebenstahl AA &amp; Klein MD &amp; Yi H" box="[810,1058,1924,1948]" pageId="18" pageNumber="372" pagination="2463 - 556" refId="ref32564" refString="Ruebenstahl AA, Klein MD, Yi H et al. Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros. The Anatomical Record 2022; 305: 2463 - 556." type="journal article" year="2022">
Ruebenstahl
<emphasis id="B97F4F2B310F9F0642ACF8043BEAF849" box="[948,998,1924,1948]" italics="true" pageId="18" pageNumber="372">et al.</emphasis>
2022
</bibRefCitation>
). In SMNS 12591a, the squamosal is considerably wider than the supratemporal fenestra in dorsal view (
<bibRefCitation id="EF9AEEC8310E9F0741DDFF1039B6FF7D" author="Sereno PC &amp; Wild R" box="[197,442,144,168]" pageId="19" pageNumber="373" pagination="435 - 58" refId="ref32722" refString="Sereno PC, Wild R. Procompsognathus: theropod, ' thecodont' or both? Journal of Vertebrate Paleontology 1992; 12: 435 - 58." type="journal article" year="1992">Sereno and Wild 1992</bibRefCitation>
). The plate is transversely distinctly convex on its dorsolateral surface and conversely concave on its ventromedial surface. The dorsolateral surface is mostly smooth but possesses a distinct longitudinal crest along most of its length, which is predominantly oriented anteroposteriorly but which curves distinctly medially near the posterior end of the bone. The dorsal surface of the squamosal of SMNS 12591a is too broken and incomplete to establish the presence of this crest. A similar crest is present in other non-crocodyliform crocodylomorphs, except
<taxonomicName id="4C0BE8BA310E9F074333FE2A3B06FE17" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[555,778,426,450]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="19" pageNumber="373" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B310E9F074333FE2A3B06FE17" box="[555,778,426,450]" italics="true" pageId="19" pageNumber="373">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0BE8BA310E9F0741A8FE4939C5FE34" box="[176,457,457,481]" class="Reptilia" family="Sphenosuchidae" genus="Litargosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="19" pageNumber="373" phylum="Chordata" rank="species" species="leptorhynchus">
<emphasis id="B97F4F2B310E9F0741A8FE4939C5FE34" box="[176,457,457,481]" italics="true" pageId="19" pageNumber="373">Litargosuchus leptorhynchus</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9AEEC8310E9F0740C4FE493A55FE34" author="Crush PJ" box="[476,601,457,481]" pageId="19" pageNumber="373" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310E9F074370FE4938B9FDD4" author="Clark JM &amp; Sues H-D" pageId="19" pageNumber="373" pagination="77 - 95" refId="ref30970" refString="Clark JM, Sues H-D. Two new basal crocodylomorph archosaurs from the Lower Jurassic and the monophyly of the Sphenosuchia. Zoological Journal of the Linnean Society 2002; 136: 77 - 95." type="journal article" year="2002">Clark and Sues 2002</bibRefCitation>
). It represents the demarcation of the contribution of the squamosal to the supratemporal fossa and thus the dorsolateral extent of the jaw adductor musculature, which is placed medial to the crest (
<bibRefCitation id="EF9AEEC8310E9F074041FDC73A4AFD8A" author="Sereno PC &amp; Wild R" box="[345,582,582,607]" pageId="19" pageNumber="373" pagination="435 - 58" refId="ref32722" refString="Sereno PC, Wild R. Procompsognathus: theropod, ' thecodont' or both? Journal of Vertebrate Paleontology 1992; 12: 435 - 58." type="journal article" year="1992">Sereno and Wild 1992</bibRefCitation>
). This medial surface would have predominantly faced dorsally, and the surface lateral to the crest predominantly laterally, with the former surface being most prominent. The outer surface of the squamosal also faces predominantly dorsally in
<taxonomicName id="4C0BE8BA310E9F0740E5FD443AD0FD09" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[509,732,708,732]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="19" pageNumber="373" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B310E9F0740E5FD443AD0FD09" box="[509,732,708,732]" italics="true" pageId="19" pageNumber="373">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0BE8BA310E9F074199FD633999FD2E" box="[129,405,739,763]" class="Reptilia" family="Sphenosuchidae" genus="Litargosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="19" pageNumber="373" phylum="Chordata" rank="species" species="leptorhynchus">
<emphasis id="B97F4F2B310E9F074199FD633999FD2E" box="[129,405,739,763]" italics="true" pageId="19" pageNumber="373">Litargosuchus leptorhynchus</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9AEEC8310E9F0740BCFD633A10FD2E" author="Crush PJ" box="[420,540,739,763]" pageId="19" pageNumber="373" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310E9F07433FFD633AF4FD2E" author="Clark JM &amp; Sues H-D" box="[551,760,739,763]" pageId="19" pageNumber="373" pagination="77 - 95" refId="ref30970" refString="Clark JM, Sues H-D. Two new basal crocodylomorph archosaurs from the Lower Jurassic and the monophyly of the Sphenosuchia. Zoological Journal of the Linnean Society 2002; 136: 77 - 95." type="journal article" year="2002">Clark and Sues 2002</bibRefCitation>
). The medial margin of the squamosal drops steeply ventrally, as can best be seen in SMNS 12596a, similar to the condition in
<taxonomicName id="4C0BE8BA310E9F074199FCC13968FC8C" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[129,356,833,857]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="19" pageNumber="373" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B310E9F074199FCC13968FC8C" box="[129,356,833,857]" italics="true" pageId="19" pageNumber="373">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
(NHMUK PV R 7591). The anterior portion of the dorsolateral surface of the squamosal of SMNS 12596a is incomplete, but a very large articular surface for the postorbital can nevertheless be observed.
</paragraph>
<paragraph id="8BB49339310E9F074184FC3E392CF814" blockId="19.[128,778,144,1985]" pageId="19" pageNumber="373">
Three processes extend from the posterior end of the less squamosal of SMNS 12596a and the right squamosal of SMNS 12597b (
<figureCitation id="13308FBC310E9F0741C7FC7D395CFBC0" box="[223,336,1021,1045]" captionStart="Figure 11" captionStartId="17.[127,192,1672,1696]" captionTargetBox="[131,1404,147,1640]" captionTargetId="figure-6@17.[127,1407,144,1644]" captionTargetPageId="17" captionText="Figure 11. Details of the cranial anatomy of Saltoposuchus connectens. A, frontals of SMNS 12597a in ventral view; B, less squamosal of SMNS 12596a in dorsal view; C, right squamosal of SMNS 12597b in ventral view; D, less quadrate of SMNS 12597b in posterolateral view; E, less quadrate of SMNS 12596a in anterolateral view; F, enlarged anterior maxillary tooth of SMNS 12352a in angled lateral view.Abbreviations: adp, anterodorsal process; ap, anterior process; dh, dorsal head; fr, frontal; iqf, internal quadrate fenestra; l., less; lcr, longitudinal crest; lr, longitudinal ridge; olb, olfactory bulb; otfl, otoccipital flange; paf, parietal facet; pmp, posteromedial process; pof, postorbital facet; ppeg, posterior peg; prff, prefrontal facet; ptfl, post-temporal flange; ptp, pterygoid process; ptra, pterygoid ramus; qf, quadrate facet; r., right; stfo, supratemporal fossa; vdp, ventrally directed process; vee, ventral end." figureDoi="http://doi.org/10.5281/zenodo.8426173" httpUri="https://zenodo.org/record/8426173/files/figure.png" pageId="19" pageNumber="373">Fig. 11B, C</figureCitation>
). Despite the disarticulation of these elements, the nature of these processes can be established based on observations on
<taxonomicName id="4C0BE8BA310E9F074047FBBC3A74FB81" box="[351,632,1084,1108]" class="Reptilia" family="Sphenosuchidae" genus="Litargosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="19" pageNumber="373" phylum="Chordata" rank="species" species="leptorhynchus">
<emphasis id="B97F4F2B310E9F074047FBBC3A74FB81" box="[351,632,1084,1108]" italics="true" pageId="19" pageNumber="373">Litargosuchus leptorhynchus</emphasis>
</taxonomicName>
(BP/I/5237, unpublished CT data) and
<taxonomicName id="4C0BE8BA310E9F0740B6FBDB3A83FBA6" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[430,655,1115,1139]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="19" pageNumber="373" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B310E9F0740B6FBDB3A83FBA6" box="[430,655,1115,1139]" italics="true" pageId="19" pageNumber="373">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
(NHMUK PV R 7591), which both have a similar squamosal morphology. The first process extends from the posteromedial end of the squamosal and is dubbed the posteromedial process. It is hook-shaped, and would have formed the articulation with the posterolateral process of the parietal. The second process is positioned directly ventrolaterally to the posteromedial process, and both are separated by a rounded opening, the post-temporal fenestra, whichwassmallandsubcircularin
<emphasis id="B97F4F2B310E9F07430AFAD63B08FABB" box="[530,772,1366,1390]" italics="true" pageId="19" pageNumber="373">Saltoposuchusconnectens</emphasis>
. This second process is named the post-temporal flange and formed the lateral margin of the fenestra. It is projected ventrally and slightly posteriorly at its base, but distally it curves distinctly medially. It is incomplete in both SMNS 12596a and SMNS 12597b but would probably have tapered to a point distally, as in
<taxonomicName id="4C0BE8BA310E9F074198F9923950F9FF" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[128,348,1554,1578]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="19" pageNumber="373" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B310E9F074198F9923950F9FF" box="[128,348,1554,1578]" italics="true" pageId="19" pageNumber="373">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0BE8BA310E9F074088F9923AAFF9FF" box="[400,675,1554,1578]" class="Reptilia" family="Sphenosuchidae" genus="Litargosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="19" pageNumber="373" phylum="Chordata" rank="species" species="leptorhynchus">
<emphasis id="B97F4F2B310E9F074088F9923AAFF9FF" box="[400,675,1554,1578]" italics="true" pageId="19" pageNumber="373">Litargosuchus leptorhynchus</emphasis>
</taxonomicName>
. The final process represents the largest of the three. It is positioned lateral to the post-temporal flange and is directed posteroventrally.Since it gradually tapers distally in dorsal view, it is roughly triangular in outline and has a flat, posterodorsally directed surface. This represents the articular surface for the paroccipital process of the otoccipital, and this process is therefore named the otoccipital flange. It is thickened on its lateral side but very dorsoventrally thin on its medial side. The ventral surface in SMNS 12597b possesses a clear concavity at the base of the posteromedial process, which represents the articular surface for the dorsal head of the quadrate. Thus, as in other crocodylomorphs, the dorsal head of the quadrate was positioned medially relative to the ventral end of the quadrate.
</paragraph>
<paragraph id="8BB49339310E9F074221FF0F3C7EFDCA" blockId="19.[825,1475,143,1828]" pageId="19" pageNumber="373">
<emphasis id="B97F4F2B310E9F074221FF0F3B92FF72" box="[825,926,143,167]" italics="true" pageId="19" pageNumber="373">Quadrate:</emphasis>
A partial less quadrate is preserved in both SMNS 12597b and SMNS 12596a (
<figureCitation id="13308FBC310E9F074546FF2F3CA0FF12" box="[1118,1196,175,199]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="19" pageNumber="373">Figs 2C</figureCitation>
,
<figureCitation id="13308FBC310E9F0745AEFF2F3CD8FF12" box="[1206,1236,175,199]" captionStart="Figure 3" captionStartId="4.[113,178,1499,1523]" captionTargetBox="[147,1426,144,1471]" captionTargetId="figure-217@4.[146,1426,144,1471]" captionTargetPageId="4" captionText="Figure 3. Part of Saltoposuchus connectens referred specimen SMNS 12596. A, SMNS 12596a, cranial block; B, SMNS 12596b, two dentaries exposed in ventrolateral view; C, SMNS 12596c exposing a less femur in anterolateral view; D, exposing the opposite side of SMNS 12596c, Abbreviations:de, dentary; dr, dorsal rib; dv, dorsal vertebra; fe, femur; ga, gastralia; l., less; mx, maxilla; na, nasal; os, osteoderm; q, quadrate; r., right; sq, squamosal; sr, sacral rib; sv, sacral vertebra; tp, transverse process." figureDoi="http://doi.org/10.5281/zenodo.8426153" httpUri="https://zenodo.org/record/8426153/files/figure.png" pageId="19" pageNumber="373">3A</figureCitation>
,
<figureCitation id="13308FBC310E9F0745F8FF2F3D28FF13" box="[1248,1316,175,199]" captionStart="Figure 11" captionStartId="17.[127,192,1672,1696]" captionTargetBox="[131,1404,147,1640]" captionTargetId="figure-6@17.[127,1407,144,1644]" captionTargetPageId="17" captionText="Figure 11. Details of the cranial anatomy of Saltoposuchus connectens. A, frontals of SMNS 12597a in ventral view; B, less squamosal of SMNS 12596a in dorsal view; C, right squamosal of SMNS 12597b in ventral view; D, less quadrate of SMNS 12597b in posterolateral view; E, less quadrate of SMNS 12596a in anterolateral view; F, enlarged anterior maxillary tooth of SMNS 12352a in angled lateral view.Abbreviations: adp, anterodorsal process; ap, anterior process; dh, dorsal head; fr, frontal; iqf, internal quadrate fenestra; l., less; lcr, longitudinal crest; lr, longitudinal ridge; olb, olfactory bulb; otfl, otoccipital flange; paf, parietal facet; pmp, posteromedial process; pof, postorbital facet; ppeg, posterior peg; prff, prefrontal facet; ptfl, post-temporal flange; ptp, pterygoid process; ptra, pterygoid ramus; qf, quadrate facet; r., right; stfo, supratemporal fossa; vdp, ventrally directed process; vee, ventral end." figureDoi="http://doi.org/10.5281/zenodo.8426173" httpUri="https://zenodo.org/record/8426173/files/figure.png" pageId="19" pageNumber="373">11D, E</figureCitation>
). The former is exposed in posterolateral view and was originally interpreted as a right quadrate by
<bibRefCitation id="EF9AEEC8310E9F0742E7FF6E3C98FED3" author="Huene FR" box="[1023,1172,238,262]" pageId="19" pageNumber="373" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene (1921)</bibRefCitation>
, whereas the laưer is exposed in anterior view and was originally interpreted as a right palatine. The interpretation offered here corroborates that suggested by
<bibRefCitation id="EF9AEEC8310E9F074221FECB3BC6FEB1" author="Crush PJ" box="[825,970,331,356]" pageId="19" pageNumber="373" refId="ref31157" refString="Crush PJ. An early, terrestrial, crocodile from South Wales. Unpublished Ph. D. Thesis, University College London, 1980." type="book" year="1980">Crush (1980)</bibRefCitation>
. SMNS 12591a also preserves distorted remains of both quadrates (
<figureCitation id="13308FBC310E9F0742E6FEEB3C3AFE56" box="[1022,1078,363,387]" captionStart="Figure 8" captionStartId="9.[129,194,1772,1796]" captionTargetBox="[178,1429,144,1744]" captionTargetId="figure-6@9.[174,1429,144,1744]" captionTargetPageId="9" captionText="Figure 8. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in right lateral view; B, interpretative drawing of A; C, less lateral view; D, interpretative drawing of (C). Abbreviations:an, angular; cc, crista cranii; de, dentary; fr, frontal; j, jugal; la, lacrimal; ls, laterosphenoid; mx, maxilla; na, nasal; pa, parietal; pl, palatine; po, postorbital; prf, prefrontal; pt, pterygoid; q, quadrate; qj, quadratojugal; sa, surangular; sq, squamosal." figureDoi="http://doi.org/10.5281/zenodo.8426167" httpUri="https://zenodo.org/record/8426167/files/figure.png" pageId="19" pageNumber="373">Fig. 8</figureCitation>
). Only the less quadrate was identified by
<bibRefCitation id="EF9AEEC8310E9F074242FE0A3C50FE77" author="Sereno PC &amp; Wild R" box="[858,1116,394,418]" pageId="19" pageNumber="373" pagination="435 - 58" refId="ref32722" refString="Sereno PC, Wild R. Procompsognathus: theropod, ' thecodont' or both? Journal of Vertebrate Paleontology 1992; 12: 435 - 58." type="journal article" year="1992">Sereno and Wild (1992)</bibRefCitation>
, probably because the right quadrate, which is distinctly displaced medially and anteriorly, was still embedded in matrix at that time. The less quadrate is heavily distorted, whereas the dorsal portion of the right quadrate is in comparatively good condition.
</paragraph>
<paragraph id="8BB49339310E9F07424DFDA73D5DFB86" blockId="19.[825,1475,143,1828]" pageId="19" pageNumber="373">
The shass of the quadrate has a mildly but continuously concave posterior margin. The posterolateral surface of the shass is characterized by a distinct and deep excavation in SMNS 12597b and the right quadrate of SMNS 12591a. This excavation fully pierces the shass in SMNS 12597b, but this is almost certainly the result of the shass being partially broken. In SMNS 12597b, the excavation is oval in posterior view, being dorsoventrally taller than mediolaterally wide (
<figureCitation id="13308FBC310E9F0745A0FC823D1CFCCF" box="[1208,1296,770,794]" captionStart="Figure 11" captionStartId="17.[127,192,1672,1696]" captionTargetBox="[131,1404,147,1640]" captionTargetId="figure-6@17.[127,1407,144,1644]" captionTargetPageId="17" captionText="Figure 11. Details of the cranial anatomy of Saltoposuchus connectens. A, frontals of SMNS 12597a in ventral view; B, less squamosal of SMNS 12596a in dorsal view; C, right squamosal of SMNS 12597b in ventral view; D, less quadrate of SMNS 12597b in posterolateral view; E, less quadrate of SMNS 12596a in anterolateral view; F, enlarged anterior maxillary tooth of SMNS 12352a in angled lateral view.Abbreviations: adp, anterodorsal process; ap, anterior process; dh, dorsal head; fr, frontal; iqf, internal quadrate fenestra; l., less; lcr, longitudinal crest; lr, longitudinal ridge; olb, olfactory bulb; otfl, otoccipital flange; paf, parietal facet; pmp, posteromedial process; pof, postorbital facet; ppeg, posterior peg; prff, prefrontal facet; ptfl, post-temporal flange; ptp, pterygoid process; ptra, pterygoid ramus; qf, quadrate facet; r., right; stfo, supratemporal fossa; vdp, ventrally directed process; vee, ventral end." figureDoi="http://doi.org/10.5281/zenodo.8426173" httpUri="https://zenodo.org/record/8426173/files/figure.png" pageId="19" pageNumber="373">Fig. 11D</figureCitation>
). This excavation represents the internal quadrate fenestra, a pneumatic feature of the crocodylomorph quadrate, which is connected to a system of internal diverticula in hallopodids (
<bibRefCitation id="EF9AEEC8310E9F0745D8FCE03D7DFCAD" author="Leardi JM &amp; Pol D &amp; Clark JM" box="[1216,1393,864,888]" pageId="19" pageNumber="373" pagination="2801" refId="ref31922" refString="Leardi JM, Pol D, Clark JM. Detailed anatomy of the braincase of Macelognathus vagans Marsh, 1884 (Archosauria, Crocodylomorpha) using high resolution tomography and new insights on basal crocodylomorph phylogeny. PeerJ 2017; 5: e 2801." type="journal article" year="2017">
Leardi
<emphasis id="B97F4F2B310E9F074411FCE13D34FCAD" box="[1289,1336,864,888]" italics="true" pageId="19" pageNumber="373">et al.</emphasis>
2017
</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310E9F074465FCE03DBDFCAD" author="Leardi JM &amp; Pol D &amp; Clark JM" box="[1405,1457,864,888]" pageId="19" pageNumber="373" pagination="48 - 73" refId="ref31967" refString="Leardi JM, Pol D, Clark JM. Braincase anatomy of Almadasuchus figarii (Archosauria, Crocodylomorpha) and a review of the cranial pneumaticity in the origins of Crocodylomorpha. Journal of Anatomy 2020; 237: 48 - 73." type="journal article" year="2020">2020</bibRefCitation>
). Both in the shape of the fenestra and the lack of internal quadrate diverticula, the morphology of SMNS 12597b is very similar to
<taxonomicName id="4C0BE8BA310E9F074221FC3E3C15FC03" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[825,1049,958,982]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="19" pageNumber="373" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B310E9F074221FC3E3C15FC03" box="[825,1049,958,982]" italics="true" pageId="19" pageNumber="373">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0BE8BA310E9F07454DFC3E3D44FC03" box="[1109,1352,958,982]" class="Reptilia" family="Sphenosuchidae" genus="Dromicosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="19" pageNumber="373" phylum="Chordata" rank="species" species="grallator">
<emphasis id="B97F4F2B310E9F07454DFC3E3D44FC03" box="[1109,1352,958,982]" italics="true" pageId="19" pageNumber="373">Dromicosuchus grallator</emphasis>
</taxonomicName>
(NHMUK PV R 7591,
<bibRefCitation id="EF9AEEC8310E9F0742A9FC5D3C46FC20" author="Sues H-D &amp; Olsen PE &amp; Carter JG" box="[945,1098,989,1013]" pageId="19" pageNumber="373" pagination="329 - 43" refId="ref32828" refString="Sues H-D, Olsen PE, Carter JG et al. A new crocodylomorph archosaur from the Upper Triassic of North Carolina. Journal of Vertebrate Paleontology 2003; 23: 329 - 43." type="journal article" year="2003">
Sues
<emphasis id="B97F4F2B310E9F0742FDFC5E3C1FFC20" box="[997,1043,989,1013]" italics="true" pageId="19" pageNumber="373">et al.</emphasis>
2003
</bibRefCitation>
). The remainder of the posterolateral surface of the quadrate is roughly flat to slightly concave. The posterior margin of the quadrate curves strongly posterodorsally towards its dorsal head, which is transversely narrow.
</paragraph>
<paragraph id="8BB49339310E9F07424DFBDB3BB1F8F1" blockId="19.[825,1475,143,1828]" pageId="19" pageNumber="373">
In lateral view, the anterodorsal portion of the quadrate is expanded and terminates in a roughly rectangular end that is confluent with the more posteromedially placed dorsal head in both SMNS 12597b and the right quadrate of SMNS 12591a. This expansion corresponds to the anterodorsal process described by
<bibRefCitation id="EF9AEEC8310E9F074240FB773BE3FADA" author="Walker AD" box="[856,1007,1271,1295]" pageId="19" pageNumber="373" pagination="1 - 120" refId="ref33031" refString="Walker AD. A revision of Sphenosuchus acutus Haughton, a crocodylomorph reptile from the Elliot Formation (Late Triassic or Early Jurassic) of South Africa. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1990; 330: 1 - 120." type="journal article" year="1990">Walker (1990)</bibRefCitation>
for
<taxonomicName id="4C0BE8BA310E9F074500FB773CEDFADA" authorityName="Haughton" authorityYear="1915" box="[1048,1249,1271,1295]" class="Reptilia" genus="Sphenosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="19" pageNumber="373" phylum="Chordata" rank="species" species="acutus">
<emphasis id="B97F4F2B310E9F074500FB773CEDFADA" box="[1048,1249,1271,1295]" italics="true" pageId="19" pageNumber="373">Sphenosuchus acutus</emphasis>
</taxonomicName>
, although it is considerably anteroposteriorly shorter in
<taxonomicName id="4C0BE8BA310E9F0745BDFA963D97FAFB" authorityName="Huene" authorityYear="1921" box="[1189,1435,1302,1326]" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="19" pageNumber="373" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B310E9F0745BDFA963D97FAFB" box="[1189,1435,1302,1326]" italics="true" pageId="19" pageNumber="373">Saltoposuchus connectens</emphasis>
</taxonomicName>
. Its ventral end of the quadrate is transversely expanded compared to the shass. Its ventral margin is slightly convex in posterior or anterior view. The anterior surface, exposed in SMNS 12596a (
<figureCitation id="13308FBC310E9F07425CFA143B94FA79" box="[836,920,1428,1452]" captionStart="Figure 11" captionStartId="17.[127,192,1672,1696]" captionTargetBox="[131,1404,147,1640]" captionTargetId="figure-6@17.[127,1407,144,1644]" captionTargetPageId="17" captionText="Figure 11. Details of the cranial anatomy of Saltoposuchus connectens. A, frontals of SMNS 12597a in ventral view; B, less squamosal of SMNS 12596a in dorsal view; C, right squamosal of SMNS 12597b in ventral view; D, less quadrate of SMNS 12597b in posterolateral view; E, less quadrate of SMNS 12596a in anterolateral view; F, enlarged anterior maxillary tooth of SMNS 12352a in angled lateral view.Abbreviations: adp, anterodorsal process; ap, anterior process; dh, dorsal head; fr, frontal; iqf, internal quadrate fenestra; l., less; lcr, longitudinal crest; lr, longitudinal ridge; olb, olfactory bulb; otfl, otoccipital flange; paf, parietal facet; pmp, posteromedial process; pof, postorbital facet; ppeg, posterior peg; prff, prefrontal facet; ptfl, post-temporal flange; ptp, pterygoid process; ptra, pterygoid ramus; qf, quadrate facet; r., right; stfo, supratemporal fossa; vdp, ventrally directed process; vee, ventral end." figureDoi="http://doi.org/10.5281/zenodo.8426173" httpUri="https://zenodo.org/record/8426173/files/figure.png" pageId="19" pageNumber="373">Fig. 11E</figureCitation>
), is mostly transversely concave. The pterygoid ramus is directed anteriorly, medially, and ventrally. Although it is incomplete in SMNS 12596a, it is clear that it is mediolaterally thin and that it gradually reduces in dorsoventral height distally. Its ventral margin is strongly concave proximally, whereas its broken dorsal margin is slightly convex. The ventral margin is thickened relative to the rest of the ramus and its margin is distinctly curved medially. A small ventral projection is present near the distal end of the ramus (
<figureCitation id="13308FBC310E9F074542F90E3CA1F973" box="[1114,1197,1678,1702]" captionStart="Figure 11" captionStartId="17.[127,192,1672,1696]" captionTargetBox="[131,1404,147,1640]" captionTargetId="figure-6@17.[127,1407,144,1644]" captionTargetPageId="17" captionText="Figure 11. Details of the cranial anatomy of Saltoposuchus connectens. A, frontals of SMNS 12597a in ventral view; B, less squamosal of SMNS 12596a in dorsal view; C, right squamosal of SMNS 12597b in ventral view; D, less quadrate of SMNS 12597b in posterolateral view; E, less quadrate of SMNS 12596a in anterolateral view; F, enlarged anterior maxillary tooth of SMNS 12352a in angled lateral view.Abbreviations: adp, anterodorsal process; ap, anterior process; dh, dorsal head; fr, frontal; iqf, internal quadrate fenestra; l., less; lcr, longitudinal crest; lr, longitudinal ridge; olb, olfactory bulb; otfl, otoccipital flange; paf, parietal facet; pmp, posteromedial process; pof, postorbital facet; ppeg, posterior peg; prff, prefrontal facet; ptfl, post-temporal flange; ptp, pterygoid process; ptra, pterygoid ramus; qf, quadrate facet; r., right; stfo, supratemporal fossa; vdp, ventrally directed process; vee, ventral end." figureDoi="http://doi.org/10.5281/zenodo.8426173" httpUri="https://zenodo.org/record/8426173/files/figure.png" pageId="19" pageNumber="373">Fig. 11E</figureCitation>
), which might be homologous with the ‘pterygoid process’ of
<taxonomicName id="4C0BE8BA310E9F0745BAF92E3B75F930" authority="(Walker 1990)" baseAuthorityName="Walker" baseAuthorityYear="1990" class="Reptilia" genus="Sphenosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="19" pageNumber="373" phylum="Chordata" rank="species" species="acutus">
<emphasis id="B97F4F2B310E9F0745BAF92E3D66F910" box="[1186,1386,1710,1734]" italics="true" pageId="19" pageNumber="373">Sphenosuchus acutus</emphasis>
(
<bibRefCitation id="EF9AEEC8310E9F074462F92E3B61F930" author="Walker AD" pageId="19" pageNumber="373" pagination="1 - 120" refId="ref33031" refString="Walker AD. A revision of Sphenosuchus acutus Haughton, a crocodylomorph reptile from the Elliot Formation (Late Triassic or Early Jurassic) of South Africa. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1990; 330: 1 - 120." type="journal article" year="1990">Walker 1990</bibRefCitation>
)
</taxonomicName>
. A pterygoid process is also present in the hallopodids
<taxonomicName id="4C0BE8BA310E9F074221F96C3C16F8D1" authorityName="Marsh" authorityYear="1884" box="[825,1050,1772,1796]" class="Reptilia" family="Hypsilophodontidae" genus="Macelognathus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="19" pageNumber="373" phylum="Chordata" rank="species" species="vagans">
<emphasis id="B97F4F2B310E9F074221F96C3C16F8D1" box="[825,1050,1772,1796]" italics="true" pageId="19" pageNumber="373">Macelognathus vagans</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0BE8BA310E9F07454FF96C3D26F8D1" box="[1111,1322,1772,1796]" genus="Almadasuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="19" pageNumber="373" phylum="Chordata" rank="species" species="figarii">
<emphasis id="B97F4F2B310E9F07454FF96C3D26F8D1" box="[1111,1322,1772,1796]" italics="true" pageId="19" pageNumber="373">Almadasuchus figarii</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9AEEC8310E9F074458F96C3B61F8F1" author="Leardi JM &amp; Pol D &amp; Clark JM" pageId="19" pageNumber="373" pagination="2801" refId="ref31922" refString="Leardi JM, Pol D, Clark JM. Detailed anatomy of the braincase of Macelognathus vagans Marsh, 1884 (Archosauria, Crocodylomorpha) using high resolution tomography and new insights on basal crocodylomorph phylogeny. PeerJ 2017; 5: e 2801." type="journal article" year="2017">
Leardi
<emphasis id="B97F4F2B310E9F074496F96D3DCFF8D1" box="[1422,1475,1772,1796]" italics="true" pageId="19" pageNumber="373">et al.</emphasis>
2017
</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310E9F074260F88C3BA0F8F1" author="Leardi JM &amp; Pol D &amp; Clark JM" box="[888,940,1804,1828]" pageId="19" pageNumber="373" pagination="48 - 73" refId="ref31967" refString="Leardi JM, Pol D, Clark JM. Braincase anatomy of Almadasuchus figarii (Archosauria, Crocodylomorpha) and a review of the cranial pneumaticity in the origins of Crocodylomorpha. Journal of Anatomy 2020; 237: 48 - 73." type="journal article" year="2020">2020</bibRefCitation>
).
</paragraph>
<paragraph id="8BB49339310E9F004221F8CE38BAFEF0" blockId="19.[825,1475,1870,1988]" lastBlockId="20.[113,762,144,293]" lastPageId="20" lastPageNumber="374" pageId="19" pageNumber="373">
<emphasis id="B97F4F2B310E9F074221F8CE3BDCF8B3" box="[825,976,1870,1894]" italics="true" pageId="19" pageNumber="373">Quadratojugal:</emphasis>
Poorly preserved remains are tentatively identified as the remnants of the less quadratojugal in SMNS 12591a. This is an anteroposteriorly thin bone that is strongly curved so that its anterior margin is strongly convex, and its posterior margin concave, in lateral view (
<figureCitation id="13308FBC31099F0040D1FF103A34FF72" box="[457,568,144,168]" captionStart="Figure 8" captionStartId="9.[129,194,1772,1796]" captionTargetBox="[178,1429,144,1744]" captionTargetId="figure-6@9.[174,1429,144,1744]" captionTargetPageId="9" captionText="Figure 8. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in right lateral view; B, interpretative drawing of A; C, less lateral view; D, interpretative drawing of (C). Abbreviations:an, angular; cc, crista cranii; de, dentary; fr, frontal; j, jugal; la, lacrimal; ls, laterosphenoid; mx, maxilla; na, nasal; pa, parietal; pl, palatine; po, postorbital; prf, prefrontal; pt, pterygoid; q, quadrate; qj, quadratojugal; sa, surangular; sq, squamosal." figureDoi="http://doi.org/10.5281/zenodo.8426167" httpUri="https://zenodo.org/record/8426167/files/figure.png" pageId="20" pageNumber="374">Fig. 8C, D</figureCitation>
). However, this is probably the result of the distortion if this indeed represents the quadratojugal, as this strongly deviates from the straighter quadratojugals observed in other crocodylomorphs (e.g.
<bibRefCitation id="EF9AEEC831099F0043AAFF6E38A9FEF0" author="Walker AD" pageId="20" pageNumber="374" pagination="1 - 120" refId="ref33031" refString="Walker AD. A revision of Sphenosuchus acutus Haughton, a crocodylomorph reptile from the Elliot Formation (Late Triassic or Early Jurassic) of South Africa. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1990; 330: 1 - 120." type="journal article" year="1990">Walker 1990</bibRefCitation>
).
</paragraph>
<paragraph id="8BB4933931099F004169FEC7394FFCE1" blockId="20.[113,763,326,820]" pageId="20" pageNumber="374">
<emphasis id="B97F4F2B31099F004169FEC738B4FE8B" box="[113,184,327,350]" italics="true" pageId="20" pageNumber="374">Vomer:</emphasis>
The additional preparation of SMNS 12591a allows for a more extensive description of the palate than was previously possible (
<bibRefCitation id="EF9AEEC831099F0041CEFE05396EFE48" author="Ostrom JH" box="[214,354,389,413]" pageId="20" pageNumber="374" pagination="179 - 95" refId="ref32208" refString="Ostrom JH. Procompsognathus - Theropod or Thecodont? Palaeontographica A 1981; 175: 179 - 95." type="journal article" year="1981">Ostrom 1981</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831099F004077FE053A55FE48" author="Sereno PC &amp; Wild R" box="[367,601,389,413]" pageId="20" pageNumber="374" pagination="435 - 58" refId="ref32722" refString="Sereno PC, Wild R. Procompsognathus: theropod, ' thecodont' or both? Journal of Vertebrate Paleontology 1992; 12: 435 - 58." type="journal article" year="1992">Sereno and Wild 1992</bibRefCitation>
), including the first observations of the vomers, which are partially exposed in ventral view and badly cracked (
<figureCitation id="13308FBC31099F0040AFFE443A12FE0E" box="[439,542,452,476]" captionStart="Figure 9" captionStartId="10.[113,178,1772,1796]" captionTargetBox="[222,1352,144,1744]" captionTargetId="figure-6@10.[220,1352,144,1744]" captionTargetPageId="10" captionText="Figure 9. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in dorsal view; B, interpretative drawing of (A); C, ventral view; D, interpretative drawing of (C). Abbreviations:an, angular; antfo, antorbital fossa; bpp, basipterygoid process; crb, ceratobranchial I; de, dentary; ect, ectopterygoid; fr, frontal; j, jugal; la, lacrimal; mx, maxilla; na, nasal; pa, parietal; pbs, parabasisphenoid; pl, palatine; plf, palatine fossa; po, postorbital; pra, pre-articular; prf, prefrontal; pt, pterygoid; ptmr, pterygoid medial rim; sa, surangular; spl, splenial; sq, squamosal; vo, vomer." figureDoi="http://doi.org/10.5281/zenodo.8426169" httpUri="https://zenodo.org/record/8426169/files/figure.png" pageId="20" pageNumber="374">Fig. 9C, D</figureCitation>
). They are apparently unfused and form elongate, transversely thin, rod-like structures that connect to the palatine and the anterior end of the pterygoid posteriorly and to the palatal contribution of the maxilla anteriorly. Because the vomers are so narrow, the choanae, of which they form the medial margin, are correspondingly wide, similar to the condition seen in
<taxonomicName id="4C0BE8BA31099F004098FD003AFAFD4D" authority="(Walker 1990)" baseAuthorityName="Walker" baseAuthorityYear="1990" box="[384,758,640,664]" class="Reptilia" genus="Sphenosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="20" pageNumber="374" phylum="Chordata" rank="species" species="acutus">
<emphasis id="B97F4F2B31099F004098FD003A42FD4D" box="[384,590,640,664]" italics="true" pageId="20" pageNumber="374">Sphenosuchus acutus</emphasis>
(
<bibRefCitation id="EF9AEEC831099F00437BFD003AE5FD4D" author="Walker AD" box="[611,745,640,664]" pageId="20" pageNumber="374" pagination="1 - 120" refId="ref33031" refString="Walker AD. A revision of Sphenosuchus acutus Haughton, a crocodylomorph reptile from the Elliot Formation (Late Triassic or Early Jurassic) of South Africa. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1990; 330: 1 - 120." type="journal article" year="1990">Walker 1990</bibRefCitation>
)
</taxonomicName>
. In contrast,
<bibRefCitation id="EF9AEEC831099F0041E8FD1F3991FD62" author="Gauthier JA" box="[240,413,671,695]" pageId="20" pageNumber="374" pagination="1 - 55" refId="ref31317" refString="Gauthier JA. Saurischian monophyly and the origin of birds. Memoirs of the California Academy of Sciences 1986; 8: 1 - 55." type="journal article" year="1986">Gauthier (1986)</bibRefCitation>
considered the vomers of SMNS 12591a to be fused anteriorly, which supported a theropod interpretation of this specimen. It is unclear on which this interpretation was based, since it predates the additional preparation that revealed the vomers.
</paragraph>
<paragraph id="8BB4933931099F004169FCE139F7F8B1" blockId="20.[113,763,865,1892]" pageId="20" pageNumber="374">
<emphasis id="B97F4F2B31099F004169FCE138C4FCAC" box="[113,200,865,889]" italics="true" pageId="20" pageNumber="374">Palatine:</emphasis>
Both palatines are present but badly preserved in SMNS 12591a (
<figureCitation id="13308FBC31099F004032FC0039AFFC4D" box="[298,419,896,921]" captionStart="Figure 9" captionStartId="10.[113,178,1772,1796]" captionTargetBox="[222,1352,144,1744]" captionTargetId="figure-6@10.[220,1352,144,1744]" captionTargetPageId="10" captionText="Figure 9. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in dorsal view; B, interpretative drawing of (A); C, ventral view; D, interpretative drawing of (C). Abbreviations:an, angular; antfo, antorbital fossa; bpp, basipterygoid process; crb, ceratobranchial I; de, dentary; ect, ectopterygoid; fr, frontal; j, jugal; la, lacrimal; mx, maxilla; na, nasal; pa, parietal; pbs, parabasisphenoid; pl, palatine; plf, palatine fossa; po, postorbital; pra, pre-articular; prf, prefrontal; pt, pterygoid; ptmr, pterygoid medial rim; sa, surangular; spl, splenial; sq, squamosal; vo, vomer." figureDoi="http://doi.org/10.5281/zenodo.8426169" httpUri="https://zenodo.org/record/8426169/files/figure.png" pageId="20" pageNumber="374">Fig. 9C, D</figureCitation>
). The palatine articulates with the pterygoid along its medial margin for most of its length. Additionally, it also connects to the maxilla, and possibly the prefrontal and lacrimal, laterally and the vomer anteromedially. As indicated above, together with the rest of the anterior palatal region, the palatines have been diagenetically compressed dorsally against the nasals and, consequently, they are visible through the antorbital fenestra in lateral view on both sides. The less palatine is best preserved; it is a plate-like bone with a distinctly concave anterior margin in ventral view, which forms the posterior margin of the choana. A distinct rim on the choanal margin, as described for
<taxonomicName id="4C0BE8BA31099F004035FB5939F1FB24" baseAuthorityName="Walker" baseAuthorityYear="1990" box="[301,509,1241,1265]" class="Reptilia" genus="Sphenosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="20" pageNumber="374" phylum="Chordata" rank="species" species="acutus">
<emphasis id="B97F4F2B31099F004035FB5939F1FB24" box="[301,509,1241,1265]" italics="true" pageId="20" pageNumber="374">Sphenosuchus acutus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0BE8BA31099F004308FB593AF9FB24" authorityName="Clark &amp; Sues" authorityYear="2002" box="[528,757,1241,1265]" class="Reptilia" family="Sphenosuchidae" genus="Kayentasuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="20" pageNumber="374" phylum="Chordata" rank="species" species="walkeri">
<emphasis id="B97F4F2B31099F004308FB593AF9FB24" box="[528,757,1241,1265]" italics="true" pageId="20" pageNumber="374">Kayentasuchus walkeri</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0BE8BA31099F004169FB78396CFAC5" authorityName="Simmons" authorityYear="1965" box="[113,352,1272,1296]" class="Reptilia" family="Sphenosuchidae" genus="Dibothrosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="20" pageNumber="374" phylum="Chordata" rank="species" species="elaphros">
<emphasis id="B97F4F2B31099F004169FB78396CFAC5" box="[113,352,1272,1296]" italics="true" pageId="20" pageNumber="374">Dibothrosuchus elaphros</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C0BE8BA31099F004080FB783A6DFAC5" box="[408,609,1272,1296]" class="Reptilia" genus="Junggarsuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="20" pageNumber="374" phylum="Chordata" rank="species" species="sloani">
<emphasis id="B97F4F2B31099F004080FB783A6DFAC5" box="[408,609,1272,1296]" italics="true" pageId="20" pageNumber="374">Junggarsuchus sloani</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9AEEC831099F00436AFB783AF8FAC5" author="Walker AD" box="[626,756,1272,1296]" pageId="20" pageNumber="374" pagination="1 - 120" refId="ref33031" refString="Walker AD. A revision of Sphenosuchus acutus Haughton, a crocodylomorph reptile from the Elliot Formation (Late Triassic or Early Jurassic) of South Africa. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1990; 330: 1 - 120." type="journal article" year="1990">Walker 1990</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831099F004169FA9838FBFAE5" author="Nesbiu SJ" box="[113,247,1304,1328]" pageId="20" pageNumber="374" pagination="1 - 292" refId="ref32089" refString="Nesbiu SJ. The early evolution of archosaurs: relationships and the origin of major clades. Bulletin of the American Museum of Natural History 2011; 352: 1 - 292." type="journal article" year="2011">Nesbiư 2011</bibRefCitation>
: ch. 91,
<bibRefCitation id="EF9AEEC831099F004057FA983A32FAE5" author="Ruebenstahl AA &amp; Klein MD &amp; Yi H" box="[335,574,1304,1328]" pageId="20" pageNumber="374" pagination="2463 - 556" refId="ref32564" refString="Ruebenstahl AA, Klein MD, Yi H et al. Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros. The Anatomical Record 2022; 305: 2463 - 556." type="journal article" year="2022">
Ruebenstahl
<emphasis id="B97F4F2B31099F0040CDFA983A09FAE5" box="[469,517,1304,1328]" italics="true" pageId="20" pageNumber="374">et al.</emphasis>
2022
</bibRefCitation>
), is absent, as it is in
<taxonomicName id="4C0BE8BA31099F004194FAB73964FA9A" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[140,360,1335,1359]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="20" pageNumber="374" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B31099F004194FAB73964FA9A" box="[140,360,1335,1359]" italics="true" pageId="20" pageNumber="374">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
(NHMUK PV R 38291,
<bibRefCitation id="EF9AEEC831099F004377FAB73AE5FA9A" author="Crush PJ" box="[623,745,1335,1359]" pageId="20" pageNumber="374" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
). The exact posterior extent of the palatine is unknown but it is clear that the posterior portion is considerably longer than the anterior portion, as is also the case in other non-crocodyliform crocodylomorphs (e.g.
<bibRefCitation id="EF9AEEC831099F004071FA3439E4FA19" author="Crush PJ" box="[361,488,1460,1484]" pageId="20" pageNumber="374" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831099F0040E1FA343A8CFA19" author="Walker AD" box="[505,640,1460,1484]" pageId="20" pageNumber="374" pagination="1 - 120" refId="ref33031" refString="Walker AD. A revision of Sphenosuchus acutus Haughton, a crocodylomorph reptile from the Elliot Formation (Late Triassic or Early Jurassic) of South Africa. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1990; 330: 1 - 120." type="journal article" year="1990">Walker 1990</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831099F004388FA3438D5FA39" author="Clark JM &amp; Sues H-D" pageId="20" pageNumber="374" pagination="77 - 95" refId="ref30970" refString="Clark JM, Sues H-D. Two new basal crocodylomorph archosaurs from the Lower Jurassic and the monophyly of the Sphenosuchia. Zoological Journal of the Linnean Society 2002; 136: 77 - 95." type="journal article" year="2002">Clark and Sues 2002</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831099F0041FBFA5439C1FA39" author="Ruebenstahl AA &amp; Klein MD &amp; Yi H" box="[227,461,1492,1516]" pageId="20" pageNumber="374" pagination="2463 - 556" refId="ref32564" refString="Ruebenstahl AA, Klein MD, Yi H et al. Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros. The Anatomical Record 2022; 305: 2463 - 556." type="journal article" year="2022">
Ruebenstahl
<emphasis id="B97F4F2B31099F00407FFA543999FA39" box="[359,405,1492,1516]" italics="true" pageId="20" pageNumber="374">et al.</emphasis>
2022
</bibRefCitation>
). The posterolateral margin is probably incomplete on both sides. However, the preserved portion of this margin, which forms the anterior and anteromedial margins of the suborbital fenestra, is continuously and widely concave in ventral view. The nature of the lateral suture with the maxilla is unclear, while the medial suture with the pterygoid is straight. The ventral surface of the less palatine possesses a very large, oval fossa, which is outlined by a radially striated surface medial to it (
<figureCitation id="13308FBC31099F0041ECF94E3957F933" box="[244,347,1742,1766]" captionStart="Figure 9" captionStartId="10.[113,178,1772,1796]" captionTargetBox="[222,1352,144,1744]" captionTargetId="figure-6@10.[220,1352,144,1744]" captionTargetPageId="10" captionText="Figure 9. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in dorsal view; B, interpretative drawing of (A); C, ventral view; D, interpretative drawing of (C). Abbreviations:an, angular; antfo, antorbital fossa; bpp, basipterygoid process; crb, ceratobranchial I; de, dentary; ect, ectopterygoid; fr, frontal; j, jugal; la, lacrimal; mx, maxilla; na, nasal; pa, parietal; pbs, parabasisphenoid; pl, palatine; plf, palatine fossa; po, postorbital; pra, pre-articular; prf, prefrontal; pt, pterygoid; ptmr, pterygoid medial rim; sa, surangular; spl, splenial; sq, squamosal; vo, vomer." figureDoi="http://doi.org/10.5281/zenodo.8426169" httpUri="https://zenodo.org/record/8426169/files/figure.png" pageId="20" pageNumber="374">Fig. 9C, D</figureCitation>
). This morphology is unknown for other crocodylomorphs and appears to be unique to
<taxonomicName id="4C0BE8BA31099F00436AF96E38DBF8F0" authorityName="Huene" authorityYear="1921" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="20" pageNumber="374" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B31099F00436AF96E38DBF8F0" italics="true" pageId="20" pageNumber="374">Saltoposuchus connectens</emphasis>
</taxonomicName>
. The dorsal surface of the palatine as seen through the antorbital fenestra is thickened but otherwise too incompletely preserved to comment on (
<figureCitation id="13308FBC31099F00409FF8CB39E6F8B6" box="[391,490,1867,1892]" captionStart="Figure 9" captionStartId="10.[113,178,1772,1796]" captionTargetBox="[222,1352,144,1744]" captionTargetId="figure-6@10.[220,1352,144,1744]" captionTargetPageId="10" captionText="Figure 9. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in dorsal view; B, interpretative drawing of (A); C, ventral view; D, interpretative drawing of (C). Abbreviations:an, angular; antfo, antorbital fossa; bpp, basipterygoid process; crb, ceratobranchial I; de, dentary; ect, ectopterygoid; fr, frontal; j, jugal; la, lacrimal; mx, maxilla; na, nasal; pa, parietal; pbs, parabasisphenoid; pl, palatine; plf, palatine fossa; po, postorbital; pra, pre-articular; prf, prefrontal; pt, pterygoid; ptmr, pterygoid medial rim; sa, surangular; spl, splenial; sq, squamosal; vo, vomer." figureDoi="http://doi.org/10.5281/zenodo.8426169" httpUri="https://zenodo.org/record/8426169/files/figure.png" pageId="20" pageNumber="374">Fig. 9A, B</figureCitation>
).
