531 lines
79 KiB
XML
531 lines
79 KiB
XML
<document ID-DOI="10.1093/zoolinnean/zlac027" ID-GBIF-Dataset="97a5e580-c3c7-4542-8022-c0500c0368b4" ID-ISSN="0024-4082" ID-Zenodo-Dep="7695267" checkinTime="1677743926356" checkinUser="plazi" docAuthor="Cowgill, Thomas, Young Fls, Mark T, Schwab, Julia A, Walsh, Stig, Witmer, Lawrence M, Herrera, Yanina, Dollman, Kathleen N, Turner, Alan H & Brusatte, Stephen L" docDate="2023" docId="B45FFE69DC5C51564237D3063103FDA5" docLanguage="en" docName="zlac027.pdf" docOrigin="Zoological Journal of the Linnean Society 197 (3)" docSource="https://academic.oup.com/zoolinnean/article/197/3/812/6593330" docStyle="DocumentStyle:36B3BD6A90C22AB4F7F465C853188CC8.7:ZoolJLinnSoc.2017-.journal_article" docStyleId="36B3BD6A90C22AB4F7F465C853188CC8" docStyleName="ZoolJLinnSoc.2017-.journal_article" docStyleVersion="7" docTitle="Protosuchus haughtoni" docType="treatment" docVersion="3" lastPageNumber="823" masterDocId="48668611DC5A515D430CD14D3547FF89" masterDocTitle="Cephalic salt gland evolution in Mesozoic pelagic crocodylomorphs" masterLastPageNumber="835" masterPageNumber="812" pageNumber="818" updateTime="1677839180419" updateUser="ExternalLinkService">
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<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:titleInfo>
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<mods:title>Cephalic salt gland evolution in Mesozoic pelagic crocodylomorphs</mods:title>
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</mods:titleInfo>
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<mods:name type="personal">
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Cowgill, Thomas</mods:namePart>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Young Fls, Mark T</mods:namePart>
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<mods:name type="personal">
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Schwab, Julia A</mods:namePart>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Walsh, Stig</mods:namePart>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Witmer, Lawrence M</mods:namePart>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Herrera, Yanina</mods:namePart>
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<mods:name type="personal">
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Dollman, Kathleen N</mods:namePart>
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<mods:name type="personal">
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Turner, Alan H</mods:namePart>
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<mods:name type="personal">
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Brusatte, Stephen L</mods:namePart>
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<mods:date>2023</mods:date>
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<mods:number>2023-03-01</mods:number>
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<mods:number>197</mods:number>
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<mods:number>3</mods:number>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.7695269" ID-Zenodo-Dep="7695269" LSID="urn:lsid:plazi:treatment:B45FFE69DC5C51564237D3063103FDA5" httpUri="http://treatment.plazi.org/id/B45FFE69DC5C51564237D3063103FDA5" lastPageId="11" lastPageNumber="823" pageId="6" pageNumber="818">
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<subSubSection box="[315,626,587,610]" pageId="6" pageNumber="818" type="nomenclature">
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<paragraph blockId="6.[315,626,587,610]" box="[315,626,587,610]" pageId="6" pageNumber="818">
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<heading box="[315,626,587,610]" centered="true" fontSize="9" level="2" pageId="6" pageNumber="818" reason="2">
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<taxonomicName baseAuthorityName="Busbey & Gow" baseAuthorityYear="1984" box="[315,626,587,610]" class="Reptilia" family="Protosuchidae" genus="Protosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="6" pageNumber="833" phylum="Chordata" rank="species" species="haughtoni">
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<emphasis box="[315,626,587,610]" italics="true" pageId="6" pageNumber="818">
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<emphasis box="[315,485,587,610]" italics="true" pageId="6" pageNumber="818">PROTOSUCHUS</emphasis>
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HAUGHTONI
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</emphasis>
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</taxonomicName>
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</heading>
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</paragraph>
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</subSubSection>
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<subSubSection pageId="6" pageNumber="818" type="description">
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<paragraph blockId="6.[163,779,626,1047]" lastBlockId="6.[827,1444,197,894]" pageId="6" pageNumber="818">
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The nasal cavity in
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<taxonomicName baseAuthorityName="Busbey & Gow" baseAuthorityYear="1984" box="[380,641,626,647]" class="Reptilia" family="Protosuchidae" genus="Protosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="6" pageNumber="818" phylum="Chordata" rank="species" species="haughtoni">
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<emphasis box="[380,641,626,647]" italics="true" pageId="6" pageNumber="818">Protosuchus haughtoni</emphasis>
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</taxonomicName>
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(BP/1/4770) is almost complete, with only the right dorsolateral portion missing (
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<figureCitation box="[396,473,688,710]" captionStart="Figure 2" captionStartId="6.[164,242,1801,1823]" captionTargetBox="[328,1280,1092,1761]" captionTargetId="figure-462@6.[321,1284,1089,1762]" captionTargetPageId="6" captionText="Figure 2. Internal rostral anatomy of Protosuchus haughtoni (BP/1/4770) showing the nasal cavity and antorbital fenestra. A, B, dorsal (A) and left lateral (B) views with skull rendered transparent. C, D, transverse CT slices of nasal cavity showing the concavities where the nasal glands are inferred (C) and small dorsal expansions of the olfactory region (D). Internal reconstruction colours: nasal cavity, yellow; antorbital fenestra, green. Scale bars equal 10 mm (A, B) and 5 mm (C, D)." figureDoi="http://doi.org/10.5281/zenodo.7695273" httpUri="https://zenodo.org/record/7695273/files/figure.png" pageId="6" pageNumber="818">Fig. 2</figureCitation>
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). The naris and nasal vestibule are oriented anteriorly, and the nasal cavity proper is relatively short compared to more derived crocodyliforms because of the shorter, dorsoventrally tall rostrum. The olfactory region comprises a relatively larger portion of the nasal cavity and shows greater dorsoventral than mediolateral expansion. Ventral to the olfactory region the nasal cavity opens ventrally into a single choana, homologous with the primary choana in mesoeucrocodylians. Lateral to the choana, the antorbital cavity opens laterally from the nasal cavity to create an external antorbital fenestra surrounded by the maxilla dorsally and ventrally and the lacrimal posteriorly (
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<figureCitation box="[1291,1375,228,250]" captionStart="Figure 2" captionStartId="6.[164,242,1801,1823]" captionTargetBox="[328,1280,1092,1761]" captionTargetId="figure-462@6.[321,1284,1089,1762]" captionTargetPageId="6" captionText="Figure 2. Internal rostral anatomy of Protosuchus haughtoni (BP/1/4770) showing the nasal cavity and antorbital fenestra. A, B, dorsal (A) and left lateral (B) views with skull rendered transparent. C, D, transverse CT slices of nasal cavity showing the concavities where the nasal glands are inferred (C) and small dorsal expansions of the olfactory region (D). Internal reconstruction colours: nasal cavity, yellow; antorbital fenestra, green. Scale bars equal 10 mm (A, B) and 5 mm (C, D)." figureDoi="http://doi.org/10.5281/zenodo.7695273" httpUri="https://zenodo.org/record/7695273/files/figure.png" pageId="6" pageNumber="818">Fig. 2B</figureCitation>
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). The dorsal surface of the nasal cavity exhibits a shallow depression ventral to a descending process of the nasals that medially separates two shallow dorsal expansions; at the olfactory region these expansions are significantly deeper (
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<figureCitation box="[1109,1190,381,403]" captionStart="Figure 2" captionStartId="6.[164,242,1801,1823]" captionTargetBox="[328,1280,1092,1761]" captionTargetId="figure-462@6.[321,1284,1089,1762]" captionTargetPageId="6" captionText="Figure 2. Internal rostral anatomy of Protosuchus haughtoni (BP/1/4770) showing the nasal cavity and antorbital fenestra. A, B, dorsal (A) and left lateral (B) views with skull rendered transparent. C, D, transverse CT slices of nasal cavity showing the concavities where the nasal glands are inferred (C) and small dorsal expansions of the olfactory region (D). Internal reconstruction colours: nasal cavity, yellow; antorbital fenestra, green. Scale bars equal 10 mm (A, B) and 5 mm (C, D)." figureDoi="http://doi.org/10.5281/zenodo.7695273" httpUri="https://zenodo.org/record/7695273/files/figure.png" pageId="6" pageNumber="818">Fig. 2C</figureCitation>
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).