</paragraph>
<paragraph id="8BB4933931099F004169F8053C5CFDA8" blockId="20.[113,762,1925,2012]" lastBlockId="20.[810,1460,143,637]" pageId="20" pageNumber="374">
<emphasis id="B97F4F2B31099F004169F80538F2F848" box="[113,254,1925,1949]" italics="true" pageId="20" pageNumber="374">Ectopterygoid:</emphasis>
Both ectopterygoids are present but incomplete and poorly preserved in SMNS 12591a, with the right element being best preserved (
<figureCitation id="13308FBC31099F0040CBF8433A19F809" box="[467,533,1987,2012]" captionStart="Figure 9" captionStartId="10.[113,178,1772,1796]" captionTargetBox="[222,1352,144,1744]" captionTargetId="figure-6@10.[220,1352,144,1744]" captionTargetPageId="10" captionText="Figure 9. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in dorsal view; B, interpretative drawing of (A); C, ventral view; D, interpretative drawing of (C). Abbreviations:an, angular; antfo, antorbital fossa; bpp, basipterygoid process; crb, ceratobranchial I; de, dentary; ect, ectopterygoid; fr, frontal; j, jugal; la, lacrimal; mx, maxilla; na, nasal; pa, parietal; pbs, parabasisphenoid; pl, palatine; plf, palatine fossa; po, postorbital; pra, pre-articular; prf, prefrontal; pt, pterygoid; ptmr, pterygoid medial rim; sa, surangular; spl, splenial; sq, squamosal; vo, vomer." figureDoi="http://doi.org/10.5281/zenodo.8426169" httpUri="https://zenodo.org/record/8426169/files/figure.png" pageId="20" pageNumber="374">Fig. 9</figureCitation>
). The ectopterygoid contacts the pterygoid posteromedially and the jugal laterally. The ectopterygoid is characterized by a distinctly curved shass that possesses a convex anterior and concave posterior margin in dorsal or ventral view. Furthermore, the lateral side of the ectopterygoid is positioned considerably further dorsally than its posteromedial side. The articular facet with the jugal is composed of an anterior and posterior process, and as such, it is considerably anteroposteriorly longer than dorsoventrally tall, as is also the case in other non-crocodyliform crocodylomorphs (e.g.
<bibRefCitation id="EF9AEEC831099F004232FE293BA2FE17" author="Walker AD" box="[810,942,425,450]" pageId="20" pageNumber="374" pagination="1 - 120" refId="ref33031" refString="Walker AD. A revision of Sphenosuchus acutus Haughton, a crocodylomorph reptile from the Elliot Formation (Late Triassic or Early Jurassic) of South Africa. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1990; 330: 1 - 120." type="journal article" year="1990">Walker 1990</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831099F0042A3FE293CA2FE17" author="Ruebenstahl AA &amp; Klein MD &amp; Yi H" box="[955,1198,425,450]" pageId="20" pageNumber="374" pagination="2463 - 556" refId="ref32564" refString="Ruebenstahl AA, Klein MD, Yi H et al. Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros. The Anatomical Record 2022; 305: 2463 - 556." type="journal article" year="2022">
Ruebenstahl
<emphasis id="B97F4F2B31099F00455BFE2A3C78FE14" box="[1091,1140,425,449]" italics="true" pageId="20" pageNumber="374">et al.</emphasis>
2022
</bibRefCitation>
). The anterior process is displaced ventrally relative to rest of the lateral end, whereas the posterior process curves posterodorsally. The exact articulation with the pterygoid is somewhat unclear, but it is obvious that the ectopterygoid expands lateromedially near its posteromedial end and it articulates on the anterolateral surface of the transverse flange of the pterygoid.
</paragraph>
<paragraph id="8BB4933931099F004232FD1F3B97FA73" blockId="20.[810,1459,671,2011]" pageId="20" pageNumber="374">
<emphasis id="B97F4F2B31099F004232FD1F3B82FD62" box="[810,910,671,695]" italics="true" pageId="20" pageNumber="374">Pterygoid:</emphasis>
Both pterygoids are present but incomplete and somewhat poorly preserved in SMNS 12591a (
<figureCitation id="13308FBC31099F004436FD3E3DAFFD03" box="[1326,1443,702,726]" captionStart="Figure 9" captionStartId="10.[113,178,1772,1796]" captionTargetBox="[222,1352,144,1744]" captionTargetId="figure-6@10.[220,1352,144,1744]" captionTargetPageId="10" captionText="Figure 9. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in dorsal view; B, interpretative drawing of (A); C, ventral view; D, interpretative drawing of (C). Abbreviations:an, angular; antfo, antorbital fossa; bpp, basipterygoid process; crb, ceratobranchial I; de, dentary; ect, ectopterygoid; fr, frontal; j, jugal; la, lacrimal; mx, maxilla; na, nasal; pa, parietal; pbs, parabasisphenoid; pl, palatine; plf, palatine fossa; po, postorbital; pra, pre-articular; prf, prefrontal; pt, pterygoid; ptmr, pterygoid medial rim; sa, surangular; spl, splenial; sq, squamosal; vo, vomer." figureDoi="http://doi.org/10.5281/zenodo.8426169" httpUri="https://zenodo.org/record/8426169/files/figure.png" pageId="20" pageNumber="374">Fig. 9C, D</figureCitation>
). The anterior portion of the pterygoids is deformed dorsally and crushed against the dorsal roof of the snout. The pterygoid articulates anteriorly with the vomer and anterolaterally with the palatine. Laterally it articulates with the ectopterygoid; the posterolateral articulation with the quadrate is not preserved or exposed. Posterodorsally, the pterygoid articulates with the parabasisphenoid. The palatal ramus of the pterygoid is very elongate and reached approximately until the anterior end of the antorbital fenestra. Its lateral margin gradually tapers anteriorly and is partially ventrally overlapped by the palatine. The medial margin is characterized by a thickened, ventrally directed rim along its entire length, as is also the case in
<taxonomicName id="4C0BE8BA31099F00443AFBB63BA5FBB8" authority=", CM" authorityName="CM" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="20" pageNumber="374" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B31099F00443AFBB63B62FBB8" italics="true" pageId="20" pageNumber="374">Terrestrisuchus gracilis</emphasis>
, CM
</taxonomicName>
29894,
<taxonomicName id="4C0BE8BA31099F00451FFBD53CD4FBB8" baseAuthorityName="Walker" baseAuthorityYear="1990" box="[1031,1240,1109,1133]" class="Reptilia" genus="Sphenosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="20" pageNumber="374" phylum="Chordata" rank="species" species="acutus">
<emphasis id="B97F4F2B31099F00451FFBD53CD4FBB8" box="[1031,1240,1109,1133]" italics="true" pageId="20" pageNumber="374">Sphenosuchus acutus</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C0BE8BA31099F004404FBD53B77FB58" authorityName="Simmons" authorityYear="1965" class="Reptilia" family="Sphenosuchidae" genus="Dibothrosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="20" pageNumber="374" phylum="Chordata" rank="species" species="elaphros">
<emphasis id="B97F4F2B31099F004404FBD53B77FB58" italics="true" pageId="20" pageNumber="374">Dibothrosuchus elaphros</emphasis>
</taxonomicName>
(e.g.
<bibRefCitation id="EF9AEEC831099F0042D9FBF53C4EFB58" author="Crush PJ" box="[961,1090,1141,1165]" pageId="20" pageNumber="374" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831099F00454CFBF53CD2FB58" author="Walker AD" box="[1108,1246,1141,1165]" pageId="20" pageNumber="374" pagination="1 - 120" refId="ref33031" refString="Walker AD. A revision of Sphenosuchus acutus Haughton, a crocodylomorph reptile from the Elliot Formation (Late Triassic or Early Jurassic) of South Africa. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1990; 330: 1 - 120." type="journal article" year="1990">Walker 1990</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831099F0045E9FBF53DA1FB58" author="Clark JM &amp; Sues H-D &amp; Berman DS" box="[1265,1453,1141,1165]" pageId="20" pageNumber="374" pagination="683 - 704" refId="ref31006" refString="Clark JM, Sues H-D, Berman DS. A new specimen of Hesperosuchus agilis from the Upper Triassic of New Mexico and the interrelationships of basal crocodylomorph archosaurs. Journal of Vertebrate Paleontology 2000; 20: 683 - 704." type="journal article" year="2000">
Clark
<emphasis id="B97F4F2B31099F00442EFBF53D61FB58" box="[1334,1389,1141,1165]" italics="true" pageId="20" pageNumber="374">et al.</emphasis>
2000
</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831099F004232FB143C2CFB79" author="Ruebenstahl AA &amp; Klein MD &amp; Yi H" box="[810,1056,1172,1196]" pageId="20" pageNumber="374" pagination="2463 - 556" refId="ref32564" refString="Ruebenstahl AA, Klein MD, Yi H et al. Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros. The Anatomical Record 2022; 305: 2463 - 556." type="journal article" year="2022">
Ruebenstahl
<emphasis id="B97F4F2B31099F0042ABFB153BE9FB79" box="[947,997,1172,1196]" italics="true" pageId="20" pageNumber="374">et al.</emphasis>
2022
</bibRefCitation>
). This rim is mostly transversely thin, but it thickens approximately anterior to the level of the palatine–maxilla articulation. Such a transverse thickening is also observed in
<taxonomicName id="4C0BE8BA31099F0042B0FB723D1DFADF" authority="(Walker 1990)" baseAuthorityName="Walker" baseAuthorityYear="1990" box="[936,1297,1266,1290]" class="Reptilia" genus="Sphenosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="20" pageNumber="374" phylum="Chordata" rank="species" species="acutus">
<emphasis id="B97F4F2B31099F0042B0FB723C7EFADF" box="[936,1138,1266,1290]" italics="true" pageId="20" pageNumber="374">Sphenosuchus acutus</emphasis>
(
<bibRefCitation id="EF9AEEC831099F00459BFB723D09FADF" author="Walker AD" box="[1155,1285,1266,1290]" pageId="20" pageNumber="374" pagination="1 - 120" refId="ref33031" refString="Walker AD. A revision of Sphenosuchus acutus Haughton, a crocodylomorph reptile from the Elliot Formation (Late Triassic or Early Jurassic) of South Africa. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1990; 330: 1 - 120." type="journal article" year="1990">Walker 1990</bibRefCitation>
)
</taxonomicName>
. This thickened anterior portion of the medial rim was interpreted as the vomer by
<bibRefCitation id="EF9AEEC831099F004252FAB13BE7FA9C" author="Ostrom JH" box="[842,1003,1329,1353]" pageId="20" pageNumber="374" pagination="179 - 95" refId="ref32208" refString="Ostrom JH. Procompsognathus - Theropod or Thecodont? Palaeontographica A 1981; 175: 179 - 95." type="journal article" year="1981">Ostrom (1981)</bibRefCitation>
, but later correctly identified by
<bibRefCitation id="EF9AEEC831099F004426FAB13BA1FABD" author="Sereno PC &amp; Wild R" pageId="20" pageNumber="374" pagination="435 - 58" refId="ref32722" refString="Sereno PC, Wild R. Procompsognathus: theropod, ' thecodont' or both? Journal of Vertebrate Paleontology 1992; 12: 435 - 58." type="journal article" year="1992">Sereno and Wild (1992)</bibRefCitation>
. The remainder of the ventral surface of the pterygoid is badly broken on both sides but appears to be smooth and mostly flat.
</paragraph>
<paragraph id="8BB4933931099F01425DFA2E3960FE90" blockId="20.[810,1459,671,2011]" lastBlockId="21.[129,778,144,325]" lastPageId="21" lastPageNumber="375" pageId="20" pageNumber="374">
The pterygoid connects to its antimere along the entire length of the palatal ramus and an interpterygoid vacuity was therefore absent. This corresponds to the condition in all other non-crocodyliform crocodylomorphs known from articulated palates with well preserved pterygoids (
<bibRefCitation id="EF9AEEC831099F004561F9AC3D12F996" author="Bonaparte JF" box="[1145,1310,1579,1603]" pageId="20" pageNumber="374" pagination="1 - 183" refId="ref30694" refString="Bonaparte JF. Los tetrapodos del sector superior de la Formacion Los Colorados, La Rioja, Argentina (Triasico Superior). 1 Parte. Opera Lilloana 1972; 22: 1 - 183." type="journal article" year="1972">Bonaparte 1972</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831099F004432F9AB3DA1F996" author="Walker AD" box="[1322,1453,1579,1603]" pageId="20" pageNumber="374" pagination="1 - 120" refId="ref33031" refString="Walker AD. A revision of Sphenosuchus acutus Haughton, a crocodylomorph reptile from the Elliot Formation (Late Triassic or Early Jurassic) of South Africa. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1990; 330: 1 - 120." type="journal article" year="1990">Walker 1990</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831099F004232F9CA3BD5F9B6" author="Clark JM &amp; Sues H-D &amp; Berman DS" box="[810,985,1610,1635]" pageId="20" pageNumber="374" pagination="683 - 704" refId="ref31006" refString="Clark JM, Sues H-D, Berman DS. A new specimen of Hesperosuchus agilis from the Upper Triassic of New Mexico and the interrelationships of basal crocodylomorph archosaurs. Journal of Vertebrate Paleontology 2000; 20: 683 - 704." type="journal article" year="2000">
Clark
<emphasis id="B97F4F2B31099F004273F9CB3B92F9B7" box="[875,926,1610,1634]" italics="true" pageId="20" pageNumber="374">et al.</emphasis>
2000
</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831099F0042F0F9CA3CD2F9B6" author="Ruebenstahl AA &amp; Klein MD &amp; Yi H" box="[1000,1246,1610,1635]" pageId="20" pageNumber="374" pagination="2463 - 556" refId="ref32564" refString="Ruebenstahl AA, Klein MD, Yi H et al. Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros. The Anatomical Record 2022; 305: 2463 - 556." type="journal article" year="2022">
Ruebenstahl
<emphasis id="B97F4F2B31099F004568F9CB3CAFF9B7" box="[1136,1187,1610,1634]" italics="true" pageId="20" pageNumber="374">et al.</emphasis>
2022
</bibRefCitation>
). An interpterygoid vacuity was considered to be present in
<taxonomicName id="4C0BE8BA31099F0045F7F9EA3B42F974" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="20" pageNumber="374" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B31099F0045F7F9EA3B42F974" italics="true" pageId="20" pageNumber="374">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9AEEC831099F004279F9093BD2F974" author="Crush PJ" box="[865,990,1673,1697]" pageId="20" pageNumber="374" refId="ref31157" refString="Crush PJ. An early, terrestrial, crocodile from South Wales. Unpublished Ph. D. Thesis, University College London, 1980." type="book" year="1980">Crush 1980</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831099F0042F4F9093C2CF974" author="Crush PJ" box="[1004,1056,1673,1697]" pageId="20" pageNumber="374" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">1984</bibRefCitation>
), but this was based on disarticulated elements and this interpretation is therefore uncertain, as was also discussed by
<bibRefCitation id="EF9AEEC831099F0042F0F9483CA3F935" author="Clark JM &amp; Sues H-D &amp; Berman DS" box="[1000,1199,1736,1760]" pageId="20" pageNumber="374" pagination="683 - 704" refId="ref31006" refString="Clark JM, Sues H-D, Berman DS. A new specimen of Hesperosuchus agilis from the Upper Triassic of New Mexico and the interrelationships of basal crocodylomorph archosaurs. Journal of Vertebrate Paleontology 2000; 20: 683 - 704." type="journal article" year="2000">
Clark
<emphasis id="B97F4F2B31099F004532F9483C50F935" box="[1066,1116,1736,1760]" italics="true" pageId="20" pageNumber="374">et al.</emphasis>
(2000)
</bibRefCitation>
. The shape of the transverse flange of the pterygoid is hard to ascertain but it appears to be elongated posterolaterally, as in other non-crocodyliform crocodylomorphs (e.g.
<bibRefCitation id="EF9AEEC831099F004503F8A63C9AF8EB" author="Crush PJ" box="[1051,1174,1830,1854]" pageId="20" pageNumber="374" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831099F0045BAF8A63D2AF8EB" author="Walker AD" box="[1186,1318,1830,1854]" pageId="20" pageNumber="374" pagination="1 - 120" refId="ref33031" refString="Walker AD. A revision of Sphenosuchus acutus Haughton, a crocodylomorph reptile from the Elliot Formation (Late Triassic or Early Jurassic) of South Africa. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1990; 330: 1 - 120." type="journal article" year="1990">Walker 1990</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831099F00442BF8A63B9DF888" author="Ruebenstahl AA &amp; Klein MD &amp; Yi H" pageId="20" pageNumber="374" pagination="2463 - 556" refId="ref32564" refString="Ruebenstahl AA, Klein MD, Yi H et al. Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros. The Anatomical Record 2022; 305: 2463 - 556." type="journal article" year="2022">
Ruebenstahl
<emphasis id="B97F4F2B31099F004232F8C63B55F888" box="[810,857,1861,1885]" italics="true" pageId="20" pageNumber="374">et al.</emphasis>
2022
</bibRefCitation>
). Posterior to the palatal ramus, the ventral rim of the pterygoid curves posterolaterally where it forms a process that is slightly posteroventrally directed, the medial pterygoid flange (
<emphasis id="B97F4F2B31099F00422DF8243B64F86E" box="[821,872,1956,1979]" italics="true" pageId="20" pageNumber="374">sensu</emphasis>
<bibRefCitation id="EF9AEEC831099F004277F8233BFDF86E" author="Walker AD" box="[879,1009,1955,1979]" pageId="20" pageNumber="374" pagination="1 - 120" refId="ref33031" refString="Walker AD. A revision of Sphenosuchus acutus Haughton, a crocodylomorph reptile from the Elliot Formation (Late Triassic or Early Jurassic) of South Africa. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1990; 330: 1 - 120." type="journal article" year="1990">Walker 1990</bibRefCitation>
). Its posterior end is covered by the ceratobranchial I ventrally on both sides. A thin, transversely concave lamina is present on the ventral surface medial to this flange in the right pterygoid, and its posteromedial corner clearly articulates with the basipterygoid process of the parabasisphenoid. A short portion of the posterolaterally directed quadrate flange of the right pterygoid can be observed in ventral view, but its morphology is unclear.
</paragraph>
<paragraph id="8BB4933931089F014199FEE73A8AFC25" blockId="21.[128,779,358,1886]" pageId="21" pageNumber="375">
<emphasis id="B97F4F2B31089F014199FEE738EAFEAB" box="[129,230,359,382]" italics="true" pageId="21" pageNumber="375">Braincase:</emphasis>
Incomplete braincase remains are certainly preserved in SMNS 12591a. Because the skull is abruptly broken posteriorly and the elements near this break are severely distorted, most bones cannot be identified. The only braincase element to exhibit sufficient preservation is the parabasisphenoid, which is partially exposed in ventral view (
<figureCitation id="13308FBC31089F0140F8FD833A40FDCF" box="[480,588,515,539]" captionStart="Figure 9" captionStartId="10.[113,178,1772,1796]" captionTargetBox="[222,1352,144,1744]" captionTargetId="figure-6@10.[220,1352,144,1744]" captionTargetPageId="10" captionText="Figure 9. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in dorsal view; B, interpretative drawing of (A); C, ventral view; D, interpretative drawing of (C). Abbreviations:an, angular; antfo, antorbital fossa; bpp, basipterygoid process; crb, ceratobranchial I; de, dentary; ect, ectopterygoid; fr, frontal; j, jugal; la, lacrimal; mx, maxilla; na, nasal; pa, parietal; pbs, parabasisphenoid; pl, palatine; plf, palatine fossa; po, postorbital; pra, pre-articular; prf, prefrontal; pt, pterygoid; ptmr, pterygoid medial rim; sa, surangular; spl, splenial; sq, squamosal; vo, vomer." figureDoi="http://doi.org/10.5281/zenodo.8426169" httpUri="https://zenodo.org/record/8426169/files/figure.png" pageId="21" pageNumber="375">Fig. 9C, D</figureCitation>
). It was identified by
<bibRefCitation id="EF9AEEC831089F014187FDA2393FFDEF" author="Huene FR" box="[159,307,546,570]" pageId="21" pageNumber="375" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene (1921)</bibRefCitation>
and also briefly described by
<bibRefCitation id="EF9AEEC831089F014347FDA238C7FD8C" author="Sereno PC &amp; Wild R" pageId="21" pageNumber="375" pagination="435 - 58" refId="ref32722" refString="Sereno PC, Wild R. Procompsognathus: theropod, ' thecodont' or both? Journal of Vertebrate Paleontology 1992; 12: 435 - 58." type="journal article" year="1992">Sereno and Wild (1992)</bibRefCitation>
. The exposed portion of the parabasisphenoid represents the ventral surface, including remains of both basipterygoid processes, which are incomplete. The ventral surface is inclined slightly posterodorsally to anteroventrally relative to the main axis of the skull. Its surface is transversely concave along its length, forming a shallow longitudinal trough that represents the anterior portion of the median pharyngeal or sub-basisphenoid recess. The posterior part of this recess, which is not preserved, is deeply excavated either directly anterior to, or confluent with, the basioccipital in other non-crocodyliform crocodyliforms and non-crocodylomorph loricatans (
<bibRefCitation id="EF9AEEC831089F01433AFCFA3AA2FC46" author="Walker AD" box="[546,686,890,915]" pageId="21" pageNumber="375" pagination="1 - 120" refId="ref33031" refString="Walker AD. A revision of Sphenosuchus acutus Haughton, a crocodylomorph reptile from the Elliot Formation (Late Triassic or Early Jurassic) of South Africa. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1990; 330: 1 - 120." type="journal article" year="1990">Walker 1990</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831089F0143DBFCFB38B9FC67" author="Gower DJ" pageId="21" pageNumber="375" pagination="49 - 76" refId="ref31485" refString="Gower DJ. Braincase evolution in suchian archosaurs (Reptilia: Diapsida): evidence from the rauisuchian Batrachotomus kupferzellensis. Zoological Journal of the Linnean Society 2002; 136: 49 - 76." type="journal article" year="2002">Gower 2002</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831089F0141A7FC1A3961FC67" author="Leardi JM &amp; Pol D &amp; Clark JM" box="[191,365,922,946]" pageId="21" pageNumber="375" pagination="48 - 73" refId="ref31967" refString="Leardi JM, Pol D, Clark JM. Braincase anatomy of Almadasuchus figarii (Archosauria, Crocodylomorpha) and a review of the cranial pneumaticity in the origins of Crocodylomorpha. Journal of Anatomy 2020; 237: 48 - 73." type="journal article" year="2020">
Leardi
<emphasis id="B97F4F2B31089F01401EFC1B3939FC67" box="[262,309,922,946]" italics="true" pageId="21" pageNumber="375">et al.</emphasis>
2020
</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831089F01406FFC1A3A6FFC67" author="Ruebenstahl AA &amp; Klein MD &amp; Yi H" box="[375,611,922,946]" pageId="21" pageNumber="375" pagination="2463 - 556" refId="ref32564" refString="Ruebenstahl AA, Klein MD, Yi H et al. Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros. The Anatomical Record 2022; 305: 2463 - 556." type="journal article" year="2022">
Ruebenstahl
<emphasis id="B97F4F2B31089F0140E5FC1B3A27FC67" box="[509,555,922,946]" italics="true" pageId="21" pageNumber="375">et al.</emphasis>
2022
</bibRefCitation>
). The preserved ventral body is roughly hourglass-shaped in ventral view, being somewhat constricted in its anteroposterior centre.
</paragraph>
<paragraph id="8BB4933931089F014184FC783B06F9F1" blockId="21.[128,779,358,1886]" pageId="21" pageNumber="375">
The right basipterygoid process is clearly exposed adjacent to its articular surface on the right pterygoid. As preserved, it is composed of two knob-like processes and a large and well excavated concavity lateral to these processes (
<figureCitation id="13308FBC31089F014354FBD63ABBFBB8" box="[588,695,1110,1134]" captionStart="Figure 9" captionStartId="10.[113,178,1772,1796]" captionTargetBox="[222,1352,144,1744]" captionTargetId="figure-6@10.[220,1352,144,1744]" captionTargetPageId="10" captionText="Figure 9. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in dorsal view; B, interpretative drawing of (A); C, ventral view; D, interpretative drawing of (C). Abbreviations:an, angular; antfo, antorbital fossa; bpp, basipterygoid process; crb, ceratobranchial I; de, dentary; ect, ectopterygoid; fr, frontal; j, jugal; la, lacrimal; mx, maxilla; na, nasal; pa, parietal; pbs, parabasisphenoid; pl, palatine; plf, palatine fossa; po, postorbital; pra, pre-articular; prf, prefrontal; pt, pterygoid; ptmr, pterygoid medial rim; sa, surangular; spl, splenial; sq, squamosal; vo, vomer." figureDoi="http://doi.org/10.5281/zenodo.8426169" httpUri="https://zenodo.org/record/8426169/files/figure.png" pageId="21" pageNumber="375">Fig. 9C, D</figureCitation>
). In extant crocodylians, as well as crocodyliforms and the hallopodid
<taxonomicName id="4C0BE8BA31089F014198FB143941FB79" box="[128,333,1172,1196]" genus="Almadasuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="21" pageNumber="375" phylum="Chordata" rank="species" species="figarii">
<emphasis id="B97F4F2B31089F014198FB143941FB79" box="[128,333,1172,1196]" italics="true" pageId="21" pageNumber="375">Almadasuchus figarii</emphasis>
</taxonomicName>
, the parabasisphenoid is tightly sutured to the pterygoids and the basipterygoid processes are lost (
<bibRefCitation id="EF9AEEC831089F0143D0FB3438E5FB3E" author="Leardi JM &amp; Pol D &amp; Clark JM" pageId="21" pageNumber="375" pagination="48 - 73" refId="ref31967" refString="Leardi JM, Pol D, Clark JM. Braincase anatomy of Almadasuchus figarii (Archosauria, Crocodylomorpha) and a review of the cranial pneumaticity in the origins of Crocodylomorpha. Journal of Anatomy 2020; 237: 48 - 73." type="journal article" year="2020">
Leardi
<emphasis id="B97F4F2B31089F014198FB5438BCFB3E" box="[128,176,1235,1259]" italics="true" pageId="21" pageNumber="375">et al.</emphasis>
2020
</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831089F0141EEFB5339BAFB3E" author="Kuzmin IT &amp; Boitsova EA &amp; Gombolevskiy VA" box="[246,438,1235,1259]" pageId="21" pageNumber="375" pagination="983 - 1038" refId="ref31880" refString="Kuzmin IT, Boitsova EA, Gombolevskiy VA et al. Braincase anatomy of extant Crocodylia, with new insights into the development and evolution of the neurocranium in crocodylomorphs. Journal of Anatomy 2021; 239: 983 - 1038." type="journal article" year="2021">
Kuzmin
<emphasis id="B97F4F2B31089F014056FB543972FB3E" box="[334,382,1235,1259]" italics="true" pageId="21" pageNumber="375">et al.</emphasis>
2021
</bibRefCitation>
). However, in several other non-crocodyliform crocodylomorphs the basipterygoid processes form very prominent but hollow protuberances that cover much of the anterior portion of the ventral body in ventral view (
<bibRefCitation id="EF9AEEC831089F0143DAFAB138BFFABD" author="Walker AD" pageId="21" pageNumber="375" pagination="1 - 120" refId="ref33031" refString="Walker AD. A revision of Sphenosuchus acutus Haughton, a crocodylomorph reptile from the Elliot Formation (Late Triassic or Early Jurassic) of South Africa. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1990; 330: 1 - 120." type="journal article" year="1990">Walker 1990</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831089F0141D9FAD0397FFABD" author="Leardi JM &amp; Pol D &amp; Clark JM" box="[193,371,1360,1384]" pageId="21" pageNumber="375" pagination="2801" refId="ref31922" refString="Leardi JM, Pol D, Clark JM. Detailed anatomy of the braincase of Macelognathus vagans Marsh, 1884 (Archosauria, Crocodylomorpha) using high resolution tomography and new insights on basal crocodylomorph phylogeny. PeerJ 2017; 5: e 2801." type="journal article" year="2017">
Leardi
<emphasis id="B97F4F2B31089F014011FAD13936FABD" box="[265,314,1360,1384]" italics="true" pageId="21" pageNumber="375">et al.</emphasis>
2017
</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831089F014098FAD03A7CFABD" author="Ruebenstahl AA &amp; Klein MD &amp; Yi H" box="[384,624,1360,1384]" pageId="21" pageNumber="375" pagination="2463 - 556" refId="ref32564" refString="Ruebenstahl AA, Klein MD, Yi H et al. Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros. The Anatomical Record 2022; 305: 2463 - 556." type="journal article" year="2022">
Ruebenstahl
<emphasis id="B97F4F2B31089F01431EFAD13A3BFABD" box="[518,567,1360,1384]" italics="true" pageId="21" pageNumber="375">et al.</emphasis>
2022
</bibRefCitation>
). It is possible that much of the ventral surface of the basipterygoid processes of SMNS 12591a are incomplete, and that the large concavity lateral to the two knob-like processes represents the opened internal cavity of the right basipterygoid process. The presence of a cultriform process could not be established, since this region of the parabasisphenoid is covered in ventral view by the pterygoids.
</paragraph>
<paragraph id="8BB4933931089F014184F9AC3A8CF88B" blockId="21.[128,779,358,1886]" pageId="21" pageNumber="375">
Additional remains of the braincase are visible through the orbits, particularly the right orbit (
<figureCitation id="13308FBC31089F0140C3F9CB3A30F9B7" box="[475,572,1611,1635]" captionStart="Figure 8" captionStartId="9.[129,194,1772,1796]" captionTargetBox="[178,1429,144,1744]" captionTargetId="figure-6@9.[174,1429,144,1744]" captionTargetPageId="9" captionText="Figure 8. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in right lateral view; B, interpretative drawing of A; C, less lateral view; D, interpretative drawing of (C). Abbreviations:an, angular; cc, crista cranii; de, dentary; fr, frontal; j, jugal; la, lacrimal; ls, laterosphenoid; mx, maxilla; na, nasal; pa, parietal; pl, palatine; po, postorbital; prf, prefrontal; pt, pterygoid; q, quadrate; qj, quadratojugal; sa, surangular; sq, squamosal." figureDoi="http://doi.org/10.5281/zenodo.8426167" httpUri="https://zenodo.org/record/8426167/files/figure.png" pageId="21" pageNumber="375">Fig. 8A, B</figureCitation>
). They form a transversely thin, plate-like structure that connects to the skull roof dorsally and to the prefrontal anteriorly. The structure is notched along its ventral margin. The remains are poorly preserved and incomplete. Posteriorly, they probably represent part of the laterosphenoid, whereas anteriorly they might be the cristae cranii, which extend far ventrally as a thin lamina from both margins of the olfactory tract of the frontal in
<taxonomicName id="4C0BE8BA31089F014305F8A63B06F8EB" authorityName="Simmons" authorityYear="1965" box="[541,778,1830,1854]" class="Reptilia" family="Sphenosuchidae" genus="Dibothrosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="21" pageNumber="375" phylum="Chordata" rank="species" species="elaphros">
<emphasis id="B97F4F2B31089F014305F8A63B06F8EB" box="[541,778,1830,1854]" italics="true" pageId="21" pageNumber="375">Dibothrosuchus elaphros</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0BE8BA31089F0141B4F8C53A70F88B" authority="(Ruebenstahl et al. 2022)" baseAuthorityName="Ruebenstahl" baseAuthorityYear="2022" box="[172,636,1861,1886]" class="Reptilia" genus="Junggarsuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="21" pageNumber="375" phylum="Chordata" rank="species" species="sloani">
<emphasis id="B97F4F2B31089F0141B4F8C53978F888" box="[172,372,1861,1885]" italics="true" pageId="21" pageNumber="375">Junggarsuchus sloani</emphasis>
(
<bibRefCitation id="EF9AEEC831089F01409CF8C53A7CF88B" author="Ruebenstahl AA &amp; Klein MD &amp; Yi H" box="[388,624,1861,1886]" pageId="21" pageNumber="375" pagination="2463 - 556" refId="ref32564" refString="Ruebenstahl AA, Klein MD, Yi H et al. Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros. The Anatomical Record 2022; 305: 2463 - 556." type="journal article" year="2022">
Ruebenstahl
<emphasis id="B97F4F2B31089F014311F8C63A34F888" box="[521,568,1861,1885]" italics="true" pageId="21" pageNumber="375">et al.</emphasis>
2022
</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
<paragraph id="8BB4933931089F01408FF8F339FEF858" blockId="21.[407,498,1907,1933]" box="[407,498,1907,1933]" pageId="21" pageNumber="375">
<emphasis id="B97F4F2B31089F01408FF8F339FEF858" box="[407,498,1907,1933]" italics="true" pageId="21" pageNumber="375">Mandible</emphasis>
</paragraph>
<paragraph id="8BB4933931089F014199F81B3B05F804" blockId="21.[129,777,1946,2001]" pageId="21" pageNumber="375">
<emphasis id="B97F4F2B31089F014199F81B38D4F867" box="[129,216,1947,1970]" italics="true" pageId="21" pageNumber="375">Dentary:</emphasis>
Remains of the dentary are preserved in SMNS 12596b, SMNS 12352a, and SMNS 12591a. In SMNS 12596b,
</paragraph>
<paragraph id="8BB4933931089F014221FF103BB6FE34" blockId="21.[825,1475,144,1139]" pageId="21" pageNumber="375">
the anterior end of the less dentary is preserved together with a small sliver of the right dentary and they are exposed in ventrolateral view (
<figureCitation id="13308FBC31089F0142A7FF4F3C0AFF32" box="[959,1030,207,231]" captionStart="Figure 3" captionStartId="4.[113,178,1499,1523]" captionTargetBox="[147,1426,144,1471]" captionTargetId="figure-217@4.[146,1426,144,1471]" captionTargetPageId="4" captionText="Figure 3. Part of Saltoposuchus connectens referred specimen SMNS 12596. A, SMNS 12596a, cranial block; B, SMNS 12596b, two dentaries exposed in ventrolateral view; C, SMNS 12596c exposing a less femur in anterolateral view; D, exposing the opposite side of SMNS 12596c, Abbreviations:de, dentary; dr, dorsal rib; dv, dorsal vertebra; fe, femur; ga, gastralia; l., less; mx, maxilla; na, nasal; os, osteoderm; q, quadrate; r., right; sq, squamosal; sr, sacral rib; sv, sacral vertebra; tp, transverse process." figureDoi="http://doi.org/10.5281/zenodo.8426153" httpUri="https://zenodo.org/record/8426153/files/figure.png" pageId="21" pageNumber="375">Fig. 3B</figureCitation>
). In SMNS 12352a, most of the right dentary is preserved, except for the anteriormost and posteriormost parts. A fragment of the less dentary is also preserved. They are exposed in lateral view and their dorsal margins, as well as their teeth, are mostly obscured by the maxillae (
<figureCitation id="13308FBC31089F014418FECC3D7BFEB1" box="[1280,1399,332,356]" captionStart="Figure 10" captionStartId="14.[113,178,1772,1796]" captionTargetBox="[197,1376,144,1744]" captionTargetId="figure-5@14.[197,1376,144,1744]" captionTargetPageId="14" captionText="Figure 10. Saltoposuchus connectens referred specimen SMNS 12352.A, SMNS 12352a with the snout region exposed in angled right lateral view; B, interpretative drawing of (A); C, opposite side of SMNS 12352a exposing the posterior portion of the less maxilla and lacrimal in angled lateral view; D, interpretative drawing of (C); E, SMNS 12352b, carpus and manus in dorsal view; F, interpretative drawing of (E); G, SMNS 12352b in medial view; H, interpretative drawing of (G). Abbreviations; antfe, antorbital fenestra; antfo, antorbital fossa; dc, distal carpal; de, dentary; exp, extensor pit; fr, frontal; j, jugal; la, lacrimal; lad, lacrimal duct; mc, metacarpal; mx, maxilla; na, nasal; ph, phalanx; pm, premaxilla; prf, prefrontal; ra, radiale; ul, ulnare." figureDoi="http://doi.org/10.5281/zenodo.8426171" httpUri="https://zenodo.org/record/8426171/files/figure.png" pageId="21" pageNumber="375">Fig. 10A–D</figureCitation>
). Both dentaries of SMNS 12591a are virtually completely preserved, only missing the anterior ends (
<figureCitation id="13308FBC31089F01459BFE0B3CCCFE76" box="[1155,1216,395,419]" captionStart="Figure 8" captionStartId="9.[129,194,1772,1796]" captionTargetBox="[178,1429,144,1744]" captionTargetId="figure-6@9.[174,1429,144,1744]" captionTargetPageId="9" captionText="Figure 8. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in right lateral view; B, interpretative drawing of A; C, less lateral view; D, interpretative drawing of (C). Abbreviations:an, angular; cc, crista cranii; de, dentary; fr, frontal; j, jugal; la, lacrimal; ls, laterosphenoid; mx, maxilla; na, nasal; pa, parietal; pl, palatine; po, postorbital; prf, prefrontal; pt, pterygoid; q, quadrate; qj, quadratojugal; sa, surangular; sq, squamosal." figureDoi="http://doi.org/10.5281/zenodo.8426167" httpUri="https://zenodo.org/record/8426167/files/figure.png" pageId="21" pageNumber="375">Figs 8</figureCitation>
,
<figureCitation id="13308FBC31089F0145D4FE0B3D06FE77" box="[1228,1290,395,419]" captionStart="Figure 9" captionStartId="10.[113,178,1772,1796]" captionTargetBox="[222,1352,144,1744]" captionTargetId="figure-6@10.[220,1352,144,1744]" captionTargetPageId="10" captionText="Figure 9. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in dorsal view; B, interpretative drawing of (A); C, ventral view; D, interpretative drawing of (C). Abbreviations:an, angular; antfo, antorbital fossa; bpp, basipterygoid process; crb, ceratobranchial I; de, dentary; ect, ectopterygoid; fr, frontal; j, jugal; la, lacrimal; mx, maxilla; na, nasal; pa, parietal; pbs, parabasisphenoid; pl, palatine; plf, palatine fossa; po, postorbital; pra, pre-articular; prf, prefrontal; pt, pterygoid; ptmr, pterygoid medial rim; sa, surangular; spl, splenial; sq, squamosal; vo, vomer." figureDoi="http://doi.org/10.5281/zenodo.8426169" httpUri="https://zenodo.org/record/8426169/files/figure.png" pageId="21" pageNumber="375">9C, D</figureCitation>
). However, as for the rest of this skull, the dentaries are strongly dorsoventrally compressed.
</paragraph>
<paragraph id="8BB4933931089F01424DFE693BFCFBA6" blockId="21.[825,1475,144,1139]" pageId="21" pageNumber="375">
The dentary is straight for most of its length, but it widens dorsoventrally and curves slightly dorsally at its anterior end in both SMNS 12596b and SMNS 12352a. This is similar to the condition of
<taxonomicName id="4C0BE8BA31089F0142D1FDC63C98FD8B" box="[969,1172,582,606]" class="Reptilia" family="Sphenosuchidae" genus="Hesperosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="21" pageNumber="375" phylum="Chordata" rank="species" species="agilis">
<emphasis id="B97F4F2B31089F0142D1FDC63C98FD8B" box="[969,1172,582,606]" italics="true" pageId="21" pageNumber="375">Hesperosuchus agilis</emphasis>
</taxonomicName>
(AMNH FARB 6758) and
<taxonomicName id="4C0BE8BA31089F014221FDE63BD4FDAB" authorityName="Reig" authorityYear="1963" box="[825,984,614,638]" class="Polyplacophora" genus="Trialestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodilomorphi" pageId="21" pageNumber="375" phylum="Mollusca" rank="species" species="romeri">
<emphasis id="B97F4F2B31089F014221FDE63BD4FDAB" box="[825,984,614,638]" italics="true" pageId="21" pageNumber="375">Trialestes romeri</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9AEEC831089F0142FEFDE63C61FDAB" author="Colbert EH" box="[998,1133,614,638]" pageId="21" pageNumber="375" pagination="561 - 92" refId="ref31083" refString="Colbert EH. A pseudosuchian reptile from Arizona. Bulletin of the American Museum of Natural History 1952; 99: 561 - 92." type="journal article" year="1952">Colbert 1952</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831089F01456EFDE63D3AFDAB" author="Lecuona A &amp; Ezcurra MD &amp; Irmis RB" box="[1142,1334,614,638]" pageId="21" pageNumber="375" pagination="585 - 622" refId="ref32010" refString="Lecuona A, Ezcurra MD, Irmis RB. Revision of the early crocodylomorph Trialestes romeri (Archosauria, Suchia) from the lower Upper Triassic Ischigualasto Formation of Argentina: one of the oldest-known crocodylomorphs. Papers in Palaeontology 2016; 2: 585 - 622." type="journal article" year="2016">
Lecuona
<emphasis id="B97F4F2B31089F0145CBFDE73D0CFDAB" box="[1235,1280,614,638]" italics="true" pageId="21" pageNumber="375">et al.</emphasis>
2016
</bibRefCitation>
), whereas the anterior ends of the dentary are not expanded in
<taxonomicName id="4C0BE8BA31089F014429FD053B71FD69" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="21" pageNumber="375" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B31089F014429FD053B71FD69" italics="true" pageId="21" pageNumber="375">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0BE8BA31089F01428BFD243CA3FD69" box="[915,1199,676,700]" class="Reptilia" family="Sphenosuchidae" genus="Litargosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="21" pageNumber="375" phylum="Chordata" rank="species" species="leptorhynchus">
<emphasis id="B97F4F2B31089F01428BFD243CA3FD69" box="[915,1199,676,700]" italics="true" pageId="21" pageNumber="375">Litargosuchus leptorhynchus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0BE8BA31089F0145DDFD243DB1FD69" box="[1221,1469,676,700]" class="Reptilia" family="Sphenosuchidae" genus="Dromicosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="21" pageNumber="375" phylum="Chordata" rank="species" species="grallator">
<emphasis id="B97F4F2B31089F0145DDFD243DB1FD69" box="[1221,1469,676,700]" italics="true" pageId="21" pageNumber="375">Dromicosuchus grallator</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0BE8BA31089F014221FD443C08FD09" baseAuthorityName="Walker" baseAuthorityYear="1990" box="[825,1028,708,732]" class="Reptilia" genus="Sphenosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="21" pageNumber="375" phylum="Chordata" rank="species" species="acutus">
<emphasis id="B97F4F2B31089F014221FD443C08FD09" box="[825,1028,708,732]" italics="true" pageId="21" pageNumber="375">Sphenosuchus acutus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0BE8BA31089F014508FD443CF3FD09" authorityName="Simmons" authorityYear="1965" box="[1040,1279,708,732]" class="Reptilia" family="Sphenosuchidae" genus="Dibothrosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="21" pageNumber="375" phylum="Chordata" rank="species" species="elaphros">
<emphasis id="B97F4F2B31089F014508FD443CF3FD09" box="[1040,1279,708,732]" italics="true" pageId="21" pageNumber="375">Dibothrosuchus elaphros</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C0BE8BA31089F014420FD443B7EFD2E" class="Reptilia" genus="Junggarsuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="21" pageNumber="375" phylum="Chordata" rank="species" species="sloani">
<emphasis id="B97F4F2B31089F014420FD443B7EFD2E" italics="true" pageId="21" pageNumber="375">Junggarsuchus sloani</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9AEEC831089F014290FD633C0BFD2E" author="Crush PJ" box="[904,1031,739,763]" pageId="21" pageNumber="375" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831089F014500FD633C93FD2E" author="Walker AD" box="[1048,1183,739,763]" pageId="21" pageNumber="375" pagination="1 - 120" refId="ref33031" refString="Walker AD. A revision of Sphenosuchus acutus Haughton, a crocodylomorph reptile from the Elliot Formation (Late Triassic or Early Jurassic) of South Africa. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1990; 330: 1 - 120." type="journal article" year="1990">Walker 1990</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831089F0145A8FD643DB0FD2E" author="Wu X-C &amp; Chauerjee S" box="[1200,1468,739,763]" pageId="21" pageNumber="375" pagination="58 - 89" refId="ref33269" refString="Wu X-C, Chauerjee S. Dibothrosuchus elaphros, a crocodylomorph from the Lower Jurassic of China and the phylogeny of the Sphenosuchia. Journal of Vertebrate Paleontology 1993; 13: 58 - 89." type="journal article" year="1993">Wu and Chaưerjee 1993</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831089F014221FC823C01FCCE" author="Clark JM &amp; Sues H-D" box="[825,1037,770,795]" pageId="21" pageNumber="375" pagination="77 - 95" refId="ref30970" refString="Clark JM, Sues H-D. Two new basal crocodylomorph archosaurs from the Lower Jurassic and the monophyly of the Sphenosuchia. Zoological Journal of the Linnean Society 2002; 136: 77 - 95." type="journal article" year="2002">Clark and Sues 2002</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831089F014501FC833CBBFCCE" author="Sues H-D &amp; Olsen PE &amp; Carter JG" box="[1049,1207,770,795]" pageId="21" pageNumber="375" pagination="329 - 43" refId="ref32828" refString="Sues H-D, Olsen PE, Carter JG et al. A new crocodylomorph archosaur from the Upper Triassic of North Carolina. Journal of Vertebrate Paleontology 2003; 23: 329 - 43." type="journal article" year="2003">
Sues
<emphasis id="B97F4F2B31089F014556FC833C72FCCF" box="[1102,1150,770,794]" italics="true" pageId="21" pageNumber="375">et al.</emphasis>
2003
</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831089F0145DAFC823DBDFCCE" author="Ruebenstahl AA &amp; Klein MD &amp; Yi H" box="[1218,1457,770,795]" pageId="21" pageNumber="375" pagination="2463 - 556" refId="ref32564" refString="Ruebenstahl AA, Klein MD, Yi H et al. Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros. The Anatomical Record 2022; 305: 2463 - 556." type="journal article" year="2022">
Ruebenstahl
<emphasis id="B97F4F2B31089F014451FC833D74FCCF" box="[1353,1400,770,794]" italics="true" pageId="21" pageNumber="375">et al.</emphasis>
2022
</bibRefCitation>
). A single, large tooth is preserved on the anterior portion of the less dentary of SMNS 12596b. Two further, more posteriorly positioned, teeth are preserved in the fragmentary right dentary. The dentaries of SMNS 12591a meet at the anterior end of the specimen in ventral view. Since their anteriormost parts are not preserved, the symphysis formed by the dentaries was considerable. The dentary tooth row extended posteriorly until the posterior end of the maxilla but the tooth count cannot be discerned. The posterior articulations of the dentary are unknown because this region is heavily cracked in both mandibular rami in SMNS 12591a.
</paragraph>
<paragraph id="8BB4933931089F014221FB143C81FADE" blockId="21.[825,1474,1172,1291]" pageId="21" pageNumber="375">
<emphasis id="B97F4F2B31089F014221FB143B82FB79" box="[825,910,1172,1196]" italics="true" pageId="21" pageNumber="375">Splenial:</emphasis>
Both splenials are present in SMNS 12591a. The splenial is a thin, plate-like bone that anteriorly contributes to the mandibular symphysis, and posteriorly reaches until the external mandibular fenestra (
<figureCitation id="13308FBC31089F01450DFB723C71FADF" box="[1045,1149,1266,1290]" captionStart="Figure 9" captionStartId="10.[113,178,1772,1796]" captionTargetBox="[222,1352,144,1744]" captionTargetId="figure-6@10.[220,1352,144,1744]" captionTargetPageId="10" captionText="Figure 9. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in dorsal view; B, interpretative drawing of (A); C, ventral view; D, interpretative drawing of (C). Abbreviations:an, angular; antfo, antorbital fossa; bpp, basipterygoid process; crb, ceratobranchial I; de, dentary; ect, ectopterygoid; fr, frontal; j, jugal; la, lacrimal; mx, maxilla; na, nasal; pa, parietal; pbs, parabasisphenoid; pl, palatine; plf, palatine fossa; po, postorbital; pra, pre-articular; prf, prefrontal; pt, pterygoid; ptmr, pterygoid medial rim; sa, surangular; spl, splenial; sq, squamosal; vo, vomer." figureDoi="http://doi.org/10.5281/zenodo.8426169" httpUri="https://zenodo.org/record/8426169/files/figure.png" pageId="21" pageNumber="375">Fig. 9C, D</figureCitation>
).
</paragraph>
<paragraph id="8BB4933931089F014221FAAC3D1CF988" blockId="21.[825,1474,1324,1630]" pageId="21" pageNumber="375">
<emphasis id="B97F4F2B31089F014221FAAC3BA2FA91" box="[825,942,1324,1348]" italics="true" pageId="21" pageNumber="375">Surangular:</emphasis>
Remains of the surangular are exclusively preserved on both mandibular rami of SMNS 12591a (
<figureCitation id="13308FBC31089F0145E6FACB3D35FAB6" box="[1278,1337,1355,1379]" captionStart="Figure 8" captionStartId="9.[129,194,1772,1796]" captionTargetBox="[178,1429,144,1744]" captionTargetId="figure-6@9.[174,1429,144,1744]" captionTargetPageId="9" captionText="Figure 8. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in right lateral view; B, interpretative drawing of A; C, less lateral view; D, interpretative drawing of (C). Abbreviations:an, angular; cc, crista cranii; de, dentary; fr, frontal; j, jugal; la, lacrimal; ls, laterosphenoid; mx, maxilla; na, nasal; pa, parietal; pl, palatine; po, postorbital; prf, prefrontal; pt, pterygoid; q, quadrate; qj, quadratojugal; sa, surangular; sq, squamosal." figureDoi="http://doi.org/10.5281/zenodo.8426167" httpUri="https://zenodo.org/record/8426167/files/figure.png" pageId="21" pageNumber="375">Figs 8</figureCitation>
,
<figureCitation id="13308FBC31089F01445CFACB3D8DFAB6" box="[1348,1409,1355,1379]" captionStart="Figure 9" captionStartId="10.[113,178,1772,1796]" captionTargetBox="[222,1352,144,1744]" captionTargetId="figure-6@10.[220,1352,144,1744]" captionTargetPageId="10" captionText="Figure 9. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in dorsal view; B, interpretative drawing of (A); C, ventral view; D, interpretative drawing of (C). Abbreviations:an, angular; antfo, antorbital fossa; bpp, basipterygoid process; crb, ceratobranchial I; de, dentary; ect, ectopterygoid; fr, frontal; j, jugal; la, lacrimal; mx, maxilla; na, nasal; pa, parietal; pbs, parabasisphenoid; pl, palatine; plf, palatine fossa; po, postorbital; pra, pre-articular; prf, prefrontal; pt, pterygoid; ptmr, pterygoid medial rim; sa, surangular; spl, splenial; sq, squamosal; vo, vomer." figureDoi="http://doi.org/10.5281/zenodo.8426169" httpUri="https://zenodo.org/record/8426169/files/figure.png" pageId="21" pageNumber="375">9C, D</figureCitation>
). Due to extensive crushing its exact outline is unclear but it forms the dorsal margin of the external mandibular fenestra, which is framed ventrally by the angular.The external mandibular fenestra of this specimen was not identified by
<bibRefCitation id="EF9AEEC831089F0145ADFA483D5FFA35" author="Ostrom JH" box="[1205,1363,1480,1504]" pageId="21" pageNumber="375" pagination="179 - 95" refId="ref32208" refString="Ostrom JH. Procompsognathus - Theropod or Thecodont? Palaeontographica A 1981; 175: 179 - 95." type="journal article" year="1981">Ostrom (1981)</bibRefCitation>
, whereas it was recognized by
<bibRefCitation id="EF9AEEC831089F0142E1FA683C82F9D5" author="Huene FR" box="[1017,1166,1512,1536]" pageId="21" pageNumber="375" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene (1921)</bibRefCitation>
and
<bibRefCitation id="EF9AEEC831089F0145D9FA683DB0F9D5" author="Sereno PC &amp; Wild R" box="[1217,1468,1512,1536]" pageId="21" pageNumber="375" pagination="435 - 58" refId="ref32722" refString="Sereno PC, Wild R. Procompsognathus: theropod, ' thecodont' or both? Journal of Vertebrate Paleontology 1992; 12: 435 - 58." type="journal article" year="1992">Sereno and Wild (1992)</bibRefCitation>
. It is badly broken but clearly oval, being anteroposteriorly longer than dorsoventrally tall, as in other crocodylomorphs. The dorsal margin of the surangular is very gently convex.
</paragraph>
<paragraph id="8BB4933931089F024221F9FF38B2FEB1" blockId="21.[825,1474,1663,2000]" lastBlockId="22.[113,763,144,1045]" lastPageId="22" lastPageNumber="376" pageId="21" pageNumber="375">
<emphasis id="B97F4F2B31089F014221F9FF3B9CF942" box="[825,912,1663,1687]" italics="true" pageId="21" pageNumber="375">Angular:</emphasis>
Several remains of angulars are preserved in SMNS 12597. Remains of two angulars were identified on SMNS 12597a by
<bibRefCitation id="EF9AEEC831089F0142B2F93E3C4CF903" author="Huene FR" box="[938,1088,1726,1750]" pageId="21" pageNumber="375" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene (1921)</bibRefCitation>
, which he considered to probably belong to
<taxonomicName id="4C0BE8BA31089F01429CF95D3C9AF920" authorityName="Fraas" authorityYear="1913" box="[900,1174,1757,1781]" class="Reptilia" family="Coelophysidae" genus="Procompsognathus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="21" pageNumber="375" phylum="Chordata" rank="species" species="triassicus">
<emphasis id="B97F4F2B31089F01429CF95D3C9AF920" box="[900,1174,1757,1781]" italics="true" pageId="21" pageNumber="375">Procompsognathus triassicus</emphasis>
</taxonomicName>
.
<bibRefCitation id="EF9AEEC831089F0145B8F95D3D22F920" author="Crush PJ" box="[1184,1326,1757,1781]" pageId="21" pageNumber="375" refId="ref31157" refString="Crush PJ. An early, terrestrial, crocodile from South Wales. Unpublished Ph. D. Thesis, University College London, 1980." type="book" year="1980">Crush (1980)</bibRefCitation>
also described these elements and referred them to
<taxonomicName id="4C0BE8BA31089F0145DDF97C3DB0F8C1" authorityName="Huene" authorityYear="1921" box="[1221,1468,1788,1812]" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="21" pageNumber="375" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B31089F0145DDF97C3DB0F8C1" box="[1221,1468,1788,1812]" italics="true" pageId="21" pageNumber="375">Saltoposuchus connectens</emphasis>
</taxonomicName>
. Both elements are incomplete and preserved in close association. The less angular is most complete and exposed in ventral and lateral views, whereas the right angular is partially overlapped by the less element and exposed in medial view (
<figureCitation id="13308FBC31089F014471F8FA3DBDF847" box="[1385,1457,1914,1938]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="21" pageNumber="375">Fig. 2A</figureCitation>
). A nicely preserved, isolated angular is also preserved in SMNS 12597e (
<figureCitation id="13308FBC31089F014281F8383BEEF805" box="[921,994,1976,2000]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="21" pageNumber="375">Fig. 2F</figureCitation>
). This element was originally interpreted as a clavicle by
<bibRefCitation id="EF9AEEC8310B9F0241F8FF103978FF7D" author="Huene FR" box="[224,372,144,168]" pageId="22" pageNumber="376" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene (1921)</bibRefCitation>
, but it was recognized as an angular by
<bibRefCitation id="EF9AEEC8310B9F024169FF2F390FFF12" author="Crush PJ" box="[113,259,175,199]" pageId="22" pageNumber="376" refId="ref31157" refString="Crush PJ. An early, terrestrial, crocodile from South Wales. Unpublished Ph. D. Thesis, University College London, 1980." type="book" year="1980">Crush (1980)</bibRefCitation>
. It represents a largely complete less angular exposed in medial view, and belongs to a different individual than the angulars of SMNS 12597a. Its assignment to
<taxonomicName id="4C0BE8BA310B9F02436AFF6E38DBFEF0" authorityName="Huene" authorityYear="1921" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="22" pageNumber="376" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B310B9F02436AFF6E38DBFEF0" italics="true" pageId="22" pageNumber="376">Saltoposuchus connectens</emphasis>
</taxonomicName>
can therefore only be made tentatively. Both angulars are also present but poorly preserved in SMNS 12591a (
<figureCitation id="13308FBC310B9F0243A0FEAD3AF9FE90" box="[696,757,301,325]" captionStart="Figure 8" captionStartId="9.[129,194,1772,1796]" captionTargetBox="[178,1429,144,1744]" captionTargetId="figure-6@9.[174,1429,144,1744]" captionTargetPageId="9" captionText="Figure 8. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in right lateral view; B, interpretative drawing of A; C, less lateral view; D, interpretative drawing of (C). Abbreviations:an, angular; cc, crista cranii; de, dentary; fr, frontal; j, jugal; la, lacrimal; ls, laterosphenoid; mx, maxilla; na, nasal; pa, parietal; pl, palatine; po, postorbital; prf, prefrontal; pt, pterygoid; q, quadrate; qj, quadratojugal; sa, surangular; sq, squamosal." figureDoi="http://doi.org/10.5281/zenodo.8426167" httpUri="https://zenodo.org/record/8426167/files/figure.png" pageId="22" pageNumber="376">Figs 8</figureCitation>
,
<figureCitation id="13308FBC310B9F024169FECC38A2FEB6" box="[113,174,332,356]" captionStart="Figure 9" captionStartId="10.[113,178,1772,1796]" captionTargetBox="[222,1352,144,1744]" captionTargetId="figure-6@10.[220,1352,144,1744]" captionTargetPageId="10" captionText="Figure 9. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in dorsal view; B, interpretative drawing of (A); C, ventral view; D, interpretative drawing of (C). Abbreviations:an, angular; antfo, antorbital fossa; bpp, basipterygoid process; crb, ceratobranchial I; de, dentary; ect, ectopterygoid; fr, frontal; j, jugal; la, lacrimal; mx, maxilla; na, nasal; pa, parietal; pbs, parabasisphenoid; pl, palatine; plf, palatine fossa; po, postorbital; pra, pre-articular; prf, prefrontal; pt, pterygoid; ptmr, pterygoid medial rim; sa, surangular; spl, splenial; sq, squamosal; vo, vomer." figureDoi="http://doi.org/10.5281/zenodo.8426169" httpUri="https://zenodo.org/record/8426169/files/figure.png" pageId="22" pageNumber="376">9C, D</figureCitation>
).
</paragraph>
<paragraph id="8BB49339310B9F024195FEEB38CBFBC0" blockId="22.[113,763,144,1045]" pageId="22" pageNumber="376">
The less angular of SMNS 12597a shows that the ventral margin of the angular was gently convex in lateral view and that the angular would have curved around the ventral margin of the mandibular ramus, and thus was visible both in lateral and medial view. The dorsal margin of the element is also visible in lateral view. It was slightly concave, particularly in its central portion, where the angular formed the posteroventral margin of the external mandibular fenestra. Posterior to this, the angular widens somewhat in dorsoventral height, before tapering into a slender posterior process, which would have articulated dorsally with the articular, as in
<taxonomicName id="4C0BE8BA310B9F024044FD243AC5FD68" authority="(Crush 1984)" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[348,713,676,701]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="22" pageNumber="376" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B310B9F024044FD243A3AFD69" box="[348,566,676,700]" italics="true" pageId="22" pageNumber="376">Terrestrisuchus gracilis</emphasis>
(
<bibRefCitation id="EF9AEEC8310B9F02435DFD243AB1FD68" author="Crush PJ" box="[581,701,676,701]" pageId="22" pageNumber="376" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
)
</taxonomicName>
. The lateral surface of the angular is smooth and slightly transversely convex. The medial surface, as seen the isolated less angular of SMNS 12597e, is markedly transversely concave and characterized by a longitudinal groove approximately in its dorsoventral centre. This groove probably forms the Meckelian canal anteriorly, whereas further posteriorly it formed the articular facet for the pre-articular, as in
<taxonomicName id="4C0BE8BA310B9F02407AFC003A4CFC4D" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[354,576,896,920]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="22" pageNumber="376" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B310B9F02407AFC003A4CFC4D" box="[354,576,896,920]" italics="true" pageId="22" pageNumber="376">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
(NHMUK PV R 7593a,
<bibRefCitation id="EF9AEEC8310B9F0241A3FC1F393FFC62" author="Crush PJ" box="[187,307,927,951]" pageId="22" pageNumber="376" refId="ref31157" refString="Crush PJ. An early, terrestrial, crocodile from South Wales. Unpublished Ph. D. Thesis, University College London, 1980." type="book" year="1980">Crush 1980</bibRefCitation>
). The slender posterior process, which forms the articulation facet for the articular, curves posterodorsally and medially, and its medial surface is tilted to face predominantly dorsally.
</paragraph>
<paragraph id="8BB49339310B9F024169FBB63A32FA13" blockId="22.[113,763,1078,1478]" pageId="22" pageNumber="376">
<emphasis id="B97F4F2B310B9F024169FBB638F9FB9B" box="[113,245,1078,1102]" italics="true" pageId="22" pageNumber="376">Pre-articular:</emphasis>
On either side of SMNS 12591a, near the posterior end of the mandible, an element is projected posteromedially from the medial side of the mandible. Based on their relative positions, these represent pre-articulars that have been somewhat displaced posteromedially (
<figureCitation id="13308FBC310B9F0240C8FB343A4DFB1E" box="[464,577,1204,1228]" captionStart="Figure 9" captionStartId="10.[113,178,1772,1796]" captionTargetBox="[222,1352,144,1744]" captionTargetId="figure-6@10.[220,1352,144,1744]" captionTargetPageId="10" captionText="Figure 9. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in dorsal view; B, interpretative drawing of (A); C, ventral view; D, interpretative drawing of (C). Abbreviations:an, angular; antfo, antorbital fossa; bpp, basipterygoid process; crb, ceratobranchial I; de, dentary; ect, ectopterygoid; fr, frontal; j, jugal; la, lacrimal; mx, maxilla; na, nasal; pa, parietal; pbs, parabasisphenoid; pl, palatine; plf, palatine fossa; po, postorbital; pra, pre-articular; prf, prefrontal; pt, pterygoid; ptmr, pterygoid medial rim; sa, surangular; spl, splenial; sq, squamosal; vo, vomer." figureDoi="http://doi.org/10.5281/zenodo.8426169" httpUri="https://zenodo.org/record/8426169/files/figure.png" pageId="22" pageNumber="376">Fig. 9C, D</figureCitation>
). They were also identified as such by
<bibRefCitation id="EF9AEEC8310B9F02407CFB543A0CFB3E" author="Huene FR" box="[356,512,1235,1259]" pageId="22" pageNumber="376" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene (1921)</bibRefCitation>
and
<bibRefCitation id="EF9AEEC8310B9F024358FB5338B0FADF" author="Sereno PC &amp; Wild R" pageId="22" pageNumber="376" pagination="435 - 58" refId="ref32722" refString="Sereno PC, Wild R. Procompsognathus: theropod, ' thecodont' or both? Journal of Vertebrate Paleontology 1992; 12: 435 - 58." type="journal article" year="1992">Sereno and Wild (1992)</bibRefCitation>
. They are incomplete on both sides, but form elongate elements with distinctly transversely convex medial surfaces. Due to the compression of SMNS 12591a, the dorsomedial portions of both mandibular rami are obscured and not externally visible. Therefore, the coronoid could not be confidently identified. Furthermore, because their posterior ends are missing, neither mandibular ramus preserves an articular.
</paragraph>
<paragraph id="8BB49339310B9F024169FA683C1DFF12" blockId="22.[113,762,1512,2006]" lastBlockId="22.[810,1459,143,606]" pageId="22" pageNumber="376">
<emphasis id="B97F4F2B310B9F024169FA6838DAFA2A" box="[113,214,1512,1535]" italics="true" pageId="22" pageNumber="376">Dentition:</emphasis>
The dental distribution and, where possible, tooth count estimates of the premaxilla, maxilla, and dentary are outlined above for those respective bones. All teeth of
<taxonomicName id="4C0BE8BA310B9F02436AF9A63AF6F9EB" authorityName="von Huene" authorityYear="1921" box="[626,762,1574,1598]" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="22" pageNumber="376" phylum="Chordata" rank="genus">
<emphasis id="B97F4F2B310B9F02436AF9A63AF6F9EB" box="[626,762,1574,1598]" italics="true" pageId="22" pageNumber="376">Saltoposuchus</emphasis>
</taxonomicName>
are recurved and labiolingually compressed with readily distinguishable carinae. Serrations are present on both the mesial and distal carinae. They are most obvious on the distal (posterior) carina, where they are present along the entire crown. The entire mesial (anterior) carina of smaller teeth is also serrated, but in larger teeth the proximal half of the crown lacks serrations in the maxillary and dentary teeth in SMNS 12352a (
<figureCitation id="13308FBC310B9F02436AF8823AC8F8CF" box="[626,708,1794,1818]" captionStart="Figure 11" captionStartId="17.[127,192,1672,1696]" captionTargetBox="[131,1404,147,1640]" captionTargetId="figure-6@17.[127,1407,144,1644]" captionTargetPageId="17" captionText="Figure 11. Details of the cranial anatomy of Saltoposuchus connectens. A, frontals of SMNS 12597a in ventral view; B, less squamosal of SMNS 12596a in dorsal view; C, right squamosal of SMNS 12597b in ventral view; D, less quadrate of SMNS 12597b in posterolateral view; E, less quadrate of SMNS 12596a in anterolateral view; F, enlarged anterior maxillary tooth of SMNS 12352a in angled lateral view.Abbreviations: adp, anterodorsal process; ap, anterior process; dh, dorsal head; fr, frontal; iqf, internal quadrate fenestra; l., less; lcr, longitudinal crest; lr, longitudinal ridge; olb, olfactory bulb; otfl, otoccipital flange; paf, parietal facet; pmp, posteromedial process; pof, postorbital facet; ppeg, posterior peg; prff, prefrontal facet; ptfl, post-temporal flange; ptp, pterygoid process; ptra, pterygoid ramus; qf, quadrate facet; r., right; stfo, supratemporal fossa; vdp, ventrally directed process; vee, ventral end." figureDoi="http://doi.org/10.5281/zenodo.8426173" httpUri="https://zenodo.org/record/8426173/files/figure.png" pageId="22" pageNumber="376">Fig. 11F</figureCitation>
) and SMNS 12591a, which is also the case in
<taxonomicName id="4C0BE8BA310B9F024306F8A13AF6F8EC" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[542,762,1825,1849]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="22" pageNumber="376" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B310B9F024306F8A13AF6F8EC" box="[542,762,1825,1849]" italics="true" pageId="22" pageNumber="376">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
(NHMUK PV R 7557). Some of the larger teeth, as well as some of the smaller posteriormost teeth, have a very widely rounded labial (lateral) surface. Serrations are clearly present on the distal carinae of the maxillary teeth and the large anterior dentary tooth of SMNS 12596b, but poorly preserved serrations on the mesial carina can only be observed in the second maxillary tooth counted from anterior.
</paragraph>
<paragraph id="8BB49339310B9F02425DFF4F3CDAFD8B" blockId="22.[810,1459,143,606]" pageId="22" pageNumber="376">
In SMNS 12352a, the largest maxillary tooth appears to be the third or fourth counted from anterior, positioned on the most convex portion of the maxillary alveolar margin in lateral view (
<figureCitation id="13308FBC310B9F02422DFEAC3BA7FE91" box="[821,939,300,324]" captionStart="Figure 10" captionStartId="14.[113,178,1772,1796]" captionTargetBox="[197,1376,144,1744]" captionTargetId="figure-5@14.[197,1376,144,1744]" captionTargetPageId="14" captionText="Figure 10. Saltoposuchus connectens referred specimen SMNS 12352.A, SMNS 12352a with the snout region exposed in angled right lateral view; B, interpretative drawing of (A); C, opposite side of SMNS 12352a exposing the posterior portion of the less maxilla and lacrimal in angled lateral view; D, interpretative drawing of (C); E, SMNS 12352b, carpus and manus in dorsal view; F, interpretative drawing of (E); G, SMNS 12352b in medial view; H, interpretative drawing of (G). Abbreviations; antfe, antorbital fenestra; antfo, antorbital fossa; dc, distal carpal; de, dentary; exp, extensor pit; fr, frontal; j, jugal; la, lacrimal; lad, lacrimal duct; mc, metacarpal; mx, maxilla; na, nasal; ph, phalanx; pm, premaxilla; prf, prefrontal; ra, radiale; ul, ulnare." figureDoi="http://doi.org/10.5281/zenodo.8426171" httpUri="https://zenodo.org/record/8426171/files/figure.png" pageId="22" pageNumber="376">Figs 10A, B</figureCitation>
,
<figureCitation id="13308FBC310B9F0242AEFEAC3BD2FE91" box="[950,990,300,324]" captionStart="Figure 11" captionStartId="17.[127,192,1672,1696]" captionTargetBox="[131,1404,147,1640]" captionTargetId="figure-6@17.[127,1407,144,1644]" captionTargetPageId="17" captionText="Figure 11. Details of the cranial anatomy of Saltoposuchus connectens. A, frontals of SMNS 12597a in ventral view; B, less squamosal of SMNS 12596a in dorsal view; C, right squamosal of SMNS 12597b in ventral view; D, less quadrate of SMNS 12597b in posterolateral view; E, less quadrate of SMNS 12596a in anterolateral view; F, enlarged anterior maxillary tooth of SMNS 12352a in angled lateral view.Abbreviations: adp, anterodorsal process; ap, anterior process; dh, dorsal head; fr, frontal; iqf, internal quadrate fenestra; l., less; lcr, longitudinal crest; lr, longitudinal ridge; olb, olfactory bulb; otfl, otoccipital flange; paf, parietal facet; pmp, posteromedial process; pof, postorbital facet; ppeg, posterior peg; prff, prefrontal facet; ptfl, post-temporal flange; ptp, pterygoid process; ptra, pterygoid ramus; qf, quadrate facet; r., right; stfo, supratemporal fossa; vdp, ventrally directed process; vee, ventral end." figureDoi="http://doi.org/10.5281/zenodo.8426173" httpUri="https://zenodo.org/record/8426173/files/figure.png" pageId="22" pageNumber="376">11F</figureCitation>
). The maxillary teeth anterior to this are considerably smaller, whereas posteriorly the size of the maxillary teeth reduces gradually. Although less obvious, this condition is also present in SMNS 12591a (
<figureCitation id="13308FBC310B9F024588FE0A3CC7FE77" box="[1168,1227,394,418]" captionStart="Figure 8" captionStartId="9.[129,194,1772,1796]" captionTargetBox="[178,1429,144,1744]" captionTargetId="figure-6@9.[174,1429,144,1744]" captionTargetPageId="9" captionText="Figure 8. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in right lateral view; B, interpretative drawing of A; C, less lateral view; D, interpretative drawing of (C). Abbreviations:an, angular; cc, crista cranii; de, dentary; fr, frontal; j, jugal; la, lacrimal; ls, laterosphenoid; mx, maxilla; na, nasal; pa, parietal; pl, palatine; po, postorbital; prf, prefrontal; pt, pterygoid; q, quadrate; qj, quadratojugal; sa, surangular; sq, squamosal." figureDoi="http://doi.org/10.5281/zenodo.8426167" httpUri="https://zenodo.org/record/8426167/files/figure.png" pageId="22" pageNumber="376">Fig. 8</figureCitation>
). The presence of enlarged teeth on the anterior portion of the maxilla, as well as the presence of an enlarged dentary tooth near the anterior end of the bone, as seen in the less dentary of SMNS 12596b (
<figureCitation id="13308FBC310B9F024444FE683DAEFDD5" box="[1372,1442,488,512]" captionStart="Figure 3" captionStartId="4.[113,178,1499,1523]" captionTargetBox="[147,1426,144,1471]" captionTargetId="figure-217@4.[146,1426,144,1471]" captionTargetPageId="4" captionText="Figure 3. Part of Saltoposuchus connectens referred specimen SMNS 12596. A, SMNS 12596a, cranial block; B, SMNS 12596b, two dentaries exposed in ventrolateral view; C, SMNS 12596c exposing a less femur in anterolateral view; D, exposing the opposite side of SMNS 12596c, Abbreviations:de, dentary; dr, dorsal rib; dv, dorsal vertebra; fe, femur; ga, gastralia; l., less; mx, maxilla; na, nasal; os, osteoderm; q, quadrate; r., right; sq, squamosal; sr, sacral rib; sv, sacral vertebra; tp, transverse process." figureDoi="http://doi.org/10.5281/zenodo.8426153" httpUri="https://zenodo.org/record/8426153/files/figure.png" pageId="22" pageNumber="376">Fig. 3B</figureCitation>
), also occurs in the crocodylomorphs CM 29894,
<taxonomicName id="4C0BE8BA310B9F024404FD873B77FDEA" authorityName="Simmons" authorityYear="1965" class="Reptilia" family="Sphenosuchidae" genus="Dibothrosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="22" pageNumber="376" phylum="Chordata" rank="species" species="elaphros">
<emphasis id="B97F4F2B310B9F024404FD873B77FDEA" italics="true" pageId="22" pageNumber="376">Dibothrosuchus elaphros</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C0BE8BA310B9F0242A5FDA73C80FDEA" box="[957,1164,551,575]" class="Reptilia" genus="Junggarsuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="22" pageNumber="376" phylum="Chordata" rank="species" species="sloani">
<emphasis id="B97F4F2B310B9F0242A5FDA73C80FDEA" box="[957,1164,551,575]" italics="true" pageId="22" pageNumber="376">Junggarsuchus sloani</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9AEEC8310B9F0245BAFDA73DA1FDEA" author="Wu X-C &amp; Chauerjee S" box="[1186,1453,551,575]" pageId="22" pageNumber="376" pagination="58 - 89" refId="ref33269" refString="Wu X-C, Chauerjee S. Dibothrosuchus elaphros, a crocodylomorph from the Lower Jurassic of China and the phylogeny of the Sphenosuchia. Journal of Vertebrate Paleontology 1993; 13: 58 - 89." type="journal article" year="1993">Wu and Chaưerjee 1993</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310B9F024232FDC63BC3FD8B" author="Clark JM &amp; Sues H-D &amp; Berman DS" box="[810,975,582,606]" pageId="22" pageNumber="376" pagination="683 - 704" refId="ref31006" refString="Clark JM, Sues H-D, Berman DS. A new specimen of Hesperosuchus agilis from the Upper Triassic of New Mexico and the interrelationships of basal crocodylomorph archosaurs. Journal of Vertebrate Paleontology 2000; 20: 683 - 704." type="journal article" year="2000">
Clark
<emphasis id="B97F4F2B310B9F024270FDC73B9BFD8B" box="[872,919,582,606]" italics="true" pageId="22" pageNumber="376">et al.</emphasis>
2000
</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310B9F0242C2FDC63CCAFD8B" author="Ruebenstahl AA &amp; Klein MD &amp; Yi H" box="[986,1222,582,606]" pageId="22" pageNumber="376" pagination="2463 - 556" refId="ref32564" refString="Ruebenstahl AA, Klein MD, Yi H et al. Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros. The Anatomical Record 2022; 305: 2463 - 556." type="journal article" year="2022">
Ruebenstahl
<emphasis id="B97F4F2B310B9F024547FDC73C82FD8B" box="[1119,1166,582,606]" italics="true" pageId="22" pageNumber="376">et al.</emphasis>
2022
</bibRefCitation>
).
</paragraph>
<paragraph id="8BB49339310B9F024232FDFF3D9FFCA6" blockId="22.[810,1459,639,883]" pageId="22" pageNumber="376">
<emphasis id="B97F4F2B310B9F024232FDFF3C11FD42" box="[810,1053,639,663]" italics="true" pageId="22" pageNumber="376">Hyobranchial apparatus:</emphasis>
Two slender, rod-like elements are preserved lateroventral to each side of the parabasisphenoid in ventral view in SMNS 12591a (
<figureCitation id="13308FBC310B9F02454BFD3E3CB3FD03" box="[1107,1215,702,726]" captionStart="Figure 9" captionStartId="10.[113,178,1772,1796]" captionTargetBox="[222,1352,144,1744]" captionTargetId="figure-6@10.[220,1352,144,1744]" captionTargetPageId="10" captionText="Figure 9. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in dorsal view; B, interpretative drawing of (A); C, ventral view; D, interpretative drawing of (C). Abbreviations:an, angular; antfo, antorbital fossa; bpp, basipterygoid process; crb, ceratobranchial I; de, dentary; ect, ectopterygoid; fr, frontal; j, jugal; la, lacrimal; mx, maxilla; na, nasal; pa, parietal; pbs, parabasisphenoid; pl, palatine; plf, palatine fossa; po, postorbital; pra, pre-articular; prf, prefrontal; pt, pterygoid; ptmr, pterygoid medial rim; sa, surangular; spl, splenial; sq, squamosal; vo, vomer." figureDoi="http://doi.org/10.5281/zenodo.8426169" httpUri="https://zenodo.org/record/8426169/files/figure.png" pageId="22" pageNumber="376">Fig. 9C, D</figureCitation>
). These slightly curved bones represent the paired ceratobranchial I. They are similar in morphology to those of other non- mesoeucrocodylian crocodylomorphs that preserve this feature of the hyobranchial apparatus in some detail:
<taxonomicName id="4C0BE8BA310B9F024529FCBB3D00FC86" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[1073,1292,827,851]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="22" pageNumber="376" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B310B9F024529FCBB3D00FC86" box="[1073,1292,827,851]" italics="true" pageId="22" pageNumber="376">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0BE8BA310B9F024427FCBB3B9BFCA6" baseAuthorityName="Brown" baseAuthorityYear="1933" class="Reptilia" family="Protosuchidae" genus="Protosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="22" pageNumber="376" phylum="Chordata" rank="species" species="richardsoni">
<emphasis id="B97F4F2B310B9F024427FCBB3B9BFCA6" italics="true" pageId="22" pageNumber="376">Protosuchus richardsoni</emphasis>
</taxonomicName>
(NHMUK PV R 7591,
<bibRefCitation id="EF9AEEC8310B9F024593FCDB3D8EFCA7" author="Colbert EH &amp; Mook CC" box="[1163,1410,858,883]" pageId="22" pageNumber="376" pagination="143 - 82" refId="ref31130" refString="Colbert EH, Mook CC. The ancestral crocodilian Protosuchus. Bulletin of the American Museum of Natural History 1951; 97: 143 - 82." type="journal article" year="1951">Colbert and Mook 1951</bibRefCitation>
).
</paragraph>
<paragraph id="8BB49339310B9F02452CFC0C3CA5FC70" blockId="22.[1076,1193,908,933]" box="[1076,1193,908,933]" pageId="22" pageNumber="376">
<emphasis id="B97F4F2B310B9F02452CFC0C3CA5FC70" box="[1076,1193,908,933]" italics="true" pageId="22" pageNumber="376">Postcranium</emphasis>
</paragraph>
<paragraph id="8BB49339310B9F024232FC323C54F8A3" blockId="22.[810,1459,946,2004]" pageId="22" pageNumber="376">
<emphasis id="B97F4F2B310B9F024232FC323BECFC1F" box="[810,992,946,970]" italics="true" pageId="22" pageNumber="376">Cervical vertebrae:</emphasis>
<bibRefCitation id="EF9AEEC8310B9F0242F4FC323C8DFC1F" author="Huene FR" box="[1004,1153,946,970]" pageId="22" pageNumber="376" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene (1921)</bibRefCitation>
and
<bibRefCitation id="EF9AEEC8310B9F0245ABFC323D4EFC1F" author="Crush PJ" box="[1203,1346,946,970]" pageId="22" pageNumber="376" refId="ref31157" refString="Crush PJ. An early, terrestrial, crocodile from South Wales. Unpublished Ph. D. Thesis, University College London, 1980." type="book" year="1980">Crush (1980)</bibRefCitation>
both identified two articulated vertebrae preserved in SMNS 12597c as cervical vertebrae. Both are exposed in dorsal view and the anterior and beưer preserved vertebra exhibits the pre- and postzygapophyses, neural arch, and right diapophysis (
<figureCitation id="13308FBC310B9F024443FBAF3DAEFB92" box="[1371,1442,1071,1095]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="22" pageNumber="376">Fig. 2E</figureCitation>
). The anterior position of the diapophysis indeed suggests that this element represents a posterior cervical vertebra. However, these elements are not very informative, and the best representatives of the cervical vertebrae are two isolated vertebrae, SMNS 12596d–e (
<figureCitation id="13308FBC310B9F0242EBFB4C3C6CFB31" box="[1011,1120,1228,1252]" captionStart="Figure 4" captionStartId="5.[129,194,1154,1178]" captionTargetBox="[164,1441,148,1126]" captionTargetId="figure-363@5.[161,1441,144,1126]" captionTargetPageId="5" captionText="Figure 4. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, SMNS 12596d, cervical vertebra exposed in less lateral view; B, interpretative drawing of (A); C, SMNS 12596e, posterior cervical or anterior dorsal vertebra exposed in an angled right anterolateral view; D, interpretative drawing of (C); E, SMNS 12596c, close-up of second sacral vertebra in right lateral view and osteoderm in dorsal view; F, interpretative drawing of (E); G, SMNS 12596f, proximal caudal vertebra block exposed in less lateral view; H, interpretative drawing of (G). Abbreviations: accs, accessory anterior spine; dp, diapophysis; izl, interzygapophyseal lamina; ncs, neurocentral suture; ns, neural spine; os, osteoderm; pap, parapophysis; pcdl, posterior centrodiapophyseal lamina; pofl, posterior flange; poz, postzygaphysis; prdl, prezygodiapophyseal lamina; prz, prezygapophysis; sr, sacral rib; tp, transverse process." figureDoi="http://doi.org/10.5281/zenodo.8426157" httpUri="https://zenodo.org/record/8426157/files/figure.png" pageId="22" pageNumber="376">Fig. 4A–D</figureCitation>
). Based on the relative position of the dia- and parapophysis, both represent approximately mid-cervical vertebrae.
<bibRefCitation id="EF9AEEC8310B9F02453CFA8B3CB2FAF6" author="Huene FR" box="[1060,1214,1291,1315]" pageId="22" pageNumber="376" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene (1921)</bibRefCitation>
provided a similar interpretation for SMNS 12596d, but considered SMNS 12596e to represent a transitional vertebra between the cervical and dorsal series.
<bibRefCitation id="EF9AEEC8310B9F0242ABFAE83C4FFA54" author="Crush PJ" box="[947,1091,1384,1409]" pageId="22" pageNumber="376" refId="ref31157" refString="Crush PJ. An early, terrestrial, crocodile from South Wales. Unpublished Ph. D. Thesis, University College London, 1980." type="book" year="1980">Crush (1980)</bibRefCitation>
identified both elements as cervical vertebrae. The anterior surface of the centrum is exposed in SMNS 12596e (
<figureCitation id="13308FBC310B9F0242CBFA273C31FA6A" box="[979,1085,1447,1471]" captionStart="Figure 4" captionStartId="5.[129,194,1154,1178]" captionTargetBox="[164,1441,148,1126]" captionTargetId="figure-363@5.[161,1441,144,1126]" captionTargetPageId="5" captionText="Figure 4. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, SMNS 12596d, cervical vertebra exposed in less lateral view; B, interpretative drawing of (A); C, SMNS 12596e, posterior cervical or anterior dorsal vertebra exposed in an angled right anterolateral view; D, interpretative drawing of (C); E, SMNS 12596c, close-up of second sacral vertebra in right lateral view and osteoderm in dorsal view; F, interpretative drawing of (E); G, SMNS 12596f, proximal caudal vertebra block exposed in less lateral view; H, interpretative drawing of (G). Abbreviations: accs, accessory anterior spine; dp, diapophysis; izl, interzygapophyseal lamina; ncs, neurocentral suture; ns, neural spine; os, osteoderm; pap, parapophysis; pcdl, posterior centrodiapophyseal lamina; pofl, posterior flange; poz, postzygaphysis; prdl, prezygodiapophyseal lamina; prz, prezygapophysis; sr, sacral rib; tp, transverse process." figureDoi="http://doi.org/10.5281/zenodo.8426157" httpUri="https://zenodo.org/record/8426157/files/figure.png" pageId="22" pageNumber="376">Fig. 4C, D</figureCitation>
) and the posterior surface in SMNS 12596d (
<figureCitation id="13308FBC310B9F024294FA463BF9FA0B" box="[908,1013,1478,1503]" captionStart="Figure 4" captionStartId="5.[129,194,1154,1178]" captionTargetBox="[164,1441,148,1126]" captionTargetId="figure-363@5.[161,1441,144,1126]" captionTargetPageId="5" captionText="Figure 4. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, SMNS 12596d, cervical vertebra exposed in less lateral view; B, interpretative drawing of (A); C, SMNS 12596e, posterior cervical or anterior dorsal vertebra exposed in an angled right anterolateral view; D, interpretative drawing of (C); E, SMNS 12596c, close-up of second sacral vertebra in right lateral view and osteoderm in dorsal view; F, interpretative drawing of (E); G, SMNS 12596f, proximal caudal vertebra block exposed in less lateral view; H, interpretative drawing of (G). Abbreviations: accs, accessory anterior spine; dp, diapophysis; izl, interzygapophyseal lamina; ncs, neurocentral suture; ns, neural spine; os, osteoderm; pap, parapophysis; pcdl, posterior centrodiapophyseal lamina; pofl, posterior flange; poz, postzygaphysis; prdl, prezygodiapophyseal lamina; prz, prezygapophysis; sr, sacral rib; tp, transverse process." figureDoi="http://doi.org/10.5281/zenodo.8426157" httpUri="https://zenodo.org/record/8426157/files/figure.png" pageId="22" pageNumber="376">Fig. 4A, B</figureCitation>
). Both are concave; the cervical vertebrae were therefore amphicoelous. The ventral margin is strongly concave in lateral view; a clear ventral keel is absent. This corresponds to the condition in the mid- to posterior cervical vertebrae of
<taxonomicName id="4C0BE8BA310B9F0242ADF9C43C99F989" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[949,1173,1604,1628]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="22" pageNumber="376" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B310B9F0242ADF9C43C99F989" box="[949,1173,1604,1628]" italics="true" pageId="22" pageNumber="376">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
(NHMUK PV R 7593b), whereas all cervical vertebrae are keeled in
<taxonomicName id="4C0BE8BA310B9F024407F9E33C4FF94E" authority="(Sues et al. 2003)" baseAuthorityName="Sues" baseAuthorityYear="2003" class="Reptilia" family="Sphenosuchidae" genus="Dromicosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="22" pageNumber="376" phylum="Chordata" rank="species" species="grallator">
<emphasis id="B97F4F2B310B9F024407F9E33B72F94F" italics="true" pageId="22" pageNumber="376">Dromicosuchus grallator</emphasis>
(
<bibRefCitation id="EF9AEEC8310B9F024289F9033C3BF94E" author="Sues H-D &amp; Olsen PE &amp; Carter JG" box="[913,1079,1666,1691]" pageId="22" pageNumber="376" pagination="329 - 43" refId="ref32828" refString="Sues H-D, Olsen PE, Carter JG et al. A new crocodylomorph archosaur from the Upper Triassic of North Carolina. Journal of Vertebrate Paleontology 2003; 23: 329 - 43." type="journal article" year="2003">
Sues
<emphasis id="B97F4F2B310B9F0242D1F9033BF7F94F" box="[969,1019,1666,1690]" italics="true" pageId="22" pageNumber="376">et al.</emphasis>
2003
</bibRefCitation>
)
</taxonomicName>
. Hypapophyses, as occur in extant crocodylians and
<taxonomicName id="4C0BE8BA310B9F0242C2F9223CAEF96F" box="[986,1186,1698,1722]" class="Reptilia" genus="Junggarsuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="22" pageNumber="376" phylum="Chordata" rank="species" species="sloani">
<emphasis id="B97F4F2B310B9F0242C2F9223CAEF96F" box="[986,1186,1698,1722]" italics="true" pageId="22" pageNumber="376">Junggarsuchus sloani</emphasis>
</taxonomicName>
among non-crocodyliform crocodylomorphs (Hoffsteưer and Gasc 1969,
<bibRefCitation id="EF9AEEC8310B9F024419F9413DAEF90C" author="Clark JM &amp; Xu X &amp; Forster CA" box="[1281,1442,1729,1753]" pageId="22" pageNumber="376" pagination="1021 - 4" refId="ref31048" refString="Clark JM, Xu X, Forster CA et al. A Middle Jurassic ' sphenosuchian' from China and the origin of the crocodylian skull. Nature 2004; 430: 1021 - 4." type="journal article" year="2004">
Clark
<emphasis id="B97F4F2B310B9F024425F9423D67F90C" box="[1341,1387,1729,1753]" italics="true" pageId="22" pageNumber="376">et al.</emphasis>
2004
</bibRefCitation>
), are also absent. In both elements, the parapophysis is positioned near the anteroventral corner of the lateral side of the centrum, whereas the diapophysis is positioned entirely on the neural arch at approximately the dorsoventral level of the posterodorsal end of the centrum in lateral view.
</paragraph>
<paragraph id="8BB49339310B9F03425DF8FD3A92FDAB" blockId="22.[810,1459,946,2004]" lastBlockId="23.[129,778,144,638]" lastPageId="23" lastPageNumber="377" pageId="22" pageNumber="376">
The neurocentral suture can be observed in SMNS 12596e (
<figureCitation id="13308FBC310B9F02422DF81C3B84F861" box="[821,904,1948,1972]" captionStart="Figure 4" captionStartId="5.[129,194,1154,1178]" captionTargetBox="[164,1441,148,1126]" captionTargetId="figure-363@5.[161,1441,144,1126]" captionTargetPageId="5" captionText="Figure 4. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, SMNS 12596d, cervical vertebra exposed in less lateral view; B, interpretative drawing of (A); C, SMNS 12596e, posterior cervical or anterior dorsal vertebra exposed in an angled right anterolateral view; D, interpretative drawing of (C); E, SMNS 12596c, close-up of second sacral vertebra in right lateral view and osteoderm in dorsal view; F, interpretative drawing of (E); G, SMNS 12596f, proximal caudal vertebra block exposed in less lateral view; H, interpretative drawing of (G). Abbreviations: accs, accessory anterior spine; dp, diapophysis; izl, interzygapophyseal lamina; ncs, neurocentral suture; ns, neural spine; os, osteoderm; pap, parapophysis; pcdl, posterior centrodiapophyseal lamina; pofl, posterior flange; poz, postzygaphysis; prdl, prezygodiapophyseal lamina; prz, prezygapophysis; sr, sacral rib; tp, transverse process." figureDoi="http://doi.org/10.5281/zenodo.8426157" httpUri="https://zenodo.org/record/8426157/files/figure.png" pageId="22" pageNumber="376">Fig. 4D</figureCitation>
), whereas no suture is visible in SMNS 12596d, which could be the result of poor preservation. Two clear laminae are preserved on the neural arches in both specimens. A prezygodiapophyseal lamina (
<emphasis id="B97F4F2B310A9F0340C4FF303A03FF12" box="[476,527,176,199]" italics="true" pageId="23" pageNumber="377">sensu</emphasis>
<bibRefCitation id="EF9AEEC8310A9F034302FF2F3AA8FF12" author="Wilson JA" box="[538,676,175,199]" pageId="23" pageNumber="377" pagination="639 - 53" refId="ref33242" refString="Wilson JA. A nomenclature for vertebral laminae in sauropods and other saurischian dinosaurs. Journal of Vertebrate Paleontology 1999; 19: 639 - 53." type="journal article" year="1999">Wilson 1999</bibRefCitation>
) is positioned between the prezygapophysis and diapophysis, and an interzygapophyseal lamina is formed between the pre- and postzygapophyses with a very clear and deeply excavated fossa positioned directly dorsomedial to it (
<figureCitation id="13308FBC310A9F034316FEAD3A75FE90" box="[526,633,301,325]" captionStart="Figure 4" captionStartId="5.[129,194,1154,1178]" captionTargetBox="[164,1441,148,1126]" captionTargetId="figure-363@5.[161,1441,144,1126]" captionTargetPageId="5" captionText="Figure 4. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, SMNS 12596d, cervical vertebra exposed in less lateral view; B, interpretative drawing of (A); C, SMNS 12596e, posterior cervical or anterior dorsal vertebra exposed in an angled right anterolateral view; D, interpretative drawing of (C); E, SMNS 12596c, close-up of second sacral vertebra in right lateral view and osteoderm in dorsal view; F, interpretative drawing of (E); G, SMNS 12596f, proximal caudal vertebra block exposed in less lateral view; H, interpretative drawing of (G). Abbreviations: accs, accessory anterior spine; dp, diapophysis; izl, interzygapophyseal lamina; ncs, neurocentral suture; ns, neural spine; os, osteoderm; pap, parapophysis; pcdl, posterior centrodiapophyseal lamina; pofl, posterior flange; poz, postzygaphysis; prdl, prezygodiapophyseal lamina; prz, prezygapophysis; sr, sacral rib; tp, transverse process." figureDoi="http://doi.org/10.5281/zenodo.8426157" httpUri="https://zenodo.org/record/8426157/files/figure.png" pageId="23" pageNumber="377">Fig. 4A–D</figureCitation>
). In addition, a poorly developed lamina extends posteroventrally from the diapophysis, the posterior centrodiapophyseal lamina. The elongate pre- and postzygapophyses of both vertebrae are directed anterodorsally and posterodorsally, respectively, with a strong dorsal inclination. The neural spines are tall and slender. In SMNS 12596e, the neural spine expands very slightly distally in lateral view. In SMNS 12596d, the neural spine appears to taper slightly distally, but this is because its posterior margin is incomplete. The distal ends of the neural spines are flat to slightly convex in lateral view, and not transversely expanded.