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</paragraph>
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</subSubSection>
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<subSubSection lastPageId="11" lastPageNumber="823" pageId="6" pageNumber="818" type="description">
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<paragraph blockId="6.[827,1444,197,894]" pageId="6" pageNumber="818">
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Using the Extant Phylogenetic Bracket method,
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<bibRefCitation author="Witmer LM" box="[827,1017,442,464]" pageId="6" pageNumber="818" pagination="269 - 327" refId="ref17586" refString="Witmer LM. 1995. Homology of facial structures in extant archosaurs (birds and crocodilians), with special reference to paranasal pneumaticity and nasal conchae. Journal of Morphology 225: 269 - 327." type="journal article" year="1995">Witmer (1995</bibRefCitation>
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,
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<bibRefCitation author="Witmer LM" box="[1042,1105,443,464]" pageId="6" pageNumber="818" pagination="1 - 76" refId="ref17623" refString="Witmer LM. 1997. The evolution of the antorbital cavity in archosaurs: a study in soft-tissue reconstruction in the fossil record with an analysis of the function of pneumaticity. Journal of Vertebrate Palaeontology 17: 1 - 76." type="journal article" year="1997">1997</bibRefCitation>
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) hypothesized that the plesiomorphic archosaur morphology of the nasal gland was positioned dorsally or dorsolaterally within the rostrum and excavated grooved internal surfaces of the bones close to the nasomaxillary suture. A shallow concave internal surface around the nasomaxillary suture is present in
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<taxonomicName authorityName="Brown" authorityYear="1934" box="[1065,1207,626,647]" class="Reptilia" family="Protosuchidae" genus="Protosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="6" pageNumber="818" phylum="Chordata" rank="genus">
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<emphasis box="[1065,1207,626,647]" italics="true" pageId="6" pageNumber="818">Protosuchus</emphasis>
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</taxonomicName>
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(
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<figureCitation box="[1223,1308,626,648]" captionStart="Figure 2" captionStartId="6.[164,242,1801,1823]" captionTargetBox="[328,1280,1092,1761]" captionTargetId="figure-462@6.[321,1284,1089,1762]" captionTargetPageId="6" captionText="Figure 2. Internal rostral anatomy of Protosuchus haughtoni (BP/1/4770) showing the nasal cavity and antorbital fenestra. A, B, dorsal (A) and left lateral (B) views with skull rendered transparent. C, D, transverse CT slices of nasal cavity showing the concavities where the nasal glands are inferred (C) and small dorsal expansions of the olfactory region (D). Internal reconstruction colours: nasal cavity, yellow; antorbital fenestra, green. Scale bars equal 10 mm (A, B) and 5 mm (C, D)." figureDoi="http://doi.org/10.5281/zenodo.7695273" httpUri="https://zenodo.org/record/7695273/files/figure.png" pageId="6" pageNumber="818">Fig. 2D</figureCitation>
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) across the posterior half of the rostrum but is more poorly defined than in the extant crocodylians. The dorsal expansions of the nasal cavity are located dorsomedially to the nasomaxillary suture and are unlikely to be associated with the nasal gland. Instead, they most likely indicate the dorsal surface of the cartilaginous nasal capsule with the descending nasal process inferring the location of the nasal septum.
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</paragraph>
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<paragraph blockId="6.[1033,1236,954,979]" box="[1033,1236,954,979]" pageId="6" pageNumber="818">THALATTOSUCHIA</paragraph>
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<paragraph blockId="6.[827,1442,994,1047]" lastBlockId="7.[145,761,197,771]" lastPageId="7" lastPageNumber="819" pageId="6" pageNumber="818">
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The nasal cavity reconstructions in the thalattosuchian skulls comprise only the posterior half of the nasal cavity, with the exception of
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<taxonomicName baseAuthorityName="Fernandez & Herrera" baseAuthorityYear="2009" box="[466,761,198,219]" class="Reptilia" family="Metriorhynchidae" genus="Cricosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="7" pageNumber="819" phylum="Chordata" rank="species" species="araucanensis">
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<emphasis box="[466,761,198,219]" italics="true" pageId="7" pageNumber="819">Cricosaurus araucanensis</emphasis>
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</taxonomicName>
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(MLP 72-IV-7-1) where the whole rostrum and nasal cavity is preserved.
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</paragraph>
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<caption ID-DOI="http://doi.org/10.5281/zenodo.7695273" ID-Zenodo-Dep="7695273" httpUri="https://zenodo.org/record/7695273/files/figure.png" pageId="6" pageNumber="818" startId="6.[164,242,1801,1823]" targetBox="[328,1280,1092,1761]" targetPageId="6">
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<paragraph blockId="6.[163,1444,1801,1911]" pageId="6" pageNumber="818">
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<emphasis bold="true" box="[164,267,1801,1823]" pageId="6" pageNumber="818">Figure 2.</emphasis>
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Internal rostral anatomy of
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<taxonomicName baseAuthorityName="Busbey & Gow" baseAuthorityYear="1984" box="[565,804,1801,1823]" class="Reptilia" family="Protosuchidae" genus="Protosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="6" pageNumber="818" phylum="Chordata" rank="species" species="haughtoni">
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<emphasis box="[565,804,1801,1823]" italics="true" pageId="6" pageNumber="818">Protosuchus haughtoni</emphasis>
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</taxonomicName>
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(BP/1/4770) showing the nasal cavity and antorbital fenestra. A, B, dorsal (A) and left lateral (B) views with skull rendered transparent. C, D, transverse CT slices of nasal cavity showing the concavities where the nasal glands are inferred (C) and small dorsal expansions of the olfactory region (D). Internal reconstruction colours: nasal cavity, yellow; antorbital fenestra, green. Scale bars equal 10 mm (A, B) and 5 mm (C, D).