</paragraph>
<paragraph id="8BB49339310A9F034199FD1F3999FABD" blockId="23.[128,778,671,1979]" pageId="23" pageNumber="377">
<emphasis id="B97F4F2B310A9F034199FD1F3926FD62" box="[129,298,671,695]" italics="true" pageId="23" pageNumber="377">Dorsal vertebrae:</emphasis>
An articulated series of at least seven dorsal vertebrae is preserved in SMNS 12597a but they are largely embedded in matrix and covered by overlying osteoderms, and only the less sides of their neural arches are exposed. Additionally, two broken but largely complete dorsal vertebrae exposed in right lateral view are preserved in SMNS 12597c (
<figureCitation id="13308FBC310A9F034389FCBC3AF4FC86" box="[657,760,828,852]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="23" pageNumber="377">Fig. 2A, E</figureCitation>
). The right side of a well preserved neural arch of a dorsal vertebrae is exposed in ventral view in SMNS 12596c (
<figureCitation id="13308FBC310A9F03436EFCFA3AC8FC46" box="[630,708,890,915]" captionStart="Figure 3" captionStartId="4.[113,178,1499,1523]" captionTargetBox="[147,1426,144,1471]" captionTargetId="figure-217@4.[146,1426,144,1471]" captionTargetPageId="4" captionText="Figure 3. Part of Saltoposuchus connectens referred specimen SMNS 12596. A, SMNS 12596a, cranial block; B, SMNS 12596b, two dentaries exposed in ventrolateral view; C, SMNS 12596c exposing a less femur in anterolateral view; D, exposing the opposite side of SMNS 12596c, Abbreviations:de, dentary; dr, dorsal rib; dv, dorsal vertebra; fe, femur; ga, gastralia; l., less; mx, maxilla; na, nasal; os, osteoderm; q, quadrate; r., right; sq, squamosal; sr, sacral rib; sv, sacral vertebra; tp, transverse process." figureDoi="http://doi.org/10.5281/zenodo.8426153" httpUri="https://zenodo.org/record/8426153/files/figure.png" pageId="23" pageNumber="377">Figs 3D</figureCitation>
,
<figureCitation id="13308FBC310A9F0343D6FCFB3AF5FC46" box="[718,761,891,915]" captionStart="Figure 12" captionStartId="24.[113,178,1772,1796]" captionTargetBox="[151,1424,144,1744]" captionTargetId="figure-6@24.[148,1424,144,1744]" captionTargetPageId="24" captionText="Figure 12. Details of the postcranial anatomy of Saltoposuchus connectens. A, dorsal vertebral neural arch and dorsal rib of SMNS 12596c in ventral view; B, associated osteoderms of SMNS 12596c in dorsal view; C, sacral ribs and associated elements of SMNS 12597a in dorsal view; D, distal portion of right scapular blade of SMNS 12597a in lateral view; E, proximal end of right tibia of SMNS 12597a in proximal view; F, less tarsus and pes of SMNS 12597a with the calcaneum exposed in lateral view; G, close-up of less tarsus of SMNS 12597a; H, close-up of less pes of SMNS 12597a. Abbreviations: ane, anterior expansion; cal, calcaneum; calt, calcaneal tuber; cp, capitulum; dp, diapophysis; dt, distal tarsal; exp, extensor pit; fe, femur; fi, fibula; il, ilium; lam, lamina; latc, lateral condyle; le, lateral edge; lr, longitudinal ridge; me, medial edge; mt, metatarsal; no, notch; pap, parapophysis; ph, phalanx; pme, posterior extension; poe, posterior expansion; pofl, posterior flange; poz, postzygapophysis; os, osteoderm; r., right; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426175" httpUri="https://zenodo.org/record/8426175/files/figure.png" pageId="23" pageNumber="377">12A</figureCitation>
). Finally, two isolated vertebral centra are preserved in SMNS 55009. Based on the relative elongation of these centra, SMNS 55009a represents a dorsal centrum, whereas SMNS 55009d is a proximal caudal centrum (
<figureCitation id="13308FBC310A9F034088FC7839F8FBDA" box="[400,500,1016,1040]" captionStart="Figure 6" captionStartId="7.[129,194,1605,1629]" captionTargetBox="[161,1441,657,1577]" captionTargetId="figure-310@7.[161,1441,657,1577]" captionTargetPageId="7" captionText="Figure 6. Part of Saltoposuchus connectens referred specimen SMNS 55009. A, SMNS 55009a, dorsal vertebral centrum in less lateral view; B, SMNS 55009d, proximal caudal vertebral centrum in angled lateral view; C, SMNS 55009b, first sacral vertebra in less lateral view; D, interpretative drawing of (C); E, SMNS 55009c, proximal caudal vertebra in posterior view; F, interpretative drawing of (E). Abbreviations: car, caudal rib; ns, neural spine; pas, posterior articular surface; poz, postzygapophysis; prz, prezygapophysis; sr, sacral rib; sut, suture." figureDoi="http://doi.org/10.5281/zenodo.8426161" httpUri="https://zenodo.org/record/8426161/files/figure.png" pageId="23" pageNumber="377">Fig. 6A, B</figureCitation>
). However, these elements are largely undiagnostic, and their association with the other remains assigned to SMNS 55009 is unclear, and therefore their assignment to
<taxonomicName id="4C0BE8BA310A9F03403AFBD63A11FBB8" authorityName="Huene" authorityYear="1921" box="[290,541,1110,1134]" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="23" pageNumber="377" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B310A9F03403AFBD63A11FBB8" box="[290,541,1110,1134]" italics="true" pageId="23" pageNumber="377">Saltoposuchus connectens</emphasis>
</taxonomicName>
should be treated as tentative. Nevertheless, the dorsal centrum has a concave ventral margin in lateral view, and a slightly transversely concave lateral surface, which corresponds to the condition seen in the dorsal centrum of SMNS 12597c and that of
<taxonomicName id="4C0BE8BA310A9F034360FB5338C9FADF" baseAuthorityName="Crush" baseAuthorityYear="1984" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="23" pageNumber="377" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B310A9F034360FB5338C9FADF" italics="true" pageId="23" pageNumber="377">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0BE8BA310A9F0341CDFB7239ADFADF" box="[213,417,1266,1290]" class="Reptilia" family="Sphenosuchidae" genus="Hesperosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="23" pageNumber="377" phylum="Chordata" rank="species" species="agilis">
<emphasis id="B97F4F2B310A9F0341CDFB7239ADFADF" box="[213,417,1266,1290]" italics="true" pageId="23" pageNumber="377">Hesperosuchus agilis</emphasis>
</taxonomicName>
(AMNH FARB 6758),
<taxonomicName id="4C0BE8BA310A9F0343B6FB7238CDFAFC" authorityName="Reig" authorityYear="1963" class="Polyplacophora" genus="Trialestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodilomorphi" pageId="23" pageNumber="377" phylum="Mollusca" rank="species" species="romeri">
<emphasis id="B97F4F2B310A9F0343B6FB7238CDFAFC" italics="true" pageId="23" pageNumber="377">Trialestes romeri</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0BE8BA310A9F0341D4FA9239E9FAFF" authorityName="Bonaparte" authorityYear="1969" box="[204,485,1298,1322]" class="Polyplacophora" genus="Pseudhesperosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="23" pageNumber="377" phylum="Mollusca" rank="species" species="jachaleri">
<emphasis id="B97F4F2B310A9F0341D4FA9239E9FAFF" box="[204,485,1298,1322]" italics="true" pageId="23" pageNumber="377">Pseudhesperosuchus jachaleri</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C0BE8BA310A9F034304FA923B06FAFF" authorityName="Simmons" authorityYear="1965" box="[540,778,1298,1322]" class="Reptilia" family="Sphenosuchidae" genus="Dibothrosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="23" pageNumber="377" phylum="Chordata" rank="species" species="elaphros">
<emphasis id="B97F4F2B310A9F034304FA923B06FAFF" box="[540,778,1298,1322]" italics="true" pageId="23" pageNumber="377">Dibothrosuchus elaphros</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9AEEC8310A9F034193FAB1391FFA9C" author="Colbert EH" box="[139,275,1329,1353]" pageId="23" pageNumber="377" pagination="561 - 92" refId="ref31083" refString="Colbert EH. A pseudosuchian reptile from Arizona. Bulletin of the American Museum of Natural History 1952; 99: 561 - 92." type="journal article" year="1952">Colbert 1952</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310A9F034005FAB139B3FA9C" author="Bonaparte JF" box="[285,447,1329,1353]" pageId="23" pageNumber="377" pagination="1 - 183" refId="ref30694" refString="Bonaparte JF. Los tetrapodos del sector superior de la Formacion Los Colorados, La Rioja, Argentina (Triasico Superior). 1 Parte. Opera Lilloana 1972; 22: 1 - 183." type="journal article" year="1972">Bonaparte 1972</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310A9F0340D1FAB13A4DFA9C" author="Crush PJ" box="[457,577,1329,1353]" pageId="23" pageNumber="377" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310A9F034353FAB138B9FABD" author="Wu X-C &amp; Chauerjee S" pageId="23" pageNumber="377" pagination="58 - 89" refId="ref33269" refString="Wu X-C, Chauerjee S. Dibothrosuchus elaphros, a crocodylomorph from the Lower Jurassic of China and the phylogeny of the Sphenosuchia. Journal of Vertebrate Paleontology 1993; 13: 58 - 89." type="journal article" year="1993">Wu and Chaưerjee 1993</bibRefCitation>
,
<bibRefCitation id="EF9AEEC8310A9F0341A7FAD13989FABD" author="Lecuona A &amp; Ezcurra MD &amp; Irmis RB" box="[191,389,1360,1384]" pageId="23" pageNumber="377" pagination="585 - 622" refId="ref32010" refString="Lecuona A, Ezcurra MD, Irmis RB. Revision of the early crocodylomorph Trialestes romeri (Archosauria, Suchia) from the lower Upper Triassic Ischigualasto Formation of Argentina: one of the oldest-known crocodylomorphs. Papers in Palaeontology 2016; 2: 585 - 622." type="journal article" year="2016">
Lecuona
<emphasis id="B97F4F2B310A9F034006FAD13940FABD" box="[286,332,1360,1384]" italics="true" pageId="23" pageNumber="377">et al.</emphasis>
2016
</bibRefCitation>
).
</paragraph>
<paragraph id="8BB49339310A9F034184FAF0390DF86E" blockId="23.[128,778,671,1979]" pageId="23" pageNumber="377">
The isolated centrum SMNS 55009a is the only element to expose the articular surfaces of the centrum. The centrum is amphicoelous despite being strongly compressed mediolaterally. The most complete dorsal vertebra of SMNS 12597c has a clearly separated dia- and parapophysis (
<figureCitation id="13308FBC310A9F0340FFFA6D3A3EF9D0" box="[487,562,1517,1541]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="23" pageNumber="377">Fig. 2E</figureCitation>
). The laưer is positioned directly dorsal to the neurocentral suture, just anterior to the anteroposterior mid-level of the vertebra, identifying this element as an anterior dorsal vertebra. The diapophysis is badly broken and no clear laminae can be discerned. SMNS 12596c preserves the morphology of the neural arch of a more posterior dorsal vertebra (
<figureCitation id="13308FBC310A9F034031F929398CF914" box="[297,384,1705,1729]" captionStart="Figure 12" captionStartId="24.[113,178,1772,1796]" captionTargetBox="[151,1424,144,1744]" captionTargetId="figure-6@24.[148,1424,144,1744]" captionTargetPageId="24" captionText="Figure 12. Details of the postcranial anatomy of Saltoposuchus connectens. A, dorsal vertebral neural arch and dorsal rib of SMNS 12596c in ventral view; B, associated osteoderms of SMNS 12596c in dorsal view; C, sacral ribs and associated elements of SMNS 12597a in dorsal view; D, distal portion of right scapular blade of SMNS 12597a in lateral view; E, proximal end of right tibia of SMNS 12597a in proximal view; F, less tarsus and pes of SMNS 12597a with the calcaneum exposed in lateral view; G, close-up of less tarsus of SMNS 12597a; H, close-up of less pes of SMNS 12597a. Abbreviations: ane, anterior expansion; cal, calcaneum; calt, calcaneal tuber; cp, capitulum; dp, diapophysis; dt, distal tarsal; exp, extensor pit; fe, femur; fi, fibula; il, ilium; lam, lamina; latc, lateral condyle; le, lateral edge; lr, longitudinal ridge; me, medial edge; mt, metatarsal; no, notch; pap, parapophysis; ph, phalanx; pme, posterior extension; poe, posterior expansion; pofl, posterior flange; poz, postzygapophysis; os, osteoderm; r., right; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426175" httpUri="https://zenodo.org/record/8426175/files/figure.png" pageId="23" pageNumber="377">Fig. 12A</figureCitation>
). The diapophysis is extended directly laterally from the neural arch and it is widely rounded distally in ventral view. Its ventral surface is thickened centrally, whereas the anterior and posterior margins are formed by thin laminae. The parapophysis is positioned at roughly the same dorsoventral level as the diapophysis, suggesting this is a mid- to posterior dorsal vertebra. The parapophysis is separated from the diapophysis and its transverse process by a distinct concavity in ventral view.
</paragraph>
<paragraph id="8BB49339310A9F034221FF0F3C82FD2E" blockId="23.[825,1475,143,763]" pageId="23" pageNumber="377">
<emphasis id="B97F4F2B310A9F034221FF0F3BA7FF72" box="[825,939,143,167]" italics="true" pageId="23" pageNumber="377">Dorsal ribs:</emphasis>
Some rib fragments are preserved in SMNS 12597a and SMNS 12596c. Both blocks also possess a well preserved example of a dorsal rib, including its proximal end. The well preserved rib of SMNS 12597a is exposed in anterior view, whereas that of SMNS 12596c is exposed in posteroventral view (
<figureCitation id="13308FBC310A9F03425CFEAC3B9AFE91" box="[836,918,300,324]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="23" pageNumber="377">Figs 2A</figureCitation>
,
<figureCitation id="13308FBC310A9F0342B0FEAC3BCBFE91" box="[936,967,300,324]" captionStart="Figure 3" captionStartId="4.[113,178,1499,1523]" captionTargetBox="[147,1426,144,1471]" captionTargetId="figure-217@4.[146,1426,144,1471]" captionTargetPageId="4" captionText="Figure 3. Part of Saltoposuchus connectens referred specimen SMNS 12596. A, SMNS 12596a, cranial block; B, SMNS 12596b, two dentaries exposed in ventrolateral view; C, SMNS 12596c exposing a less femur in anterolateral view; D, exposing the opposite side of SMNS 12596c, Abbreviations:de, dentary; dr, dorsal rib; dv, dorsal vertebra; fe, femur; ga, gastralia; l., less; mx, maxilla; na, nasal; os, osteoderm; q, quadrate; r., right; sq, squamosal; sr, sacral rib; sv, sacral vertebra; tp, transverse process." figureDoi="http://doi.org/10.5281/zenodo.8426153" httpUri="https://zenodo.org/record/8426153/files/figure.png" pageId="23" pageNumber="377">3D</figureCitation>
,
<figureCitation id="13308FBC310A9F0342CFFEAC3C0EFE91" box="[983,1026,300,324]" captionStart="Figure 12" captionStartId="24.[113,178,1772,1796]" captionTargetBox="[151,1424,144,1744]" captionTargetId="figure-6@24.[148,1424,144,1744]" captionTargetPageId="24" captionText="Figure 12. Details of the postcranial anatomy of Saltoposuchus connectens. A, dorsal vertebral neural arch and dorsal rib of SMNS 12596c in ventral view; B, associated osteoderms of SMNS 12596c in dorsal view; C, sacral ribs and associated elements of SMNS 12597a in dorsal view; D, distal portion of right scapular blade of SMNS 12597a in lateral view; E, proximal end of right tibia of SMNS 12597a in proximal view; F, less tarsus and pes of SMNS 12597a with the calcaneum exposed in lateral view; G, close-up of less tarsus of SMNS 12597a; H, close-up of less pes of SMNS 12597a. Abbreviations: ane, anterior expansion; cal, calcaneum; calt, calcaneal tuber; cp, capitulum; dp, diapophysis; dt, distal tarsal; exp, extensor pit; fe, femur; fi, fibula; il, ilium; lam, lamina; latc, lateral condyle; le, lateral edge; lr, longitudinal ridge; me, medial edge; mt, metatarsal; no, notch; pap, parapophysis; ph, phalanx; pme, posterior extension; poe, posterior expansion; pofl, posterior flange; poz, postzygapophysis; os, osteoderm; r., right; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426175" httpUri="https://zenodo.org/record/8426175/files/figure.png" pageId="23" pageNumber="377">12A</figureCitation>
). Both ribs are dichocephalous, although the tuberculum of the rib of SMNS 12596c is poorly developed, whereas that of SMNS 12597a is elongate and separated widely from the capitulum. This suggests that the laưer represents an anterior dorsal rib and the former a posterior dorsal rib. It seems likely that almost all dorsal ribs were dichocephalous, with the possible exception of the posteriormost few elements, as in
<taxonomicName id="4C0BE8BA310A9F034274FD873C4AFDCA" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[876,1094,519,543]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="23" pageNumber="377" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B310A9F034274FD873C4AFDCA" box="[876,1094,519,543]" italics="true" pageId="23" pageNumber="377">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
(NHMUK PV R 7562). The anterior dorsal rib of SMNS 12597a bears a thin lamina that extends from its anterior surface. Proximally, this lamina originates where the capitulum and tuberculum meet, and it tapers out a few mm further distally. It is incomplete. The posterior dorsal rib of SMNS 12596c exhibits a more prominent anterior lamina. It originates at the base of the capitulum, and gradually taper distally towards the end of the preserved rib shass.
</paragraph>
<paragraph id="8BB49339310A9F034221FC9C3DBAF996" blockId="23.[825,1475,796,1979]" pageId="23" pageNumber="377">
<emphasis id="B97F4F2B310A9F034221FC9C3C20FCE1" box="[825,1068,796,820]" italics="true" pageId="23" pageNumber="377">Sacral vertebrae and ribs:</emphasis>
Remains of two poorly preserved sacral vertebrae are associated with the pelvic girdle and dorsal vertebral series in SMNS 12597a (
<figureCitation id="13308FBC310A9F03459FFCDB3CDAFCA6" box="[1159,1238,859,883]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="23" pageNumber="377">Figs 2A</figureCitation>
,
<figureCitation id="13308FBC310A9F0345FDFCDB3D1DFCA6" box="[1253,1297,859,883]" captionStart="Figure 12" captionStartId="24.[113,178,1772,1796]" captionTargetBox="[151,1424,144,1744]" captionTargetId="figure-6@24.[148,1424,144,1744]" captionTargetPageId="24" captionText="Figure 12. Details of the postcranial anatomy of Saltoposuchus connectens. A, dorsal vertebral neural arch and dorsal rib of SMNS 12596c in ventral view; B, associated osteoderms of SMNS 12596c in dorsal view; C, sacral ribs and associated elements of SMNS 12597a in dorsal view; D, distal portion of right scapular blade of SMNS 12597a in lateral view; E, proximal end of right tibia of SMNS 12597a in proximal view; F, less tarsus and pes of SMNS 12597a with the calcaneum exposed in lateral view; G, close-up of less tarsus of SMNS 12597a; H, close-up of less pes of SMNS 12597a. Abbreviations: ane, anterior expansion; cal, calcaneum; calt, calcaneal tuber; cp, capitulum; dp, diapophysis; dt, distal tarsal; exp, extensor pit; fe, femur; fi, fibula; il, ilium; lam, lamina; latc, lateral condyle; le, lateral edge; lr, longitudinal ridge; me, medial edge; mt, metatarsal; no, notch; pap, parapophysis; ph, phalanx; pme, posterior extension; poe, posterior expansion; pofl, posterior flange; poz, postzygapophysis; os, osteoderm; r., right; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426175" httpUri="https://zenodo.org/record/8426175/files/figure.png" pageId="23" pageNumber="377">12C</figureCitation>
).
<bibRefCitation id="EF9AEEC8310A9F034432FCDB3DCEFCA6" author="Huene FR" box="[1322,1474,859,883]" pageId="23" pageNumber="377" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene (1921)</bibRefCitation>
suggested three sacral vertebrae were present here, but
<bibRefCitation id="EF9AEEC8310A9F03449AFCFA3B88FC64" author="Crush PJ" pageId="23" pageNumber="377" refId="ref31157" refString="Crush PJ. An early, terrestrial, crocodile from South Wales. Unpublished Ph. D. Thesis, University College London, 1980." type="book" year="1980">Crush (1980)</bibRefCitation>
correctly identified the presence of only two sacrals, as in all other crocodylomorphs, except for rare cases in which a dorsal or caudal vertebra is incorporated in the sacrum (
<bibRefCitation id="EF9AEEC8310A9F034469FC583BA8FBDA" author="Scheyer TM &amp; Hutchinson JR &amp; Strauss O" pageId="23" pageNumber="377" pagination="49972" refId="ref32600" refString="Scheyer TM, Hutchinson JR, Strauss O et al. Giant extinct caiman breaks constraint on the axial skeleton of extant crocodylians. eLife 2019; 8: e 49972." type="journal article" year="2019">
Scheyer
<emphasis id="B97F4F2B310A9F034221FC783B66FBDA" box="[825,874,1015,1039]" italics="true" pageId="23" pageNumber="377">et al.</emphasis>
2019
</bibRefCitation>
). A single, disarticulated sacral vertebra is preserved in SMNS 12596c and in SMNS 55009b (
<figureCitation id="13308FBC310A9F0345FCFB973D5DFBFB" box="[1252,1361,1047,1071]" captionStart="Figure 3" captionStartId="4.[113,178,1499,1523]" captionTargetBox="[147,1426,144,1471]" captionTargetId="figure-217@4.[146,1426,144,1471]" captionTargetPageId="4" captionText="Figure 3. Part of Saltoposuchus connectens referred specimen SMNS 12596. A, SMNS 12596a, cranial block; B, SMNS 12596b, two dentaries exposed in ventrolateral view; C, SMNS 12596c exposing a less femur in anterolateral view; D, exposing the opposite side of SMNS 12596c, Abbreviations:de, dentary; dr, dorsal rib; dv, dorsal vertebra; fe, femur; ga, gastralia; l., less; mx, maxilla; na, nasal; os, osteoderm; q, quadrate; r., right; sq, squamosal; sr, sacral rib; sv, sacral vertebra; tp, transverse process." figureDoi="http://doi.org/10.5281/zenodo.8426153" httpUri="https://zenodo.org/record/8426153/files/figure.png" pageId="23" pageNumber="377">Figs 3C, D</figureCitation>
,
<figureCitation id="13308FBC310A9F034445FB973D9EFBFB" box="[1373,1426,1047,1071]" captionStart="Figure 4" captionStartId="5.[129,194,1154,1178]" captionTargetBox="[164,1441,148,1126]" captionTargetId="figure-363@5.[161,1441,144,1126]" captionTargetPageId="5" captionText="Figure 4. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, SMNS 12596d, cervical vertebra exposed in less lateral view; B, interpretative drawing of (A); C, SMNS 12596e, posterior cervical or anterior dorsal vertebra exposed in an angled right anterolateral view; D, interpretative drawing of (C); E, SMNS 12596c, close-up of second sacral vertebra in right lateral view and osteoderm in dorsal view; F, interpretative drawing of (E); G, SMNS 12596f, proximal caudal vertebra block exposed in less lateral view; H, interpretative drawing of (G). Abbreviations: accs, accessory anterior spine; dp, diapophysis; izl, interzygapophyseal lamina; ncs, neurocentral suture; ns, neural spine; os, osteoderm; pap, parapophysis; pcdl, posterior centrodiapophyseal lamina; pofl, posterior flange; poz, postzygaphysis; prdl, prezygodiapophyseal lamina; prz, prezygapophysis; sr, sacral rib; tp, transverse process." figureDoi="http://doi.org/10.5281/zenodo.8426157" httpUri="https://zenodo.org/record/8426157/files/figure.png" pageId="23" pageNumber="377">4E, F</figureCitation>
,
<figureCitation id="13308FBC310A9F034486FB963B40FB98" captionStart="Figure 6" captionStartId="7.[129,194,1605,1629]" captionTargetBox="[161,1441,657,1577]" captionTargetId="figure-310@7.[161,1441,657,1577]" captionTargetPageId="7" captionText="Figure 6. Part of Saltoposuchus connectens referred specimen SMNS 55009. A, SMNS 55009a, dorsal vertebral centrum in less lateral view; B, SMNS 55009d, proximal caudal vertebral centrum in angled lateral view; C, SMNS 55009b, first sacral vertebra in less lateral view; D, interpretative drawing of (C); E, SMNS 55009c, proximal caudal vertebra in posterior view; F, interpretative drawing of (E). Abbreviations: car, caudal rib; ns, neural spine; pas, posterior articular surface; poz, postzygapophysis; prz, prezygapophysis; sr, sacral rib; sut, suture." figureDoi="http://doi.org/10.5281/zenodo.8426161" httpUri="https://zenodo.org/record/8426161/files/figure.png" pageId="23" pageNumber="377">6C, D</figureCitation>
). The sacral remains of SMNS 12597a are represented by the right sacral ribs exposed in dorsal view and are preserved in articulation with the right ilium (
<figureCitation id="13308FBC310A9F034591FBF43CE8FB58" box="[1161,1252,1140,1165]" captionStart="Figure 12" captionStartId="24.[113,178,1772,1796]" captionTargetBox="[151,1424,144,1744]" captionTargetId="figure-6@24.[148,1424,144,1744]" captionTargetPageId="24" captionText="Figure 12. Details of the postcranial anatomy of Saltoposuchus connectens. A, dorsal vertebral neural arch and dorsal rib of SMNS 12596c in ventral view; B, associated osteoderms of SMNS 12596c in dorsal view; C, sacral ribs and associated elements of SMNS 12597a in dorsal view; D, distal portion of right scapular blade of SMNS 12597a in lateral view; E, proximal end of right tibia of SMNS 12597a in proximal view; F, less tarsus and pes of SMNS 12597a with the calcaneum exposed in lateral view; G, close-up of less tarsus of SMNS 12597a; H, close-up of less pes of SMNS 12597a. Abbreviations: ane, anterior expansion; cal, calcaneum; calt, calcaneal tuber; cp, capitulum; dp, diapophysis; dt, distal tarsal; exp, extensor pit; fe, femur; fi, fibula; il, ilium; lam, lamina; latc, lateral condyle; le, lateral edge; lr, longitudinal ridge; me, medial edge; mt, metatarsal; no, notch; pap, parapophysis; ph, phalanx; pme, posterior extension; poe, posterior expansion; pofl, posterior flange; poz, postzygapophysis; os, osteoderm; r., right; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426175" httpUri="https://zenodo.org/record/8426175/files/figure.png" pageId="23" pageNumber="377">Fig. 12C</figureCitation>
).
<bibRefCitation id="EF9AEEC8310A9F0345E6FBF43D9DFB58" author="Crush PJ" box="[1278,1425,1140,1165]" pageId="23" pageNumber="377" refId="ref31157" refString="Crush PJ. An early, terrestrial, crocodile from South Wales. Unpublished Ph. D. Thesis, University College London, 1980." type="book" year="1980">Crush (1980)</bibRefCitation>
also identified a part of the less rib of the second sacral vertebra, but this element is not recognized here. The ribs of both vertebrae are strongly developed and anteroposteriorly broaden distally. The second sacral rib is broader than the first. The first sacral rib has concave anterior and posterior margins in dorsal view, with the former being most concave. This rib is most expanded anterodistally. Its dorsal surface is mostly transversely concave, which might be the result of compression, apart from a raised, flaưened platform on its anterodistal portion. The second sacral rib is composed of an elevated anterior half and a posterior half that is deflected posteroventrally. Both portions are separated by a proximodistally directed, thin ridge, and the dorsal surfaces of both portions are gently transversely concave. The anterior margin is concave, whereas the posterior margin is incomplete.
</paragraph>
<paragraph id="8BB49339310A9F0D424DF9CA3A71FE14" blockId="23.[825,1475,796,1979]" lastBlockId="25.[128,778,144,1014]" lastPageId="25" lastPageNumber="379" pageId="23" pageNumber="377">
The sacral vertebra of SMNS 12596c is a second sacral vertebra exposed in right lateral view (
<figureCitation id="13308FBC310A9F0345BCF9EA3D0AF954" box="[1188,1286,1642,1666]" captionStart="Figure 4" captionStartId="5.[129,194,1154,1178]" captionTargetBox="[164,1441,148,1126]" captionTargetId="figure-363@5.[161,1441,144,1126]" captionTargetPageId="5" captionText="Figure 4. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, SMNS 12596d, cervical vertebra exposed in less lateral view; B, interpretative drawing of (A); C, SMNS 12596e, posterior cervical or anterior dorsal vertebra exposed in an angled right anterolateral view; D, interpretative drawing of (C); E, SMNS 12596c, close-up of second sacral vertebra in right lateral view and osteoderm in dorsal view; F, interpretative drawing of (E); G, SMNS 12596f, proximal caudal vertebra block exposed in less lateral view; H, interpretative drawing of (G). Abbreviations: accs, accessory anterior spine; dp, diapophysis; izl, interzygapophyseal lamina; ncs, neurocentral suture; ns, neural spine; os, osteoderm; pap, parapophysis; pcdl, posterior centrodiapophyseal lamina; pofl, posterior flange; poz, postzygaphysis; prdl, prezygodiapophyseal lamina; prz, prezygapophysis; sr, sacral rib; tp, transverse process." figureDoi="http://doi.org/10.5281/zenodo.8426157" httpUri="https://zenodo.org/record/8426157/files/figure.png" pageId="23" pageNumber="377">Fig. 4E, F</figureCitation>
). The neural arch, including the neural spine and right pre- and postzygapophysis are preserved, as well as the right rib. The less rib of the same element can be observed on the other side of the slab (
<figureCitation id="13308FBC310A9F034484F9483B54F92A" captionStart="Figure 3" captionStartId="4.[113,178,1499,1523]" captionTargetBox="[147,1426,144,1471]" captionTargetId="figure-217@4.[146,1426,144,1471]" captionTargetPageId="4" captionText="Figure 3. Part of Saltoposuchus connectens referred specimen SMNS 12596. A, SMNS 12596a, cranial block; B, SMNS 12596b, two dentaries exposed in ventrolateral view; C, SMNS 12596c exposing a less femur in anterolateral view; D, exposing the opposite side of SMNS 12596c, Abbreviations:de, dentary; dr, dorsal rib; dv, dorsal vertebra; fe, femur; ga, gastralia; l., less; mx, maxilla; na, nasal; os, osteoderm; q, quadrate; r., right; sq, squamosal; sr, sacral rib; sv, sacral vertebra; tp, transverse process." figureDoi="http://doi.org/10.5281/zenodo.8426153" httpUri="https://zenodo.org/record/8426153/files/figure.png" pageId="23" pageNumber="377">Fig. 3C</figureCitation>
). The postzygapophysis is placed considerably further dorsally than the prezygapophysis. The neural spine is low and its lateral surface is distinctly transversely convex. An incomplete interdigitating suture is present between the transverse process and rib; it is placed far proximally, near the base of the neural spine. The right rib broadens distally and both the anterior and posterior margins are concave. The dorsal surface of the rib is transverse concave. Distally, the rib is strongly expanded posteriorly to form a dorsoventrally thin posterior flange, similar to that seen in
<taxonomicName id="4C0BE8BA31049F0D41D3FF4F39A8FF32" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[203,420,207,231]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="25" pageNumber="379" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B31049F0D41D3FF4F39A8FF32" box="[203,420,207,231]" italics="true" pageId="25" pageNumber="379">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0BE8BA31049F0D40B6FF4F3A4DFF33" baseAuthorityName="Marsh" baseAuthorityYear="1877" box="[430,577,207,231]" class="Reptilia" family="Coeluridae" genus="Hallopus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="25" pageNumber="379" phylum="Chordata" rank="species" species="victor">
<emphasis id="B97F4F2B31049F0D40B6FF4F3A4DFF33" box="[430,577,207,231]" italics="true" pageId="25" pageNumber="379">Hallopus victor</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C0BE8BA31049F0D436EFF4F38D9FED3" class="Reptilia" family="Sphenosuchidae" genus="Dromicosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="25" pageNumber="379" phylum="Chordata" rank="species" species="grallator">
<emphasis id="B97F4F2B31049F0D436EFF4F38D9FED3" italics="true" pageId="25" pageNumber="379">Dromicosuchus grallator</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9AEEC831049F0D41FDFF6E396AFED3" author="Walker AD" box="[229,358,238,262]" pageId="25" pageNumber="379" pagination="323 - 72" refId="ref32988" refString="Walker AD. A revision of the Jurassic reptile Hallopus victor (Marsh), with remarks on the classification of crocodiles. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1970; 257: 323 - 72." type="journal article" year="1970">Walker 1970</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831049F0D4069FF6E39E7FED3" author="Crush PJ" box="[369,491,238,262]" pageId="25" pageNumber="379" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831049F0D40EEFF6E3A9EFED3" author="Sues H-D &amp; Olsen PE &amp; Carter JG" box="[502,658,238,262]" pageId="25" pageNumber="379" pagination="329 - 43" refId="ref32828" refString="Sues H-D, Olsen PE, Carter JG et al. A new crocodylomorph archosaur from the Upper Triassic of North Carolina. Journal of Vertebrate Paleontology 2003; 23: 329 - 43." type="journal article" year="2003">
Sues
<emphasis id="B97F4F2B31049F0D4332FF6F3A55FED3" box="[554,601,238,262]" italics="true" pageId="25" pageNumber="379">et al.</emphasis>
2003
</bibRefCitation>
). However, a distinct groove on the lateral surface of the rib associated with the flange, as present in
<taxonomicName id="4C0BE8BA31049F0D406BFEAD3A42FE90" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[371,590,301,325]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="25" pageNumber="379" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B31049F0D406BFEAD3A42FE90" box="[371,590,301,325]" italics="true" pageId="25" pageNumber="379">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
, is absent. The lateral surface of the rib is rugose and, except for the thin posterior flange, it is dorsoventrally thickened with a transversely gently concave lateral surface. The dorsal side of the posterior flange is convex in lateral view, and its ventral side concave.
</paragraph>
<caption id="DF74C3B131059F0C4169F96C3BC1F81D" ID-DOI="http://doi.org/10.5281/zenodo.8426175" ID-Zenodo-Dep="8426175" httpUri="https://zenodo.org/record/8426175/files/figure.png" pageId="24" pageNumber="378" startId="24.[113,178,1772,1796]" targetBox="[151,1424,144,1744]" targetPageId="24" targetType="figure">
<paragraph id="8BB4933931059F0C4169F96C3BC1F81D" blockId="24.[113,1455,1772,1992]" pageId="24" pageNumber="378">
<emphasis id="B97F4F2B31059F0C4169F96C38DAF8D1" bold="true" box="[113,214,1772,1796]" pageId="24" pageNumber="378">Figure 12.</emphasis>
Details of the postcranial anatomy of
<taxonomicName id="4C0BE8BA31059F0C4321F96C3B17F8D1" authorityName="Huene" authorityYear="1921" box="[569,795,1772,1796]" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="24" pageNumber="378" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B31059F0C4321F96C3B17F8D1" box="[569,795,1772,1796]" italics="true" pageId="24" pageNumber="378">Saltoposuchus connectens</emphasis>
</taxonomicName>
. A, dorsal vertebral neural arch and dorsal rib of SMNS 12596c in ventral view; B, associated osteoderms of SMNS 12596c in dorsal view; C, sacral ribs and associated elements of SMNS 12597a in dorsal view; D, distal portion of right scapular blade of SMNS 12597a in lateral view; E, proximal end of right tibia of SMNS 12597a in proximal view; F, less tarsus and pes of SMNS 12597a with the calcaneum exposed in lateral view; G, close-up of less tarsus of SMNS 12597a; H, close-up of less pes of SMNS 12597a. Abbreviations: ane, anterior expansion; cal, calcaneum; calt, calcaneal tuber; cp, capitulum; dp, diapophysis; dt, distal tarsal; exp, extensor pit; fe, femur; fi, fibula; il, ilium; lam, lamina; latc, lateral condyle; le, lateral edge; lr, longitudinal ridge; me, medial edge; mt, metatarsal; no, notch; pap, parapophysis; ph, phalanx; pme, posterior extension; poe, posterior expansion; pofl, posterior flange; poz, postzygapophysis; os, osteoderm; r., right; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual.
</paragraph>
</caption>
<paragraph id="8BB4933931049F0D4184FE493A9EFC23" blockId="25.[128,778,144,1014]" pageId="25" pageNumber="379">
The sacral vertebra of SMNS 55009b is strongly deformed so that the less side of the specimen, including the less rib, is displaced ventrally and the right side dorsally (
<figureCitation id="13308FBC31049F0D431BFD883A63FDCA" box="[515,623,520,544]" captionStart="Figure 6" captionStartId="7.[129,194,1605,1629]" captionTargetBox="[161,1441,657,1577]" captionTargetId="figure-310@7.[161,1441,657,1577]" captionTargetPageId="7" captionText="Figure 6. Part of Saltoposuchus connectens referred specimen SMNS 55009. A, SMNS 55009a, dorsal vertebral centrum in less lateral view; B, SMNS 55009d, proximal caudal vertebral centrum in angled lateral view; C, SMNS 55009b, first sacral vertebra in less lateral view; D, interpretative drawing of (C); E, SMNS 55009c, proximal caudal vertebra in posterior view; F, interpretative drawing of (E). Abbreviations: car, caudal rib; ns, neural spine; pas, posterior articular surface; poz, postzygapophysis; prz, prezygapophysis; sr, sacral rib; sut, suture." figureDoi="http://doi.org/10.5281/zenodo.8426161" httpUri="https://zenodo.org/record/8426161/files/figure.png" pageId="25" pageNumber="379">Fig. 6C, D</figureCitation>
). The exposed anterior articular surface is clearly concave. The neural spine has a gradual transverse expansion distally but its lateral margin is flat to slightly concave, although it is unclear whether this is the result of compression. The postzygapophysis is again positioned considerably further dorsally than the prezygapophysis and an interdigitating suture is present between the rib and transverse process near the base of the neural spine. The rib is badly broken laterally and ventrally but clearly broadens distally in dorsal view. Its dorsal surface is mostly flat. The lateral side of the rib is incomplete and largely uninformative, but a thin posterior flange appears to be lacking; this element is therefore identified as a first sacral vertebra (
<emphasis id="B97F4F2B31049F0D404FFC013986FC4D" box="[343,394,897,920]" italics="true" pageId="25" pageNumber="379">sensu</emphasis>
<bibRefCitation id="EF9AEEC831049F0D4089FC003A00FC4D" author="Crush PJ" box="[401,524,896,920]" pageId="25" pageNumber="379" refId="ref31157" refString="Crush PJ. An early, terrestrial, crocodile from South Wales. Unpublished Ph. D. Thesis, University College London, 1980." type="book" year="1980">Crush 1980</bibRefCitation>
but
<emphasis id="B97F4F2B31049F0D4326FC013A77FC4D" box="[574,635,897,920]" italics="true" pageId="25" pageNumber="379">contra</emphasis>
<bibRefCitation id="EF9AEEC831049F0D439BFC003B08FC4D" author="Huene FR" box="[643,772,896,920]" pageId="25" pageNumber="379" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene 1921</bibRefCitation>
, who identified it as a second sacral vertebra). Due to its isolated nature and lack of diagnostic features, the assignment of this element to
<taxonomicName id="4C0BE8BA31049F0D41EAFC5E39EAFC20" authorityName="Huene" authorityYear="1921" box="[242,486,990,1014]" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="25" pageNumber="379" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B31049F0D41EAFC5E39EAFC20" box="[242,486,990,1014]" italics="true" pageId="25" pageNumber="379">Saltoposuchus connectens</emphasis>
</taxonomicName>
is only tentative.
</paragraph>
<paragraph id="8BB4933931049F0D4198FB973ABDF935" blockId="25.[128,778,1047,1980]" pageId="25" pageNumber="379">
<emphasis id="B97F4F2B31049F0D4198FB97398CFBFA" box="[128,384,1047,1071]" italics="true" pageId="25" pageNumber="379">Caudal vertebrae and ribs:</emphasis>
An articulated series of both the proximal and distal caudal vertebrae is preserved in SMNS 12597. The proximal series is preserved in SMNS 12597a and the distal series in SMNS 12597d (
<figureCitation id="13308FBC31049F0D409FFBF539E3FB58" box="[391,495,1141,1165]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="25" pageNumber="379">Fig. 2B, D</figureCitation>
). The slabs do not connect directly and, based on the presence of a well developed transverse process on the posteriormost preserved caudal vertebra of SMNS 12597a, and the complete absence of a transverse process on the anteriormost caudal vertebra on SMNS 12597d, a considerable part of the tail is missing, as was also suggested by
<bibRefCitation id="EF9AEEC831049F0D43DCFA9238C6FA9C" author="Huene FR" pageId="25" pageNumber="379" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene (1921)</bibRefCitation>
. A partially preserved, isolated caudal vertebra is present in SMNS 12596f. It is exposed in less lateral view, and based on the presence of a poorly developed transverse process, this represents approximately a vertebra from the middle of the caudal series (
<figureCitation id="13308FBC31049F0D41D1FA2E393CFA13" box="[201,304,1454,1478]" captionStart="Figure 4" captionStartId="5.[129,194,1154,1178]" captionTargetBox="[164,1441,148,1126]" captionTargetId="figure-363@5.[161,1441,144,1126]" captionTargetPageId="5" captionText="Figure 4. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, SMNS 12596d, cervical vertebra exposed in less lateral view; B, interpretative drawing of (A); C, SMNS 12596e, posterior cervical or anterior dorsal vertebra exposed in an angled right anterolateral view; D, interpretative drawing of (C); E, SMNS 12596c, close-up of second sacral vertebra in right lateral view and osteoderm in dorsal view; F, interpretative drawing of (E); G, SMNS 12596f, proximal caudal vertebra block exposed in less lateral view; H, interpretative drawing of (G). Abbreviations: accs, accessory anterior spine; dp, diapophysis; izl, interzygapophyseal lamina; ncs, neurocentral suture; ns, neural spine; os, osteoderm; pap, parapophysis; pcdl, posterior centrodiapophyseal lamina; pofl, posterior flange; poz, postzygaphysis; prdl, prezygodiapophyseal lamina; prz, prezygapophysis; sr, sacral rib; tp, transverse process." figureDoi="http://doi.org/10.5281/zenodo.8426157" httpUri="https://zenodo.org/record/8426157/files/figure.png" pageId="25" pageNumber="379">Fig. 4G, H</figureCitation>
). Two further, very poorly preserved, proximal caudal vertebrae are preserved in SMNS 12596h in association with a less ischium. They are exposed in right lateral view and the complete right side of the neural arch dorsal to the transverse process is visible in the proximal element, whereas the right prezygapophysis and a small part of the centrum is visible for the distal vertebra (
<figureCitation id="13308FBC31049F0D4055F9EA3995F957" box="[333,409,1642,1666]" captionStart="Figure 5" captionStartId="6.[113,178,1777,1801]" captionTargetBox="[221,1353,149,1749]" captionTargetId="figure-5@6.[220,1353,149,1749]" captionTargetPageId="6" captionText="Figure 5. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, B, SMNS 12596g, right pubis in (A) anterior and (B) angled medial views; C, SMNS 12596h, right ischium exposed in dorsomedial view and two proximal caudal vertebrae exposed in dorsal view; D, SMNS 12596i, tentative less ulna in posteromedial view; E, F, G, H, SMNS 12596j, right femur in (E) posteromedial, (F) anterolateral, (G) proximal, and (H) distal views; I, proximal half of the right femur in posterolateral view; J, K, SMNS 12596k, right tibia and fibula, (J) with the tibia in posteromedial and the fibula in lateral(?) view and (K) with the tibia in anterolateral and the fibula in medial(?) view. Abbreviations: alcr, anterolateral crest; amt, anteromedial tuber; car, caudal rib; cav, caudal vertebra; cr, crest; crtf, crista tibiofibularis; fi, fibula; gr, groove; ilf, ilium facet; iss, ischial shass; lc, lateral condyle; ltr, lesser trochanter; mc, medial condyle; no, notch; ns, neural spine; obf, obturator foramen; olp, olecranon process; os, osteoderm; poz, postzygapophysis; pr., proximal; prz, prezygapophysis; pus, pubic shass; put, pubic tubercle; r., right; ti, tibia; tr, trochanter." figureDoi="http://doi.org/10.5281/zenodo.8426159" httpUri="https://zenodo.org/record/8426159/files/figure.png" pageId="25" pageNumber="379">Fig. 5C</figureCitation>
). SMNS 55009d represents an isolated centrum of a proximal caudal vertebra and SMNS 55009c represents a complete and isolated, proximal caudal vertebra including ribs exposed in posterior view (
<figureCitation id="13308FBC31049F0D433EF9483AACF935" box="[550,672,1736,1760]" captionStart="Figure 6" captionStartId="7.[129,194,1605,1629]" captionTargetBox="[161,1441,657,1577]" captionTargetId="figure-310@7.[161,1441,657,1577]" captionTargetPageId="7" captionText="Figure 6. Part of Saltoposuchus connectens referred specimen SMNS 55009. A, SMNS 55009a, dorsal vertebral centrum in less lateral view; B, SMNS 55009d, proximal caudal vertebral centrum in angled lateral view; C, SMNS 55009b, first sacral vertebra in less lateral view; D, interpretative drawing of (C); E, SMNS 55009c, proximal caudal vertebra in posterior view; F, interpretative drawing of (E). Abbreviations: car, caudal rib; ns, neural spine; pas, posterior articular surface; poz, postzygapophysis; prz, prezygapophysis; sr, sacral rib; sut, suture." figureDoi="http://doi.org/10.5281/zenodo.8426161" httpUri="https://zenodo.org/record/8426161/files/figure.png" pageId="25" pageNumber="379">Fig. 6B, E, F</figureCitation>
).
</paragraph>
<paragraph id="8BB4933931049F0D4184F9673BAFFDF5" blockId="25.[128,778,1047,1980]" lastBlockId="25.[825,1475,143,1640]" pageId="25" pageNumber="379">
Nine articulated, proximal caudal vertebrae are preserved on SMNS 12597a, whereas 15 distal caudal vertebrae are preserved in SMNS 12597d (
<figureCitation id="13308FBC31049F0D4097F8A639FAF8EB" box="[399,502,1830,1854]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="25" pageNumber="379">Fig. 2B, D</figureCitation>
). The end of the tail is not preserved, but the diminutive size of the posteriormost caudal vertebrae indicates that they are positioned very close to the terminus. Based on the separation between the sacrum and proximalmost exposed caudal vertebra in SMNS 12597a, as well as the presence of a chevron anterior to the anteriormost exposed caudal vertebra, at least one but possibly a few caudal vertebrae are still imbedded in matrix, as was also suggested by
<bibRefCitation id="EF9AEEC831049F0D4221FF6E3BDEFED3" author="Huene FR" box="[825,978,238,262]" pageId="25" pageNumber="379" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene (1921)</bibRefCitation>
.
<bibRefCitation id="EF9AEEC831049F0D42F9FF6E3C63FED3" author="Huene FR" box="[993,1135,238,262]" pageId="25" pageNumber="379" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene (1921</bibRefCitation>
: fig. 31) estimated that the gap between both blocks comprised at least 15 vertebrae and reconstructed
<taxonomicName id="4C0BE8BA31049F0D42A3FEAC3CBEFE91" authorityName="Huene" authorityYear="1921" box="[955,1202,300,324]" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="25" pageNumber="379" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B31049F0D42A3FEAC3CBEFE91" box="[955,1202,300,324]" italics="true" pageId="25" pageNumber="379">Saltoposuchus connectens</emphasis>
</taxonomicName>
with 53 caudal vertebrae, which is very likely an exaggeration.
<taxonomicName id="4C0BE8BA31049F0D45A7FECB3DA5FEB6" baseAuthorityName="Brown" baseAuthorityYear="1933" box="[1215,1449,331,355]" class="Reptilia" family="Protosuchidae" genus="Protosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="25" pageNumber="379" phylum="Chordata" rank="species" species="richardsoni">
<emphasis id="B97F4F2B31049F0D45A7FECB3DA5FEB6" box="[1215,1449,331,355]" italics="true" pageId="25" pageNumber="379">Protosuchus richardsoni</emphasis>
</taxonomicName>
is the only early crocodylomorph for which the caudal vertebral count is known, possessing around 35 caudal vertebrae (
<bibRefCitation id="EF9AEEC831049F0D446AFE0A3BD0FE14" author="Colbert EH &amp; Mook CC" pageId="25" pageNumber="379" pagination="143 - 82" refId="ref31130" refString="Colbert EH, Mook CC. The ancestral crocodilian Protosuchus. Bulletin of the American Museum of Natural History 1951; 97: 143 - 82." type="journal article" year="1951">Colbert and Mook 1951</bibRefCitation>
), whilst in modern crocodylians the caudal vertebral count is 30 to 40 (Hoffsteưer and Gasc 1969). It seems likely that the count of
<taxonomicName id="4C0BE8BA31049F0D4530FE683D10FDD5" authorityName="Huene" authorityYear="1921" box="[1064,1308,488,512]" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="25" pageNumber="379" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B31049F0D4530FE683D10FDD5" box="[1064,1308,488,512]" italics="true" pageId="25" pageNumber="379">Saltoposuchus connectens</emphasis>
</taxonomicName>
also falls within this range.