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</paragraph>
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</caption>
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<paragraph blockId="7.[145,761,197,771]" lastBlockId="7.[809,1425,197,771]" pageId="7" pageNumber="819">
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The basal thalattosuchian
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<taxonomicName baseAuthorityName="Westphal" baseAuthorityYear="1961" genus="Plagiophthalmosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="819" phylum="Chordata" rank="species" species="gracilirostris">
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<emphasis italics="true" pageId="7" pageNumber="819">Plagiophthalmosuchus gracilirostris</emphasis>
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</taxonomicName>
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(NHMUK PV OR 15500) shows a nasal cavity morphology comparable to that of extant crocodylians with a tubular nasal cavity proper with an expanded olfactory region at the posterior end of the rostrum (
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<figureCitation box="[301,417,442,464]" captionStart="Figure 3" captionStartId="7.[144,225,1683,1705]" captionTargetBox="[154,1428,804,1639]" captionTargetId="figure-324@7.[145,1428,795,1643]" captionTargetPageId="7" captionText="Figure 3. Internal rostral anatomy of basal thalattosuchian Plagiophthalmosuchus gracilirostris [NHMUK PV OR 15500 (A-C)] and teleosauroid Macrospondylus bollensis [NHMUK PV OR 14436 (D-F) and MCZ VPRA-1063 (G-I)]. A, B, Plagiophthalmosuchus in dorsal (A) and anterodorsal views (B). C, transverse CT slice of nasal cavity olfactory region. D, E, NHMUK PV OR 14436 in dorsal (D) and anterodorsal views (E). F, transverse CT slice of nasal cavity olfactory region showing small dorsal expansions where of the olfactory region where the nasal capsule is inferred. G, H, MCZ VPRA-1063 in dorsal (G) and anterodorsal views (H). I, transverse CT slice showing dorsal expansions of the olfactory region. All skulls are rendered transparent. Internal reconstruction colours: nasal cavity, yellow; antorbital fenestra, green. Scale bars equal 50 mm (A, B, D, E, G, H) and 10 mm (C, F, I)." figureDoi="http://doi.org/10.5281/zenodo.7695275" httpUri="https://zenodo.org/record/7695275/files/figure.png" pageId="7" pageNumber="819">Fig. 3A, B</figureCitation>
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). The antorbital cavity opens laterally into a reduced external antorbital fenestra (preserved on the right side of the skull) compared to
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<taxonomicName authorityName="Brown" authorityYear="1934" box="[178,326,534,555]" class="Reptilia" family="Protosuchidae" genus="Protosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="7" pageNumber="819" phylum="Chordata" rank="genus">
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<emphasis box="[178,326,534,555]" italics="true" pageId="7" pageNumber="819">Protosuchus</emphasis>
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</taxonomicName>
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, which is bordered by the lacrimal dorsally and maxilla ventrally. The nasals comprising the anterodorsal walls of the nasal cavity are missing and the boundaries between the fossilized bones and sedimentary matrix are poorly defined around the olfactory region (
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<figureCitation box="[347,432,688,710]" captionStart="Figure 3" captionStartId="7.[144,225,1683,1705]" captionTargetBox="[154,1428,804,1639]" captionTargetId="figure-324@7.[145,1428,795,1643]" captionTargetPageId="7" captionText="Figure 3. Internal rostral anatomy of basal thalattosuchian Plagiophthalmosuchus gracilirostris [NHMUK PV OR 15500 (A-C)] and teleosauroid Macrospondylus bollensis [NHMUK PV OR 14436 (D-F) and MCZ VPRA-1063 (G-I)]. A, B, Plagiophthalmosuchus in dorsal (A) and anterodorsal views (B). C, transverse CT slice of nasal cavity olfactory region. D, E, NHMUK PV OR 14436 in dorsal (D) and anterodorsal views (E). F, transverse CT slice of nasal cavity olfactory region showing small dorsal expansions where of the olfactory region where the nasal capsule is inferred. G, H, MCZ VPRA-1063 in dorsal (G) and anterodorsal views (H). I, transverse CT slice showing dorsal expansions of the olfactory region. All skulls are rendered transparent. Internal reconstruction colours: nasal cavity, yellow; antorbital fenestra, green. Scale bars equal 50 mm (A, B, D, E, G, H) and 10 mm (C, F, I)." figureDoi="http://doi.org/10.5281/zenodo.7695275" httpUri="https://zenodo.org/record/7695275/files/figure.png" pageId="7" pageNumber="819">Fig. 3C</figureCitation>
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), particularly on the dorsal surface, so it is not possible to identify any osteological correlates for either nasal glands comparable to crocodylians or enlarged nasal salt glands in metriorhynchids.
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</paragraph>
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<paragraph blockId="7.[809,1425,197,771]" lastBlockId="8.[163,780,197,1905]" lastPageId="8" lastPageNumber="820" pageId="7" pageNumber="819">
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The olfactory region of the nasal cavity is better preserved in the teleosauroid skulls (
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<figureCitation box="[1266,1332,289,311]" captionStart="Figure 3" captionStartId="7.[144,225,1683,1705]" captionTargetBox="[154,1428,804,1639]" captionTargetId="figure-324@7.[145,1428,795,1643]" captionTargetPageId="7" captionText="Figure 3. Internal rostral anatomy of basal thalattosuchian Plagiophthalmosuchus gracilirostris [NHMUK PV OR 15500 (A-C)] and teleosauroid Macrospondylus bollensis [NHMUK PV OR 14436 (D-F) and MCZ VPRA-1063 (G-I)]. A, B, Plagiophthalmosuchus in dorsal (A) and anterodorsal views (B). C, transverse CT slice of nasal cavity olfactory region. D, E, NHMUK PV OR 14436 in dorsal (D) and anterodorsal views (E). F, transverse CT slice of nasal cavity olfactory region showing small dorsal expansions where of the olfactory region where the nasal capsule is inferred. G, H, MCZ VPRA-1063 in dorsal (G) and anterodorsal views (H). I, transverse CT slice showing dorsal expansions of the olfactory region. All skulls are rendered transparent. Internal reconstruction colours: nasal cavity, yellow; antorbital fenestra, green. Scale bars equal 50 mm (A, B, D, E, G, H) and 10 mm (C, F, I)." figureDoi="http://doi.org/10.5281/zenodo.7695275" httpUri="https://zenodo.org/record/7695275/files/figure.png" pageId="7" pageNumber="819">Fig. 3</figureCitation>
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D-I). In
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<taxonomicName box="[809,1131,320,341]" genus="Macrospondylus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="819" phylum="Chordata" rank="species" species="bollensis">
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<emphasis box="[809,1131,320,341]" italics="true" pageId="7" pageNumber="819">Macrospondylus bollensis</emphasis>
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</taxonomicName>
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the olfactory region of both NHMUK PV OR 14436 (
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<figureCitation box="[1176,1244,350,372]" captionStart="Figure 3" captionStartId="7.[144,225,1683,1705]" captionTargetBox="[154,1428,804,1639]" captionTargetId="figure-324@7.[145,1428,795,1643]" captionTargetPageId="7" captionText="Figure 3. Internal rostral anatomy of basal thalattosuchian Plagiophthalmosuchus gracilirostris [NHMUK PV OR 15500 (A-C)] and teleosauroid Macrospondylus bollensis [NHMUK PV OR 14436 (D-F) and MCZ VPRA-1063 (G-I)]. A, B, Plagiophthalmosuchus in dorsal (A) and anterodorsal views (B). C, transverse CT slice of nasal cavity olfactory region. D, E, NHMUK PV OR 14436 in dorsal (D) and anterodorsal views (E). F, transverse CT slice of nasal cavity olfactory region showing small dorsal expansions where of the olfactory region where the nasal capsule is inferred. G, H, MCZ VPRA-1063 in dorsal (G) and anterodorsal views (H). I, transverse CT slice showing dorsal expansions of the olfactory region. All skulls are rendered transparent. Internal reconstruction colours: nasal cavity, yellow; antorbital fenestra, green. Scale bars equal 50 mm (A, B, D, E, G, H) and 10 mm (C, F, I)." figureDoi="http://doi.org/10.5281/zenodo.7695275" httpUri="https://zenodo.org/record/7695275/files/figure.png" pageId="7" pageNumber="819">Fig. 3</figureCitation>
|
|
D-F) and MCZ VPRA-1063 (
|
|
<figureCitation box="[972,1041,381,403]" captionStart="Figure 3" captionStartId="7.[144,225,1683,1705]" captionTargetBox="[154,1428,804,1639]" captionTargetId="figure-324@7.[145,1428,795,1643]" captionTargetPageId="7" captionText="Figure 3. Internal rostral anatomy of basal thalattosuchian Plagiophthalmosuchus gracilirostris [NHMUK PV OR 15500 (A-C)] and teleosauroid Macrospondylus bollensis [NHMUK PV OR 14436 (D-F) and MCZ VPRA-1063 (G-I)]. A, B, Plagiophthalmosuchus in dorsal (A) and anterodorsal views (B). C, transverse CT slice of nasal cavity olfactory region. D, E, NHMUK PV OR 14436 in dorsal (D) and anterodorsal views (E). F, transverse CT slice of nasal cavity olfactory region showing small dorsal expansions where of the olfactory region where the nasal capsule is inferred. G, H, MCZ VPRA-1063 in dorsal (G) and anterodorsal views (H). I, transverse CT slice showing dorsal expansions of the olfactory region. All skulls are rendered transparent. Internal reconstruction colours: nasal cavity, yellow; antorbital fenestra, green. Scale bars equal 50 mm (A, B, D, E, G, H) and 10 mm (C, F, I)." figureDoi="http://doi.org/10.5281/zenodo.7695275" httpUri="https://zenodo.org/record/7695275/files/figure.png" pageId="7" pageNumber="819">Fig. 3</figureCitation>
|
|
G-I) exhibits two small dorsal expansions ventral to the nasals and medial to the lacrimals, separated by a shallow midline depression (
|
|
<figureCitation box="[817,929,473,495]" captionStart="Figure 3" captionStartId="7.[144,225,1683,1705]" captionTargetBox="[154,1428,804,1639]" captionTargetId="figure-324@7.[145,1428,795,1643]" captionTargetPageId="7" captionText="Figure 3. Internal rostral anatomy of basal thalattosuchian Plagiophthalmosuchus gracilirostris [NHMUK PV OR 15500 (A-C)] and teleosauroid Macrospondylus bollensis [NHMUK PV OR 14436 (D-F) and MCZ VPRA-1063 (G-I)]. A, B, Plagiophthalmosuchus in dorsal (A) and anterodorsal views (B). C, transverse CT slice of nasal cavity olfactory region. D, E, NHMUK PV OR 14436 in dorsal (D) and anterodorsal views (E). F, transverse CT slice of nasal cavity olfactory region showing small dorsal expansions where of the olfactory region where the nasal capsule is inferred. G, H, MCZ VPRA-1063 in dorsal (G) and anterodorsal views (H). I, transverse CT slice showing dorsal expansions of the olfactory region. All skulls are rendered transparent. Internal reconstruction colours: nasal cavity, yellow; antorbital fenestra, green. Scale bars equal 50 mm (A, B, D, E, G, H) and 10 mm (C, F, I)." figureDoi="http://doi.org/10.5281/zenodo.7695275" httpUri="https://zenodo.org/record/7695275/files/figure.png" pageId="7" pageNumber="819">Fig. 3F, H</figureCitation>
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), concurring with the previous endocranial reconstruction of MCZ VPRA-1063 by
|
|
<bibRefCitation author="Wilberg EW & Beyl AR & Pierce SE & Turner AH" pageId="7" pageNumber="819" refId="ref17504" refString="Wilberg EW, Beyl AR, Pierce SE, Turner AH. 2021. Cranial and endocranial anatomy of a three-dimensionally preserved teleosauroid thalattosuchian skull. The Anatomical Record [early view]. doi: 10.1002 / ar. 24704." type="book" year="2021">
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Wilberg
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<emphasis box="[1366,1425,504,525]" italics="true" pageId="7" pageNumber="819">et al.</emphasis>
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|
(2021)
|
|
</bibRefCitation>
|
|
. This morphology is better demonstrated in MCZ VPRA-1063 where the whole olfactory region is preserved, whereas only the posterior portion is present in NHMUK PV OR 14436. Despite their occurrence in the olfactory region, when compared to metriorhynchid natural endocasts these expansions are located further dorsally in the olfactory region than the dorsolaterally positioned salt glands (
|
|
<bibRefCitation author="Fernandez M & Gasparini Z" lastPageId="8" lastPageNumber="820" pageId="7" pageNumber="819" pagination="269 - 276" refId="ref15303" refString="Fernandez M, Gasparini Z. 2000. Salt glands in a Tithonian metriorhynchid crocodyliform and their physiological significance. Lethaia 33: 269 - 276." type="journal article" year="2000">Fernández & Gasparini, 2000</bibRefCitation>
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,
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<bibRefCitation author="Fernandez M & Gasparini Z" box="[356,411,197,218]" pageId="8" pageNumber="820" pagination="79 - 84" refId="ref15330" refString="Fernandez M, Gasparini Z. 2008. Salt glands in the Jurassic metriorhynchid Geosaurus: implications for the evolution of osmoregulation in Mesozoic marine crocodyliforms. Naturaeissenschaften 95: 79 - 84." type="journal article" year="2008">2008</bibRefCitation>
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;
|
|
<bibRefCitation author="Herrera Y & Fernandez MS & Gasparini Z" box="[424,646,197,219]" pageId="8" pageNumber="820" pagination="331 - 340" refId="ref15839" refString="Herrera Y, Fernandez MS, Gasparini Z. 2013. The snout of Cricosaurus araucanensis: a case study in novel anatomy of the nasal region of metriorhynchids. Lethaia 46: 331 - 340." type="journal article" year="2013">
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Herrera
|
|
<emphasis box="[522,578,197,219]" italics="true" pageId="8" pageNumber="820">et al.</emphasis>
|
|
, 2013
|
|
</bibRefCitation>
|
|
;
|
|
<bibRefCitation author="Fernandez MS & Herrera Y" pageId="8" pageNumber="820" refId="ref15435" refString="Fernandez MS, Herrera Y. 2021. Active airflow of the paranasal sinuses in extinct crocodyliforms: evidence from a natural cast of the thalattosuchian Dakosaurus andiniensis. The Anatomical Record [early view]. doi: 10.1002 / ar. 24678." type="book" year="2021">Fernández & Herrera, 2021</bibRefCitation>
|
|
). However, their location is comparable to the small olfactory region expansions present in
|
|
<taxonomicName authorityName="Brown" authorityYear="1934" box="[163,313,289,310]" class="Reptilia" family="Protosuchidae" genus="Protosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="8" pageNumber="820" phylum="Chordata" rank="genus">
|
|
<emphasis box="[163,313,289,310]" italics="true" pageId="8" pageNumber="820">Protosuchus</emphasis>
|
|
</taxonomicName>
|
|
. It is also consistent with shallower dorsal convexities of the natural endocasts identified as part of the olfactory region of the cartilaginous nasal capsule (
|
|
<bibRefCitation author="Herrera Y & Fernandez MS & Gasparini Z" box="[347,588,381,403]" pageId="8" pageNumber="820" pagination="331 - 340" refId="ref15839" refString="Herrera Y, Fernandez MS, Gasparini Z. 2013. The snout of Cricosaurus araucanensis: a case study in novel anatomy of the nasal region of metriorhynchids. Lethaia 46: 331 - 340." type="journal article" year="2013">
|
|
Herrera
|
|
<emphasis box="[453,514,381,403]" italics="true" pageId="8" pageNumber="820">et al.</emphasis>
|
|
, 2013
|
|
</bibRefCitation>
|
|
). Therefore, we cannot confidently infer salt glands were present in
|
|
<taxonomicName box="[163,349,442,463]" genus="Macrospondylus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="820" phylum="Chordata" rank="genus">
|
|
<emphasis box="[163,349,442,463]" italics="true" pageId="8" pageNumber="820">Macrospondylus</emphasis>
|
|
</taxonomicName>
|
|
from the endocranial reconstructions, and the observed dorsal expansions likely correspond to the nasal capsule instead. The antorbital cavity of
|
|
<taxonomicName box="[192,384,534,555]" genus="Macrospondylus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="820" phylum="Chordata" rank="genus">
|
|
<emphasis box="[192,384,534,555]" italics="true" pageId="8" pageNumber="820">Macrospondylus</emphasis>
|
|
</taxonomicName>
|
|
is preserved in MCZ VPRA-1063 and, as with
|
|
<taxonomicName box="[318,587,565,586]" genus="Plagiophthalmosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="820" phylum="Chordata" rank="genus">
|
|
<emphasis box="[318,587,565,586]" italics="true" pageId="8" pageNumber="820">Plagiophthalmosuchus</emphasis>
|
|
</taxonomicName>
|
|
, opens laterally through a small antorbital fenestra. The antorbital fenestra is bordered by the lacrimal dorsally, the maxilla anteriorly and (unlike
|
|
<taxonomicName box="[511,771,657,678]" genus="Plagiophthalmosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="820" phylum="Chordata" rank="genus">
|
|
<emphasis box="[511,771,657,678]" italics="true" pageId="8" pageNumber="820">Plagiophthalmosuchus</emphasis>
|
|
</taxonomicName>
|
|
) the anterior process of the jugal ventrally.