</paragraph>
<paragraph id="8BB4933931049F0D424DFDA73BD1FB86" blockId="25.[825,1475,143,1640]" pageId="25" pageNumber="379">
The neurocentral suture is unfused in both the proximal and distal caudal vertebrae of SMNS 12597, suggesting this specimen is likely immature. The suture between the transverse process and caudal rib is visible in some but not all proximal caudal vertebrae. The ventral margin of all caudal centra is very gently concave. The posterior articular surface of SMNS 55009c is slightly concave, whereas the deeply excavated articular faces of SMNS 55009d are likely exaggerated due to compression (
<figureCitation id="13308FBC31049F0D444EFC823B4BFCEC" captionStart="Figure 6" captionStartId="7.[129,194,1605,1629]" captionTargetBox="[161,1441,657,1577]" captionTargetId="figure-310@7.[161,1441,657,1577]" captionTargetPageId="7" captionText="Figure 6. Part of Saltoposuchus connectens referred specimen SMNS 55009. A, SMNS 55009a, dorsal vertebral centrum in less lateral view; B, SMNS 55009d, proximal caudal vertebral centrum in angled lateral view; C, SMNS 55009b, first sacral vertebra in less lateral view; D, interpretative drawing of (C); E, SMNS 55009c, proximal caudal vertebra in posterior view; F, interpretative drawing of (E). Abbreviations: car, caudal rib; ns, neural spine; pas, posterior articular surface; poz, postzygapophysis; prz, prezygapophysis; sr, sacral rib; sut, suture." figureDoi="http://doi.org/10.5281/zenodo.8426161" httpUri="https://zenodo.org/record/8426161/files/figure.png" pageId="25" pageNumber="379">Fig. 6B, E, F</figureCitation>
). The relative elongation of the caudal centra increases considerably posteriorly. The transverse processes and ribs are oriented perpendicular to the axis of the vertebral column. They do not extend particularly far laterally and have a rounded to rectangular distal end. Chevrons articulate between the ventral edges of the caudal vertebrae (
<figureCitation id="13308FBC31049F0D42E9FC3E3C5AFC00" box="[1009,1110,958,982]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="25" pageNumber="379">Fig. 2B, D</figureCitation>
). They are directed strongly posteriorly and only slightly ventrally, and have a posterodorsal curvature in lateral view. They occur along the entire tail, including the distalmost vertebrae, but reduce in length gradually posteriorly along the series.
</paragraph>
<paragraph id="8BB4933931049F0D424DFBDA3C37F9BD" blockId="25.[825,1475,143,1640]" pageId="25" pageNumber="379">
The neural spine of the isolated mid-caudal vertebra SMNS 12596f is slightly directed anterodorsally in lateral view (
<figureCitation id="13308FBC31049F0D446AFBFA3B41FB64" captionStart="Figure 4" captionStartId="5.[129,194,1154,1178]" captionTargetBox="[164,1441,148,1126]" captionTargetId="figure-363@5.[161,1441,144,1126]" captionTargetPageId="5" captionText="Figure 4. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, SMNS 12596d, cervical vertebra exposed in less lateral view; B, interpretative drawing of (A); C, SMNS 12596e, posterior cervical or anterior dorsal vertebra exposed in an angled right anterolateral view; D, interpretative drawing of (C); E, SMNS 12596c, close-up of second sacral vertebra in right lateral view and osteoderm in dorsal view; F, interpretative drawing of (E); G, SMNS 12596f, proximal caudal vertebra block exposed in less lateral view; H, interpretative drawing of (G). Abbreviations: accs, accessory anterior spine; dp, diapophysis; izl, interzygapophyseal lamina; ncs, neurocentral suture; ns, neural spine; os, osteoderm; pap, parapophysis; pcdl, posterior centrodiapophyseal lamina; pofl, posterior flange; poz, postzygaphysis; prdl, prezygodiapophyseal lamina; prz, prezygapophysis; sr, sacral rib; tp, transverse process." figureDoi="http://doi.org/10.5281/zenodo.8426157" httpUri="https://zenodo.org/record/8426157/files/figure.png" pageId="25" pageNumber="379">Fig. 4G, H</figureCitation>
). Anterior to the neural spine, an additional, sharply pointed, anterodorsally directed process is present, which is separated from the spine by a gap that is widely rounded in lateral view. This is the accessory anterior spine (
<emphasis id="B97F4F2B31049F0D45AEFB783CE5FADA" box="[1206,1257,1272,1295]" italics="true" pageId="25" pageNumber="379">sensu</emphasis>
<bibRefCitation id="EF9AEEC831049F0D45E8FB773D68FADA" author="Clark JM" box="[1264,1380,1271,1295]" pageId="25" pageNumber="379" refId="ref30947" refString="Clark JM. Phylogenetic relationships of the crocodylomorphs archosaurs. Unpublished Ph. D. Thesis, University of Chicago, 1986." type="book" year="1986">Clark 1986</bibRefCitation>
: ch. 14), which is also present in
<taxonomicName id="4C0BE8BA31049F0D4559FA963D13FAFB" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[1089,1311,1302,1326]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="25" pageNumber="379" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B31049F0D4559FA963D13FAFB" box="[1089,1311,1302,1326]" italics="true" pageId="25" pageNumber="379">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
(NHMUK PV R 7561) but absent in
<taxonomicName id="4C0BE8BA31049F0D4531FAB63CCCFA98" baseAuthorityName="Marsh" baseAuthorityYear="1877" box="[1065,1216,1334,1358]" class="Reptilia" family="Coeluridae" genus="Hallopus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="25" pageNumber="379" phylum="Chordata" rank="species" species="victor">
<emphasis id="B97F4F2B31049F0D4531FAB63CCCFA98" box="[1065,1216,1334,1358]" italics="true" pageId="25" pageNumber="379">Hallopus victor</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0BE8BA31049F0D45EEFAB63DCEFA9B" baseAuthorityName="Walker" baseAuthorityYear="1990" box="[1270,1474,1334,1358]" class="Reptilia" genus="Sphenosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="25" pageNumber="379" phylum="Chordata" rank="species" species="acutus">
<emphasis id="B97F4F2B31049F0D45EEFAB63DCEFA9B" box="[1270,1474,1334,1358]" italics="true" pageId="25" pageNumber="379">Sphenosuchus acutus</emphasis>
</taxonomicName>
among non-crocodyliform crocodylomorphs (
<bibRefCitation id="EF9AEEC831049F0D4429FAD53DB1FAB8" author="Walker AD" box="[1329,1469,1365,1389]" pageId="25" pageNumber="379" pagination="323 - 72" refId="ref32988" refString="Walker AD. A revision of the Jurassic reptile Hallopus victor (Marsh), with remarks on the classification of crocodiles. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1970; 257: 323 - 72." type="journal article" year="1970">Walker 1970</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831049F0D4221FAF53B61FA58" author="Walker AD" box="[825,877,1397,1421]" pageId="25" pageNumber="379" pagination="1 - 120" refId="ref33031" refString="Walker AD. A revision of Sphenosuchus acutus Haughton, a crocodylomorph reptile from the Elliot Formation (Late Triassic or Early Jurassic) of South Africa. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1990; 330: 1 - 120." type="journal article" year="1990">1990</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831049F0D4264FAF43BF4FA58" author="Crush PJ" box="[892,1016,1396,1421]" pageId="25" pageNumber="379" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
). Neural spines are also present in the two caudal vertebrae of SMNS 12596h, but these are poorly preserved and incomplete anteriorly, and only the base of the accessory anterior spine is preserved (
<figureCitation id="13308FBC31049F0D45BDFA523CE2FA3F" box="[1189,1262,1490,1514]" captionStart="Figure 5" captionStartId="6.[113,178,1777,1801]" captionTargetBox="[221,1353,149,1749]" captionTargetId="figure-5@6.[220,1353,149,1749]" captionTargetPageId="6" captionText="Figure 5. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, B, SMNS 12596g, right pubis in (A) anterior and (B) angled medial views; C, SMNS 12596h, right ischium exposed in dorsomedial view and two proximal caudal vertebrae exposed in dorsal view; D, SMNS 12596i, tentative less ulna in posteromedial view; E, F, G, H, SMNS 12596j, right femur in (E) posteromedial, (F) anterolateral, (G) proximal, and (H) distal views; I, proximal half of the right femur in posterolateral view; J, K, SMNS 12596k, right tibia and fibula, (J) with the tibia in posteromedial and the fibula in lateral(?) view and (K) with the tibia in anterolateral and the fibula in medial(?) view. Abbreviations: alcr, anterolateral crest; amt, anteromedial tuber; car, caudal rib; cav, caudal vertebra; cr, crest; crtf, crista tibiofibularis; fi, fibula; gr, groove; ilf, ilium facet; iss, ischial shass; lc, lateral condyle; ltr, lesser trochanter; mc, medial condyle; no, notch; ns, neural spine; obf, obturator foramen; olp, olecranon process; os, osteoderm; poz, postzygapophysis; pr., proximal; prz, prezygapophysis; pus, pubic shass; put, pubic tubercle; r., right; ti, tibia; tr, trochanter." figureDoi="http://doi.org/10.5281/zenodo.8426159" httpUri="https://zenodo.org/record/8426159/files/figure.png" pageId="25" pageNumber="379">Fig. 5C</figureCitation>
). In the distal caudal vertebrae, transverse processes are absent and the neural spines are very strongly reduced, but their morphology cannot be observed because they are covered dorsally by osteoderms in SMNS 12597e (
<figureCitation id="13308FBC31049F0D42F8F9D03C26F9BD" box="[992,1066,1616,1640]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="25" pageNumber="379">Fig. 2C</figureCitation>
).
</paragraph>
<paragraph id="8BB4933931049F0E4221F9093964FDD4" blockId="25.[825,1475,1673,1979]" lastBlockId="26.[113,763,144,1171]" lastPageId="26" lastPageNumber="380" pageId="25" pageNumber="379">
<emphasis id="B97F4F2B31049F0D4221F9093BBEF974" box="[825,946,1673,1697]" italics="true" pageId="25" pageNumber="379">Osteoderms:</emphasis>
Many osteoderms are preserved in SMNS 12597a, d, e and SMNS 12596c (
<figureCitation id="13308FBC31049F0D452EF9283CD9F915" box="[1078,1237,1704,1728]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="25" pageNumber="379">Figs 2A, B, D, E</figureCitation>
,
<figureCitation id="13308FBC31049F0D45F9F9283D11F915" box="[1249,1309,1704,1728]" captionStart="Figure 3" captionStartId="4.[113,178,1499,1523]" captionTargetBox="[147,1426,144,1471]" captionTargetId="figure-217@4.[146,1426,144,1471]" captionTargetPageId="4" captionText="Figure 3. Part of Saltoposuchus connectens referred specimen SMNS 12596. A, SMNS 12596a, cranial block; B, SMNS 12596b, two dentaries exposed in ventrolateral view; C, SMNS 12596c exposing a less femur in anterolateral view; D, exposing the opposite side of SMNS 12596c, Abbreviations:de, dentary; dr, dorsal rib; dv, dorsal vertebra; fe, femur; ga, gastralia; l., less; mx, maxilla; na, nasal; os, osteoderm; q, quadrate; r., right; sq, squamosal; sr, sacral rib; sv, sacral vertebra; tp, transverse process." figureDoi="http://doi.org/10.5281/zenodo.8426153" httpUri="https://zenodo.org/record/8426153/files/figure.png" pageId="25" pageNumber="379">3C, D</figureCitation>
,
<figureCitation id="13308FBC31049F0D4430F9283D50F915" box="[1320,1372,1704,1728]" captionStart="Figure 4" captionStartId="5.[129,194,1154,1178]" captionTargetBox="[164,1441,148,1126]" captionTargetId="figure-363@5.[161,1441,144,1126]" captionTargetPageId="5" captionText="Figure 4. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, SMNS 12596d, cervical vertebra exposed in less lateral view; B, interpretative drawing of (A); C, SMNS 12596e, posterior cervical or anterior dorsal vertebra exposed in an angled right anterolateral view; D, interpretative drawing of (C); E, SMNS 12596c, close-up of second sacral vertebra in right lateral view and osteoderm in dorsal view; F, interpretative drawing of (E); G, SMNS 12596f, proximal caudal vertebra block exposed in less lateral view; H, interpretative drawing of (G). Abbreviations: accs, accessory anterior spine; dp, diapophysis; izl, interzygapophyseal lamina; ncs, neurocentral suture; ns, neural spine; os, osteoderm; pap, parapophysis; pcdl, posterior centrodiapophyseal lamina; pofl, posterior flange; poz, postzygaphysis; prdl, prezygodiapophyseal lamina; prz, prezygapophysis; sr, sacral rib; tp, transverse process." figureDoi="http://doi.org/10.5281/zenodo.8426157" httpUri="https://zenodo.org/record/8426157/files/figure.png" pageId="25" pageNumber="379">4E, F</figureCitation>
,
<figureCitation id="13308FBC31049F0D447FF9283DBEF915" box="[1383,1458,1704,1728]" captionStart="Figure 12" captionStartId="24.[113,178,1772,1796]" captionTargetBox="[151,1424,144,1744]" captionTargetId="figure-6@24.[148,1424,144,1744]" captionTargetPageId="24" captionText="Figure 12. Details of the postcranial anatomy of Saltoposuchus connectens. A, dorsal vertebral neural arch and dorsal rib of SMNS 12596c in ventral view; B, associated osteoderms of SMNS 12596c in dorsal view; C, sacral ribs and associated elements of SMNS 12597a in dorsal view; D, distal portion of right scapular blade of SMNS 12597a in lateral view; E, proximal end of right tibia of SMNS 12597a in proximal view; F, less tarsus and pes of SMNS 12597a with the calcaneum exposed in lateral view; G, close-up of less tarsus of SMNS 12597a; H, close-up of less pes of SMNS 12597a. Abbreviations: ane, anterior expansion; cal, calcaneum; calt, calcaneal tuber; cp, capitulum; dp, diapophysis; dt, distal tarsal; exp, extensor pit; fe, femur; fi, fibula; il, ilium; lam, lamina; latc, lateral condyle; le, lateral edge; lr, longitudinal ridge; me, medial edge; mt, metatarsal; no, notch; pap, parapophysis; ph, phalanx; pme, posterior extension; poe, posterior expansion; pofl, posterior flange; poz, postzygapophysis; os, osteoderm; r., right; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426175" httpUri="https://zenodo.org/record/8426175/files/figure.png" pageId="25" pageNumber="379">12A–C</figureCitation>
). In SMNS 12597, they form a paired row across both the dorsal and caudal vertebral column. Only the osteoderms on the less side of the dorsal column are well preserved, but slivers of the osteoderms of the right row can also be discerned (
<figureCitation id="13308FBC31049F0D4459F8A63D86F8EB" box="[1345,1418,1830,1854]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="25" pageNumber="379">Fig. 2A</figureCitation>
). On the tail the osteoderms reach until the posteriormost preserved vertebra (
<figureCitation id="13308FBC31049F0D42BBF8E43BFEF8A9" box="[931,1010,1892,1916]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="25" pageNumber="379">Fig. 2D</figureCitation>
). It is unclear whether the osteoderms are also paired here.
<bibRefCitation id="EF9AEEC831049F0D42F1F8043C8CF849" author="Huene FR" box="[1001,1152,1924,1948]" pageId="25" pageNumber="379" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene (1921)</bibRefCitation>
identified a tiny, lancet-shaped osteoderm associated with the two cervical vertebrae of SMNS 12597c. In contrast to
<bibRefCitation id="EF9AEEC831079F0E4077FF103A09FF7D" author="Crush PJ" box="[367,517,144,168]" pageId="26" pageNumber="380" refId="ref31157" refString="Crush PJ. An early, terrestrial, crocodile from South Wales. Unpublished Ph. D. Thesis, University College London, 1980." type="book" year="1980">Crush (1980)</bibRefCitation>
, I have recovered this element, but I do not agree with its identification. It most likely represents part of the transverse process of a vertebra. Thus, no cervical osteoderms are known for
<taxonomicName id="4C0BE8BA31079F0E40C0FF6E3AC0FED3" authorityName="Huene" authorityYear="1921" box="[472,716,238,262]" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="26" pageNumber="380" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B31079F0E40C0FF6E3AC0FED3" box="[472,716,238,262]" italics="true" pageId="26" pageNumber="380">Saltoposuchus connectens</emphasis>
</taxonomicName>
, but they were almost certainly present, as in other crocodylomorphs. Appendicular and ventral osteoderms are absent, as in all other known non-crocodyliform crocodylomorphs. The osteoderms differ in size along the vertebral column but maintain a similar morphology. The osteoderms along the dorsal vertebral series of SMNS 12597a gradually increase in width and level of ornamentation posteriorly, whereas on the tail they decrease in size posteriorly (
<figureCitation id="13308FBC31079F0E41E8FE693954FDD5" box="[240,344,489,513]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="26" pageNumber="380">Fig. 2A, D</figureCitation>
).
</paragraph>
<paragraph id="8BB4933931079F0E4195FD883AACFB47" blockId="26.[113,763,144,1171]" pageId="26" pageNumber="380">
The dorsal surface of the osteoderms is heavily sculptured by pits and ridges and, additionally, possesses a very low and narrow longitudinal ridge that is placed slightly medial to the transverse midpoint of the osteoderm (
<figureCitation id="13308FBC31079F0E430EFDE63A8EFDA8" box="[534,642,614,638]" captionStart="Figure 4" captionStartId="5.[129,194,1154,1178]" captionTargetBox="[164,1441,148,1126]" captionTargetId="figure-363@5.[161,1441,144,1126]" captionTargetPageId="5" captionText="Figure 4. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, SMNS 12596d, cervical vertebra exposed in less lateral view; B, interpretative drawing of (A); C, SMNS 12596e, posterior cervical or anterior dorsal vertebra exposed in an angled right anterolateral view; D, interpretative drawing of (C); E, SMNS 12596c, close-up of second sacral vertebra in right lateral view and osteoderm in dorsal view; F, interpretative drawing of (E); G, SMNS 12596f, proximal caudal vertebra block exposed in less lateral view; H, interpretative drawing of (G). Abbreviations: accs, accessory anterior spine; dp, diapophysis; izl, interzygapophyseal lamina; ncs, neurocentral suture; ns, neural spine; os, osteoderm; pap, parapophysis; pcdl, posterior centrodiapophyseal lamina; pofl, posterior flange; poz, postzygaphysis; prdl, prezygodiapophyseal lamina; prz, prezygapophysis; sr, sacral rib; tp, transverse process." figureDoi="http://doi.org/10.5281/zenodo.8426157" httpUri="https://zenodo.org/record/8426157/files/figure.png" pageId="26" pageNumber="380">Figs 4E, F</figureCitation>
,
<figureCitation id="13308FBC31079F0E4388FDE63AB5FDAB" box="[656,697,614,638]" captionStart="Figure 12" captionStartId="24.[113,178,1772,1796]" captionTargetBox="[151,1424,144,1744]" captionTargetId="figure-6@24.[148,1424,144,1744]" captionTargetPageId="24" captionText="Figure 12. Details of the postcranial anatomy of Saltoposuchus connectens. A, dorsal vertebral neural arch and dorsal rib of SMNS 12596c in ventral view; B, associated osteoderms of SMNS 12596c in dorsal view; C, sacral ribs and associated elements of SMNS 12597a in dorsal view; D, distal portion of right scapular blade of SMNS 12597a in lateral view; E, proximal end of right tibia of SMNS 12597a in proximal view; F, less tarsus and pes of SMNS 12597a with the calcaneum exposed in lateral view; G, close-up of less tarsus of SMNS 12597a; H, close-up of less pes of SMNS 12597a. Abbreviations: ane, anterior expansion; cal, calcaneum; calt, calcaneal tuber; cp, capitulum; dp, diapophysis; dt, distal tarsal; exp, extensor pit; fe, femur; fi, fibula; il, ilium; lam, lamina; latc, lateral condyle; le, lateral edge; lr, longitudinal ridge; me, medial edge; mt, metatarsal; no, notch; pap, parapophysis; ph, phalanx; pme, posterior extension; poe, posterior expansion; pofl, posterior flange; poz, postzygapophysis; os, osteoderm; r., right; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426175" httpUri="https://zenodo.org/record/8426175/files/figure.png" pageId="26" pageNumber="380">12B</figureCitation>
). The osteoderms are subrectangular to oval in outline in dorsal view, being considerably longer anteroposteriorly than mediolaterally wide. The medial margin, which articulated with its antimere, is straight, whereas the lateral margin is convex in dorsal view. The osteoderms of
<taxonomicName id="4C0BE8BA31079F0E4021FC823A22FCCF" authorityName="Huene" authorityYear="1921" box="[313,558,770,794]" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="26" pageNumber="380" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B31079F0E4021FC823A22FCCF" box="[313,558,770,794]" italics="true" pageId="26" pageNumber="380">Saltoposuchus connectens</emphasis>
</taxonomicName>
are similar to those seen in
<taxonomicName id="4C0BE8BA31079F0E41D2FCA23A81FCEF" authority="(Sues et al. 2003)" baseAuthorityName="Sues" baseAuthorityYear="2003" box="[202,653,802,826]" class="Reptilia" family="Sphenosuchidae" genus="Dromicosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="26" pageNumber="380" phylum="Chordata" rank="species" species="grallator">
<emphasis id="B97F4F2B31079F0E41D2FCA239B2FCEF" box="[202,446,802,826]" italics="true" pageId="26" pageNumber="380">Dromicosuchus grallator</emphasis>
(
<bibRefCitation id="EF9AEEC831079F0E40CBFCA23A8EFCEF" author="Sues H-D &amp; Olsen PE &amp; Carter JG" box="[467,642,802,826]" pageId="26" pageNumber="380" pagination="329 - 43" refId="ref32828" refString="Sues H-D, Olsen PE, Carter JG et al. A new crocodylomorph archosaur from the Upper Triassic of North Carolina. Journal of Vertebrate Paleontology 2003; 23: 329 - 43." type="journal article" year="2003">
Sues
<emphasis id="B97F4F2B31079F0E4316FCA33A4FFCEF" box="[526,579,802,826]" italics="true" pageId="26" pageNumber="380">et al.</emphasis>
2003
</bibRefCitation>
)
</taxonomicName>
, relatively much larger and much more sculpted than the osteoderms of
<taxonomicName id="4C0BE8BA31079F0E4169FCE039E1FCAC" authority="(Crush 1984)" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[113,493,864,889]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="26" pageNumber="380" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B31079F0E4169FCE0395CFCAD" box="[113,336,864,888]" italics="true" pageId="26" pageNumber="380">Terrestrisuchus gracilis</emphasis>
(
<bibRefCitation id="EF9AEEC831079F0E407CFCE039EEFCAC" author="Crush PJ" box="[356,482,864,889]" pageId="26" pageNumber="380" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
)
</taxonomicName>
, and less rectangular but rather more oval-shaped than the osteoderms of
<taxonomicName id="4C0BE8BA31079F0E4375FC0038AFFC62" class="Reptilia" family="Sphenosuchidae" genus="Hesperosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="26" pageNumber="380" phylum="Chordata" rank="species" species="agilis">
<emphasis id="B97F4F2B31079F0E4375FC0038AFFC62" italics="true" pageId="26" pageNumber="380">Hesperosuchus agilis</emphasis>
</taxonomicName>
(AMNH FARB 6758) and CM 29894 (
<bibRefCitation id="EF9AEEC831079F0E437EFC1F3AF9FC62" author="Colbert EH" box="[614,757,927,951]" pageId="26" pageNumber="380" pagination="561 - 92" refId="ref31083" refString="Colbert EH. A pseudosuchian reptile from Arizona. Bulletin of the American Museum of Natural History 1952; 99: 561 - 92." type="journal article" year="1952">Colbert 1952</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831079F0E4169FC3E3916FC02" author="Clark JM &amp; Sues H-D &amp; Berman DS" box="[113,282,958,983]" pageId="26" pageNumber="380" pagination="683 - 704" refId="ref31006" refString="Clark JM, Sues H-D, Berman DS. A new specimen of Hesperosuchus agilis from the Upper Triassic of New Mexico and the interrelationships of basal crocodylomorph archosaurs. Journal of Vertebrate Paleontology 2000; 20: 683 - 704." type="journal article" year="2000">
Clark
<emphasis id="B97F4F2B31079F0E41A8FC3F38EDFC03" box="[176,225,958,982]" italics="true" pageId="26" pageNumber="380">et al.</emphasis>
2000
</bibRefCitation>
). They possess a short, anteriorly directed peg on their anterior margin as seen in disarticulated osteoderms of SMNS 12596c. The ventral surface, exposed in an osteoderm directly adjacent to the less transverse process of the sacral vertebra in SMNS 12596c, is smooth and gently transversely concave and its posterior margin possesses a distinct notch that would have received the anterior peg of the subsequent osteoderm.
</paragraph>
<paragraph id="8BB4933931079F0E4169FB3439C7F86E" blockId="26.[113,763,1204,1979]" pageId="26" pageNumber="380">
<emphasis id="B97F4F2B31079F0E4169FB3438C5FB19" box="[113,201,1204,1228]" italics="true" pageId="26" pageNumber="380">Scapula:</emphasis>
The distal portion of a right scapular blade is preserved in SMNS 12597a (
<figureCitation id="13308FBC31079F0E4096FB5339ECFB3E" box="[398,480,1235,1259]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="26" pageNumber="380">Figs 2A</figureCitation>
,
<figureCitation id="13308FBC31079F0E40E8FB533A12FB3E" box="[496,542,1235,1259]" captionStart="Figure 12" captionStartId="24.[113,178,1772,1796]" captionTargetBox="[151,1424,144,1744]" captionTargetId="figure-6@24.[148,1424,144,1744]" captionTargetPageId="24" captionText="Figure 12. Details of the postcranial anatomy of Saltoposuchus connectens. A, dorsal vertebral neural arch and dorsal rib of SMNS 12596c in ventral view; B, associated osteoderms of SMNS 12596c in dorsal view; C, sacral ribs and associated elements of SMNS 12597a in dorsal view; D, distal portion of right scapular blade of SMNS 12597a in lateral view; E, proximal end of right tibia of SMNS 12597a in proximal view; F, less tarsus and pes of SMNS 12597a with the calcaneum exposed in lateral view; G, close-up of less tarsus of SMNS 12597a; H, close-up of less pes of SMNS 12597a. Abbreviations: ane, anterior expansion; cal, calcaneum; calt, calcaneal tuber; cp, capitulum; dp, diapophysis; dt, distal tarsal; exp, extensor pit; fe, femur; fi, fibula; il, ilium; lam, lamina; latc, lateral condyle; le, lateral edge; lr, longitudinal ridge; me, medial edge; mt, metatarsal; no, notch; pap, parapophysis; ph, phalanx; pme, posterior extension; poe, posterior expansion; pofl, posterior flange; poz, postzygapophysis; os, osteoderm; r., right; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426175" httpUri="https://zenodo.org/record/8426175/files/figure.png" pageId="26" pageNumber="380">12D</figureCitation>
). Its exposed lateral surface is smooth and flaưened to slightly transversely convex. The distal end is expanded both anteriorly and posteriorly. The posterior expansion is more gradual and positioned further dorsally than the anterior expansion and, consequently, the anterior margin is more strongly concave than the posterior one. Both expansions terminate in an acute tip, which is sharpest in the posterior expansion. The outer surface of the dorsal margin is incomplete, which according to
<bibRefCitation id="EF9AEEC831079F0E4370FA4E3AF6FA33" author="Crush PJ" box="[616,762,1486,1510]" pageId="26" pageNumber="380" refId="ref31157" refString="Crush PJ. An early, terrestrial, crocodile from South Wales. Unpublished Ph. D. Thesis, University College London, 1980." type="book" year="1980">Crush (1980)</bibRefCitation>
represents an area of partial ossification. Regardless, its outline is clearly preserved. It is distinctly convex in lateral view and from its apex it gradually curves posteroventrally towards the tip of the posterior expansion and anteroventrally under a stronger inclination towards the tip of the anterior expansion. The outline of the distal end of the scapula is very similar to that of
<taxonomicName id="4C0BE8BA31079F0E41A3F9293990F914" baseAuthorityName="Crush" baseAuthorityYear="1984" box="[187,412,1705,1729]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="26" pageNumber="380" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B31079F0E41A3F9293990F914" box="[187,412,1705,1729]" italics="true" pageId="26" pageNumber="380">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0BE8BA31079F0E40D6F9293A9BF914" box="[462,663,1705,1729]" class="Reptilia" family="Sphenosuchidae" genus="Hesperosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="26" pageNumber="380" phylum="Chordata" rank="species" species="agilis">
<emphasis id="B97F4F2B31079F0E40D6F9293A9BF914" box="[462,663,1705,1729]" italics="true" pageId="26" pageNumber="380">Hesperosuchus agilis</emphasis>
</taxonomicName>
(AMNH FARB 6758,
<bibRefCitation id="EF9AEEC831079F0E41E1F948398AF935" author="Colbert EH" box="[249,390,1736,1760]" pageId="26" pageNumber="380" pagination="561 - 92" refId="ref31083" refString="Colbert EH. A pseudosuchian reptile from Arizona. Bulletin of the American Museum of Natural History 1952; 99: 561 - 92." type="journal article" year="1952">Colbert 1952</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831079F0E408AF9483A03F935" author="Crush PJ" box="[402,527,1736,1760]" pageId="26" pageNumber="380" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
), contrasting with the anteroposteriorly wider scapular blades in
<taxonomicName id="4C0BE8BA31079F0E433FF9673AF9F92A" baseAuthorityName="Walker" baseAuthorityYear="1990" box="[551,757,1767,1791]" class="Reptilia" genus="Sphenosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="26" pageNumber="380" phylum="Chordata" rank="species" species="acutus">
<emphasis id="B97F4F2B31079F0E433FF9673AF9F92A" box="[551,757,1767,1791]" italics="true" pageId="26" pageNumber="380">Sphenosuchus acutus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0BE8BA31079F0E4169F8873910F8CA" authorityName="Reig" authorityYear="1963" box="[113,284,1799,1823]" class="Polyplacophora" genus="Trialestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodilomorphi" pageId="26" pageNumber="380" phylum="Mollusca" rank="species" species="romeri">
<emphasis id="B97F4F2B31079F0E4169F8873910F8CA" box="[113,284,1799,1823]" italics="true" pageId="26" pageNumber="380">Trialestes romeri</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0BE8BA31079F0E4037F8873A25F8CA" authorityName="Simmons" authorityYear="1965" box="[303,553,1799,1823]" class="Reptilia" family="Sphenosuchidae" genus="Dibothrosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="26" pageNumber="380" phylum="Chordata" rank="species" species="elaphros">
<emphasis id="B97F4F2B31079F0E4037F8873A25F8CA" box="[303,553,1799,1823]" italics="true" pageId="26" pageNumber="380">Dibothrosuchus elaphros</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C0BE8BA31079F0E4375F88738A7F8EB" class="Reptilia" genus="Junggarsuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="26" pageNumber="380" phylum="Chordata" rank="species" species="sloani">
<emphasis id="B97F4F2B31079F0E4375F88738A7F8EB" italics="true" pageId="26" pageNumber="380">Junggarsuchus sloani</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9AEEC831079F0E41DAF8A63941F8EB" author="Walker AD" box="[194,333,1830,1854]" pageId="26" pageNumber="380" pagination="1 - 120" refId="ref33031" refString="Walker AD. A revision of Sphenosuchus acutus Haughton, a crocodylomorph reptile from the Elliot Formation (Late Triassic or Early Jurassic) of South Africa. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1990; 330: 1 - 120." type="journal article" year="1990">Walker 1990</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831079F0E4046F8A73A63F8EB" author="Wu X-C &amp; Chauerjee S" box="[350,623,1830,1854]" pageId="26" pageNumber="380" pagination="58 - 89" refId="ref33269" refString="Wu X-C, Chauerjee S. Dibothrosuchus elaphros, a crocodylomorph from the Lower Jurassic of China and the phylogeny of the Sphenosuchia. Journal of Vertebrate Paleontology 1993; 13: 58 - 89." type="journal article" year="1993">Wu and Chaưerjee 1993</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831079F0E4398F8A638AAF88B" author="Clark JM &amp; Xu X &amp; Forster CA" pageId="26" pageNumber="380" pagination="1021 - 4" refId="ref31048" refString="Clark JM, Xu X, Forster CA et al. A Middle Jurassic ' sphenosuchian' from China and the origin of the crocodylian skull. Nature 2004; 430: 1021 - 4." type="journal article" year="2004">
Clark
<emphasis id="B97F4F2B31079F0E43DDF8A73AF7F8EB" box="[709,763,1830,1854]" italics="true" pageId="26" pageNumber="380">et al.</emphasis>
2004
</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831079F0E41ADF8C6398BF888" author="Lecuona A &amp; Ezcurra MD &amp; Irmis RB" box="[181,391,1861,1885]" pageId="26" pageNumber="380" pagination="585 - 622" refId="ref32010" refString="Lecuona A, Ezcurra MD, Irmis RB. Revision of the early crocodylomorph Trialestes romeri (Archosauria, Suchia) from the lower Upper Triassic Ischigualasto Formation of Argentina: one of the oldest-known crocodylomorphs. Papers in Palaeontology 2016; 2: 585 - 622." type="journal article" year="2016">
Lecuona
<emphasis id="B97F4F2B31079F0E4000F8C63946F888" box="[280,330,1861,1885]" italics="true" pageId="26" pageNumber="380">et al.</emphasis>
2016
</bibRefCitation>
). A small fragment preserved dorsally to the dorsal end of the scapula was tentatively identified as part of the scapulacoracoid by
<bibRefCitation id="EF9AEEC831079F0E40D0F8053A53F849" author="Huene FR" box="[456,607,1924,1948]" pageId="26" pageNumber="380" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene (1921)</bibRefCitation>
. This fragment is considered unidentifiable here.
</paragraph>
<paragraph id="8BB4933931079F0E4232FF0F3DB8FD2F" blockId="26.[810,1460,143,763]" pageId="26" pageNumber="380">
<emphasis id="B97F4F2B31079F0E4232FF0F3B68FF72" box="[810,868,143,167]" italics="true" pageId="26" pageNumber="380">Ulna:</emphasis>
A long bone present among SMNS 12596 was identified by
<bibRefCitation id="EF9AEEC831079F0E4251FF2F3BD4FF12" author="Crush PJ" box="[841,984,175,199]" pageId="26" pageNumber="380" refId="ref31157" refString="Crush PJ. An early, terrestrial, crocodile from South Wales. Unpublished Ph. D. Thesis, University College London, 1980." type="book" year="1980">Crush (1980)</bibRefCitation>
and
<bibRefCitation id="EF9AEEC831079F0E4512FF2F3CA5FF12" author="Hughes B" box="[1034,1193,175,199]" pageId="26" pageNumber="380" pagination="221 - 41" refId="ref31617" refString="Hughes B. The earliest archosaurian reptiles. South Affican Journal of Science 1963; 59: 221 - 41." type="journal article" year="1963">Hughes (1963)</bibRefCitation>
as an ulna, asser originally being interpreted as a metatarsal by
<bibRefCitation id="EF9AEEC831079F0E45B3FF4F3D48FF33" author="Huene FR" box="[1195,1348,206,230]" pageId="26" pageNumber="380" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene (1921)</bibRefCitation>
. Based on its length (Supporting Information,
<tableCitation id="C689A68231079F0E45B9FF6D3CF7FED3" box="[1185,1275,237,262]" captionStart="Table 1" captionStartId="2.[114,169,143,167]" captionText="Table 1. Summary of the taxonomic assignments in relevant studies of the specimens that now comprise Saltoposuchus connectens" pageId="26" pageNumber="380">Table S1</tableCitation>
), it is too long to represent a metatarsal. One end of the element is asymmetrical, with one of its sides reaching further away from the shass, which could correspond to the olecranon process on the proximal end of an ulna (
<figureCitation id="13308FBC31079F0E42BEFEEB3BF8FE56" box="[934,1012,363,387]" captionStart="Figure 5" captionStartId="6.[113,178,1777,1801]" captionTargetBox="[221,1353,149,1749]" captionTargetId="figure-5@6.[220,1353,149,1749]" captionTargetPageId="6" captionText="Figure 5. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, B, SMNS 12596g, right pubis in (A) anterior and (B) angled medial views; C, SMNS 12596h, right ischium exposed in dorsomedial view and two proximal caudal vertebrae exposed in dorsal view; D, SMNS 12596i, tentative less ulna in posteromedial view; E, F, G, H, SMNS 12596j, right femur in (E) posteromedial, (F) anterolateral, (G) proximal, and (H) distal views; I, proximal half of the right femur in posterolateral view; J, K, SMNS 12596k, right tibia and fibula, (J) with the tibia in posteromedial and the fibula in lateral(?) view and (K) with the tibia in anterolateral and the fibula in medial(?) view. Abbreviations: alcr, anterolateral crest; amt, anteromedial tuber; car, caudal rib; cav, caudal vertebra; cr, crest; crtf, crista tibiofibularis; fi, fibula; gr, groove; ilf, ilium facet; iss, ischial shass; lc, lateral condyle; ltr, lesser trochanter; mc, medial condyle; no, notch; ns, neural spine; obf, obturator foramen; olp, olecranon process; os, osteoderm; poz, postzygapophysis; pr., proximal; prz, prezygapophysis; pus, pubic shass; put, pubic tubercle; r., right; ti, tibia; tr, trochanter." figureDoi="http://doi.org/10.5281/zenodo.8426159" httpUri="https://zenodo.org/record/8426159/files/figure.png" pageId="26" pageNumber="380">Fig. 5D</figureCitation>
). A small process projecting from the side of the head might correspond to the anteromedial tuber, which would suggest that the element represents a less ulna. However, other aspects of the element differ from known long bones of early crocodylomorphs; the shass is curved, flaưened, and widened towards its distal end, which could potentially be the result of a pathology or diagenetic compression, and the inferred distal end of the bone is very thin. The presence of one asymmetrical end and another flaưened and widened end also does not correspond with the morphology of a metatarsal. Its identification as a less ulna of
<taxonomicName id="4C0BE8BA31079F0E42EFFD243CE0FD69" authorityName="Huene" authorityYear="1921" box="[1015,1260,676,700]" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="26" pageNumber="380" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B31079F0E42EFFD243CE0FD69" box="[1015,1260,676,700]" italics="true" pageId="26" pageNumber="380">Saltoposuchus connectens</emphasis>
</taxonomicName>
is tentatively maintained here, although, due to the disassociation of the material, it cannot be excluded that the element belongs to a different taxon.
</paragraph>
<paragraph id="8BB4933931079F0E4232FC9C3C35F974" blockId="26.[810,1460,796,1979]" pageId="26" pageNumber="380">
<emphasis id="B97F4F2B31079F0E4232FC9C3BE2FCE1" box="[810,1006,796,820]" italics="true" pageId="26" pageNumber="380">Carpus and manus:</emphasis>
An articulated less manus is preserved and exposed in dorsal and medial views in SMNS 12352b (
<figureCitation id="13308FBC31079F0E4495FCBB3B65FCA6" captionStart="Figure 8" captionStartId="9.[129,194,1772,1796]" captionTargetBox="[178,1429,144,1744]" captionTargetId="figure-6@9.[174,1429,144,1744]" captionTargetPageId="9" captionText="Figure 8. Saltoposuchus connectens referred specimen SMNS 12591a. A, skull and mandible in right lateral view; B, interpretative drawing of A; C, less lateral view; D, interpretative drawing of (C). Abbreviations:an, angular; cc, crista cranii; de, dentary; fr, frontal; j, jugal; la, lacrimal; ls, laterosphenoid; mx, maxilla; na, nasal; pa, parietal; pl, palatine; po, postorbital; prf, prefrontal; pt, pterygoid; q, quadrate; qj, quadratojugal; sa, surangular; sq, squamosal." figureDoi="http://doi.org/10.5281/zenodo.8426167" httpUri="https://zenodo.org/record/8426167/files/figure.png" pageId="26" pageNumber="380">Fig. 8E–H</figureCitation>
). Directly proximal to metacarpals I and II, the distal half of an elongate element is preserved. This element was previously identified as the less radius (
<bibRefCitation id="EF9AEEC831079F0E4540FC1A3CD7FC64" author="Huene FR" box="[1112,1243,921,945]" pageId="26" pageNumber="380" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene 1921</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831079F0E45F1FC193D7BFC64" author="Ostrom JH" box="[1257,1399,921,945]" pageId="26" pageNumber="380" pagination="179 - 95" refId="ref32208" refString="Ostrom JH. Procompsognathus - Theropod or Thecodont? Palaeontographica A 1981; 175: 179 - 95." type="journal article" year="1981">Ostrom 1981</bibRefCitation>
), but
<bibRefCitation id="EF9AEEC831079F0E4232FC393C20FC04" author="Sereno PC &amp; Wild R" box="[810,1068,953,977]" pageId="26" pageNumber="380" pagination="435 - 58" refId="ref32722" refString="Sereno PC, Wild R. Procompsognathus: theropod, ' thecodont' or both? Journal of Vertebrate Paleontology 1992; 12: 435 - 58." type="journal article" year="1992">Sereno and Wild (1992)</bibRefCitation>
later identified this element as a partial, elongate less radiale. They also identified a wide but thin piece of bone between the radiale and metacarpal II as a distal carpal, and a tiny bone sliver lateral to the distal end of the radiale as the distalmost end of the less ulnare. This configuration of a single distal carpal and an elongate radiale and ulnare proximal to this distal carpal is corroborated here, and forms a diagnostic configuration for early crocodylomorphs (
<bibRefCitation id="EF9AEEC831079F0E45C4FB143D5AFB79" author="Crush PJ" box="[1244,1366,1172,1196]" pageId="26" pageNumber="380" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831079F0E447AFB143BDEFB1E" author="Wu X-C &amp; Chauerjee S" pageId="26" pageNumber="380" pagination="58 - 89" refId="ref33269" refString="Wu X-C, Chauerjee S. Dibothrosuchus elaphros, a crocodylomorph from the Lower Jurassic of China and the phylogeny of the Sphenosuchia. Journal of Vertebrate Paleontology 1993; 13: 58 - 89." type="journal article" year="1993">Wu and Chaưerjee 1993</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831079F0E42F8FB333C64FB1E" author="Nesbiu SJ" box="[992,1128,1203,1227]" pageId="26" pageNumber="380" pagination="1 - 292" refId="ref32089" refString="Nesbiu SJ. The early evolution of archosaurs: relationships and the origin of major clades. Bulletin of the American Museum of Natural History 2011; 352: 1 - 292." type="journal article" year="2011">Nesbiư 2011</bibRefCitation>
: ch. 242,
<bibRefCitation id="EF9AEEC831079F0E45CBFB343DAEFB1E" author="Lecuona A &amp; Ezcurra MD &amp; Irmis RB" box="[1235,1442,1203,1227]" pageId="26" pageNumber="380" pagination="585 - 622" refId="ref32010" refString="Lecuona A, Ezcurra MD, Irmis RB. Revision of the early crocodylomorph Trialestes romeri (Archosauria, Suchia) from the lower Upper Triassic Ischigualasto Formation of Argentina: one of the oldest-known crocodylomorphs. Papers in Palaeontology 2016; 2: 585 - 622." type="journal article" year="2016">
Lecuona
<emphasis id="B97F4F2B31079F0E442DFB343D6BFB1E" box="[1333,1383,1203,1227]" italics="true" pageId="26" pageNumber="380">et al.</emphasis>
2016
</bibRefCitation>
). In contrast, it does not correspond to the carpal configuration of early theropods, in which the radiale and ulnare are very wide and proximodistally very short elements, and in which three distal carpals are generally present (e.g.
<bibRefCitation id="EF9AEEC831079F0E4406FAB03DA1FA9C" author="Colbert EH" box="[1310,1453,1328,1353]" pageId="26" pageNumber="380" pagination="1 - 160" refId="ref31108" refString="Colbert EH. The Triassic dinosaur Coelophysis. Museum of Northern Arizona, Bulletin 1989; 57: 1 - 160." type="journal article" year="1989">Colbert 1989</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831079F0E4232FAD03BBFFABD" author="Nesbiu SJ" box="[810,947,1360,1384]" pageId="26" pageNumber="380" pagination="1 - 292" refId="ref32089" refString="Nesbiu SJ. The early evolution of archosaurs: relationships and the origin of major clades. Bulletin of the American Museum of Natural History 2011; 352: 1 - 292." type="journal article" year="2011">Nesbiư 2011</bibRefCitation>
). The manus and associated elements therefore clearly belong to a non-crocodyliform crocodylomorph (
<emphasis id="B97F4F2B31079F0E4498FAF03DBFFA52" box="[1408,1459,1392,1415]" italics="true" pageId="26" pageNumber="380">sensu</emphasis>
<bibRefCitation id="EF9AEEC831079F0E4232FA0F3C1CFA72" author="Sereno PC &amp; Wild R" box="[810,1040,1422,1447]" pageId="26" pageNumber="380" pagination="435 - 58" refId="ref32722" refString="Sereno PC, Wild R. Procompsognathus: theropod, ' thecodont' or both? Journal of Vertebrate Paleontology 1992; 12: 435 - 58." type="journal article" year="1992">Sereno and Wild 1992</bibRefCitation>
), and they can reasonably be assigned to
<taxonomicName id="4C0BE8BA31079F0E4232FA2E3C11FA13" authorityName="Huene" authorityYear="1921" box="[810,1053,1454,1478]" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="26" pageNumber="380" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B31079F0E4232FA2E3C11FA13" box="[810,1053,1454,1478]" italics="true" pageId="26" pageNumber="380">Saltoposuchus connectens</emphasis>
</taxonomicName>
based on their association with the skull of SMNS 12352a. The shass of the radiale has a flaưened anterior face and is slightly wider mediolaterally than anteroposteriorly. The distal end of the radiale is slightly mediolaterally expanded compared to the shass. The ulnare preserves no discernible features, whereas the distal carpal is also poorly preserved but roughly ovoid in outline. It is only slightly wider than the proximal head of metacarpal II.