|
|
</paragraph>
|
|
<caption ID-DOI="http://doi.org/10.5281/zenodo.7695275" ID-Zenodo-Dep="7695275" httpUri="https://zenodo.org/record/7695275/files/figure.png" pageId="7" pageNumber="819" startId="7.[144,225,1683,1705]" targetBox="[154,1428,804,1639]" targetPageId="7">
|
|
<paragraph blockId="7.[144,1426,1683,1910]" pageId="7" pageNumber="819">
|
|
<emphasis bold="true" box="[144,254,1683,1705]" pageId="7" pageNumber="819">Figure 3.</emphasis>
|
|
Internal rostral anatomy of basal thalattosuchian
|
|
<taxonomicName baseAuthorityName="Westphal" baseAuthorityYear="1961" box="[827,1227,1683,1705]" genus="Plagiophthalmosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="819" phylum="Chordata" rank="species" species="gracilirostris">
|
|
<emphasis box="[827,1227,1683,1705]" italics="true" pageId="7" pageNumber="819">Plagiophthalmosuchus gracilirostris</emphasis>
|
|
</taxonomicName>
|
|
[NHMUK PV OR 15500 (A-C)] and teleosauroid
|
|
<taxonomicName box="[469,741,1712,1734]" genus="Macrospondylus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="819" phylum="Chordata" rank="species" species="bollensis">
|
|
<emphasis box="[469,741,1712,1734]" italics="true" pageId="7" pageNumber="819">Macrospondylus bollensis</emphasis>
|
|
</taxonomicName>
|
|
[NHMUK PV OR 14436 (D-F) and MCZ VPRA-1063 (G-I)]. A, B,
|
|
<taxonomicName box="[145,386,1742,1763]" genus="Plagiophthalmosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="819" phylum="Chordata" rank="genus">
|
|
<emphasis box="[145,386,1742,1763]" italics="true" pageId="7" pageNumber="819">Plagiophthalmosuchus</emphasis>
|
|
</taxonomicName>
|
|
in dorsal (A) and anterodorsal views (B). C, transverse CT slice of nasal cavity olfactory region. D, E, NHMUK PV OR 14436 in dorsal (D) and anterodorsal views (E). F, transverse CT slice of nasal cavity olfactory region showing small dorsal expansions where of the olfactory region where the nasal capsule is inferred. G, H, MCZ VPRA-1063 in dorsal (G) and anterodorsal views (H). I, transverse CT slice showing dorsal expansions of the olfactory region. All skulls are rendered transparent. Internal reconstruction colours: nasal cavity, yellow; antorbital fenestra, green. Scale bars equal 50 mm (A, B, D, E, G, H) and 10 mm (C, F, I).
|
|
</paragraph>
|
|
</caption>
|
|
<paragraph blockId="8.[163,780,197,1905]" lastBlockId="8.[827,1444,197,1905]" pageId="8" pageNumber="820">
|
|
In the basal metriorhynchoid
|
|
<emphasis box="[548,779,718,740]" italics="true" pageId="8" pageNumber="820">
|
|
<taxonomicName baseAuthorityName="Witmer" baseAuthorityYear="1997" box="[548,706,718,739]" class="Reptilia" family="Metriorhynchidae" genus="Pelagosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="8" pageNumber="820" phylum="Chordata" rank="genus">Pelagosaurus</taxonomicName>
|
|
<typeStatus box="[715,779,719,740]" pageId="8" pageNumber="820">typus</typeStatus>
|
|
</emphasis>
|
|
(NHMUK PV OR 32599), the olfactory region differs from crocodylians and teleosauroids by possessing a deep ridge on the ventral surface of the nasals, creating a V-shaped depression on the dorsal surface of the olfactory region that separates the shallow dorsolateral expansions on the internal surface of the prefrontals and lacrimals (
|
|
<figureCitation box="[520,584,933,955]" captionStart="Figure 4" captionStartId="9.[145,225,1611,1633]" captionTargetBox="[305,1266,199,1570]" captionTargetId="figure-26@9.[303,1266,195,1571]" captionTargetPageId="9" captionText="Figure 4. Internal rostral anatomy of basal metriorhynchoids Pelagosaurus typus [NHMUK PV OR 32599 (A-E)] and Eoneustes gaudryi [NHMUK PV R 3263 (F-H)]. A-C, Pelagosaurus in dorsal (A), anterodorsal (B) and left lateral views (C) with the skull rendered transparent. D, E, transverse CT slices of nasal cavity olfactory region showing dorsal expansions where salt glands are inferred, the dorsolateral groove separating the inferred positions of salt glands and the nasal capsule (D) and the nasal ridge creating a V-shaped depression of the olfactory region ventral to the nasals (E). F-G, Eoneustes in dorsal (F) and left lateral views (G) with the skull rendered transparent. H, transverse CT slice of nasal cavity olfactory region showing the left salt duct, larger dorsolateral expansions where salt glands are inferred and ridge on the ventral surface of the nasals. Internal reconstruction colours: nasal cavity, yellow; nasopharyngeal ducts, pale yellow; antorbital fenestra, green; preobrital fenestra, light green. Scale bars equal 50 mm (A-C, F, G) and 10 mm (D, E, H)." figureDoi="http://doi.org/10.5281/zenodo.7695277" httpUri="https://zenodo.org/record/7695277/files/figure.png" pageId="8" pageNumber="820">Fig. 4</figureCitation>
|
|
A-E). Such nasal cavity expansions have previously been reported in
|
|
<taxonomicName baseAuthorityName="Witmer" baseAuthorityYear="1997" box="[163,326,994,1015]" class="Reptilia" family="Metriorhynchidae" genus="Pelagosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="8" pageNumber="820" phylum="Chordata" rank="genus">
|
|
<emphasis box="[163,326,994,1015]" italics="true" pageId="8" pageNumber="820">Pelagosaurus</emphasis>
|
|
</taxonomicName>
|
|
endocranial reconstructions (
|
|
<bibRefCitation author="Pierce SE & Williams M & Benton MJ" pageId="8" pageNumber="820" pagination="3225" refId="ref16732" refString="Pierce SE, Williams M, Benton MJ. 2017. Virtual reconstruction of the endocranial anatomy of the Early Jurassic marine crocodylomorph Pelagosaurus typus (Thalattosuchia). PeerJ 5: e 3225." type="journal article" year="2017">
|
|
Pierce
|
|
<emphasis box="[163,223,1025,1047]" italics="true" pageId="8" pageNumber="820">et al.</emphasis>
|
|
, 2017
|
|
</bibRefCitation>
|
|
) and, unlike the more dorsally oriented expansions in teleosauroids, the morphology present in
|
|
<taxonomicName baseAuthorityName="Witmer" baseAuthorityYear="1997" box="[198,364,1086,1107]" class="Reptilia" family="Metriorhynchidae" genus="Pelagosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="8" pageNumber="820" phylum="Chordata" rank="genus">
|
|
<emphasis box="[198,364,1086,1107]" italics="true" pageId="8" pageNumber="820">Pelagosaurus</emphasis>
|
|
</taxonomicName>
|
|
more closely corresponds to the dorsolateral position of salt glands preserved in natural endocasts of metriorhynchids (
|
|
<bibRefCitation author="Fernandez M & Gasparini Z" pageId="8" pageNumber="820" pagination="269 - 276" refId="ref15303" refString="Fernandez M, Gasparini Z. 2000. Salt glands in a Tithonian metriorhynchid crocodyliform and their physiological significance. Lethaia 33: 269 - 276." type="journal article" year="2000">Fernández & Gasparini, 2000</bibRefCitation>
|
|
,
|
|
<bibRefCitation author="Fernandez M & Gasparini Z" box="[356,411,1178,1199]" pageId="8" pageNumber="820" pagination="79 - 84" refId="ref15330" refString="Fernandez M, Gasparini Z. 2008. Salt glands in the Jurassic metriorhynchid Geosaurus: implications for the evolution of osmoregulation in Mesozoic marine crocodyliforms. Naturaeissenschaften 95: 79 - 84." type="journal article" year="2008">2008</bibRefCitation>
|
|
;
|
|
<bibRefCitation author="Herrera Y & Fernandez MS & Gasparini Z" box="[424,646,1178,1200]" pageId="8" pageNumber="820" pagination="331 - 340" refId="ref15839" refString="Herrera Y, Fernandez MS, Gasparini Z. 2013. The snout of Cricosaurus araucanensis: a case study in novel anatomy of the nasal region of metriorhynchids. Lethaia 46: 331 - 340." type="journal article" year="2013">
|
|
Herrera
|
|
<emphasis box="[522,578,1178,1200]" italics="true" pageId="8" pageNumber="820">et al.</emphasis>