</paragraph>
<paragraph id="8BB4933931079F0F425EF92838E0FD69" blockId="26.[810,1460,796,1979]" lastBlockId="27.[128,778,144,700]" lastPageId="27" lastPageNumber="381" pageId="26" pageNumber="380">
All metacarpals are virtually completely preserved.Metacarpal I is considerably shorter, but not narrower, than the other metacarpals (
<figureCitation id="13308FBC31079F0E429CF9673BF7F92A" box="[900,1019,1767,1791]" captionStart="Figure 10" captionStartId="14.[113,178,1772,1796]" captionTargetBox="[197,1376,144,1744]" captionTargetId="figure-5@14.[197,1376,144,1744]" captionTargetPageId="14" captionText="Figure 10. Saltoposuchus connectens referred specimen SMNS 12352.A, SMNS 12352a with the snout region exposed in angled right lateral view; B, interpretative drawing of (A); C, opposite side of SMNS 12352a exposing the posterior portion of the less maxilla and lacrimal in angled lateral view; D, interpretative drawing of (C); E, SMNS 12352b, carpus and manus in dorsal view; F, interpretative drawing of (E); G, SMNS 12352b in medial view; H, interpretative drawing of (G). Abbreviations; antfe, antorbital fenestra; antfo, antorbital fossa; dc, distal carpal; de, dentary; exp, extensor pit; fr, frontal; j, jugal; la, lacrimal; lad, lacrimal duct; mc, metacarpal; mx, maxilla; na, nasal; ph, phalanx; pm, premaxilla; prf, prefrontal; ra, radiale; ul, ulnare." figureDoi="http://doi.org/10.5281/zenodo.8426171" httpUri="https://zenodo.org/record/8426171/files/figure.png" pageId="26" pageNumber="380">Fig. 10E–H</figureCitation>
), as was also observed previously (
<bibRefCitation id="EF9AEEC831079F0E4475F9683B52F8CB" author="Huene FR" pageId="26" pageNumber="380" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene 1921</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831079F0E4274F8863BF4F8CB" author="Ostrom JH" box="[876,1016,1798,1822]" pageId="26" pageNumber="380" pagination="179 - 95" refId="ref32208" refString="Ostrom JH. Procompsognathus - Theropod or Thecodont? Palaeontographica A 1981; 175: 179 - 95." type="journal article" year="1981">Ostrom 1981</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831079F0E451DF8863CFDF8CB" author="Sereno PC &amp; Wild R" box="[1029,1265,1798,1822]" pageId="26" pageNumber="380" pagination="435 - 58" refId="ref32722" refString="Sereno PC, Wild R. Procompsognathus: theropod, ' thecodont' or both? Journal of Vertebrate Paleontology 1992; 12: 435 - 58." type="journal article" year="1992">Sereno and Wild 1992</bibRefCitation>
). The other metacarpals are all elongate, with metacarpal V being the shortest, followed by metacarpals IV and then II, and with metacarpal III being the longest (Supporting Information,
<tableCitation id="C689A68231079F0E45FAF8E43D34F8A9" box="[1250,1336,1892,1916]" captionStart="Table 1" captionStartId="2.[114,169,143,167]" captionText="Table 1. Summary of the taxonomic assignments in relevant studies of the specimens that now comprise Saltoposuchus connectens" pageId="26" pageNumber="380">Table S1</tableCitation>
). The metacarpals are subequal in width, except for metacarpal V, which is more slender. The metacarpals are all tightly bunched and metacarpals II–V are all oriented in the same lateromedial plane, whereas metacarpal I is positioned ventromedial to metacarpal II (
<figureCitation id="13308FBC31069F0F41BDFF4F392CFF33" box="[165,288,207,231]" captionStart="Figure 10" captionStartId="14.[113,178,1772,1796]" captionTargetBox="[197,1376,144,1744]" captionTargetId="figure-5@14.[197,1376,144,1744]" captionTargetPageId="14" captionText="Figure 10. Saltoposuchus connectens referred specimen SMNS 12352.A, SMNS 12352a with the snout region exposed in angled right lateral view; B, interpretative drawing of (A); C, opposite side of SMNS 12352a exposing the posterior portion of the less maxilla and lacrimal in angled lateral view; D, interpretative drawing of (C); E, SMNS 12352b, carpus and manus in dorsal view; F, interpretative drawing of (E); G, SMNS 12352b in medial view; H, interpretative drawing of (G). Abbreviations; antfe, antorbital fenestra; antfo, antorbital fossa; dc, distal carpal; de, dentary; exp, extensor pit; fr, frontal; j, jugal; la, lacrimal; lad, lacrimal duct; mc, metacarpal; mx, maxilla; na, nasal; ph, phalanx; pm, premaxilla; prf, prefrontal; ra, radiale; ul, ulnare." figureDoi="http://doi.org/10.5281/zenodo.8426171" httpUri="https://zenodo.org/record/8426171/files/figure.png" pageId="27" pageNumber="381">Fig. 10G, H</figureCitation>
). The tightly bunched carpus is indicative of a digitigrade posture, as was previously also pointed out by
<bibRefCitation id="EF9AEEC831069F0F43DAFF6E3922FEF0" author="Sereno PC &amp; Wild R" pageId="27" pageNumber="381" pagination="435 - 58" refId="ref32722" refString="Sereno PC, Wild R. Procompsognathus: theropod, ' thecodont' or both? Journal of Vertebrate Paleontology 1992; 12: 435 - 58." type="journal article" year="1992">Sereno and Wild (1992)</bibRefCitation>
for
<taxonomicName id="4C0BE8BA31069F0F404EFE8D3A45FEF0" authorityName="Huene" authorityYear="1921" box="[342,585,269,293]" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="27" pageNumber="381" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B31069F0F404EFE8D3A45FEF0" box="[342,585,269,293]" italics="true" pageId="27" pageNumber="381">Saltoposuchus connectens</emphasis>
</taxonomicName>
. A digitigrade posture of the front limb has also been described for
<taxonomicName id="4C0BE8BA31069F0F4360FEAD38F1FEB1" authority=", CM" authorityName="CM" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="27" pageNumber="381" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B31069F0F4360FEAD38C8FEB1" italics="true" pageId="27" pageNumber="381">Terrestrisuchus gracilis</emphasis>
, CM
</taxonomicName>
29894,
<taxonomicName id="4C0BE8BA31069F0F404DFECC3A2EFEB1" box="[341,546,332,356]" class="Reptilia" genus="Junggarsuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="27" pageNumber="381" phylum="Chordata" rank="species" species="sloani">
<emphasis id="B97F4F2B31069F0F404DFECC3A2EFEB1" box="[341,546,332,356]" italics="true" pageId="27" pageNumber="381">Junggarsuchus sloani</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C0BE8BA31069F0F4347FECC3B08FEB1" authorityName="Reig" authorityYear="1963" box="[607,772,332,356]" class="Polyplacophora" genus="Trialestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodilomorphi" pageId="27" pageNumber="381" phylum="Mollusca" rank="species" species="romeri">
<emphasis id="B97F4F2B31069F0F4347FECC3B08FEB1" box="[607,772,332,356]" italics="true" pageId="27" pageNumber="381">Trialestes romeri</emphasis>
</taxonomicName>
, albeit tentatively for this final taxon (NHMUK PV R 7562, NHMUK PV R 7557b,
<bibRefCitation id="EF9AEEC831069F0F4064FE0B3A25FE76" author="Clark JM &amp; Sues H-D &amp; Berman DS" box="[380,553,395,419]" pageId="27" pageNumber="381" pagination="683 - 704" refId="ref31006" refString="Clark JM, Sues H-D, Berman DS. A new specimen of Hesperosuchus agilis from the Upper Triassic of New Mexico and the interrelationships of basal crocodylomorph archosaurs. Journal of Vertebrate Paleontology 2000; 20: 683 - 704." type="journal article" year="2000">
Clark
<emphasis id="B97F4F2B31069F0F40A4FE0B39E2FE76" box="[444,494,395,419]" italics="true" pageId="27" pageNumber="381">et al.</emphasis>
2000
</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831069F0F432EFE0B3A66FE76" author="Clark JM &amp; Xu X &amp; Forster CA" box="[566,618,395,419]" pageId="27" pageNumber="381" pagination="1021 - 4" refId="ref31048" refString="Clark JM, Xu X, Forster CA et al. A Middle Jurassic ' sphenosuchian' from China and the origin of the crocodylian skull. Nature 2004; 430: 1021 - 4." type="journal article" year="2004">2004</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831069F0F4360FE0B38B8FE17" author="Lecuona A &amp; Ezcurra MD &amp; Irmis RB" pageId="27" pageNumber="381" pagination="585 - 622" refId="ref32010" refString="Lecuona A, Ezcurra MD, Irmis RB. Revision of the early crocodylomorph Trialestes romeri (Archosauria, Suchia) from the lower Upper Triassic Ischigualasto Formation of Argentina: one of the oldest-known crocodylomorphs. Papers in Palaeontology 2016; 2: 585 - 622." type="journal article" year="2016">
Lecuona
<emphasis id="B97F4F2B31069F0F43C1FE0B3B06FE76" box="[729,778,395,419]" italics="true" pageId="27" pageNumber="381">et al.</emphasis>
2016
</bibRefCitation>
). The metacarpals are slightly expanded at their proximal and distal ends, and the dorsal surface of the distal ends possess concave extensor pits (
<figureCitation id="13308FBC31069F0F407FFE6939DEFDD5" box="[359,466,489,513]" captionStart="Figure 10" captionStartId="14.[113,178,1772,1796]" captionTargetBox="[197,1376,144,1744]" captionTargetId="figure-5@14.[197,1376,144,1744]" captionTargetPageId="14" captionText="Figure 10. Saltoposuchus connectens referred specimen SMNS 12352.A, SMNS 12352a with the snout region exposed in angled right lateral view; B, interpretative drawing of (A); C, opposite side of SMNS 12352a exposing the posterior portion of the less maxilla and lacrimal in angled lateral view; D, interpretative drawing of (C); E, SMNS 12352b, carpus and manus in dorsal view; F, interpretative drawing of (E); G, SMNS 12352b in medial view; H, interpretative drawing of (G). Abbreviations; antfe, antorbital fenestra; antfo, antorbital fossa; dc, distal carpal; de, dentary; exp, extensor pit; fr, frontal; j, jugal; la, lacrimal; lad, lacrimal duct; mc, metacarpal; mx, maxilla; na, nasal; ph, phalanx; pm, premaxilla; prf, prefrontal; ra, radiale; ul, ulnare." figureDoi="http://doi.org/10.5281/zenodo.8426171" httpUri="https://zenodo.org/record/8426171/files/figure.png" pageId="27" pageNumber="381">Fig. 10E, F</figureCitation>
). Two phalanges are preserved in digit I, of which the distalmost represents a very small ungual (
<figureCitation id="13308FBC31069F0F4193FDA7390DFDEA" box="[139,257,551,575]" captionStart="Figure 10" captionStartId="14.[113,178,1772,1796]" captionTargetBox="[197,1376,144,1744]" captionTargetId="figure-5@14.[197,1376,144,1744]" captionTargetPageId="14" captionText="Figure 10. Saltoposuchus connectens referred specimen SMNS 12352.A, SMNS 12352a with the snout region exposed in angled right lateral view; B, interpretative drawing of (A); C, opposite side of SMNS 12352a exposing the posterior portion of the less maxilla and lacrimal in angled lateral view; D, interpretative drawing of (C); E, SMNS 12352b, carpus and manus in dorsal view; F, interpretative drawing of (E); G, SMNS 12352b in medial view; H, interpretative drawing of (G). Abbreviations; antfe, antorbital fenestra; antfo, antorbital fossa; dc, distal carpal; de, dentary; exp, extensor pit; fr, frontal; j, jugal; la, lacrimal; lad, lacrimal duct; mc, metacarpal; mx, maxilla; na, nasal; ph, phalanx; pm, premaxilla; prf, prefrontal; ra, radiale; ul, ulnare." figureDoi="http://doi.org/10.5281/zenodo.8426171" httpUri="https://zenodo.org/record/8426171/files/figure.png" pageId="27" pageNumber="381">Fig. 10G, H</figureCitation>
). No further unguals are preserved and the phalangeal count of the other digits is incomplete. Digits II, IV, and V each preserve two phalanges and digit III preserves three (
<figureCitation id="13308FBC31069F0F43FBFDE638C8FD48" captionStart="Figure 10" captionStartId="14.[113,178,1772,1796]" captionTargetBox="[197,1376,144,1744]" captionTargetId="figure-5@14.[197,1376,144,1744]" captionTargetPageId="14" captionText="Figure 10. Saltoposuchus connectens referred specimen SMNS 12352.A, SMNS 12352a with the snout region exposed in angled right lateral view; B, interpretative drawing of (A); C, opposite side of SMNS 12352a exposing the posterior portion of the less maxilla and lacrimal in angled lateral view; D, interpretative drawing of (C); E, SMNS 12352b, carpus and manus in dorsal view; F, interpretative drawing of (E); G, SMNS 12352b in medial view; H, interpretative drawing of (G). Abbreviations; antfe, antorbital fenestra; antfo, antorbital fossa; dc, distal carpal; de, dentary; exp, extensor pit; fr, frontal; j, jugal; la, lacrimal; lad, lacrimal duct; mc, metacarpal; mx, maxilla; na, nasal; ph, phalanx; pm, premaxilla; prf, prefrontal; ra, radiale; ul, ulnare." figureDoi="http://doi.org/10.5281/zenodo.8426171" httpUri="https://zenodo.org/record/8426171/files/figure.png" pageId="27" pageNumber="381">Fig. 10E, F</figureCitation>
). Extensor pits are also present on the distal ends of the phalanges.
</paragraph>
<paragraph id="8BB4933931069F0F4198FD5E38DFFA9D" blockId="27.[128,778,734,1352]" pageId="27" pageNumber="381">
<emphasis id="B97F4F2B31069F0F4198FD5E38B1FD23" box="[128,189,734,758]" italics="true" pageId="27" pageNumber="381">Ilium:</emphasis>
A single, right ilium is preserved in articulation with the sacral ribs and femur in SMNS 12597a (
<figureCitation id="13308FBC31069F0F431BFD7D3A43FCC0" box="[515,591,765,789]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="27" pageNumber="381">Figs 2A</figureCitation>
,
<figureCitation id="13308FBC31069F0F4343FD7D3A8AFCC0" box="[603,646,765,789]" captionStart="Figure 12" captionStartId="24.[113,178,1772,1796]" captionTargetBox="[151,1424,144,1744]" captionTargetId="figure-6@24.[148,1424,144,1744]" captionTargetPageId="24" captionText="Figure 12. Details of the postcranial anatomy of Saltoposuchus connectens. A, dorsal vertebral neural arch and dorsal rib of SMNS 12596c in ventral view; B, associated osteoderms of SMNS 12596c in dorsal view; C, sacral ribs and associated elements of SMNS 12597a in dorsal view; D, distal portion of right scapular blade of SMNS 12597a in lateral view; E, proximal end of right tibia of SMNS 12597a in proximal view; F, less tarsus and pes of SMNS 12597a with the calcaneum exposed in lateral view; G, close-up of less tarsus of SMNS 12597a; H, close-up of less pes of SMNS 12597a. Abbreviations: ane, anterior expansion; cal, calcaneum; calt, calcaneal tuber; cp, capitulum; dp, diapophysis; dt, distal tarsal; exp, extensor pit; fe, femur; fi, fibula; il, ilium; lam, lamina; latc, lateral condyle; le, lateral edge; lr, longitudinal ridge; me, medial edge; mt, metatarsal; no, notch; pap, parapophysis; ph, phalanx; pme, posterior extension; poe, posterior expansion; pofl, posterior flange; poz, postzygapophysis; os, osteoderm; r., right; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426175" httpUri="https://zenodo.org/record/8426175/files/figure.png" pageId="27" pageNumber="381">12C</figureCitation>
). It is poorly preserved and exposed in dorsal view. The pre-acetabular process is largely missing but, although its dorsal margin is missing, the post-acetabular process is preserved across its entire length. The dorsal margin of the ilium has a sinusoidal outline in dorsal view, with the post-acetabular process curving first medially and subsequently laterally from anterior to posterior (
<figureCitation id="13308FBC31069F0F436AFC393AC4FC04" box="[626,712,953,977]" captionStart="Figure 12" captionStartId="24.[113,178,1772,1796]" captionTargetBox="[151,1424,144,1744]" captionTargetId="figure-6@24.[148,1424,144,1744]" captionTargetPageId="24" captionText="Figure 12. Details of the postcranial anatomy of Saltoposuchus connectens. A, dorsal vertebral neural arch and dorsal rib of SMNS 12596c in ventral view; B, associated osteoderms of SMNS 12596c in dorsal view; C, sacral ribs and associated elements of SMNS 12597a in dorsal view; D, distal portion of right scapular blade of SMNS 12597a in lateral view; E, proximal end of right tibia of SMNS 12597a in proximal view; F, less tarsus and pes of SMNS 12597a with the calcaneum exposed in lateral view; G, close-up of less tarsus of SMNS 12597a; H, close-up of less pes of SMNS 12597a. Abbreviations: ane, anterior expansion; cal, calcaneum; calt, calcaneal tuber; cp, capitulum; dp, diapophysis; dt, distal tarsal; exp, extensor pit; fe, femur; fi, fibula; il, ilium; lam, lamina; latc, lateral condyle; le, lateral edge; lr, longitudinal ridge; me, medial edge; mt, metatarsal; no, notch; pap, parapophysis; ph, phalanx; pme, posterior extension; poe, posterior expansion; pofl, posterior flange; poz, postzygapophysis; os, osteoderm; r., right; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426175" httpUri="https://zenodo.org/record/8426175/files/figure.png" pageId="27" pageNumber="381">Fig. 12C</figureCitation>
). This is probably due to diagenetic deformation. The lateral surface of the ilium slants ventrolaterally to form the supracetabular crest, which extends far laterally beyond the rest of the ilium. Its lateral margin is incomplete. The acetabulum itself is covered by the crest and completely embedded in matrix. The medial side of the ilium is largely covered in matrix, but it is apparent that the first sacral rib aưaches on the anterior portion of the main body of the ilium, with its anterior end connecting to the now missing pre-acetabular process. The second sacral rib connects on its anterior end to the posterior part of the main body of the ilium and on its posterior part to the proximal base of the post-acetabular process.
</paragraph>
<paragraph id="8BB4933931069F0F4199FAEA3D9FFE56" blockId="27.[129,778,1386,1974]" lastBlockId="27.[825,1474,143,387]" pageId="27" pageNumber="381">
<emphasis id="B97F4F2B31069F0F4199FAEA38D8FA57" box="[129,212,1386,1410]" italics="true" pageId="27" pageNumber="381">Ischium:</emphasis>
A large right ischium is exposed in medial or dorsal view in SMNS 12596h (
<figureCitation id="13308FBC31069F0F406FFA0A39CCFA74" box="[375,448,1417,1442]" captionStart="Figure 5" captionStartId="6.[113,178,1777,1801]" captionTargetBox="[221,1353,149,1749]" captionTargetId="figure-5@6.[220,1353,149,1749]" captionTargetPageId="6" captionText="Figure 5. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, B, SMNS 12596g, right pubis in (A) anterior and (B) angled medial views; C, SMNS 12596h, right ischium exposed in dorsomedial view and two proximal caudal vertebrae exposed in dorsal view; D, SMNS 12596i, tentative less ulna in posteromedial view; E, F, G, H, SMNS 12596j, right femur in (E) posteromedial, (F) anterolateral, (G) proximal, and (H) distal views; I, proximal half of the right femur in posterolateral view; J, K, SMNS 12596k, right tibia and fibula, (J) with the tibia in posteromedial and the fibula in lateral(?) view and (K) with the tibia in anterolateral and the fibula in medial(?) view. Abbreviations: alcr, anterolateral crest; amt, anteromedial tuber; car, caudal rib; cav, caudal vertebra; cr, crest; crtf, crista tibiofibularis; fi, fibula; gr, groove; ilf, ilium facet; iss, ischial shass; lc, lateral condyle; ltr, lesser trochanter; mc, medial condyle; no, notch; ns, neural spine; obf, obturator foramen; olp, olecranon process; os, osteoderm; poz, postzygapophysis; pr., proximal; prz, prezygapophysis; pus, pubic shass; put, pubic tubercle; r., right; ti, tibia; tr, trochanter." figureDoi="http://doi.org/10.5281/zenodo.8426159" httpUri="https://zenodo.org/record/8426159/files/figure.png" pageId="27" pageNumber="381">Fig. 5C</figureCitation>
). It is well preserved but its ventral, anterior, and probably posterior margins are incomplete and therefore the proximodistal length of the ischial shass cannot be determined. The ischial shass curves posteriorly so that its distal end is directed considerably more posteriorly than ventrally. The ischial shass gradually reduces in height distally. The dorsal margin is distinctly concave in medial view, thickened and curved proximally, but thin distally. From the base of the dorsal end of the ischial shass a crest is projected that curves anterodorsally and medially. At its dorsal end, this crest forms the posterior end of a large, concave surface that faces mediodorsally and somewhat anteriorly and that is triangular in outline. This forms the articular surface with the ilium. The medial surface directly ventral to the iliac peduncle is concave. A small, sloped surface is visible directly lateral to the peduncle, which is directed dorsolaterally and slightly anteriorly. This probably represents the incompletely exposed antitrochanter of the ischium (
<emphasis id="B97F4F2B31069F0F4399F8003AB2F842" box="[641,702,1920,1943]" italics="true" pageId="27" pageNumber="381">contra</emphasis>
<bibRefCitation id="EF9AEEC831069F0F43DAF8FF3917F863" author="Sereno PC &amp; Wild R" pageId="27" pageNumber="381" pagination="435 - 58" refId="ref32722" refString="Sereno PC, Wild R. Procompsognathus: theropod, ' thecodont' or both? Journal of Vertebrate Paleontology 1992; 12: 435 - 58." type="journal article" year="1992">Sereno and Wild 1992</bibRefCitation>
), which is also present in
<taxonomicName id="4C0BE8BA31069F0F4335F81E3B06F863" box="[557,778,1950,1974]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="27" pageNumber="381" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B31069F0F4335F81E3B06F863" box="[557,778,1950,1974]" italics="true" pageId="27" pageNumber="381">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
but absent in
<taxonomicName id="4C0BE8BA31069F0F42DCFF0F3C69FF72" authorityName="Reig" authorityYear="1963" box="[964,1125,143,167]" class="Polyplacophora" genus="Trialestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodilomorphi" pageId="27" pageNumber="381" phylum="Mollusca" rank="species" species="romeri">
<emphasis id="B97F4F2B31069F0F42DCFF0F3C69FF72" box="[964,1125,143,167]" italics="true" pageId="27" pageNumber="381">Trialestes romeri</emphasis>
</taxonomicName>
(NHMUK PV R 37728,
<bibRefCitation id="EF9AEEC831069F0F4471FF103BACFF12" author="Lecuona A &amp; Ezcurra MD &amp; Irmis RB" pageId="27" pageNumber="381" pagination="585 - 622" refId="ref32010" refString="Lecuona A, Ezcurra MD, Irmis RB. Revision of the early crocodylomorph Trialestes romeri (Archosauria, Suchia) from the lower Upper Triassic Ischigualasto Formation of Argentina: one of the oldest-known crocodylomorphs. Papers in Palaeontology 2016; 2: 585 - 622." type="journal article" year="2016">
Lecuona
<emphasis id="B97F4F2B31069F0F4221FF303B64FF12" box="[825,872,175,199]" italics="true" pageId="27" pageNumber="381">et al.</emphasis>
2016
</bibRefCitation>
). A clear concavity anterior to the iliac peduncle, as is found in
<taxonomicName id="4C0BE8BA31069F0F4281FF4E3C79FF33" box="[921,1141,206,230]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="27" pageNumber="381" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B31069F0F4281FF4E3C79FF33" box="[921,1141,206,230]" italics="true" pageId="27" pageNumber="381">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
(NHMUK PV R 37728), is absent, which could suggest that the acetabular fenestra was comparatively smaller in
<taxonomicName id="4C0BE8BA31069F0F450FFE8D3D03FEF0" authorityName="Huene" authorityYear="1921" box="[1047,1295,269,293]" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="27" pageNumber="381" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B31069F0F450FFE8D3D03FEF0" box="[1047,1295,269,293]" italics="true" pageId="27" pageNumber="381">Saltoposuchus connectens</emphasis>
</taxonomicName>
. The incomplete anterior margin is roughly straight and vertically oriented. This formed the articular surface with the pubis. The incomplete ventral margin formed the articular surface for the less ischium.
</paragraph>
<paragraph id="8BB4933931069F0F4221FE243D60FABD" blockId="27.[825,1475,420,1822]" pageId="27" pageNumber="381">
<emphasis id="B97F4F2B31069F0F4221FE243B7AFE69" box="[825,886,420,444]" italics="true" pageId="27" pageNumber="381">Pubis:</emphasis>
The pubis is preserved in SMNS 12597a and SMNS
<quantity id="4CF33EDC31069F0F4221FE433B85FE0E" box="[825,905,451,475]" metricMagnitude="1" metricUnit="kg" metricValue="1.2596" pageId="27" pageNumber="381" unit="g" value="12596.0">12596g</quantity>
. The right pubis of SMNS 12597a is disarticulated from, but still associated with, the right ilium and sacral vertebrae. Most of the shass is exposed in lateral and anterior views (
<figureCitation id="13308FBC31069F0F4484FD823B5BFDEF" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="27" pageNumber="381">Fig. 2A</figureCitation>
). SMNS
<quantity id="4CF33EDC31069F0F42D8FDA13C01FDEC" box="[960,1037,545,569]" metricMagnitude="1" metricUnit="kg" metricValue="1.2596" pageId="27" pageNumber="381" unit="g" value="12596.0">12596g</quantity>
preserves a virtually complete, disarticulated right pubis that is exposed in anterior and medial views (
<figureCitation id="13308FBC31069F0F4484FDC13B7EFDAD" captionStart="Figure 5" captionStartId="6.[113,178,1777,1801]" captionTargetBox="[221,1353,149,1749]" captionTargetId="figure-5@6.[220,1353,149,1749]" captionTargetPageId="6" captionText="Figure 5. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, B, SMNS 12596g, right pubis in (A) anterior and (B) angled medial views; C, SMNS 12596h, right ischium exposed in dorsomedial view and two proximal caudal vertebrae exposed in dorsal view; D, SMNS 12596i, tentative less ulna in posteromedial view; E, F, G, H, SMNS 12596j, right femur in (E) posteromedial, (F) anterolateral, (G) proximal, and (H) distal views; I, proximal half of the right femur in posterolateral view; J, K, SMNS 12596k, right tibia and fibula, (J) with the tibia in posteromedial and the fibula in lateral(?) view and (K) with the tibia in anterolateral and the fibula in medial(?) view. Abbreviations: alcr, anterolateral crest; amt, anteromedial tuber; car, caudal rib; cav, caudal vertebra; cr, crest; crtf, crista tibiofibularis; fi, fibula; gr, groove; ilf, ilium facet; iss, ischial shass; lc, lateral condyle; ltr, lesser trochanter; mc, medial condyle; no, notch; ns, neural spine; obf, obturator foramen; olp, olecranon process; os, osteoderm; poz, postzygapophysis; pr., proximal; prz, prezygapophysis; pus, pubic shass; put, pubic tubercle; r., right; ti, tibia; tr, trochanter." figureDoi="http://doi.org/10.5281/zenodo.8426159" httpUri="https://zenodo.org/record/8426159/files/figure.png" pageId="27" pageNumber="381">Fig. 5A, B</figureCitation>
). Its proximal portion is composed of a plate-like surface that is directed ventromedially to dorsolaterally, with the dorsal end being curved so that it is oriented directly dorsally. The plate is characterized by a very large and oval-shaped obturator foramen, which has a long axis that is directed anteroventrally to posterodorsally. The posterior margin of the pubic plate is composed of a dorsal portion, which forms approximately two-thirds of the posterior margin, and a ventral portion. The dorsal portion of the posterior margin is concave in medial view, and forms the articular surface with the ischium, whereas the ventral portion is straight and did not articulate with another element, thus forming an open notch. In contrast, the straight ventromedial margin of the plate is aligned with the medial margin of the pubic shass, and both would have articulated with the opposing pubis. The dorsal margin of the pubic plate in medial view is incomplete but would have articulated with the ilium. Near its proximal end, the pubis possesses a clear laterally and slightly posteriorly directed process, which has a triangular outline in anterolateral view. This is the pubic tubercle, which is also present in
<taxonomicName id="4C0BE8BA31069F0F4299FB333C56FB1E" box="[897,1114,1203,1227]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="27" pageNumber="381" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B31069F0F4299FB333C56FB1E" box="[897,1114,1203,1227]" italics="true" pageId="27" pageNumber="381">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0BE8BA31069F0F4591FB333D25FB1E" authorityName="Reig" authorityYear="1963" box="[1161,1321,1203,1227]" class="Polyplacophora" genus="Trialestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodilomorphi" pageId="27" pageNumber="381" phylum="Mollusca" rank="species" species="romeri">
<emphasis id="B97F4F2B31069F0F4591FB333D25FB1E" box="[1161,1321,1203,1227]" italics="true" pageId="27" pageNumber="381">Trialestes romeri</emphasis>
</taxonomicName>
(NHMUK PV R 7562,
<bibRefCitation id="EF9AEEC831069F0F428BFB533C53FB3F" author="Lecuona A &amp; Ezcurra MD &amp; Irmis RB" box="[915,1119,1234,1258]" pageId="27" pageNumber="381" pagination="585 - 622" refId="ref32010" refString="Lecuona A, Ezcurra MD, Irmis RB. Revision of the early crocodylomorph Trialestes romeri (Archosauria, Suchia) from the lower Upper Triassic Ischigualasto Formation of Argentina: one of the oldest-known crocodylomorphs. Papers in Palaeontology 2016; 2: 585 - 622." type="journal article" year="2016">
Lecuona
<emphasis id="B97F4F2B31069F0F42ECFB533C29FB3F" box="[1012,1061,1234,1258]" italics="true" pageId="27" pageNumber="381">et al.</emphasis>
2016
</bibRefCitation>
), and which forms an aưachment site for M. obliquus abdominus, M. ambiens, and ligaments (
<bibRefCitation id="EF9AEEC831069F0F425CFA913C0BFAFC" author="Hutchinson JR" box="[836,1031,1297,1321]" pageId="27" pageNumber="381" pagination="123 - 68" refId="ref31711" refString="Hutchinson JR. The evolution of pelvic osteology and soss tissues on the line to extant birds (Neornithes). Zoological Journal of the Linnean Society 2001 b; 131: 123 - 68." type="journal article" year="2001">Hutchinson 2001b</bibRefCitation>
). The portion posterodorsal to the pubic tubercle represents the iliac peduncle, which is embedded in matrix and largely obscured by an overlying osteoderm.
</paragraph>
<paragraph id="8BB4933931069F0F424DFAEF3BB4F8CB" blockId="27.[825,1475,420,1822]" pageId="27" pageNumber="381">
The pubis possesses an elongate, flaưened pubic shass that has wide anterodorsal and posteroventral surfaces and a thin and incomplete medial margin and a somewhat thickened lateral margin. The torsion between the proximal body of the pubis and the pubic shass is greater than in
<taxonomicName id="4C0BE8BA31069F0F45F0FA6C3DCEF9D1" box="[1256,1474,1516,1540]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="27" pageNumber="381" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B31069F0F45F0FA6C3DCEF9D1" box="[1256,1474,1516,1540]" italics="true" pageId="27" pageNumber="381">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
(NHMUK PV R 7561b,
<bibRefCitation id="EF9AEEC831069F0F4526F98C3CB5F9F1" author="Crush PJ" box="[1086,1209,1548,1572]" pageId="27" pageNumber="381" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
), such that the obturator foramen cannot be observed in anterior view. There is also some torsion in the pubic shass itself, so that its anterior surface curves very slightly laterally towards the distal end. The pubic shass of SMNS
<quantity id="4CF33EDC31069F0F429EF9093BDFF974" box="[902,979,1673,1697]" metricMagnitude="1" metricUnit="kg" metricValue="1.2596" pageId="27" pageNumber="381" unit="g" value="12596.0">12596g</quantity>
is incomplete distally. The lateral margin of the shass gently curves mediodistally so that it very gradually reduces in transverse width distally. The shass is also gently curved anteroventrally, so that its anterior margin is convex in lateral or medial view.
</paragraph>
<paragraph id="8BB4933931069F084221F8C0397CFDD4" blockId="27.[825,1474,1856,1974]" lastBlockId="28.[113,763,144,1985]" lastPageId="28" lastPageNumber="382" pageId="27" pageNumber="381">
<emphasis id="B97F4F2B31069F0F4221F8C03B73F882" box="[825,895,1856,1879]" italics="true" pageId="27" pageNumber="381">Femur:</emphasis>
Both femora are virtually completely preserved but damaged in SMNS 12597a (
<figureCitation id="13308FBC31069F0F452FF8DF3C8EF8A2" box="[1079,1154,1887,1911]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="27" pageNumber="381">Fig. 2A</figureCitation>
). The right femur is preserved in association with the right ilium and sacral vertebrae proximally, and the right tibia and fibula distally, although the laưer elements have been displaced distally. The proximal end of the less femur has been displaced from the vertebral column and is positioned adjacent to the angulars, but distally it is found in complete articulation with the less tibia and fibula. Both femora of SMNS 12596 (c, j) are also preserved (
<figureCitation id="13308FBC31019F084333FE8D3A70FEF0" box="[555,636,269,293]" captionStart="Figure 3" captionStartId="4.[113,178,1499,1523]" captionTargetBox="[147,1426,144,1471]" captionTargetId="figure-217@4.[146,1426,144,1471]" captionTargetPageId="4" captionText="Figure 3. Part of Saltoposuchus connectens referred specimen SMNS 12596. A, SMNS 12596a, cranial block; B, SMNS 12596b, two dentaries exposed in ventrolateral view; C, SMNS 12596c exposing a less femur in anterolateral view; D, exposing the opposite side of SMNS 12596c, Abbreviations:de, dentary; dr, dorsal rib; dv, dorsal vertebra; fe, femur; ga, gastralia; l., less; mx, maxilla; na, nasal; os, osteoderm; q, quadrate; r., right; sq, squamosal; sr, sacral rib; sv, sacral vertebra; tp, transverse process." figureDoi="http://doi.org/10.5281/zenodo.8426153" httpUri="https://zenodo.org/record/8426153/files/figure.png" pageId="28" pageNumber="382">Figs 3C</figureCitation>
,
<figureCitation id="13308FBC31019F084391FE8D3AB7FEF0" box="[649,699,269,293]" captionStart="Figure 5" captionStartId="6.[113,178,1777,1801]" captionTargetBox="[221,1353,149,1749]" captionTargetId="figure-5@6.[220,1353,149,1749]" captionTargetPageId="6" captionText="Figure 5. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, B, SMNS 12596g, right pubis in (A) anterior and (B) angled medial views; C, SMNS 12596h, right ischium exposed in dorsomedial view and two proximal caudal vertebrae exposed in dorsal view; D, SMNS 12596i, tentative less ulna in posteromedial view; E, F, G, H, SMNS 12596j, right femur in (E) posteromedial, (F) anterolateral, (G) proximal, and (H) distal views; I, proximal half of the right femur in posterolateral view; J, K, SMNS 12596k, right tibia and fibula, (J) with the tibia in posteromedial and the fibula in lateral(?) view and (K) with the tibia in anterolateral and the fibula in medial(?) view. Abbreviations: alcr, anterolateral crest; amt, anteromedial tuber; car, caudal rib; cav, caudal vertebra; cr, crest; crtf, crista tibiofibularis; fi, fibula; gr, groove; ilf, ilium facet; iss, ischial shass; lc, lateral condyle; ltr, lesser trochanter; mc, medial condyle; no, notch; ns, neural spine; obf, obturator foramen; olp, olecranon process; os, osteoderm; poz, postzygapophysis; pr., proximal; prz, prezygapophysis; pus, pubic shass; put, pubic tubercle; r., right; ti, tibia; tr, trochanter." figureDoi="http://doi.org/10.5281/zenodo.8426159" httpUri="https://zenodo.org/record/8426159/files/figure.png" pageId="28" pageNumber="382">5E–I</figureCitation>
), and they are in beưer condition than those in SMNS 12597a. The less femur of SMNS 12596c is preserved in association with disarticulated vertebrae, ribs, and osteoderms. The right femur is completely freed from matrix, and based on its length and congruent morphology belongs to the same individual (Supporting Information,
<tableCitation id="C689A68231019F084018FE49396CFE34" box="[256,352,457,481]" captionStart="Table 1" captionStartId="2.[114,169,143,167]" captionText="Table 1. Summary of the taxonomic assignments in relevant studies of the specimens that now comprise Saltoposuchus connectens" pageId="28" pageNumber="382">Table S1</tableCitation>
). All femora have been considerably transversely compressed.
</paragraph>
<paragraph id="8BB4933931019F084195FD8839D6FC62" blockId="28.[113,763,144,1985]" pageId="28" pageNumber="382">
As noted by previous studies (
<bibRefCitation id="EF9AEEC831019F0840DAFD883A37FDF5" author="Crush PJ" box="[450,571,520,544]" pageId="28" pageNumber="382" refId="ref31157" refString="Crush PJ. An early, terrestrial, crocodile from South Wales. Unpublished Ph. D. Thesis, University College London, 1980." type="book" year="1980">Crush 1980</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831019F08435EFD883AE5FDF5" author="Irmis RB &amp; Nesbiu SJ &amp; Sues H-D" box="[582,745,520,544]" pageId="28" pageNumber="382" pagination="275 - 302" refId="ref31747" refString="Irmis RB, Nesbiu SJ, Sues H-D. Early Crocodylomorpha. Geological Society, London, Special Publications 2013; 379: 275 - 302." type="journal article" year="2013">
Irmis
<emphasis id="B97F4F2B31019F08439AFD893ABDFDF5" box="[642,689,520,544]" italics="true" pageId="28" pageNumber="382">et al.</emphasis>
2013
</bibRefCitation>
), the femur of
<taxonomicName id="4C0BE8BA31019F0841E5FDA739FFFDEA" authorityName="Huene" authorityYear="1921" box="[253,499,551,575]" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="28" pageNumber="382" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B31019F0841E5FDA739FFFDEA" box="[253,499,551,575]" italics="true" pageId="28" pageNumber="382">Saltoposuchus connectens</emphasis>
</taxonomicName>
is considerably more robust (i.e. with a comparatively broad circumference relative to total length) than that of
<taxonomicName id="4C0BE8BA31019F084083FDE638EBFD48" authority="(Irmis et al. 2013: 284)" baseAuthorityName="Irmis" baseAuthorityPageNumber="284" baseAuthorityYear="2013" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="28" pageNumber="382" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B31019F084083FDE63A75FDAB" box="[411,633,614,638]" italics="true" pageId="28" pageNumber="382">Terrestrisuchus gracilis</emphasis>
(
<bibRefCitation id="EF9AEEC831019F084393FDE638D3FD48" author="Irmis RB &amp; Nesbiu SJ &amp; Sues H-D" pageId="28" pageNumber="382" pagination="275 - 302" refId="ref31747" refString="Irmis RB, Nesbiu SJ, Sues H-D. Early Crocodylomorpha. Geological Society, London, Special Publications 2013; 379: 275 - 302." type="journal article" year="2013">
Irmis
<emphasis id="B97F4F2B31019F0843D1FDE73AF7FDAB" box="[713,763,614,638]" italics="true" pageId="28" pageNumber="382">et al.</emphasis>
2013: 284
</bibRefCitation>
)
</taxonomicName>
. The femur is longer than the tibia and fibula (
<tableCitation id="C689A68231019F0843DAFD053872FD68" captionStart="Table 2" captionStartId="7.[131,186,143,167]" captionText="Table 2. Comparison of measurements among the Saltoposuchus connectensspecimens. A full list of measurements is presented in the Supporting Information, Table S1" pageId="28" pageNumber="382">Table 2</tableCitation>
;
<bibRefCitation id="EF9AEEC831019F084188FD253988FD68" author="Sereno PC &amp; Wild R" box="[144,388,676,701]" pageId="28" pageNumber="382" pagination="435 - 58" refId="ref32722" refString="Sereno PC, Wild R. Procompsognathus: theropod, ' thecodont' or both? Journal of Vertebrate Paleontology 1992; 12: 435 - 58." type="journal article" year="1992">Sereno and Wild 1992</bibRefCitation>
) as in
<taxonomicName id="4C0BE8BA31019F0840C0FD243AC6FD69" box="[472,714,676,700]" class="Reptilia" family="Sphenosuchidae" genus="Dromicosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="28" pageNumber="382" phylum="Chordata" rank="species" species="grallator">
<emphasis id="B97F4F2B31019F0840C0FD243AC6FD69" box="[472,714,676,700]" italics="true" pageId="28" pageNumber="382">Dromicosuchus grallator</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0BE8BA31019F084169FD443986FD09" authorityName="Bonaparte" authorityYear="1969" box="[113,394,708,732]" class="Polyplacophora" genus="Pseudhesperosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="28" pageNumber="382" phylum="Mollusca" rank="species" species="jachaleri">
<emphasis id="B97F4F2B31019F084169FD443986FD09" box="[113,394,708,732]" italics="true" pageId="28" pageNumber="382">Pseudhesperosuchus jachaleri</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9AEEC831019F084083FD443A34FD09" author="Sues H-D &amp; Olsen PE &amp; Carter JG" box="[411,568,708,732]" pageId="28" pageNumber="382" pagination="329 - 43" refId="ref32828" refString="Sues H-D, Olsen PE, Carter JG et al. A new crocodylomorph archosaur from the Upper Triassic of North Carolina. Journal of Vertebrate Paleontology 2003; 23: 329 - 43." type="journal article" year="2003">
Sues
<emphasis id="B97F4F2B31019F0840C8FD4539F3FD09" box="[464,511,708,732]" italics="true" pageId="28" pageNumber="382">et al.</emphasis>
2003
</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831019F08435BFD443AFFFD09" author="Leardi JM &amp; Pol D &amp; Clark JM" box="[579,755,708,732]" pageId="28" pageNumber="382" pagination="2801" refId="ref31922" refString="Leardi JM, Pol D, Clark JM. Detailed anatomy of the braincase of Macelognathus vagans Marsh, 1884 (Archosauria, Crocodylomorpha) using high resolution tomography and new insights on basal crocodylomorph phylogeny. PeerJ 2017; 5: e 2801." type="journal article" year="2017">
Leardi
<emphasis id="B97F4F2B31019F084393FD453AB6FD09" box="[651,698,708,732]" italics="true" pageId="28" pageNumber="382">et al.</emphasis>
2017
</bibRefCitation>
: supplementary information therein), whereas in
<taxonomicName id="4C0BE8BA31019F084371FD6338B9FCCF" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="28" pageNumber="382" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B31019F084371FD6338B9FCCF" italics="true" pageId="28" pageNumber="382">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0BE8BA31019F0841DAFC823954FCCF" baseAuthorityName="Marsh" baseAuthorityYear="1877" box="[194,344,770,794]" class="Reptilia" family="Coeluridae" genus="Hallopus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="28" pageNumber="382" phylum="Chordata" rank="species" species="victor">
<emphasis id="B97F4F2B31019F0841DAFC823954FCCF" box="[194,344,770,794]" italics="true" pageId="28" pageNumber="382">Hallopus victor</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C0BE8BA31019F08408BFC823A63FCCF" authorityName="Marsh" authorityYear="1884" box="[403,623,770,794]" class="Reptilia" family="Hypsilophodontidae" genus="Macelognathus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="28" pageNumber="382" phylum="Chordata" rank="species" species="vagans">
<emphasis id="B97F4F2B31019F08408BFC823A63FCCF" box="[403,623,770,794]" italics="true" pageId="28" pageNumber="382">Macelognathus vagans</emphasis>
</taxonomicName>
the tibia and fibula are longer than the femur (NHMUK PV R 7551,
<bibRefCitation id="EF9AEEC831019F0843AAFCA238A9FC8C" author="Walker AD" pageId="28" pageNumber="382" pagination="323 - 72" refId="ref32988" refString="Walker AD. A revision of the Jurassic reptile Hallopus victor (Marsh), with remarks on the classification of crocodiles. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1970; 257: 323 - 72." type="journal article" year="1970">Walker 1970</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831019F0841A8FCC13963FC8C" author="Gohlich UB &amp; Chiappe LM &amp; Clark JM" box="[176,367,833,857]" pageId="28" pageNumber="382" pagination="307 - 21" refId="ref31388" refString="Gohlich UB, Chiappe LM, Clark JM et al. The systematic position of the Late Jurassic alleged dinosaur Macelognathus (Crocodylomorpha: Sphenosuchia). Canadian Journal of Earth Sciences 2005; 42: 307 - 21." type="journal article" year="2005">
Göhlich
<emphasis id="B97F4F2B31019F084011FCC2393BFC8C" box="[265,311,833,857]" italics="true" pageId="28" pageNumber="382">et al.</emphasis>
2005
</bibRefCitation>
). The femoral shass is distinctly curved in anterolateral or posteromedial view, so that the anteromedial margin of the shass is clearly convex and the posterolateral margin clearly concave (
<figureCitation id="13308FBC31019F084071FC1F39C5FC62" box="[361,457,927,951]" captionStart="Figure 5" captionStartId="6.[113,178,1777,1801]" captionTargetBox="[221,1353,149,1749]" captionTargetId="figure-5@6.[220,1353,149,1749]" captionTargetPageId="6" captionText="Figure 5. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, B, SMNS 12596g, right pubis in (A) anterior and (B) angled medial views; C, SMNS 12596h, right ischium exposed in dorsomedial view and two proximal caudal vertebrae exposed in dorsal view; D, SMNS 12596i, tentative less ulna in posteromedial view; E, F, G, H, SMNS 12596j, right femur in (E) posteromedial, (F) anterolateral, (G) proximal, and (H) distal views; I, proximal half of the right femur in posterolateral view; J, K, SMNS 12596k, right tibia and fibula, (J) with the tibia in posteromedial and the fibula in lateral(?) view and (K) with the tibia in anterolateral and the fibula in medial(?) view. Abbreviations: alcr, anterolateral crest; amt, anteromedial tuber; car, caudal rib; cav, caudal vertebra; cr, crest; crtf, crista tibiofibularis; fi, fibula; gr, groove; ilf, ilium facet; iss, ischial shass; lc, lateral condyle; ltr, lesser trochanter; mc, medial condyle; no, notch; ns, neural spine; obf, obturator foramen; olp, olecranon process; os, osteoderm; poz, postzygapophysis; pr., proximal; prz, prezygapophysis; pus, pubic shass; put, pubic tubercle; r., right; ti, tibia; tr, trochanter." figureDoi="http://doi.org/10.5281/zenodo.8426159" httpUri="https://zenodo.org/record/8426159/files/figure.png" pageId="28" pageNumber="382">Fig. 5E, F</figureCitation>
).
</paragraph>
<paragraph id="8BB4933931019F084195FC3E3A40F9DF" blockId="28.[113,763,144,1985]" pageId="28" pageNumber="382">
The proximal femoral head is strongly inturned and directed anteromedially to articulate with the acetabulum. The proximal surface of the proximal head is convex and gradually narrows anteromedially to posterolaterally (
<figureCitation id="13308FBC31019F0840CFFB9C3A2EFBE1" box="[471,546,1052,1076]" captionStart="Figure 5" captionStartId="6.[113,178,1777,1801]" captionTargetBox="[221,1353,149,1749]" captionTargetId="figure-5@6.[220,1353,149,1749]" captionTargetPageId="6" captionText="Figure 5. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, B, SMNS 12596g, right pubis in (A) anterior and (B) angled medial views; C, SMNS 12596h, right ischium exposed in dorsomedial view and two proximal caudal vertebrae exposed in dorsal view; D, SMNS 12596i, tentative less ulna in posteromedial view; E, F, G, H, SMNS 12596j, right femur in (E) posteromedial, (F) anterolateral, (G) proximal, and (H) distal views; I, proximal half of the right femur in posterolateral view; J, K, SMNS 12596k, right tibia and fibula, (J) with the tibia in posteromedial and the fibula in lateral(?) view and (K) with the tibia in anterolateral and the fibula in medial(?) view. Abbreviations: alcr, anterolateral crest; amt, anteromedial tuber; car, caudal rib; cav, caudal vertebra; cr, crest; crtf, crista tibiofibularis; fi, fibula; gr, groove; ilf, ilium facet; iss, ischial shass; lc, lateral condyle; ltr, lesser trochanter; mc, medial condyle; no, notch; ns, neural spine; obf, obturator foramen; olp, olecranon process; os, osteoderm; poz, postzygapophysis; pr., proximal; prz, prezygapophysis; pus, pubic shass; put, pubic tubercle; r., right; ti, tibia; tr, trochanter." figureDoi="http://doi.org/10.5281/zenodo.8426159" httpUri="https://zenodo.org/record/8426159/files/figure.png" pageId="28" pageNumber="382">Fig. 5G</figureCitation>
). Of the three tubera typically present on the proximal head of the femur in archosaurs (
<bibRefCitation id="EF9AEEC831019F0841A2FBDB394FFBA6" author="Nesbiu SJ" box="[186,323,1115,1139]" pageId="28" pageNumber="382" pagination="1 - 292" refId="ref32089" refString="Nesbiu SJ. The early evolution of archosaurs: relationships and the origin of major clades. Bulletin of the American Museum of Natural History 2011; 352: 1 - 292." type="journal article" year="2011">Nesbiư 2011</bibRefCitation>
: chs 300–302), only a poorly developed anteromedial tuber can be observed on the right femur SMNS 12596j due to compression. SMNS 12597a exposes a clearly defined posteromedial tuber on its less femur and a similarly defined anterolateral tuber on its right femur. The posterolateral margin bears a longitudinal, rugose thickening directly distal to the proximal head. It is positioned on the lateral side of the margin and represents the lesser or dorsolateral trochanter (
<figureCitation id="13308FBC31019F084349FAB63AABFA9B" box="[593,679,1334,1358]" captionStart="Figure 5" captionStartId="6.[113,178,1777,1801]" captionTargetBox="[221,1353,149,1749]" captionTargetId="figure-5@6.[220,1353,149,1749]" captionTargetPageId="6" captionText="Figure 5. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, B, SMNS 12596g, right pubis in (A) anterior and (B) angled medial views; C, SMNS 12596h, right ischium exposed in dorsomedial view and two proximal caudal vertebrae exposed in dorsal view; D, SMNS 12596i, tentative less ulna in posteromedial view; E, F, G, H, SMNS 12596j, right femur in (E) posteromedial, (F) anterolateral, (G) proximal, and (H) distal views; I, proximal half of the right femur in posterolateral view; J, K, SMNS 12596k, right tibia and fibula, (J) with the tibia in posteromedial and the fibula in lateral(?) view and (K) with the tibia in anterolateral and the fibula in medial(?) view. Abbreviations: alcr, anterolateral crest; amt, anteromedial tuber; car, caudal rib; cav, caudal vertebra; cr, crest; crtf, crista tibiofibularis; fi, fibula; gr, groove; ilf, ilium facet; iss, ischial shass; lc, lateral condyle; ltr, lesser trochanter; mc, medial condyle; no, notch; ns, neural spine; obf, obturator foramen; olp, olecranon process; os, osteoderm; poz, postzygapophysis; pr., proximal; prz, prezygapophysis; pus, pubic shass; put, pubic tubercle; r., right; ti, tibia; tr, trochanter." figureDoi="http://doi.org/10.5281/zenodo.8426159" httpUri="https://zenodo.org/record/8426159/files/figure.png" pageId="28" pageNumber="382">Fig. 5F, I</figureCitation>
), which occurs widely among non-crocodyliform crocodylomorphs and represents an aưachment site for M. puboischiofemoralis internus 2 (NHMUK PV R 7562,
<bibRefCitation id="EF9AEEC831019F0840F9FA143AAFFA79" author="Hutchinson JR" box="[481,675,1428,1452]" pageId="28" pageNumber="382" pagination="169 - 97" refId="ref31675" refString="Hutchinson JR. The evolution of femoral osteology and soss tissues on the line to extant birds (Neornithes). Zoological Journal of the Linnean Society 2001 a; 131: 169 - 97." type="journal article" year="2001">Hutchinson 2001a</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831019F0843A9FA1438C9FA19" author="Sues H-D &amp; Olsen PE &amp; Carter JG" pageId="28" pageNumber="382" pagination="329 - 43" refId="ref32828" refString="Sues H-D, Olsen PE, Carter JG et al. A new crocodylomorph archosaur from the Upper Triassic of North Carolina. Journal of Vertebrate Paleontology 2003; 23: 329 - 43." type="journal article" year="2003">
Sues
<emphasis id="B97F4F2B31019F0843F1FA153886FA19" italics="true" pageId="28" pageNumber="382">et al.</emphasis>
2003
</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831019F0841CBFA343991FA1E" author="Gohlich UB &amp; Chiappe LM &amp; Clark JM" box="[211,413,1459,1484]" pageId="28" pageNumber="382" pagination="307 - 21" refId="ref31388" refString="Gohlich UB, Chiappe LM, Clark JM et al. The systematic position of the Late Jurassic alleged dinosaur Macelognathus (Crocodylomorpha: Sphenosuchia). Canadian Journal of Earth Sciences 2005; 42: 307 - 21." type="journal article" year="2005">
Göhlich
<emphasis id="B97F4F2B31019F084037FA34396DFA19" box="[303,353,1460,1484]" italics="true" pageId="28" pageNumber="382">et al.</emphasis>
2005
</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831019F0840B3FA343A4EFA19" author="Pol D &amp; Rauhut OWM &amp; Lecuona A" box="[427,578,1460,1484]" pageId="28" pageNumber="382" pagination="862 - 72" refId="ref32359" refString="Pol D, Rauhut OWM, Lecuona A et al. A new fossil from the Jurassic of Patagonia reveals the early basicranial evolution and the origins of Crocodyliformes. Biological Reviews 2013; 88: 862 - 72." type="journal article" year="2013">
Pol
<emphasis id="B97F4F2B31019F0840CDFA343A0BFA19" box="[469,519,1460,1484]" italics="true" pageId="28" pageNumber="382">et al.</emphasis>
2013
</bibRefCitation>
). The fourth trochanter is low and mount-like and the surface of the shass at its anteromedial base is quite concave (
<figureCitation id="13308FBC31019F0840F9FA723A30F9DF" box="[481,572,1522,1546]" captionStart="Figure 5" captionStartId="6.[113,178,1777,1801]" captionTargetBox="[221,1353,149,1749]" captionTargetId="figure-5@6.[220,1353,149,1749]" captionTargetPageId="6" captionText="Figure 5. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, B, SMNS 12596g, right pubis in (A) anterior and (B) angled medial views; C, SMNS 12596h, right ischium exposed in dorsomedial view and two proximal caudal vertebrae exposed in dorsal view; D, SMNS 12596i, tentative less ulna in posteromedial view; E, F, G, H, SMNS 12596j, right femur in (E) posteromedial, (F) anterolateral, (G) proximal, and (H) distal views; I, proximal half of the right femur in posterolateral view; J, K, SMNS 12596k, right tibia and fibula, (J) with the tibia in posteromedial and the fibula in lateral(?) view and (K) with the tibia in anterolateral and the fibula in medial(?) view. Abbreviations: alcr, anterolateral crest; amt, anteromedial tuber; car, caudal rib; cav, caudal vertebra; cr, crest; crtf, crista tibiofibularis; fi, fibula; gr, groove; ilf, ilium facet; iss, ischial shass; lc, lateral condyle; ltr, lesser trochanter; mc, medial condyle; no, notch; ns, neural spine; obf, obturator foramen; olp, olecranon process; os, osteoderm; poz, postzygapophysis; pr., proximal; prz, prezygapophysis; pus, pubic shass; put, pubic tubercle; r., right; ti, tibia; tr, trochanter." figureDoi="http://doi.org/10.5281/zenodo.8426159" httpUri="https://zenodo.org/record/8426159/files/figure.png" pageId="28" pageNumber="382">Fig. 5E, I</figureCitation>
).
</paragraph>
<paragraph id="8BB4933931019F084195F9923C11FF12" blockId="28.[113,763,144,1985]" lastBlockId="28.[810,1459,143,199]" pageId="28" pageNumber="382">
The compressed distal end of the right femur of SMNS 12596j bears distinct fibular and tibial condyles, which are positioned on the lateral and medial side of the distal end, respectively (
<figureCitation id="13308FBC31019F0841ABF9EF390DF952" box="[179,257,1647,1671]" captionStart="Figure 5" captionStartId="6.[113,178,1777,1801]" captionTargetBox="[221,1353,149,1749]" captionTargetId="figure-5@6.[220,1353,149,1749]" captionTargetPageId="6" captionText="Figure 5. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, B, SMNS 12596g, right pubis in (A) anterior and (B) angled medial views; C, SMNS 12596h, right ischium exposed in dorsomedial view and two proximal caudal vertebrae exposed in dorsal view; D, SMNS 12596i, tentative less ulna in posteromedial view; E, F, G, H, SMNS 12596j, right femur in (E) posteromedial, (F) anterolateral, (G) proximal, and (H) distal views; I, proximal half of the right femur in posterolateral view; J, K, SMNS 12596k, right tibia and fibula, (J) with the tibia in posteromedial and the fibula in lateral(?) view and (K) with the tibia in anterolateral and the fibula in medial(?) view. Abbreviations: alcr, anterolateral crest; amt, anteromedial tuber; car, caudal rib; cav, caudal vertebra; cr, crest; crtf, crista tibiofibularis; fi, fibula; gr, groove; ilf, ilium facet; iss, ischial shass; lc, lateral condyle; ltr, lesser trochanter; mc, medial condyle; no, notch; ns, neural spine; obf, obturator foramen; olp, olecranon process; os, osteoderm; poz, postzygapophysis; pr., proximal; prz, prezygapophysis; pus, pubic shass; put, pubic tubercle; r., right; ti, tibia; tr, trochanter." figureDoi="http://doi.org/10.5281/zenodo.8426159" httpUri="https://zenodo.org/record/8426159/files/figure.png" pageId="28" pageNumber="382">Fig. 5H</figureCitation>
). They are separated by a distinct groove on the distal surface that curves proximally onto the anterior surface of the shass where it continues proximally for a considerable length. The two condyles are separated by a strong concavity on the posterior margin of the distal end. In distal view, the lateral condyle is considerably narrower and reaches further posteriorly than the medial condyle. Although this is strongly exaggerated by compression, this would have probably also been the case in life to a lesser extent, as can be deduced from the well preserved femur of
<taxonomicName id="4C0BE8BA31019F0841D5F80939A4F874" box="[205,424,1929,1953]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="28" pageNumber="382" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B31019F0841D5F80939A4F874" box="[205,424,1929,1953]" italics="true" pageId="28" pageNumber="382">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
(NHMUK PV R 37685). A large crista tibiofibularis is projected from the lateral side of the lateral condyle. It is separated from the posterior end of the lateral condyle by a distinct notch.
</paragraph>
<paragraph id="8BB4933931019F084232FF6B3BA2FCA3" blockId="28.[810,1460,235,1982]" pageId="28" pageNumber="382">
<emphasis id="B97F4F2B31019F084232FF6B3BDEFED6" box="[810,978,235,259]" italics="true" pageId="28" pageNumber="382">Tibia and fibula:</emphasis>
Both pairs of tibiae and fibulae are preserved in articulation in SMNS 12597a. The right pair lacks the distal ends of both elements, whereas the less pair is complete, although the central portion of the fibular shass is completely covered in matrix. The less pair is preserved in articulation with the less femur proximally and less tarsus distally (
<figureCitation id="13308FBC31019F08443CFE083D86FE75" box="[1316,1418,392,416]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="28" pageNumber="382">Fig. 2A, B</figureCitation>
). A less tibia and fibula are fully freed from matrix and preserved in partial articulation in SMNS 12596k (
<figureCitation id="13308FBC31019F0845B5FE473D05FE0B" box="[1197,1289,455,479]" captionStart="Figure 5" captionStartId="6.[113,178,1777,1801]" captionTargetBox="[221,1353,149,1749]" captionTargetId="figure-5@6.[220,1353,149,1749]" captionTargetPageId="6" captionText="Figure 5. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, B, SMNS 12596g, right pubis in (A) anterior and (B) angled medial views; C, SMNS 12596h, right ischium exposed in dorsomedial view and two proximal caudal vertebrae exposed in dorsal view; D, SMNS 12596i, tentative less ulna in posteromedial view; E, F, G, H, SMNS 12596j, right femur in (E) posteromedial, (F) anterolateral, (G) proximal, and (H) distal views; I, proximal half of the right femur in posterolateral view; J, K, SMNS 12596k, right tibia and fibula, (J) with the tibia in posteromedial and the fibula in lateral(?) view and (K) with the tibia in anterolateral and the fibula in medial(?) view. Abbreviations: alcr, anterolateral crest; amt, anteromedial tuber; car, caudal rib; cav, caudal vertebra; cr, crest; crtf, crista tibiofibularis; fi, fibula; gr, groove; ilf, ilium facet; iss, ischial shass; lc, lateral condyle; ltr, lesser trochanter; mc, medial condyle; no, notch; ns, neural spine; obf, obturator foramen; olp, olecranon process; os, osteoderm; poz, postzygapophysis; pr., proximal; prz, prezygapophysis; pus, pubic shass; put, pubic tubercle; r., right; ti, tibia; tr, trochanter." figureDoi="http://doi.org/10.5281/zenodo.8426159" httpUri="https://zenodo.org/record/8426159/files/figure.png" pageId="28" pageNumber="382">Fig. 5J, K</figureCitation>
). Finally, SMNS 55009e is a complete, well preserved right tibia that is fully freed from matrix (
<figureCitation id="13308FBC31019F0842A2FD853C2EFDC8" box="[954,1058,517,541]" captionStart="Figure 6" captionStartId="7.[129,194,1605,1629]" captionTargetBox="[161,1441,657,1577]" captionTargetId="figure-310@7.[161,1441,657,1577]" captionTargetPageId="7" captionText="Figure 6. Part of Saltoposuchus connectens referred specimen SMNS 55009. A, SMNS 55009a, dorsal vertebral centrum in less lateral view; B, SMNS 55009d, proximal caudal vertebral centrum in angled lateral view; C, SMNS 55009b, first sacral vertebra in less lateral view; D, interpretative drawing of (C); E, SMNS 55009c, proximal caudal vertebra in posterior view; F, interpretative drawing of (E). Abbreviations: car, caudal rib; ns, neural spine; pas, posterior articular surface; poz, postzygapophysis; prz, prezygapophysis; sr, sacral rib; sut, suture." figureDoi="http://doi.org/10.5281/zenodo.8426161" httpUri="https://zenodo.org/record/8426161/files/figure.png" pageId="28" pageNumber="382">Fig. 6A–F</figureCitation>
). The tibia and fibula are subequal in length, but the tibia is considerably more robust (Supporting Information,
<tableCitation id="C689A68231019F0842AAFDC43C06FD89" box="[946,1034,580,604]" captionStart="Table 1" captionStartId="2.[114,169,143,167]" captionText="Table 1. Summary of the taxonomic assignments in relevant studies of the specimens that now comprise Saltoposuchus connectens" pageId="28" pageNumber="382">Table S1</tableCitation>
). The proximal ends of both elements are curved relative to the shass, so that the posterior margin directly distal to the proximal end is concave, and the opposite anterior end convex. The fibula of SMNS 12596k is apparently shissed relative to the tibia, so that its concave posterior margin faces the equivalent margin on the tibia (
<figureCitation id="13308FBC31019F08456BFD613CDEFD2D" box="[1139,1234,736,761]" captionStart="Figure 5" captionStartId="6.[113,178,1777,1801]" captionTargetBox="[221,1353,149,1749]" captionTargetId="figure-5@6.[220,1353,149,1749]" captionTargetPageId="6" captionText="Figure 5. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, B, SMNS 12596g, right pubis in (A) anterior and (B) angled medial views; C, SMNS 12596h, right ischium exposed in dorsomedial view and two proximal caudal vertebrae exposed in dorsal view; D, SMNS 12596i, tentative less ulna in posteromedial view; E, F, G, H, SMNS 12596j, right femur in (E) posteromedial, (F) anterolateral, (G) proximal, and (H) distal views; I, proximal half of the right femur in posterolateral view; J, K, SMNS 12596k, right tibia and fibula, (J) with the tibia in posteromedial and the fibula in lateral(?) view and (K) with the tibia in anterolateral and the fibula in medial(?) view. Abbreviations: alcr, anterolateral crest; amt, anteromedial tuber; car, caudal rib; cav, caudal vertebra; cr, crest; crtf, crista tibiofibularis; fi, fibula; gr, groove; ilf, ilium facet; iss, ischial shass; lc, lateral condyle; ltr, lesser trochanter; mc, medial condyle; no, notch; ns, neural spine; obf, obturator foramen; olp, olecranon process; os, osteoderm; poz, postzygapophysis; pr., proximal; prz, prezygapophysis; pus, pubic shass; put, pubic tubercle; r., right; ti, tibia; tr, trochanter." figureDoi="http://doi.org/10.5281/zenodo.8426159" httpUri="https://zenodo.org/record/8426159/files/figure.png" pageId="28" pageNumber="382">Fig. 5J, K</figureCitation>
). Distally, both elements also gradually curve anteriorly and as such have a somewhat sigmoidal outline. The curvature of the tibia SMNS 55009e is much reduced due to its strong anteroposterior compression (
<figureCitation id="13308FBC31019F08422DFCDE3B91FCA3" box="[821,925,862,886]" captionStart="Figure 7" captionStartId="8.[113,178,1656,1680]" captionTargetBox="[146,1426,144,1628]" captionTargetId="figure-102@8.[146,1426,144,1628]" captionTargetPageId="8" captionText="Figure 7. Saltoposuchus connectens referred specimen SMNS 55009, continued. A, B, C, D, E, F, SMNS 55009e, right tibia in (A) posterior, (B) anterior, (C) medial, (D) lateral, (E) proximal, and (F) distal views; G, H, SMNS 55009h, metatarsal block in (G) dorsal(?) and (H) opposing views; I, J, K, L, SMNS 55009g, right calcaneum in (I) medial, (J) lateral, (K) proximal, and (L) distal views. Abbreviations: als, anterolateral side; asf, astragalus facet; calt, calcaneal tuber; gr, groove; latc, lateral condyle; lg, longitudinal groove; mt, metatarsal; no, notch; pme, posterior extension; pms, posteromedial side; tr, trough." figureDoi="http://doi.org/10.5281/zenodo.8426163" httpUri="https://zenodo.org/record/8426163/files/figure.png" pageId="28" pageNumber="382">Fig. 7A–D</figureCitation>
).
</paragraph>
<paragraph id="8BB4933931019F08425DFCFD3C4CF936" blockId="28.[810,1460,235,1982]" pageId="28" pageNumber="382">
The proximal surface of the tibia is slightly concave. The anterior margin of the proximal head is widely rounded and thus convex in proximal view, whereas the posterior side possesses two projections, which represent the posterior expansion on the posteromedial end and the lateral condyle on the posterolateral end respectively (
<figureCitation id="13308FBC31019F0842C4FB9A3C29FBE4" box="[988,1061,1050,1074]" captionStart="Figure 7" captionStartId="8.[113,178,1656,1680]" captionTargetBox="[146,1426,144,1628]" captionTargetId="figure-102@8.[146,1426,144,1628]" captionTargetPageId="8" captionText="Figure 7. Saltoposuchus connectens referred specimen SMNS 55009, continued. A, B, C, D, E, F, SMNS 55009e, right tibia in (A) posterior, (B) anterior, (C) medial, (D) lateral, (E) proximal, and (F) distal views; G, H, SMNS 55009h, metatarsal block in (G) dorsal(?) and (H) opposing views; I, J, K, L, SMNS 55009g, right calcaneum in (I) medial, (J) lateral, (K) proximal, and (L) distal views. Abbreviations: als, anterolateral side; asf, astragalus facet; calt, calcaneal tuber; gr, groove; latc, lateral condyle; lg, longitudinal groove; mt, metatarsal; no, notch; pme, posterior extension; pms, posteromedial side; tr, trough." figureDoi="http://doi.org/10.5281/zenodo.8426163" httpUri="https://zenodo.org/record/8426163/files/figure.png" pageId="28" pageNumber="382">Figs 7E</figureCitation>
,
<figureCitation id="13308FBC31019F084537FB9A3C5BFBE7" box="[1071,1111,1050,1074]" captionStart="Figure 12" captionStartId="24.[113,178,1772,1796]" captionTargetBox="[151,1424,144,1744]" captionTargetId="figure-6@24.[148,1424,144,1744]" captionTargetPageId="24" captionText="Figure 12. Details of the postcranial anatomy of Saltoposuchus connectens. A, dorsal vertebral neural arch and dorsal rib of SMNS 12596c in ventral view; B, associated osteoderms of SMNS 12596c in dorsal view; C, sacral ribs and associated elements of SMNS 12597a in dorsal view; D, distal portion of right scapular blade of SMNS 12597a in lateral view; E, proximal end of right tibia of SMNS 12597a in proximal view; F, less tarsus and pes of SMNS 12597a with the calcaneum exposed in lateral view; G, close-up of less tarsus of SMNS 12597a; H, close-up of less pes of SMNS 12597a. Abbreviations: ane, anterior expansion; cal, calcaneum; calt, calcaneal tuber; cp, capitulum; dp, diapophysis; dt, distal tarsal; exp, extensor pit; fe, femur; fi, fibula; il, ilium; lam, lamina; latc, lateral condyle; le, lateral edge; lr, longitudinal ridge; me, medial edge; mt, metatarsal; no, notch; pap, parapophysis; ph, phalanx; pme, posterior extension; poe, posterior expansion; pofl, posterior flange; poz, postzygapophysis; os, osteoderm; r., right; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426175" httpUri="https://zenodo.org/record/8426175/files/figure.png" pageId="28" pageNumber="382">12E</figureCitation>
). The posterior expansion extends quite far beyond the shass in all tibiae of
<taxonomicName id="4C0BE8BA31019F0845A2FBB93DA1FB84" authorityName="Huene" authorityYear="1921" box="[1210,1453,1081,1105]" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="28" pageNumber="382" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B31019F0845A2FBB93DA1FB84" box="[1210,1453,1081,1105]" italics="true" pageId="28" pageNumber="382">Saltoposuchus connectens</emphasis>
</taxonomicName>
, and a similar projection is also present in
<taxonomicName id="4C0BE8BA31019F0845C0FBD83DBFFBA5" box="[1240,1459,1112,1136]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="28" pageNumber="382" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B31019F0845C0FBD83DBFFBA5" box="[1240,1459,1112,1136]" italics="true" pageId="28" pageNumber="382">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0BE8BA31019F084241FBF83C44FB45" box="[857,1096,1144,1168]" class="Reptilia" family="Sphenosuchidae" genus="Dromicosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="28" pageNumber="382" phylum="Chordata" rank="species" species="grallator">
<emphasis id="B97F4F2B31019F084241FBF83C44FB45" box="[857,1096,1144,1168]" italics="true" pageId="28" pageNumber="382">Dromicosuchus grallator</emphasis>
</taxonomicName>
(NHMUK PV R 7566,
<bibRefCitation id="EF9AEEC831019F084453FBF83B52FB7A" author="Sues H-D &amp; Olsen PE &amp; Carter JG" pageId="28" pageNumber="382" pagination="329 - 43" refId="ref32828" refString="Sues H-D, Olsen PE, Carter JG et al. A new crocodylomorph archosaur from the Upper Triassic of North Carolina. Journal of Vertebrate Paleontology 2003; 23: 329 - 43." type="journal article" year="2003">
Sues
<emphasis id="B97F4F2B31019F08449AFBF93DB8FB45" box="[1410,1460,1144,1168]" italics="true" pageId="28" pageNumber="382">et al.</emphasis>
2003
</bibRefCitation>
). The markedly exaggerated expansion in SMNS 55009e is the result of compression. The posterior expansion and lateral condyle are separated by a deep concavity in proximal view, and this concavity continues distally along the posterior surface of the shass for some distance as a trough (
<figureCitation id="13308FBC31019F0845A4FA943D08FAF8" box="[1212,1284,1300,1325]" captionStart="Figure 7" captionStartId="8.[113,178,1656,1680]" captionTargetBox="[146,1426,144,1628]" captionTargetId="figure-102@8.[146,1426,144,1628]" captionTargetPageId="8" captionText="Figure 7. Saltoposuchus connectens referred specimen SMNS 55009, continued. A, B, C, D, E, F, SMNS 55009e, right tibia in (A) posterior, (B) anterior, (C) medial, (D) lateral, (E) proximal, and (F) distal views; G, H, SMNS 55009h, metatarsal block in (G) dorsal(?) and (H) opposing views; I, J, K, L, SMNS 55009g, right calcaneum in (I) medial, (J) lateral, (K) proximal, and (L) distal views. Abbreviations: als, anterolateral side; asf, astragalus facet; calt, calcaneal tuber; gr, groove; latc, lateral condyle; lg, longitudinal groove; mt, metatarsal; no, notch; pme, posterior extension; pms, posteromedial side; tr, trough." figureDoi="http://doi.org/10.5281/zenodo.8426163" httpUri="https://zenodo.org/record/8426163/files/figure.png" pageId="28" pageNumber="382">Fig. 7A</figureCitation>
). The lateral side of the proximal head also possesses a small but distinct notch in proximal view (
<bibRefCitation id="EF9AEEC831019F0842FFFAD33C6EFABE" author="Crush PJ" box="[999,1122,1363,1387]" pageId="28" pageNumber="382" refId="ref31157" refString="Crush PJ. An early, terrestrial, crocodile from South Wales. Unpublished Ph. D. Thesis, University College London, 1980." type="book" year="1980">Crush 1980</bibRefCitation>
;
<figureCitation id="13308FBC31019F084568FAD33CB0FABE" box="[1136,1212,1363,1387]" captionStart="Figure 7" captionStartId="8.[113,178,1656,1680]" captionTargetBox="[146,1426,144,1628]" captionTargetId="figure-102@8.[146,1426,144,1628]" captionTargetPageId="8" captionText="Figure 7. Saltoposuchus connectens referred specimen SMNS 55009, continued. A, B, C, D, E, F, SMNS 55009e, right tibia in (A) posterior, (B) anterior, (C) medial, (D) lateral, (E) proximal, and (F) distal views; G, H, SMNS 55009h, metatarsal block in (G) dorsal(?) and (H) opposing views; I, J, K, L, SMNS 55009g, right calcaneum in (I) medial, (J) lateral, (K) proximal, and (L) distal views. Abbreviations: als, anterolateral side; asf, astragalus facet; calt, calcaneal tuber; gr, groove; latc, lateral condyle; lg, longitudinal groove; mt, metatarsal; no, notch; pme, posterior extension; pms, posteromedial side; tr, trough." figureDoi="http://doi.org/10.5281/zenodo.8426163" httpUri="https://zenodo.org/record/8426163/files/figure.png" pageId="28" pageNumber="382">Figs 7E</figureCitation>
,
<figureCitation id="13308FBC31019F0845D1FAD33CFDFABE" box="[1225,1265,1363,1387]" captionStart="Figure 12" captionStartId="24.[113,178,1772,1796]" captionTargetBox="[151,1424,144,1744]" captionTargetId="figure-6@24.[148,1424,144,1744]" captionTargetPageId="24" captionText="Figure 12. Details of the postcranial anatomy of Saltoposuchus connectens. A, dorsal vertebral neural arch and dorsal rib of SMNS 12596c in ventral view; B, associated osteoderms of SMNS 12596c in dorsal view; C, sacral ribs and associated elements of SMNS 12597a in dorsal view; D, distal portion of right scapular blade of SMNS 12597a in lateral view; E, proximal end of right tibia of SMNS 12597a in proximal view; F, less tarsus and pes of SMNS 12597a with the calcaneum exposed in lateral view; G, close-up of less tarsus of SMNS 12597a; H, close-up of less pes of SMNS 12597a. Abbreviations: ane, anterior expansion; cal, calcaneum; calt, calcaneal tuber; cp, capitulum; dp, diapophysis; dt, distal tarsal; exp, extensor pit; fe, femur; fi, fibula; il, ilium; lam, lamina; latc, lateral condyle; le, lateral edge; lr, longitudinal ridge; me, medial edge; mt, metatarsal; no, notch; pap, parapophysis; ph, phalanx; pme, posterior extension; poe, posterior expansion; pofl, posterior flange; poz, postzygapophysis; os, osteoderm; r., right; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426175" httpUri="https://zenodo.org/record/8426175/files/figure.png" pageId="28" pageNumber="382">12E</figureCitation>
) and such a notch is absent in
<taxonomicName id="4C0BE8BA31019F0842BAFAF23C71FA5F" box="[930,1149,1394,1418]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="28" pageNumber="382" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B31019F0842BAFAF23C71FA5F" box="[930,1149,1394,1418]" italics="true" pageId="28" pageNumber="382">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
(NHMUK PV R 7566,
<bibRefCitation id="EF9AEEC831019F08446BFAF23B52FA7F" author="Crush PJ" pageId="28" pageNumber="382" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
). The distal end of the tibia is composed of two distal projections, one on the anterolateral side and the other on the posteromedial side (
<figureCitation id="13308FBC31019F08451DFA503C7EFA3D" box="[1029,1138,1488,1513]" captionStart="Figure 7" captionStartId="8.[113,178,1656,1680]" captionTargetBox="[146,1426,144,1628]" captionTargetId="figure-102@8.[146,1426,144,1628]" captionTargetPageId="8" captionText="Figure 7. Saltoposuchus connectens referred specimen SMNS 55009, continued. A, B, C, D, E, F, SMNS 55009e, right tibia in (A) posterior, (B) anterior, (C) medial, (D) lateral, (E) proximal, and (F) distal views; G, H, SMNS 55009h, metatarsal block in (G) dorsal(?) and (H) opposing views; I, J, K, L, SMNS 55009g, right calcaneum in (I) medial, (J) lateral, (K) proximal, and (L) distal views. Abbreviations: als, anterolateral side; asf, astragalus facet; calt, calcaneal tuber; gr, groove; latc, lateral condyle; lg, longitudinal groove; mt, metatarsal; no, notch; pme, posterior extension; pms, posteromedial side; tr, trough." figureDoi="http://doi.org/10.5281/zenodo.8426163" httpUri="https://zenodo.org/record/8426163/files/figure.png" pageId="28" pageNumber="382">Fig. 7A, B</figureCitation>
). In other non-crocodyliform crocodylomorphs the posteromedial side projects further distally than the anterolateral side (NHMUK PV R 7566,
<bibRefCitation id="EF9AEEC831019F08447BF98F3B52F993" author="Colbert EH" pageId="28" pageNumber="382" pagination="561 - 92" refId="ref31083" refString="Colbert EH. A pseudosuchian reptile from Arizona. Bulletin of the American Museum of Natural History 1952; 99: 561 - 92." type="journal article" year="1952">Colbert 1952</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831019F084276F9AF3C14F992" author="Sues H-D &amp; Olsen PE &amp; Carter JG" box="[878,1048,1582,1607]" pageId="28" pageNumber="382" pagination="329 - 43" refId="ref32828" refString="Sues H-D, Olsen PE, Carter JG et al. A new crocodylomorph archosaur from the Upper Triassic of North Carolina. Journal of Vertebrate Paleontology 2003; 23: 329 - 43." type="journal article" year="2003">
Sues
<emphasis id="B97F4F2B31019F0842BFF9AF3BD7F993" box="[935,987,1582,1606]" italics="true" pageId="28" pageNumber="382">et al.</emphasis>
2003
</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831019F084530F9AE3CF4F993" author="Gohlich UB &amp; Chiappe LM &amp; Clark JM" box="[1064,1272,1582,1606]" pageId="28" pageNumber="382" pagination="307 - 21" refId="ref31388" refString="Gohlich UB, Chiappe LM, Clark JM et al. The systematic position of the Late Jurassic alleged dinosaur Macelognathus (Crocodylomorpha: Sphenosuchia). Canadian Journal of Earth Sciences 2005; 42: 307 - 21." type="journal article" year="2005">
Göhlich
<emphasis id="B97F4F2B31019F08459EF9AF3CBFF993" box="[1158,1203,1582,1606]" italics="true" pageId="28" pageNumber="382">et al</emphasis>
. 2005
</bibRefCitation>
). In both SMNS 55009e and SMNS 12596k, this condition is reversed, with the anterolateral side extending further distally (
<figureCitation id="13308FBC31019F0845F4F9ED3D3CF950" box="[1260,1328,1645,1669]" captionStart="Figure 5" captionStartId="6.[113,178,1777,1801]" captionTargetBox="[221,1353,149,1749]" captionTargetId="figure-5@6.[220,1353,149,1749]" captionTargetPageId="6" captionText="Figure 5. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, B, SMNS 12596g, right pubis in (A) anterior and (B) angled medial views; C, SMNS 12596h, right ischium exposed in dorsomedial view and two proximal caudal vertebrae exposed in dorsal view; D, SMNS 12596i, tentative less ulna in posteromedial view; E, F, G, H, SMNS 12596j, right femur in (E) posteromedial, (F) anterolateral, (G) proximal, and (H) distal views; I, proximal half of the right femur in posterolateral view; J, K, SMNS 12596k, right tibia and fibula, (J) with the tibia in posteromedial and the fibula in lateral(?) view and (K) with the tibia in anterolateral and the fibula in medial(?) view. Abbreviations: alcr, anterolateral crest; amt, anteromedial tuber; car, caudal rib; cav, caudal vertebra; cr, crest; crtf, crista tibiofibularis; fi, fibula; gr, groove; ilf, ilium facet; iss, ischial shass; lc, lateral condyle; ltr, lesser trochanter; mc, medial condyle; no, notch; ns, neural spine; obf, obturator foramen; olp, olecranon process; os, osteoderm; poz, postzygapophysis; pr., proximal; prz, prezygapophysis; pus, pubic shass; put, pubic tubercle; r., right; ti, tibia; tr, trochanter." figureDoi="http://doi.org/10.5281/zenodo.8426159" httpUri="https://zenodo.org/record/8426159/files/figure.png" pageId="28" pageNumber="382">Figs 5J</figureCitation>
,
<figureCitation id="13308FBC31019F084422F9ED3D7DF950" box="[1338,1393,1645,1669]" captionStart="Figure 7" captionStartId="8.[113,178,1656,1680]" captionTargetBox="[146,1426,144,1628]" captionTargetId="figure-102@8.[146,1426,144,1628]" captionTargetPageId="8" captionText="Figure 7. Saltoposuchus connectens referred specimen SMNS 55009, continued. A, B, C, D, E, F, SMNS 55009e, right tibia in (A) posterior, (B) anterior, (C) medial, (D) lateral, (E) proximal, and (F) distal views; G, H, SMNS 55009h, metatarsal block in (G) dorsal(?) and (H) opposing views; I, J, K, L, SMNS 55009g, right calcaneum in (I) medial, (J) lateral, (K) proximal, and (L) distal views. Abbreviations: als, anterolateral side; asf, astragalus facet; calt, calcaneal tuber; gr, groove; latc, lateral condyle; lg, longitudinal groove; mt, metatarsal; no, notch; pme, posterior extension; pms, posteromedial side; tr, trough." figureDoi="http://doi.org/10.5281/zenodo.8426163" httpUri="https://zenodo.org/record/8426163/files/figure.png" pageId="28" pageNumber="382">7A, B</figureCitation>
). This might represent an artefact of compression, since this extension is strongly exaggerated in SMNS 55009e, and the tibia of SMNS 12596k is also compressed.
</paragraph>
<paragraph id="8BB4933931019F09425DF96A3911FF32" blockId="28.[810,1460,235,1982]" lastBlockId="29.[129,777,144,231]" lastPageId="29" lastPageNumber="383" pageId="28" pageNumber="382">
The proximal and distal ends of the fibula are very slightly expanded relative to the shass and due to compression they form a thin ridge rather than a clear surface in SMNS 12596k (
<figureCitation id="13308FBC31019F084474F8A93B30F8B5" captionStart="Figure 5" captionStartId="6.[113,178,1777,1801]" captionTargetBox="[221,1353,149,1749]" captionTargetId="figure-5@6.[220,1353,149,1749]" captionTargetPageId="6" captionText="Figure 5. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, B, SMNS 12596g, right pubis in (A) anterior and (B) angled medial views; C, SMNS 12596h, right ischium exposed in dorsomedial view and two proximal caudal vertebrae exposed in dorsal view; D, SMNS 12596i, tentative less ulna in posteromedial view; E, F, G, H, SMNS 12596j, right femur in (E) posteromedial, (F) anterolateral, (G) proximal, and (H) distal views; I, proximal half of the right femur in posterolateral view; J, K, SMNS 12596k, right tibia and fibula, (J) with the tibia in posteromedial and the fibula in lateral(?) view and (K) with the tibia in anterolateral and the fibula in medial(?) view. Abbreviations: alcr, anterolateral crest; amt, anteromedial tuber; car, caudal rib; cav, caudal vertebra; cr, crest; crtf, crista tibiofibularis; fi, fibula; gr, groove; ilf, ilium facet; iss, ischial shass; lc, lateral condyle; ltr, lesser trochanter; mc, medial condyle; no, notch; ns, neural spine; obf, obturator foramen; olp, olecranon process; os, osteoderm; poz, postzygapophysis; pr., proximal; prz, prezygapophysis; pus, pubic shass; put, pubic tubercle; r., right; ti, tibia; tr, trochanter." figureDoi="http://doi.org/10.5281/zenodo.8426159" httpUri="https://zenodo.org/record/8426159/files/figure.png" pageId="28" pageNumber="382">Fig. 5J, K</figureCitation>
). The anterolateral side of the fibula possesses a thin crest on the proximal half of the shass (
<figureCitation id="13308FBC31019F084546F8E83CA4F8AA" box="[1118,1192,1895,1920]" captionStart="Figure 5" captionStartId="6.[113,178,1777,1801]" captionTargetBox="[221,1353,149,1749]" captionTargetId="figure-5@6.[220,1353,149,1749]" captionTargetPageId="6" captionText="Figure 5. Saltoposuchus connectens referred specimen SMNS 12596, continued. A, B, SMNS 12596g, right pubis in (A) anterior and (B) angled medial views; C, SMNS 12596h, right ischium exposed in dorsomedial view and two proximal caudal vertebrae exposed in dorsal view; D, SMNS 12596i, tentative less ulna in posteromedial view; E, F, G, H, SMNS 12596j, right femur in (E) posteromedial, (F) anterolateral, (G) proximal, and (H) distal views; I, proximal half of the right femur in posterolateral view; J, K, SMNS 12596k, right tibia and fibula, (J) with the tibia in posteromedial and the fibula in lateral(?) view and (K) with the tibia in anterolateral and the fibula in medial(?) view. Abbreviations: alcr, anterolateral crest; amt, anteromedial tuber; car, caudal rib; cav, caudal vertebra; cr, crest; crtf, crista tibiofibularis; fi, fibula; gr, groove; ilf, ilium facet; iss, ischial shass; lc, lateral condyle; ltr, lesser trochanter; mc, medial condyle; no, notch; ns, neural spine; obf, obturator foramen; olp, olecranon process; os, osteoderm; poz, postzygapophysis; pr., proximal; prz, prezygapophysis; pus, pubic shass; put, pubic tubercle; r., right; ti, tibia; tr, trochanter." figureDoi="http://doi.org/10.5281/zenodo.8426159" httpUri="https://zenodo.org/record/8426159/files/figure.png" pageId="28" pageNumber="382">Fig. 5K</figureCitation>
). This crest is demarcated by a marked groove on the lateral surface of the shass posterior to it, which might also represent a compression artefact. This anterolateral crest (=anterior trochanter
<emphasis id="B97F4F2B31009F094303FF113A42FF7D" box="[539,590,145,168]" italics="true" pageId="29" pageNumber="383">sensu</emphasis>
<bibRefCitation id="EF9AEEC831009F09434BFF103ADFFF7D" author="Sereno PC" box="[595,723,144,168]" pageId="29" pageNumber="383" pagination="1 - 53" refId="ref32698" refString="Sereno PC. Basal archosaurs: phylogenetic relationships and functional implications. Journal of Vertebrate Paleontology 1991; 11: 1 - 53." type="journal article" year="1991">Sereno 1991</bibRefCitation>
) was previously described by
<bibRefCitation id="EF9AEEC831009F094060FF303A01FF12" author="Huene FR" box="[376,525,175,199]" pageId="29" pageNumber="383" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene (1921)</bibRefCitation>
as an aưachment site for M. iliofibularis.
</paragraph>
<paragraph id="8BB4933931009F094199FE8939A7FBFA" blockId="29.[129,778,264,1979]" pageId="29" pageNumber="383">
<emphasis id="B97F4F2B31009F094199FE8938CBFEF5" box="[129,199,265,288]" italics="true" pageId="29" pageNumber="383">Tarsus:</emphasis>
Of the tarsus, three calcanea and a distal tarsal 4 are preserved. The less calcaneum and distal tarsal 4 are preserved in SMNS 12597a in association with the less tibia and fibula and the less pes, whereas the right calcaneum SMNS 12597f is completely freed from matrix (
<figureCitation id="13308FBC31009F094058FE0539C8FE48" box="[320,452,389,413]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="29" pageNumber="383">Figs 2B, G, H</figureCitation>
,
<figureCitation id="13308FBC31009F0940D6FE053A1DFE48" box="[462,529,389,413]" captionStart="Figure 12" captionStartId="24.[113,178,1772,1796]" captionTargetBox="[151,1424,144,1744]" captionTargetId="figure-6@24.[148,1424,144,1744]" captionTargetPageId="24" captionText="Figure 12. Details of the postcranial anatomy of Saltoposuchus connectens. A, dorsal vertebral neural arch and dorsal rib of SMNS 12596c in ventral view; B, associated osteoderms of SMNS 12596c in dorsal view; C, sacral ribs and associated elements of SMNS 12597a in dorsal view; D, distal portion of right scapular blade of SMNS 12597a in lateral view; E, proximal end of right tibia of SMNS 12597a in proximal view; F, less tarsus and pes of SMNS 12597a with the calcaneum exposed in lateral view; G, close-up of less tarsus of SMNS 12597a; H, close-up of less pes of SMNS 12597a. Abbreviations: ane, anterior expansion; cal, calcaneum; calt, calcaneal tuber; cp, capitulum; dp, diapophysis; dt, distal tarsal; exp, extensor pit; fe, femur; fi, fibula; il, ilium; lam, lamina; latc, lateral condyle; le, lateral edge; lr, longitudinal ridge; me, medial edge; mt, metatarsal; no, notch; pap, parapophysis; ph, phalanx; pme, posterior extension; poe, posterior expansion; pofl, posterior flange; poz, postzygapophysis; os, osteoderm; r., right; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426175" httpUri="https://zenodo.org/record/8426175/files/figure.png" pageId="29" pageNumber="383">12F, G</figureCitation>
). Finally, SMNS
<quantity id="4CF33EDC31009F0943A4FE053B06FE48" box="[700,778,389,413]" metricMagnitude="1" metricUnit="kg" metricValue="5.5009" pageId="29" pageNumber="383" unit="g" value="55009.0">55009g</quantity>
is another right calcaneum completely freed from matrix (
<figureCitation id="13308FBC31009F0943FBFE2538BDFE0E" captionStart="Figure 7" captionStartId="8.[113,178,1656,1680]" captionTargetBox="[146,1426,144,1628]" captionTargetId="figure-102@8.[146,1426,144,1628]" captionTargetPageId="8" captionText="Figure 7. Saltoposuchus connectens referred specimen SMNS 55009, continued. A, B, C, D, E, F, SMNS 55009e, right tibia in (A) posterior, (B) anterior, (C) medial, (D) lateral, (E) proximal, and (F) distal views; G, H, SMNS 55009h, metatarsal block in (G) dorsal(?) and (H) opposing views; I, J, K, L, SMNS 55009g, right calcaneum in (I) medial, (J) lateral, (K) proximal, and (L) distal views. Abbreviations: als, anterolateral side; asf, astragalus facet; calt, calcaneal tuber; gr, groove; latc, lateral condyle; lg, longitudinal groove; mt, metatarsal; no, notch; pme, posterior extension; pms, posteromedial side; tr, trough." figureDoi="http://doi.org/10.5281/zenodo.8426163" httpUri="https://zenodo.org/record/8426163/files/figure.png" pageId="29" pageNumber="383">Fig. 7J–L</figureCitation>
). The less calcaneum of SMNS 12597a exposes its flaưened lateral surface, including a posteriorly directed calcaneal tuber that curves posteroproximally and which has a distinctly concave proximal surface (
<figureCitation id="13308FBC31009F09405BFDA239ACFDEF" box="[323,416,546,570]" captionStart="Figure 12" captionStartId="24.[113,178,1772,1796]" captionTargetBox="[151,1424,144,1744]" captionTargetId="figure-6@24.[148,1424,144,1744]" captionTargetPageId="24" captionText="Figure 12. Details of the postcranial anatomy of Saltoposuchus connectens. A, dorsal vertebral neural arch and dorsal rib of SMNS 12596c in ventral view; B, associated osteoderms of SMNS 12596c in dorsal view; C, sacral ribs and associated elements of SMNS 12597a in dorsal view; D, distal portion of right scapular blade of SMNS 12597a in lateral view; E, proximal end of right tibia of SMNS 12597a in proximal view; F, less tarsus and pes of SMNS 12597a with the calcaneum exposed in lateral view; G, close-up of less tarsus of SMNS 12597a; H, close-up of less pes of SMNS 12597a. Abbreviations: ane, anterior expansion; cal, calcaneum; calt, calcaneal tuber; cp, capitulum; dp, diapophysis; dt, distal tarsal; exp, extensor pit; fe, femur; fi, fibula; il, ilium; lam, lamina; latc, lateral condyle; le, lateral edge; lr, longitudinal ridge; me, medial edge; mt, metatarsal; no, notch; pap, parapophysis; ph, phalanx; pme, posterior extension; poe, posterior expansion; pofl, posterior flange; poz, postzygapophysis; os, osteoderm; r., right; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426175" httpUri="https://zenodo.org/record/8426175/files/figure.png" pageId="29" pageNumber="383">Fig. 12G</figureCitation>
). A posteriorly directed calcaneal tuber is indicative of an erect posture (
<emphasis id="B97F4F2B31009F09430CFDC23A4BFD8C" box="[532,583,578,601]" italics="true" pageId="29" pageNumber="383">sensu</emphasis>
<bibRefCitation id="EF9AEEC831009F094356FDC13ADDFD8C" author="Parrish JM" box="[590,721,577,601]" pageId="29" pageNumber="383" pagination="396 - 414" refId="ref32298" refString="Parrish JM. The origin of crocodilian locomotion. Paleobiology 1987; 13: 396 - 414." type="journal article" year="1987">Parrish 1987</bibRefCitation>
) and also occurs in the non-crocodyliform crocodylomorphs
<taxonomicName id="4C0BE8BA31009F0943A9FDE138BBFD4D" baseAuthorityName="Marsh" baseAuthorityYear="1877" class="Reptilia" family="Coeluridae" genus="Hallopus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="29" pageNumber="383" phylum="Chordata" rank="species" species="victor">
<emphasis id="B97F4F2B31009F0943A9FDE138BBFD4D" italics="true" pageId="29" pageNumber="383">Hallopus victor</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0BE8BA31009F0941D3FD003978FD4D" authorityName="Reig" authorityYear="1963" box="[203,372,640,664]" class="Polyplacophora" genus="Trialestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodilomorphi" pageId="29" pageNumber="383" phylum="Mollusca" rank="species" species="romeri">
<emphasis id="B97F4F2B31009F0941D3FD003978FD4D" box="[203,372,640,664]" italics="true" pageId="29" pageNumber="383">Trialestes romeri</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0BE8BA31009F094090FD003AB8FD42" authority=", YPM" authorityName="YPM" box="[392,692,640,664]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="29" pageNumber="383" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B31009F094090FD003A60FD4D" box="[392,620,640,664]" italics="true" pageId="29" pageNumber="383">Terrestrisuchus gracilis</emphasis>
, YPM
</taxonomicName>
41198, and
<taxonomicName id="4C0BE8BA31009F0941A8FD1F3982FD62" authorityName="Marsh" authorityYear="1884" box="[176,398,671,695]" class="Reptilia" family="Hypsilophodontidae" genus="Macelognathus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="29" pageNumber="383" phylum="Chordata" rank="species" species="vagans">
<emphasis id="B97F4F2B31009F0941A8FD1F3982FD62" box="[176,398,671,695]" italics="true" pageId="29" pageNumber="383">Macelognathus vagans</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9AEEC831009F0940BAFD1F3A24FD62" author="Parrish JM" box="[418,552,671,695]" pageId="29" pageNumber="383" pagination="396 - 414" refId="ref32298" refString="Parrish JM. The origin of crocodilian locomotion. Paleobiology 1987; 13: 396 - 414." type="journal article" year="1987">Parrish 1987</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831009F09432FFD1F3B08FD62" author="Gohlich UB &amp; Chiappe LM &amp; Clark JM" box="[567,772,671,695]" pageId="29" pageNumber="383" pagination="307 - 21" refId="ref31388" refString="Gohlich UB, Chiappe LM, Clark JM et al. The systematic position of the Late Jurassic alleged dinosaur Macelognathus (Crocodylomorpha: Sphenosuchia). Canadian Journal of Earth Sciences 2005; 42: 307 - 21." type="journal article" year="2005">
Göhlich
<emphasis id="B97F4F2B31009F09438CFD203ACBFD62" box="[660,711,671,695]" italics="true" pageId="29" pageNumber="383">et al.</emphasis>
2005
</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831009F094199FD3E3904FD02" author="Nesbiu SJ" box="[129,264,702,727]" pageId="29" pageNumber="383" pagination="1 - 292" refId="ref32089" refString="Nesbiu SJ. The early evolution of archosaurs: relationships and the origin of major clades. Bulletin of the American Museum of Natural History 2011; 352: 1 - 292." type="journal article" year="2011">Nesbiư 2011</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831009F09400EFD3F39E8FD03" author="Lecuona A &amp; Ezcurra MD &amp; Irmis RB" box="[278,484,702,726]" pageId="29" pageNumber="383" pagination="585 - 622" refId="ref32010" refString="Lecuona A, Ezcurra MD, Irmis RB. Revision of the early crocodylomorph Trialestes romeri (Archosauria, Suchia) from the lower Upper Triassic Ischigualasto Formation of Argentina: one of the oldest-known crocodylomorphs. Papers in Palaeontology 2016; 2: 585 - 622." type="journal article" year="2016">
Lecuona
<emphasis id="B97F4F2B31009F09406FFD3F39A5FD03" box="[375,425,702,726]" italics="true" pageId="29" pageNumber="383">et al.</emphasis>
2016
</bibRefCitation>
). It deviates from the more posterolaterally oriented calcaneal tubera of extant crocodylians (
<bibRefCitation id="EF9AEEC831009F094193FD7D3902FCC0" author="Parrish JM" box="[139,270,765,789]" pageId="29" pageNumber="383" pagination="396 - 414" refId="ref32298" refString="Parrish JM. The origin of crocodilian locomotion. Paleobiology 1987; 13: 396 - 414." type="journal article" year="1987">Parrish 1987</bibRefCitation>
). The anterior margin of the calcaneum is curved anteriorly and proximally, whereas its anterodistal portion is flattened and in articulation with the distal tarsal 4 directly distal to it. A process is projected proximal to the main body of the calcaneum. This probably represents the heavily crushed lip from the medial side of the calcaneum, which has been shissed proximally, whereas it was projected medially in life (
<figureCitation id="13308FBC31009F094334FC393A89FC04" box="[556,645,953,977]" captionStart="Figure 12" captionStartId="24.[113,178,1772,1796]" captionTargetBox="[151,1424,144,1744]" captionTargetId="figure-6@24.[148,1424,144,1744]" captionTargetPageId="24" captionText="Figure 12. Details of the postcranial anatomy of Saltoposuchus connectens. A, dorsal vertebral neural arch and dorsal rib of SMNS 12596c in ventral view; B, associated osteoderms of SMNS 12596c in dorsal view; C, sacral ribs and associated elements of SMNS 12597a in dorsal view; D, distal portion of right scapular blade of SMNS 12597a in lateral view; E, proximal end of right tibia of SMNS 12597a in proximal view; F, less tarsus and pes of SMNS 12597a with the calcaneum exposed in lateral view; G, close-up of less tarsus of SMNS 12597a; H, close-up of less pes of SMNS 12597a. Abbreviations: ane, anterior expansion; cal, calcaneum; calt, calcaneal tuber; cp, capitulum; dp, diapophysis; dt, distal tarsal; exp, extensor pit; fe, femur; fi, fibula; il, ilium; lam, lamina; latc, lateral condyle; le, lateral edge; lr, longitudinal ridge; me, medial edge; mt, metatarsal; no, notch; pap, parapophysis; ph, phalanx; pme, posterior extension; poe, posterior expansion; pofl, posterior flange; poz, postzygapophysis; os, osteoderm; r., right; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426175" httpUri="https://zenodo.org/record/8426175/files/figure.png" pageId="29" pageNumber="383">Fig. 12G</figureCitation>
). Anteriorly, the lip is deeply excavated, and here it formed the posterior surface of the deep facet that received the laterally projected peg of the astragalus (
<bibRefCitation id="EF9AEEC831009F094002FB973997FBFA" author="Sereno PC" box="[282,411,1047,1071]" pageId="29" pageNumber="383" pagination="1 - 53" refId="ref32698" refString="Sereno PC. Basal archosaurs: phylogenetic relationships and functional implications. Journal of Vertebrate Paleontology 1991; 11: 1 - 53." type="journal article" year="1991">Sereno 1991</bibRefCitation>
).