|
|
, 2013
|
|
</bibRefCitation>
|
|
;
|
|
<bibRefCitation author="Fernandez MS & Herrera Y" pageId="8" pageNumber="820" refId="ref15435" refString="Fernandez MS, Herrera Y. 2021. Active airflow of the paranasal sinuses in extinct crocodyliforms: evidence from a natural cast of the thalattosuchian Dakosaurus andiniensis. The Anatomical Record [early view]. doi: 10.1002 / ar. 24678." type="book" year="2021">Fernández & Herrera, 2021</bibRefCitation>
|
|
). Therefore, we concur with
|
|
<bibRefCitation author="Pierce SE & Williams M & Benton MJ" pageId="8" pageNumber="820" pagination="3225" refId="ref16732" refString="Pierce SE, Williams M, Benton MJ. 2017. Virtual reconstruction of the endocranial anatomy of the Early Jurassic marine crocodylomorph Pelagosaurus typus (Thalattosuchia). PeerJ 5: e 3225." type="journal article" year="2017">
|
|
Pierce
|
|
<emphasis box="[163,219,1240,1261]" italics="true" pageId="8" pageNumber="820">et al.</emphasis>
|
|
(2017)
|
|
</bibRefCitation>
|
|
and can infer that nasal salt glands were present in
|
|
<emphasis box="[297,532,1270,1292]" italics="true" pageId="8" pageNumber="820">
|
|
<taxonomicName baseAuthorityName="Witmer" baseAuthorityYear="1997" box="[297,457,1270,1291]" class="Reptilia" family="Metriorhynchidae" genus="Pelagosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="8" pageNumber="820" phylum="Chordata" rank="genus">Pelagosaurus</taxonomicName>
|
|
<typeStatus box="[467,532,1271,1292]" pageId="8" pageNumber="820">typus</typeStatus>
|
|
</emphasis>
|
|
from such olfactory region expansions, which may have occupied the lateral portion of these expansions while the dorsal portion (as with the teleosauroid expansions), was likely occupied by part of the nasal capsule. The size of the olfactory region expansions suggests the salt glands of
|
|
<taxonomicName baseAuthorityName="Witmer" baseAuthorityYear="1997" box="[329,484,1454,1475]" class="Reptilia" family="Metriorhynchidae" genus="Pelagosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="8" pageNumber="820" phylum="Chordata" rank="genus">
|
|
<emphasis box="[329,484,1454,1475]" italics="true" pageId="8" pageNumber="820">Pelagosaurus</emphasis>
|
|
</taxonomicName>
|
|
were smaller than those observed in metriorhynchid natural endocasts. The posterior end of the nasal cavity bears a shallow dorsolateral groove ventral to the prefrontal that separates the olfactory region expansions into dorsal and lateral portions (
|
|
<figureCitation box="[416,502,1608,1630]" captionStart="Figure 4" captionStartId="9.[145,225,1611,1633]" captionTargetBox="[305,1266,199,1570]" captionTargetId="figure-26@9.[303,1266,195,1571]" captionTargetPageId="9" captionText="Figure 4. Internal rostral anatomy of basal metriorhynchoids Pelagosaurus typus [NHMUK PV OR 32599 (A-E)] and Eoneustes gaudryi [NHMUK PV R 3263 (F-H)]. A-C, Pelagosaurus in dorsal (A), anterodorsal (B) and left lateral views (C) with the skull rendered transparent. D, E, transverse CT slices of nasal cavity olfactory region showing dorsal expansions where salt glands are inferred, the dorsolateral groove separating the inferred positions of salt glands and the nasal capsule (D) and the nasal ridge creating a V-shaped depression of the olfactory region ventral to the nasals (E). F-G, Eoneustes in dorsal (F) and left lateral views (G) with the skull rendered transparent. H, transverse CT slice of nasal cavity olfactory region showing the left salt duct, larger dorsolateral expansions where salt glands are inferred and ridge on the ventral surface of the nasals. Internal reconstruction colours: nasal cavity, yellow; nasopharyngeal ducts, pale yellow; antorbital fenestra, green; preobrital fenestra, light green. Scale bars equal 50 mm (A-C, F, G) and 10 mm (D, E, H)." figureDoi="http://doi.org/10.5281/zenodo.7695277" httpUri="https://zenodo.org/record/7695277/files/figure.png" pageId="8" pageNumber="820">Fig. 4D</figureCitation>
|
|
) similar to the shallow dorsal groove separating the glands and nasal capsule in metriorhynchid natural endocasts (
|
|
<bibRefCitation author="Herrera Y & Fernandez MS & Gasparini Z" pageId="8" pageNumber="820" pagination="331 - 340" refId="ref15839" refString="Herrera Y, Fernandez MS, Gasparini Z. 2013. The snout of Cricosaurus araucanensis: a case study in novel anatomy of the nasal region of metriorhynchids. Lethaia 46: 331 - 340." type="journal article" year="2013">
|
|
Herrera
|
|
<emphasis box="[714,772,1669,1691]" italics="true" pageId="8" pageNumber="820">et al.</emphasis>
|
|
, 2013
|
|
</bibRefCitation>
|
|
) and likely indicates the same morphology here, with salt glands occupying the lateral portions of the expansions and the nasal capsule occupying the dorsal portions. The antorbital cavity in
|
|
<taxonomicName baseAuthorityName="Witmer" baseAuthorityYear="1997" box="[545,696,1791,1812]" class="Reptilia" family="Metriorhynchidae" genus="Pelagosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="8" pageNumber="820" phylum="Chordata" rank="genus">
|
|
<emphasis box="[545,696,1791,1812]" italics="true" pageId="8" pageNumber="820">Pelagosaurus</emphasis>
|
|
</taxonomicName>
|
|
shares with
|
|
<taxonomicName box="[224,492,1822,1843]" genus="Plagiophthalmosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="820" phylum="Chordata" rank="genus">
|
|
<emphasis box="[224,492,1822,1843]" italics="true" pageId="8" pageNumber="820">Plagiophthalmosuchus</emphasis>
|
|
</taxonomicName>
|
|
and the teleosauroids a lateral opening through a reduced antorbital fenestra bordered by the lacrimal and maxilla. Dorsal to the antorbital fenestra, a novel antorbital duct is present that opens anteriorly into the nasal cavity through the maxilla anterior to the fenestra (
|
|
<figureCitation box="[1281,1367,258,280]" captionStart="Figure 4" captionStartId="9.[145,225,1611,1633]" captionTargetBox="[305,1266,199,1570]" captionTargetId="figure-26@9.[303,1266,195,1571]" captionTargetPageId="9" captionText="Figure 4. Internal rostral anatomy of basal metriorhynchoids Pelagosaurus typus [NHMUK PV OR 32599 (A-E)] and Eoneustes gaudryi [NHMUK PV R 3263 (F-H)]. A-C, Pelagosaurus in dorsal (A), anterodorsal (B) and left lateral views (C) with the skull rendered transparent. D, E, transverse CT slices of nasal cavity olfactory region showing dorsal expansions where salt glands are inferred, the dorsolateral groove separating the inferred positions of salt glands and the nasal capsule (D) and the nasal ridge creating a V-shaped depression of the olfactory region ventral to the nasals (E). F-G, Eoneustes in dorsal (F) and left lateral views (G) with the skull rendered transparent. H, transverse CT slice of nasal cavity olfactory region showing the left salt duct, larger dorsolateral expansions where salt glands are inferred and ridge on the ventral surface of the nasals. Internal reconstruction colours: nasal cavity, yellow; nasopharyngeal ducts, pale yellow; antorbital fenestra, green; preobrital fenestra, light green. Scale bars equal 50 mm (A-C, F, G) and 10 mm (D, E, H)." figureDoi="http://doi.org/10.5281/zenodo.7695277" httpUri="https://zenodo.org/record/7695277/files/figure.png" pageId="8" pageNumber="820">Fig. 4E</figureCitation>
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|
). It is currently unknown what system occupied this duct, but we do not regard this duct as either an extension of the antorbital cavity, as it does not communicate with the antorbital fenestra or as a rudimentary salt duct to facilitate drainage of the salt glands because no posterior opening into the olfactory region is observed.
|
|
</paragraph>
|
|
<paragraph blockId="8.[827,1444,197,1905]" pageId="8" pageNumber="820">
|
|
In
|
|
<taxonomicName box="[893,1140,473,494]" class="Reptilia" genus="Eoneustes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="8" pageNumber="820" phylum="Chordata" rank="species" species="gaudryi">
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<emphasis box="[893,1140,473,494]" italics="true" pageId="8" pageNumber="820">Eoneustes gaudryi</emphasis>
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</taxonomicName>
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|
(NHMUK PV R3263), the expansions of the olfactory region are further enlarged compared to
|
|
<taxonomicName baseAuthorityName="Witmer" baseAuthorityYear="1997" box="[1081,1233,534,555]" class="Reptilia" family="Metriorhynchidae" genus="Pelagosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="8" pageNumber="820" phylum="Chordata" rank="genus">
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|
<emphasis box="[1081,1233,534,555]" italics="true" pageId="8" pageNumber="820">Pelagosaurus</emphasis>
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|
</taxonomicName>
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|
, with two bulbous dorsolateral concave depressions on the prefrontals and lacrimals (
|
|
<figureCitation box="[1007,1074,595,617]" captionStart="Figure 4" captionStartId="9.[145,225,1611,1633]" captionTargetBox="[305,1266,199,1570]" captionTargetId="figure-26@9.[303,1266,195,1571]" captionTargetPageId="9" captionText="Figure 4. Internal rostral anatomy of basal metriorhynchoids Pelagosaurus typus [NHMUK PV OR 32599 (A-E)] and Eoneustes gaudryi [NHMUK PV R 3263 (F-H)]. A-C, Pelagosaurus in dorsal (A), anterodorsal (B) and left lateral views (C) with the skull rendered transparent. D, E, transverse CT slices of nasal cavity olfactory region showing dorsal expansions where salt glands are inferred, the dorsolateral groove separating the inferred positions of salt glands and the nasal capsule (D) and the nasal ridge creating a V-shaped depression of the olfactory region ventral to the nasals (E). F-G, Eoneustes in dorsal (F) and left lateral views (G) with the skull rendered transparent. H, transverse CT slice of nasal cavity olfactory region showing the left salt duct, larger dorsolateral expansions where salt glands are inferred and ridge on the ventral surface of the nasals. Internal reconstruction colours: nasal cavity, yellow; nasopharyngeal ducts, pale yellow; antorbital fenestra, green; preobrital fenestra, light green. Scale bars equal 50 mm (A-C, F, G) and 10 mm (D, E, H)." figureDoi="http://doi.org/10.5281/zenodo.7695277" httpUri="https://zenodo.org/record/7695277/files/figure.png" pageId="8" pageNumber="820">Fig. 4</figureCitation>
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F-H), corresponding to the salt gland location in metriorhynchid natural endocasts (
|
|
<bibRefCitation author="Fernandez M & Gasparini Z" box="[835,1190,657,679]" pageId="8" pageNumber="820" pagination="269 - 276" refId="ref15303" refString="Fernandez M, Gasparini Z. 2000. Salt glands in a Tithonian metriorhynchid crocodyliform and their physiological significance. Lethaia 33: 269 - 276." type="journal article" year="2000">Fernández & Gasparini, 2000</bibRefCitation>
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,
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<bibRefCitation author="Fernandez M & Gasparini Z" box="[1202,1258,657,678]" pageId="8" pageNumber="820" pagination="79 - 84" refId="ref15330" refString="Fernandez M, Gasparini Z. 2008. Salt glands in the Jurassic metriorhynchid Geosaurus: implications for the evolution of osmoregulation in Mesozoic marine crocodyliforms. Naturaeissenschaften 95: 79 - 84." type="journal article" year="2008">2008</bibRefCitation>
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;
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<bibRefCitation author="Herrera Y & Fernandez MS & Gasparini Z" pageId="8" pageNumber="820" pagination="331 - 340" refId="ref15839" refString="Herrera Y, Fernandez MS, Gasparini Z. 2013. The snout of Cricosaurus araucanensis: a case study in novel anatomy of the nasal region of metriorhynchids. Lethaia 46: 331 - 340." type="journal article" year="2013">
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Herrera
|
|
<emphasis box="[1377,1436,657,678]" italics="true" pageId="8" pageNumber="820">et al.</emphasis>
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|
, 2013
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|
</bibRefCitation>
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|
) and the prefrontal depressions reported by
|
|
<bibRefCitation author="Gandola R & Buffetaut E & Monaghan N & Dyke G" box="[827,1095,718,740]" pageId="8" pageNumber="820" pagination="1009 - 1010" refId="ref15557" refString="Gandola R, Buffetaut E, Monaghan N, Dyke G. 2006. Salt glands in the fossil crocodile Metriorhynchus. Journal of Vertebrate Paleontology 26: 1009 - 1010." type="journal article" year="2006">
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Gandola
|
|
<emphasis box="[942,1006,718,740]" italics="true" pageId="8" pageNumber="820">et al.</emphasis>
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|
(2006)
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|
</bibRefCitation>
|
|
. Thus, we infer that large, hypertrophied nasal salt glands comparable in size to those reported in metriorhynchids were present in
|
|
<taxonomicName authorityName="Young, Brusatte, Ruta & Andrade" authorityYear="2010" box="[857,972,811,832]" class="Reptilia" genus="Eoneustes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="8" pageNumber="820" phylum="Chordata" rank="genus">
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|
<emphasis box="[857,972,811,832]" italics="true" pageId="8" pageNumber="820">Eoneustes</emphasis>
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|
</taxonomicName>
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|
. As in
|
|
<taxonomicName baseAuthorityName="Witmer" baseAuthorityYear="1997" box="[1053,1207,810,831]" class="Reptilia" family="Metriorhynchidae" genus="Pelagosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="8" pageNumber="820" phylum="Chordata" rank="genus">
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|
<emphasis box="[1053,1207,810,831]" italics="true" pageId="8" pageNumber="820">Pelagosaurus</emphasis>
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|
</taxonomicName>
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|
, a deep ridge on the ventral surface of the nasals and the corresponding V-shaped nasal cavity depression medially separates the two dorsolateral expansions of the olfactory region. In