</paragraph>
<paragraph id="8BB4933931009F094184FBB63901FA33" blockId="29.[129,778,264,1979]" pageId="29" pageNumber="383">
The element distal to this calcaneum was interpreted as the astragalus by
<bibRefCitation id="EF9AEEC831009F094013FBD639ADFBBB" author="Huene FR" box="[267,417,1110,1134]" pageId="29" pageNumber="383" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene (1921)</bibRefCitation>
, but it was later identified as distal tarsal 4 by
<bibRefCitation id="EF9AEEC831009F0941E8FBF5398CFB58" author="Crush PJ" box="[240,384,1141,1165]" pageId="29" pageNumber="383" refId="ref31157" refString="Crush PJ. An early, terrestrial, crocodile from South Wales. Unpublished Ph. D. Thesis, University College London, 1980." type="book" year="1980">Crush (1980)</bibRefCitation>
. It is around two-thirds the size of the calcaneum (
<figureCitation id="13308FBC31009F094018FB143956FB78" box="[256,346,1172,1197]" captionStart="Figure 12" captionStartId="24.[113,178,1772,1796]" captionTargetBox="[151,1424,144,1744]" captionTargetId="figure-6@24.[148,1424,144,1744]" captionTargetPageId="24" captionText="Figure 12. Details of the postcranial anatomy of Saltoposuchus connectens. A, dorsal vertebral neural arch and dorsal rib of SMNS 12596c in ventral view; B, associated osteoderms of SMNS 12596c in dorsal view; C, sacral ribs and associated elements of SMNS 12597a in dorsal view; D, distal portion of right scapular blade of SMNS 12597a in lateral view; E, proximal end of right tibia of SMNS 12597a in proximal view; F, less tarsus and pes of SMNS 12597a with the calcaneum exposed in lateral view; G, close-up of less tarsus of SMNS 12597a; H, close-up of less pes of SMNS 12597a. Abbreviations: ane, anterior expansion; cal, calcaneum; calt, calcaneal tuber; cp, capitulum; dp, diapophysis; dt, distal tarsal; exp, extensor pit; fe, femur; fi, fibula; il, ilium; lam, lamina; latc, lateral condyle; le, lateral edge; lr, longitudinal ridge; me, medial edge; mt, metatarsal; no, notch; pap, parapophysis; ph, phalanx; pme, posterior extension; poe, posterior expansion; pofl, posterior flange; poz, postzygapophysis; os, osteoderm; r., right; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426175" httpUri="https://zenodo.org/record/8426175/files/figure.png" pageId="29" pageNumber="383">Fig. 12G</figureCitation>
). It is widest at its posterodistal end relative to the calcaneum, and narrower at its anteroproximal end. It also possesses a small, laterally directed process near its proximal end. Its morphology is somewhat similar to that of distal tarsal 4 of pseudosuchian-line archosaurs, including YPM 41198, which also possesses a narrow and wide end, respectively (
<bibRefCitation id="EF9AEEC831009F094367FAB13B0EFA9C" author="Nesbiu SJ" box="[639,770,1329,1353]" pageId="29" pageNumber="383" pagination="1 - 292" refId="ref32089" refString="Nesbiu SJ. The early evolution of archosaurs: relationships and the origin of major clades. Bulletin of the American Museum of Natural History 2011; 352: 1 - 292." type="journal article" year="2011">Nesbiư 2011</bibRefCitation>
: ch. 349). Furthermore, the element lacks the complex structure seen in the astragalus of crocodylomorphs that includes a prominent peg for the articulation with the calcaneum (
<bibRefCitation id="EF9AEEC831009F094361FA0F3AF5FA72" author="Sereno PC" box="[633,761,1423,1447]" pageId="29" pageNumber="383" pagination="1 - 53" refId="ref32698" refString="Sereno PC. Basal archosaurs: phylogenetic relationships and functional implications. Journal of Vertebrate Paleontology 1991; 11: 1 - 53." type="journal article" year="1991">Sereno 1991</bibRefCitation>
). The interpretation of this element as a distal tarsal 4 is therefore corroborated.
</paragraph>
<paragraph id="8BB4933931009F094184FA6D3C3FFE14" blockId="29.[129,778,264,1979]" lastBlockId="29.[825,1474,143,449]" pageId="29" pageNumber="383">
The isolated right calcaneum of SMNS 12597f possesses a complete calcaneal tuber that roughly corresponds in morphology to that of the less calcaneum (
<figureCitation id="13308FBC31009F0940EEF9AC3A6FF996" box="[502,611,1580,1604]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="29" pageNumber="383">Fig. 2G, H</figureCitation>
). Its main body is badly broken, with its proximal half missing. Nevertheless, its medial surface was clearly deeply excavated, forming the facet for the astragalar peg. The right calcaneum SMNS 12597f was already isolated from the rest of the material of SMNS
<quantity id="4CF33EDC31009F0943B6F9293B05F914" box="[686,777,1705,1729]" metricMagnitude="2" metricUnit="m" metricValue="3.1996379999999998" pageId="29" pageNumber="383" unit="in" value="12597.0">12597 in</quantity>
1921 (
<bibRefCitation id="EF9AEEC831009F0941D0F9493947F935" author="Huene FR" box="[200,331,1736,1760]" pageId="29" pageNumber="383" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene 1921</bibRefCitation>
). This element is considerably larger than the associated less calcaneum of SMNS 12597a, to such an extent (Supporting Information,
<tableCitation id="C689A68231009F0940DBF8873A12F8CA" box="[451,542,1799,1823]" captionStart="Table 1" captionStartId="2.[114,169,143,167]" captionText="Table 1. Summary of the taxonomic assignments in relevant studies of the specimens that now comprise Saltoposuchus connectens" pageId="29" pageNumber="383">Table S1</tableCitation>
) that it is certain that both elements derive from separate individuals. Furthermore, the right calcaneum SMNS 12597f is also stouter and less anteroposteriorly elongate compared to the calcanea of SMNS 12597a and SMNS
<quantity id="4CF33EDC31009F094046F80439A7F849" box="[350,427,1924,1948]" metricMagnitude="1" metricUnit="kg" metricValue="5.5009" pageId="29" pageNumber="383" unit="g" value="55009.0">55009g</quantity>
(
<figureCitation id="13308FBC31009F0940DAF8043A31F849" box="[450,573,1924,1948]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="29" pageNumber="383">Figs 2G, H</figureCitation>
,
<figureCitation id="13308FBC31009F094348F8053A8EF849" box="[592,642,1925,1948]" captionStart="Figure 7" captionStartId="8.[113,178,1656,1680]" captionTargetBox="[146,1426,144,1628]" captionTargetId="figure-102@8.[146,1426,144,1628]" captionTargetPageId="8" captionText="Figure 7. Saltoposuchus connectens referred specimen SMNS 55009, continued. A, B, C, D, E, F, SMNS 55009e, right tibia in (A) posterior, (B) anterior, (C) medial, (D) lateral, (E) proximal, and (F) distal views; G, H, SMNS 55009h, metatarsal block in (G) dorsal(?) and (H) opposing views; I, J, K, L, SMNS 55009g, right calcaneum in (I) medial, (J) lateral, (K) proximal, and (L) distal views. Abbreviations: als, anterolateral side; asf, astragalus facet; calt, calcaneal tuber; gr, groove; latc, lateral condyle; lg, longitudinal groove; mt, metatarsal; no, notch; pme, posterior extension; pms, posteromedial side; tr, trough." figureDoi="http://doi.org/10.5281/zenodo.8426163" httpUri="https://zenodo.org/record/8426163/files/figure.png" pageId="29" pageNumber="383">7I–L</figureCitation>
,
<figureCitation id="13308FBC31009F09438CF8043ACDF849" box="[660,705,1924,1948]" captionStart="Figure 12" captionStartId="24.[113,178,1772,1796]" captionTargetBox="[151,1424,144,1744]" captionTargetId="figure-6@24.[148,1424,144,1744]" captionTargetPageId="24" captionText="Figure 12. Details of the postcranial anatomy of Saltoposuchus connectens. A, dorsal vertebral neural arch and dorsal rib of SMNS 12596c in ventral view; B, associated osteoderms of SMNS 12596c in dorsal view; C, sacral ribs and associated elements of SMNS 12597a in dorsal view; D, distal portion of right scapular blade of SMNS 12597a in lateral view; E, proximal end of right tibia of SMNS 12597a in proximal view; F, less tarsus and pes of SMNS 12597a with the calcaneum exposed in lateral view; G, close-up of less tarsus of SMNS 12597a; H, close-up of less pes of SMNS 12597a. Abbreviations: ane, anterior expansion; cal, calcaneum; calt, calcaneal tuber; cp, capitulum; dp, diapophysis; dt, distal tarsal; exp, extensor pit; fe, femur; fi, fibula; il, ilium; lam, lamina; latc, lateral condyle; le, lateral edge; lr, longitudinal ridge; me, medial edge; mt, metatarsal; no, notch; pap, parapophysis; ph, phalanx; pme, posterior extension; poe, posterior expansion; pofl, posterior flange; poz, postzygapophysis; os, osteoderm; r., right; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426175" httpUri="https://zenodo.org/record/8426175/files/figure.png" pageId="29" pageNumber="383">12G</figureCitation>
). This could be related to allometric variation or an anteroposterior compression of the element. Based on the prominent calcaneal tuber and deeply excavated astragalar facet that is framed posteriorly by a distinct lip, this element clearly belongs to a crocodylomorph and its assignment to
<taxonomicName id="4C0BE8BA31009F0945D5FF6D3DCEFED0" authorityName="Huene" authorityYear="1921" box="[1229,1474,237,261]" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="29" pageNumber="383" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B31009F0945D5FF6D3DCEFED0" box="[1229,1474,237,261]" italics="true" pageId="29" pageNumber="383">Saltoposuchus connectens</emphasis>
</taxonomicName>
is tentatively maintained. The isolated right calcaneum SMNS
<quantity id="4CF33EDC31009F094221FEAC3B8BFE91" box="[825,903,300,324]" metricMagnitude="1" metricUnit="kg" metricValue="5.5009" pageId="29" pageNumber="383" unit="g" value="55009.0">55009g</quantity>
is mostly complete, although the distal surface of the calcaneal tuber is missing, and it is poorly preserved, being heavily distorted (
<figureCitation id="13308FBC31009F0942ECFEEB3C5CFE57" box="[1012,1104,363,387]" captionStart="Figure 7" captionStartId="8.[113,178,1656,1680]" captionTargetBox="[146,1426,144,1628]" captionTargetId="figure-102@8.[146,1426,144,1628]" captionTargetPageId="8" captionText="Figure 7. Saltoposuchus connectens referred specimen SMNS 55009, continued. A, B, C, D, E, F, SMNS 55009e, right tibia in (A) posterior, (B) anterior, (C) medial, (D) lateral, (E) proximal, and (F) distal views; G, H, SMNS 55009h, metatarsal block in (G) dorsal(?) and (H) opposing views; I, J, K, L, SMNS 55009g, right calcaneum in (I) medial, (J) lateral, (K) proximal, and (L) distal views. Abbreviations: als, anterolateral side; asf, astragalus facet; calt, calcaneal tuber; gr, groove; latc, lateral condyle; lg, longitudinal groove; mt, metatarsal; no, notch; pme, posterior extension; pms, posteromedial side; tr, trough." figureDoi="http://doi.org/10.5281/zenodo.8426163" httpUri="https://zenodo.org/record/8426163/files/figure.png" pageId="29" pageNumber="383">Fig. 7I–L</figureCitation>
). It possesses the same morphology as that seen in the calcanea of SMNS 12597, as well as those of other crocodylomorphs.
</paragraph>
<paragraph id="8BB4933931009F094221FE633D5CFCC1" blockId="29.[825,1475,483,1447]" pageId="29" pageNumber="383">
<emphasis id="B97F4F2B31009F094221FE633B6CFE2F" box="[825,864,483,506]" italics="true" pageId="29" pageNumber="383">Pes:</emphasis>
Remains of the less pes are preserved in SMNS 12597a in articulation with the rest of the less hindlimb. They consist of very fragmentary remains of at least two metatarsals and the phalanges of the corresponding digits (
<figureCitation id="13308FBC31009F0945CAFDC13D13FD8C" box="[1234,1311,577,601]" captionStart="Figure 2" captionStartId="3.[129,193,1777,1801]" captionTargetBox="[238,1364,149,1749]" captionTargetId="figure-6@3.[238,1364,149,1749]" captionTargetPageId="3" captionText="Figure 2. Saltoposuchus connectens holotype specimen SMNS 12597a–d and referred specimens SMNS 12597e and SMNS 12597f. A, B, SMNS 12597a, main skeletal block in (A) dorsal and (B) ventral views; C, SMNS 12597b, associated cranial elements; D, SMNS 12597d, articulated caudal vertebrae exposed in less lateral view; E, SMNS 12597c with two cervical vertebrae exposed in dorsal view, two dorsal vertebrae exposed in right lateral view, and an osteoderm exposed in dorsal view; F, SMNS 12597e, less angular exposed in medial view; G, H, SMNS 12597f, isolated right calcaneum in (G) medial and (H) lateral views.Abbreviations: an, angular; asf, astragular facet; ca., caudal; cal, calcaneum; calt, calcaneal tuber; cav, caudal vertebra; cev, cervical vertebra; ch, chevron; cp, capitulum; do., dorsal; dp, diapophysis; dr, dorsal rib; ds., distal; dt, distal tarsal; dv, dorsal vertebra; exp, extensor pit; fe, femur; fi, fibula; fr, frontal; gr, groove; il, ilium; j, jugal; l., less; mt, metatarsal; na, nasal; ncs, neurocentral suture; ns, neural spine; os, osteoderm; ot, otoccipital; pa, parietal; pap, parapophysis; ph, phalanx; pop, posterior process; poz, postzygapophysis; pr., proximal; pu, pubis; q, quadrate; r., right; sc, scapula; sq, squamosal; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426151" httpUri="https://zenodo.org/record/8426151/files/figure.png" pageId="29" pageNumber="383">Figs 2B</figureCitation>
,
<figureCitation id="13308FBC31009F094434FDC13D79FD8C" box="[1324,1397,577,601]" captionStart="Figure 12" captionStartId="24.[113,178,1772,1796]" captionTargetBox="[151,1424,144,1744]" captionTargetId="figure-6@24.[148,1424,144,1744]" captionTargetPageId="24" captionText="Figure 12. Details of the postcranial anatomy of Saltoposuchus connectens. A, dorsal vertebral neural arch and dorsal rib of SMNS 12596c in ventral view; B, associated osteoderms of SMNS 12596c in dorsal view; C, sacral ribs and associated elements of SMNS 12597a in dorsal view; D, distal portion of right scapular blade of SMNS 12597a in lateral view; E, proximal end of right tibia of SMNS 12597a in proximal view; F, less tarsus and pes of SMNS 12597a with the calcaneum exposed in lateral view; G, close-up of less tarsus of SMNS 12597a; H, close-up of less pes of SMNS 12597a. Abbreviations: ane, anterior expansion; cal, calcaneum; calt, calcaneal tuber; cp, capitulum; dp, diapophysis; dt, distal tarsal; exp, extensor pit; fe, femur; fi, fibula; il, ilium; lam, lamina; latc, lateral condyle; le, lateral edge; lr, longitudinal ridge; me, medial edge; mt, metatarsal; no, notch; pap, parapophysis; ph, phalanx; pme, posterior extension; poe, posterior expansion; pofl, posterior flange; poz, postzygapophysis; os, osteoderm; r., right; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426175" httpUri="https://zenodo.org/record/8426175/files/figure.png" pageId="29" pageNumber="383">12F–H</figureCitation>
). Only the proximal ends of the metatarsals are informative. The outer element is more gracile than the element associated with distal tarsal 4, and its proximal head is smaller and bulbous (
<figureCitation id="13308FBC31009F094472FD1F3B40FD03" captionStart="Figure 12" captionStartId="24.[113,178,1772,1796]" captionTargetBox="[151,1424,144,1744]" captionTargetId="figure-6@24.[148,1424,144,1744]" captionTargetPageId="24" captionText="Figure 12. Details of the postcranial anatomy of Saltoposuchus connectens. A, dorsal vertebral neural arch and dorsal rib of SMNS 12596c in ventral view; B, associated osteoderms of SMNS 12596c in dorsal view; C, sacral ribs and associated elements of SMNS 12597a in dorsal view; D, distal portion of right scapular blade of SMNS 12597a in lateral view; E, proximal end of right tibia of SMNS 12597a in proximal view; F, less tarsus and pes of SMNS 12597a with the calcaneum exposed in lateral view; G, close-up of less tarsus of SMNS 12597a; H, close-up of less pes of SMNS 12597a. Abbreviations: ane, anterior expansion; cal, calcaneum; calt, calcaneal tuber; cp, capitulum; dp, diapophysis; dt, distal tarsal; exp, extensor pit; fe, femur; fi, fibula; il, ilium; lam, lamina; latc, lateral condyle; le, lateral edge; lr, longitudinal ridge; me, medial edge; mt, metatarsal; no, notch; pap, parapophysis; ph, phalanx; pme, posterior extension; poe, posterior expansion; pofl, posterior flange; poz, postzygapophysis; os, osteoderm; r., right; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426175" httpUri="https://zenodo.org/record/8426175/files/figure.png" pageId="29" pageNumber="383">Fig. 12F, G</figureCitation>
). This represents metatarsal V, as was also suggested by
<bibRefCitation id="EF9AEEC831009F094465FD3E3B8FFD20" author="Huene FR" pageId="29" pageNumber="383" pagination="329 - 403" refId="ref31595" refString="Huene FR. Neue Pseudosuchier und Coelurosaurier aus dem Wuruembergischen Keuper. Acta Zoologica 1921; 2: 329 - 403." type="journal article" year="1921">Huene (1921)</bibRefCitation>
. The proximal head of the second metatarsal is incomplete but markedly broader than that of metatarsal V.
</paragraph>
<paragraph id="8BB4933931009F09424DFC9C3BC5FBB8" blockId="29.[825,1475,483,1447]" pageId="29" pageNumber="383">
Three incomplete metatarsals are preserved in SMNS 55009f (
<figureCitation id="13308FBC31009F09425CFCBB3BA3FC86" box="[836,943,827,851]" captionStart="Figure 7" captionStartId="8.[113,178,1656,1680]" captionTargetBox="[146,1426,144,1628]" captionTargetId="figure-102@8.[146,1426,144,1628]" captionTargetPageId="8" captionText="Figure 7. Saltoposuchus connectens referred specimen SMNS 55009, continued. A, B, C, D, E, F, SMNS 55009e, right tibia in (A) posterior, (B) anterior, (C) medial, (D) lateral, (E) proximal, and (F) distal views; G, H, SMNS 55009h, metatarsal block in (G) dorsal(?) and (H) opposing views; I, J, K, L, SMNS 55009g, right calcaneum in (I) medial, (J) lateral, (K) proximal, and (L) distal views. Abbreviations: als, anterolateral side; asf, astragalus facet; calt, calcaneal tuber; gr, groove; latc, lateral condyle; lg, longitudinal groove; mt, metatarsal; no, notch; pme, posterior extension; pms, posteromedial side; tr, trough." figureDoi="http://doi.org/10.5281/zenodo.8426163" httpUri="https://zenodo.org/record/8426163/files/figure.png" pageId="29" pageNumber="383">Fig. 7G, H</figureCitation>
). Their shasss are straight, elongate, and tightly bundled, similar to the condition in
<taxonomicName id="4C0BE8BA31009F09459CFCDB3D52FCA6" box="[1156,1374,859,883]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="29" pageNumber="383" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B97F4F2B31009F09459CFCDB3D52FCA6" box="[1156,1374,859,883]" italics="true" pageId="29" pageNumber="383">Terrestrisuchus gracilis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0BE8BA31009F094472FCDB3B63FC47" baseAuthorityName="Marsh" baseAuthorityYear="1877" class="Reptilia" family="Coeluridae" genus="Hallopus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="29" pageNumber="383" phylum="Chordata" rank="species" species="victor">
<emphasis id="B97F4F2B31009F094472FCDB3B63FC47" italics="true" pageId="29" pageNumber="383">Hallopus victor</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C0BE8BA31009F0942BCFCFA3C72FC47" authorityName="Marsh" authorityYear="1884" box="[932,1150,890,914]" class="Reptilia" family="Hypsilophodontidae" genus="Macelognathus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="29" pageNumber="383" phylum="Chordata" rank="species" species="vagans">
<emphasis id="B97F4F2B31009F0942BCFCFA3C72FC47" box="[932,1150,890,914]" italics="true" pageId="29" pageNumber="383">Macelognathus vagans</emphasis>
</taxonomicName>
(NHMUK PV R 7551b,
<bibRefCitation id="EF9AEEC831009F094462FCFA3B61FC64" author="Walker AD" pageId="29" pageNumber="383" pagination="323 - 72" refId="ref32988" refString="Walker AD. A revision of the Jurassic reptile Hallopus victor (Marsh), with remarks on the classification of crocodiles. Philosophical Transactions of the Royal Society of London B: Biological Sciences 1970; 257: 323 - 72." type="journal article" year="1970">Walker 1970</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831009F09426FFC193BFCFC64" author="Crush PJ" box="[887,1008,921,945]" pageId="29" pageNumber="383" pagination="131 - 57" refId="ref31183" refString="Crush PJ. A late Upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 1984; 27: 131 - 57." type="journal article" year="1984">Crush 1984</bibRefCitation>
,
<bibRefCitation id="EF9AEEC831009F0942E2FC193CBBFC64" author="Gohlich UB &amp; Chiappe LM &amp; Clark JM" box="[1018,1207,921,945]" pageId="29" pageNumber="383" pagination="307 - 21" refId="ref31388" refString="Gohlich UB, Chiappe LM, Clark JM et al. The systematic position of the Late Jurassic alleged dinosaur Macelognathus (Crocodylomorpha: Sphenosuchia). Canadian Journal of Earth Sciences 2005; 42: 307 - 21." type="journal article" year="2005">
Göhlich
<emphasis id="B97F4F2B31009F09454AFC1A3C75FC64" box="[1106,1145,921,945]" italics="true" pageId="29" pageNumber="383">et al</emphasis>
. 2005
</bibRefCitation>
), whereas they are slightly more spread in
<taxonomicName id="4C0BE8BA31009F0942CCFC393D5EFC04" authority="(Lecuona et al. 2016)" baseAuthorityName="Lecuona" baseAuthorityYear="2016" box="[980,1362,953,977]" class="Polyplacophora" genus="Trialestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodilomorphi" pageId="29" pageNumber="383" phylum="Mollusca" rank="species" species="romeri">
<emphasis id="B97F4F2B31009F0942CCFC393C78FC05" box="[980,1140,953,977]" italics="true" pageId="29" pageNumber="383">Trialestes romeri</emphasis>
(
<bibRefCitation id="EF9AEEC831009F09459BFC393D4AFC04" author="Lecuona A &amp; Ezcurra MD &amp; Irmis RB" box="[1155,1350,953,977]" pageId="29" pageNumber="383" pagination="585 - 622" refId="ref32010" refString="Lecuona A, Ezcurra MD, Irmis RB. Revision of the early crocodylomorph Trialestes romeri (Archosauria, Suchia) from the lower Upper Triassic Ischigualasto Formation of Argentina: one of the oldest-known crocodylomorphs. Papers in Palaeontology 2016; 2: 585 - 622." type="journal article" year="2016">
Lecuona
<emphasis id="B97F4F2B31009F0945F9FC393D03FC04" box="[1249,1295,953,977]" italics="true" pageId="29" pageNumber="383">et al.</emphasis>
2016
</bibRefCitation>
)
</taxonomicName>
. The metatarsals differ from other non-crocodyliform crocodylomorphs in possessing a distinct longitudinal groove along the shass. This was aưributed to compression by
<bibRefCitation id="EF9AEEC831009F094596FB973D11FBFA" author="Crush PJ" box="[1166,1309,1047,1071]" pageId="29" pageNumber="383" refId="ref31157" refString="Crush PJ. An early, terrestrial, crocodile from South Wales. Unpublished Ph. D. Thesis, University College London, 1980." type="book" year="1980">Crush (1980)</bibRefCitation>
but it cannot be excluded that such a groove, possibly to a lesser extent, was also present in life.
</paragraph>
<paragraph id="8BB4933931009F09424DFBF43B74FA72" blockId="29.[825,1475,483,1447]" pageId="29" pageNumber="383">
The phalanges of SMNS 12597a have been partially disarticulated, preventing their referral to specific digits and determining the phalangeal count (
<figureCitation id="13308FBC31009F094504FB333C81FB1E" box="[1052,1165,1203,1227]" captionStart="Figure 12" captionStartId="24.[113,178,1772,1796]" captionTargetBox="[151,1424,144,1744]" captionTargetId="figure-6@24.[148,1424,144,1744]" captionTargetPageId="24" captionText="Figure 12. Details of the postcranial anatomy of Saltoposuchus connectens. A, dorsal vertebral neural arch and dorsal rib of SMNS 12596c in ventral view; B, associated osteoderms of SMNS 12596c in dorsal view; C, sacral ribs and associated elements of SMNS 12597a in dorsal view; D, distal portion of right scapular blade of SMNS 12597a in lateral view; E, proximal end of right tibia of SMNS 12597a in proximal view; F, less tarsus and pes of SMNS 12597a with the calcaneum exposed in lateral view; G, close-up of less tarsus of SMNS 12597a; H, close-up of less pes of SMNS 12597a. Abbreviations: ane, anterior expansion; cal, calcaneum; calt, calcaneal tuber; cp, capitulum; dp, diapophysis; dt, distal tarsal; exp, extensor pit; fe, femur; fi, fibula; il, ilium; lam, lamina; latc, lateral condyle; le, lateral edge; lr, longitudinal ridge; me, medial edge; mt, metatarsal; no, notch; pap, parapophysis; ph, phalanx; pme, posterior extension; poe, posterior expansion; pofl, posterior flange; poz, postzygapophysis; os, osteoderm; r., right; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426175" httpUri="https://zenodo.org/record/8426175/files/figure.png" pageId="29" pageNumber="383">Fig. 12F, H</figureCitation>
). However, based on the presence of two more robust, proximally positioned phalanges and two unguals, they likely represent remains of two digits. The non-terminal phalanges possess expanded proximal and distal ends relative to the shass. The proximal ends are flat to slightly concave, whereas the distal ends are marked by distinct extensor pits. The unguals are small in size and curved in lateral view (
<figureCitation id="13308FBC31009F094484FAEF3B6BFA72" captionStart="Figure 12" captionStartId="24.[113,178,1772,1796]" captionTargetBox="[151,1424,144,1744]" captionTargetId="figure-6@24.[148,1424,144,1744]" captionTargetPageId="24" captionText="Figure 12. Details of the postcranial anatomy of Saltoposuchus connectens. A, dorsal vertebral neural arch and dorsal rib of SMNS 12596c in ventral view; B, associated osteoderms of SMNS 12596c in dorsal view; C, sacral ribs and associated elements of SMNS 12597a in dorsal view; D, distal portion of right scapular blade of SMNS 12597a in lateral view; E, proximal end of right tibia of SMNS 12597a in proximal view; F, less tarsus and pes of SMNS 12597a with the calcaneum exposed in lateral view; G, close-up of less tarsus of SMNS 12597a; H, close-up of less pes of SMNS 12597a. Abbreviations: ane, anterior expansion; cal, calcaneum; calt, calcaneal tuber; cp, capitulum; dp, diapophysis; dt, distal tarsal; exp, extensor pit; fe, femur; fi, fibula; il, ilium; lam, lamina; latc, lateral condyle; le, lateral edge; lr, longitudinal ridge; me, medial edge; mt, metatarsal; no, notch; pap, parapophysis; ph, phalanx; pme, posterior extension; poe, posterior expansion; pofl, posterior flange; poz, postzygapophysis; os, osteoderm; r., right; sr, sacral rib; tb, tuberculum; ti, tibia; un, ungual." figureDoi="http://doi.org/10.5281/zenodo.8426175" httpUri="https://zenodo.org/record/8426175/files/figure.png" pageId="29" pageNumber="383">Fig. 12H</figureCitation>
).
</paragraph>
<paragraph id="8BB4933931009F094299FA473D77FA34" blockId="29.[897,1403,1479,1505]" box="[897,1403,1479,1505]" pageId="29" pageNumber="383">
<emphasis id="B97F4F2B31009F094299FA473D77FA34" box="[897,1403,1479,1505]" italics="true" pageId="29" pageNumber="383">Histological section of the right femur of SMNS 12596</emphasis>
</paragraph>
<paragraph id="8BB4933931009F0B4221FA6D3A8FFF32" blockId="29.[825,1475,1517,1980]" lastBlockId="31.[129,777,144,231]" lastPageId="31" lastPageNumber="385" pageId="29" pageNumber="383">
The strong anterolateral to posteromedial compression of the right femur SMNS 12596j is also reflected in the histological section of its mid-shass. The medullary cavity has been diagenetically reduced to an anteromedially to posterolaterally oriented slit-like opening (
<figureCitation id="13308FBC31009F0942E9F9EB3C45F956" box="[1009,1097,1643,1667]" captionStart="Figure 13" captionStartId="30.[114,179,1746,1770]" captionTargetBox="[146,1426,164,1719]" captionTargetId="figure-65@30.[146,1426,144,1719]" captionTargetPageId="30" captionText="Figure 13. Images of the histological cross section of the right femur SMNS 12596j.A, complete overview of the cross-section; B, C, close-up of the endosteal region under (B) normal and (C) polarized light; D, E, close-up of the periosteal region revealing the two annuli under (D) normal and (E) polarized light. Arrow heads indicate the laminae that form the annuli. Abbreviations:eb, endosteal bone; flb, fibrolamellar bone; mc, medullary cavity; vac, vascular canal." figureDoi="http://doi.org/10.5281/zenodo.8426177" httpUri="https://zenodo.org/record/8426177/files/figure.png" pageId="29" pageNumber="383">Fig. 13A</figureCitation>
). Along the preserved long axis, the medullary cavity comprises approximately half of the diameter of the shass. Previously studied long bone sections of non-crocodyliform crocodylomorphs comprise an isolated, likely immature humerus assigned to
<taxonomicName id="4C0BE8BA31009F094565F9683D3DF92A" box="[1149,1329,1768,1792]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" kingdom="Animalia" order="Prolacertiformes" pageId="29" pageNumber="383" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B97F4F2B31009F094565F9683D02F8D5" box="[1149,1294,1768,1792]" italics="true" pageId="29" pageNumber="383">Terrestrisuchus</emphasis>
sp.
</taxonomicName>
(Riqclès
<emphasis id="B97F4F2B31009F09448BF9693DCFF8D5" box="[1427,1475,1768,1792]" italics="true" pageId="29" pageNumber="383">et al.</emphasis>
2003) and a femur and long bone fragments of
<taxonomicName id="4C0BE8BA31009F09442DF8873B67F8EA" class="Reptilia" family="Sphenosuchidae" genus="Hesperosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="29" pageNumber="383" phylum="Chordata" rank="species" species="agilis">
<emphasis id="B97F4F2B31009F09442DF8873B67F8EA" italics="true" pageId="29" pageNumber="383">Hesperosuchus agilis</emphasis>
</taxonomicName>
(AMNH FARB 6767, material associated with
<typeStatus id="54B02D9B31009F09447EF8A73DCEF8EA" box="[1382,1474,1831,1855]" pageId="29" pageNumber="383">holotype</typeStatus>
specimen AMNH FARB 6758; Riqclès
<emphasis id="B97F4F2B31009F0945DFF8C73CF9F88B" box="[1223,1269,1862,1886]" italics="true" pageId="29" pageNumber="383">et al.</emphasis>
2008, Buffrénil
<emphasis id="B97F4F2B31009F09448DF8C73DCFF88B" box="[1429,1475,1862,1886]" italics="true" pageId="29" pageNumber="383">et al.</emphasis>
2021). In the former, the medullary cavity is clearly considerably larger compared to the preserved long axis in SMNS 12596j, whereas the medullary cavity of the laưer is approximately similar in width. However, it is largely filled by extensive trabeculae in
<taxonomicName id="4C0BE8BA31039F0A4169F8033930F84E" box="[113,316,1923,1947]" class="Reptilia" family="Sphenosuchidae" genus="Hesperosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="30" pageNumber="384" phylum="Chordata" rank="species" species="agilis">
<emphasis id="B97F4F2B31039F0A4169F8033930F84E" box="[113,316,1923,1947]" italics="true" pageId="30" pageNumber="384">Hesperosuchus agilis</emphasis>
</taxonomicName>
, which are absent in both
<taxonomicName id="4C0BE8BA31039F0A4371F8033881F86F" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" kingdom="Animalia" order="Prolacertiformes" pageId="30" pageNumber="384" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B97F4F2B31039F0A4371F8033AF6F84E" box="[617,762,1923,1947]" italics="true" pageId="30" pageNumber="384">Terrestrisuchus</emphasis>
sp.
</taxonomicName>
and
<taxonomicName id="4C0BE8BA31039F0A41D0F82239CDF86F" authorityName="Huene" authorityYear="1921" box="[200,449,1954,1978]" class="Reptilia" family="Saltoposuchidae" genus="Saltoposuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="30" pageNumber="384" phylum="Chordata" rank="species" species="connectens">
<emphasis id="B97F4F2B31039F0A41D0F82239CDF86F" box="[200,449,1954,1978]" italics="true" pageId="30" pageNumber="384">Saltoposuchus connectens</emphasis>
</taxonomicName>
. In SMNS 12596j, the inner margin of the periosteal tissue has been resorbed and subsequently an endosteal layer of lamellar bone with flaưened and elongated osteocyte lacunae has been deposited that is strongly birefringent in polarized light (
<figureCitation id="13308FBC31029F0B40A3FF103A20FF7D" box="[443,556,144,168]" captionStart="Figure 13" captionStartId="30.[114,179,1746,1770]" captionTargetBox="[146,1426,164,1719]" captionTargetId="figure-65@30.[146,1426,144,1719]" captionTargetPageId="30" captionText="Figure 13. Images of the histological cross section of the right femur SMNS 12596j.A, complete overview of the cross-section; B, C, close-up of the endosteal region under (B) normal and (C) polarized light; D, E, close-up of the periosteal region revealing the two annuli under (D) normal and (E) polarized light. Arrow heads indicate the laminae that form the annuli. Abbreviations:eb, endosteal bone; flb, fibrolamellar bone; mc, medullary cavity; vac, vascular canal." figureDoi="http://doi.org/10.5281/zenodo.8426177" httpUri="https://zenodo.org/record/8426177/files/figure.png" pageId="31" pageNumber="385">Fig. 13B, C</figureCitation>
). This layer is considerably thicker than the very thin endosteal layer identified in the humerus of
<taxonomicName id="4C0BE8BA31029F0B41E2FF4F39A0FF33" box="[250,428,207,231]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" kingdom="Animalia" order="Prolacertiformes" pageId="31" pageNumber="385" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B97F4F2B31029F0B41E2FF4F3987FF32" box="[250,395,207,231]" italics="true" pageId="31" pageNumber="385">Terrestrisuchus</emphasis>
sp.
</taxonomicName>
(Riqclès
<emphasis id="B97F4F2B31029F0B4314FF4F3A37FF32" box="[524,571,207,231]" italics="true" pageId="31" pageNumber="385">et al.</emphasis>
2003).
</paragraph>
<caption id="DF74C3B131039F0A416AF9523A23F8EA" ID-DOI="http://doi.org/10.5281/zenodo.8426177" ID-Zenodo-Dep="8426177" httpUri="https://zenodo.org/record/8426177/files/figure.png" pageId="30" pageNumber="384" startId="30.[114,179,1746,1770]" targetBox="[146,1426,164,1719]" targetPageId="30" targetType="figure">
<paragraph id="8BB4933931039F0A416AF9523A23F8EA" blockId="30.[113,1441,1746,1855]" pageId="30" pageNumber="384">
<emphasis id="B97F4F2B31039F0A416AF95238DBF93E" bold="true" box="[114,215,1746,1771]" pageId="30" pageNumber="384">Figure 13.</emphasis>
Images of the histological cross section of the right femur SMNS 12596j. A, complete overview of the cross-section; B, C, close-up of the endosteal region under (B) normal and (C) polarized light; D, E, close-up of the periosteal region revealing the two annuli under (D) normal and (E) polarized light. Arrow heads indicate the laminae that form the annuli. Abbreviations: eb, endosteal bone; flb, fibrolamellar bone; mc, medullary cavity; vac, vascular canal.
</paragraph>
</caption>
<paragraph id="8BB4933931029F0B4184FE8D3975FBE1" blockId="31.[129,778,269,1076]" pageId="31" pageNumber="385">
The periosteal matrix of SMNS 12596j is composed of primary fibrolamellar or woven-parallel bone with numerous, relatively round osteocyte lacunae. The cortex possesses a high density of simple vascular canals that are roughly equally distributed throughout the matrix (
<figureCitation id="13308FBC31029F0B40BCFE0B3A18FE76" box="[420,532,395,419]" captionStart="Figure 13" captionStartId="30.[114,179,1746,1770]" captionTargetBox="[146,1426,164,1719]" captionTargetId="figure-65@30.[146,1426,144,1719]" captionTargetPageId="30" captionText="Figure 13. Images of the histological cross section of the right femur SMNS 12596j.A, complete overview of the cross-section; B, C, close-up of the endosteal region under (B) normal and (C) polarized light; D, E, close-up of the periosteal region revealing the two annuli under (D) normal and (E) polarized light. Arrow heads indicate the laminae that form the annuli. Abbreviations:eb, endosteal bone; flb, fibrolamellar bone; mc, medullary cavity; vac, vascular canal." figureDoi="http://doi.org/10.5281/zenodo.8426177" httpUri="https://zenodo.org/record/8426177/files/figure.png" pageId="31" pageNumber="385">Fig. 13B, C</figureCitation>
). No secondary osteons are present, in contrast to the femoral section of
<taxonomicName id="4C0BE8BA31029F0B4364FE2A3990FE34" authority="(Buffrenil et al. 2021)" baseAuthorityName="Buffrenil" baseAuthorityYear="2021" class="Reptilia" family="Sphenosuchidae" genus="Hesperosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="31" pageNumber="385" phylum="Chordata" rank="species" species="agilis">
<emphasis id="B97F4F2B31029F0B4364FE2A38BFFE34" italics="true" pageId="31" pageNumber="385">Hesperosuchus agilis</emphasis>
(Buffrénil
<emphasis id="B97F4F2B31029F0B403DFE4A395AFE34" box="[293,342,457,481]" italics="true" pageId="31" pageNumber="385">et al.</emphasis>
2021)
</taxonomicName>
. The vascular canals are predominantly longitudinally oriented, with some forming anastomoses (
<figureCitation id="13308FBC31029F0B4193FD8838EFFDF5" box="[139,227,520,544]" captionStart="Figure 13" captionStartId="30.[114,179,1746,1770]" captionTargetBox="[146,1426,164,1719]" captionTargetId="figure-65@30.[146,1426,144,1719]" captionTargetPageId="30" captionText="Figure 13. Images of the histological cross section of the right femur SMNS 12596j.A, complete overview of the cross-section; B, C, close-up of the endosteal region under (B) normal and (C) polarized light; D, E, close-up of the periosteal region revealing the two annuli under (D) normal and (E) polarized light. Arrow heads indicate the laminae that form the annuli. Abbreviations:eb, endosteal bone; flb, fibrolamellar bone; mc, medullary cavity; vac, vascular canal." figureDoi="http://doi.org/10.5281/zenodo.8426177" httpUri="https://zenodo.org/record/8426177/files/figure.png" pageId="31" pageNumber="385">Fig. 13A</figureCitation>
). As such, the composition of the periosteal matrix is broadly similar to that seen in the humerus of
<taxonomicName id="4C0BE8BA31029F0B434FFDA73B06FDEA" box="[599,778,551,575]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" kingdom="Animalia" order="Prolacertiformes" pageId="31" pageNumber="385" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B97F4F2B31029F0B434FFDA73AE4FDEA" box="[599,744,551,575]" italics="true" pageId="31" pageNumber="385">Terrestrisuchus</emphasis>
sp.
</taxonomicName>
(Riqclès
<emphasis id="B97F4F2B31029F0B41C3FDC73906FD8A" box="[219,266,583,607]" italics="true" pageId="31" pageNumber="385">et al.</emphasis>
2003). In contrast, in
<taxonomicName id="4C0BE8BA31029F0B40F2FDC73AA2FD8A" box="[490,686,583,607]" class="Reptilia" family="Sphenosuchidae" genus="Hesperosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="31" pageNumber="385" phylum="Chordata" rank="species" species="agilis">
<emphasis id="B97F4F2B31029F0B40F2FDC73AA2FD8A" box="[490,686,583,607]" italics="true" pageId="31" pageNumber="385">Hesperosuchus agilis</emphasis>
</taxonomicName>
the periosteal matrix is formed by parallel-fibred tissue that is very distinctly lamellar for most of its width, being least organized near the medullary cavity (Riqclès
<emphasis id="B97F4F2B31029F0B40AFFD2539E4FD69" box="[439,488,676,700]" italics="true" pageId="31" pageNumber="385">et al.</emphasis>
2008, Buffrénil
<emphasis id="B97F4F2B31029F0B4396FD253AB3FD69" box="[654,703,676,700]" italics="true" pageId="31" pageNumber="385">et al.</emphasis>
2021). In the humerus of
<taxonomicName id="4C0BE8BA31029F0B4020FD4439E6FD0E" box="[312,490,708,732]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" kingdom="Animalia" order="Prolacertiformes" pageId="31" pageNumber="385" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B97F4F2B31029F0B4020FD4439C5FD09" box="[312,457,708,732]" italics="true" pageId="31" pageNumber="385">Terrestrisuchus</emphasis>
sp.
</taxonomicName>
the periosteal matrix is most organized (i.e. lamellar) near the periphery, as is also indicated by the presence of the two lines of arrested growth (LAGs) (Riqclès
<emphasis id="B97F4F2B31029F0B41C2FCA33904FCEF" box="[218,264,802,826]" italics="true" pageId="31" pageNumber="385">et al.</emphasis>
2003). Sharpey’s fibres, as present in
<taxonomicName id="4C0BE8BA31029F0B4364FCA23992FC8C" authority="(Buffrenil et al. 2021)" baseAuthorityName="Buffrenil" baseAuthorityYear="2021" class="Reptilia" family="Sphenosuchidae" genus="Hesperosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="31" pageNumber="385" phylum="Chordata" rank="species" species="agilis">
<emphasis id="B97F4F2B31029F0B4364FCA238BFFC8C" italics="true" pageId="31" pageNumber="385">Hesperosuchus agilis</emphasis>
(Buffrénil
<emphasis id="B97F4F2B31029F0B403FFCC23954FC8C" box="[295,344,833,857]" italics="true" pageId="31" pageNumber="385">et al.</emphasis>
2021)
</taxonomicName>
, are absent in SMNS 12596j. Two regions of increased organisation are present in the periosteal matrix of SMNS 12596j, most clearly visible in polarized light. Each area is composed of three closely packed lines (
<figureCitation id="13308FBC31029F0B43B2FC1F3883FC03" captionStart="Figure 13" captionStartId="30.[114,179,1746,1770]" captionTargetBox="[146,1426,164,1719]" captionTargetId="figure-65@30.[146,1426,144,1719]" captionTargetPageId="30" captionText="Figure 13. Images of the histological cross section of the right femur SMNS 12596j.A, complete overview of the cross-section; B, C, close-up of the endosteal region under (B) normal and (C) polarized light; D, E, close-up of the periosteal region revealing the two annuli under (D) normal and (E) polarized light. Arrow heads indicate the laminae that form the annuli. Abbreviations:eb, endosteal bone; flb, fibrolamellar bone; mc, medullary cavity; vac, vascular canal." figureDoi="http://doi.org/10.5281/zenodo.8426177" httpUri="https://zenodo.org/record/8426177/files/figure.png" pageId="31" pageNumber="385">Fig. 13D, E</figureCitation>
), which might reflect temporarily arrested growth. The first region of arrested growth is formed at approximately the centre of the cortex, whereas the second region is positioned close to the outer edge of the cortex.
</paragraph>
</subSubSection>
</treatment>
</document>
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