|
|
<taxonomicName authorityName="Young, Brusatte, Ruta & Andrade" authorityYear="2010" box="[858,971,933,954]" class="Reptilia" genus="Eoneustes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="8" pageNumber="820" phylum="Chordata" rank="genus">
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|
<emphasis box="[858,971,933,954]" italics="true" pageId="8" pageNumber="820">Eoneustes</emphasis>
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</taxonomicName>
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|
, there is a fenestra in the typical position for an antorbital fenestra. This region of
|
|
<taxonomicName authorityName="Young, Brusatte, Ruta & Andrade" authorityYear="2010" box="[1325,1443,964,985]" class="Reptilia" genus="Eoneustes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="8" pageNumber="820" phylum="Chordata" rank="genus">
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|
<emphasis box="[1325,1443,964,985]" italics="true" pageId="8" pageNumber="820">Eoneustes</emphasis>
|
|
</taxonomicName>
|
|
includes a long, obliquely oriented fossa on the external surface of the nasal, maxilla and lacrimal leading to an internal fenestra bordered by the nasal dorsally and lacrimal ventrally. Each fenestra is connected to the nasal cavity by short ducts running posteriorly from the fenestra to the dorsolateral expansions of the olfactory region, comparable to ducts preserved in metriorhynchid natural endocasts (
|
|
<bibRefCitation author="Fernandez MS & Herrera Y" pageId="8" pageNumber="820" pagination="702 - 714" refId="ref15396" refString="Fernandez MS, Herrera Y. 2009. Paranasal sinus system of Geosaurus araucanensis and the homology of the antorbital fenestra of metriorhynchids (Thalattosuchia: Crocodylomorpha). Journal of Vertebrate Paleontology 29: 702 - 714." type="journal article" year="2009">Fernández & Herrera, 2009</bibRefCitation>
|
|
). As was reported by
|
|
<bibRefCitation author="Cowgill T & Young MT & Schwab JA & Walsh S & Witmer LM & Herrera Y & Dollman KN & Choiniere JN & Brusatte SL" pageId="8" pageNumber="820" refId="ref15046" refString="Cowgill T, Young MT, Schwab JA, Walsh S, Witmer LM, Herrera Y, Dollman KN, Choiniere JN, Brusatte SL. 2021. Paranasal sinus system and upper respiratory tract evolution in Mesozoic pelagic crocodylomorphs. The Anatomical Record [early view] doi: 10.1002 / ar. 24727." type="book" year="2021">
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|
Cowgill
|
|
<emphasis box="[1383,1443,1239,1261]" italics="true" pageId="8" pageNumber="820">et al.</emphasis>
|
|
(2021)
|
|
</bibRefCitation>
|
|
, no antorbital cavity could be differentiated from the nasal cavity. However, given that antorbital cavities are present in both basal thalattosuchians and more derived metriorhynchids and the closest homologous location of the antorbital cavity is immediately posterior to the dorsal alveolar canal, ventral to the fenestra, we conclude that it was likely present in
|
|
<taxonomicName authorityName="Young, Brusatte, Ruta & Andrade" authorityYear="2010" box="[959,1077,1485,1506]" class="Reptilia" genus="Eoneustes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="8" pageNumber="820" phylum="Chordata" rank="genus">
|
|
<emphasis box="[959,1077,1485,1506]" italics="true" pageId="8" pageNumber="820">Eoneustes</emphasis>
|
|
</taxonomicName>
|
|
as well. The ducts connecting the fenestrae to the nasal cavity do not extend to the likely location of the antorbital cavity; thus, there is no evidence that the external fenestra and antorbital cavity are connected, and as such, we refer to it as a pre-orbital fenestra (
|
|
<bibRefCitation author="Fernandez MS & Herrera Y" box="[1103,1436,1638,1660]" pageId="8" pageNumber="820" pagination="702 - 714" refId="ref15396" refString="Fernandez MS, Herrera Y. 2009. Paranasal sinus system of Geosaurus araucanensis and the homology of the antorbital fenestra of metriorhynchids (Thalattosuchia: Crocodylomorpha). Journal of Vertebrate Paleontology 29: 702 - 714." type="journal article" year="2009">Fernández & Herrera, 2009</bibRefCitation>
|
|
;
|
|
<bibRefCitation author="Leardi JM & Pol D & Fernandez MS" box="[827,1032,1669,1691]" pageId="8" pageNumber="820" pagination="490 - 494" refId="ref16231" refString="Leardi JM, Pol D, Fernandez MS. 2012. The antorbital fenestra of Metriorhynchidae (Crocodyliformes, Thalattosuchia): testing its homology within a phylogenetic framework. Journal of Vertebrate Paleontology 32: 490 - 494." type="journal article" year="2012">
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|
Leardi
|
|
<emphasis box="[909,965,1669,1690]" italics="true" pageId="8" pageNumber="820">et al.</emphasis>
|
|
, 2012
|
|
</bibRefCitation>
|
|
).
|
|
</paragraph>
|
|
<paragraph blockId="8.[827,1444,197,1905]" lastBlockId="11.[145,762,197,1047]" lastPageId="11" lastPageNumber="823" pageId="8" pageNumber="820">
|
|
Among the metriorhynchids, the reconstructed nasal cavity morphology in
|
|
<taxonomicName baseAuthorityName="Fernandez & Herrera" baseAuthorityYear="2009" box="[1145,1443,1730,1752]" class="Reptilia" family="Metriorhynchidae" genus="Cricosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="8" pageNumber="820" phylum="Chordata" rank="species" species="araucanensis">
|
|
<emphasis box="[1145,1443,1730,1752]" italics="true" pageId="8" pageNumber="820">Cricosaurus araucanensis</emphasis>
|
|
</taxonomicName>
|
|
(MLP 72-IV-7-1) is concordant with the observed internal rostral morphology in natural endocasts from the same species (
|
|
<bibRefCitation author="Fernandez M & Gasparini Z" pageId="8" pageNumber="820" pagination="269 - 276" refId="ref15303" refString="Fernandez M, Gasparini Z. 2000. Salt glands in a Tithonian metriorhynchid crocodyliform and their physiological significance. Lethaia 33: 269 - 276." type="journal article" year="2000">Fernández & Gasparini, 2000</bibRefCitation>
|
|
,
|
|
<bibRefCitation author="Fernandez M & Gasparini Z" box="[896,951,1853,1874]" pageId="8" pageNumber="820" pagination="79 - 84" refId="ref15330" refString="Fernandez M, Gasparini Z. 2008. Salt glands in the Jurassic metriorhynchid Geosaurus: implications for the evolution of osmoregulation in Mesozoic marine crocodyliforms. Naturaeissenschaften 95: 79 - 84." type="journal article" year="2008">2008</bibRefCitation>
|
|
;
|
|
<bibRefCitation author="Herrera Y & Fernandez MS & Gasparini Z" box="[964,1190,1852,1874]" pageId="8" pageNumber="820" pagination="331 - 340" refId="ref15839" refString="Herrera Y, Fernandez MS, Gasparini Z. 2013. The snout of Cricosaurus araucanensis: a case study in novel anatomy of the nasal region of metriorhynchids. Lethaia 46: 331 - 340." type="journal article" year="2013">
|
|
Herrera
|
|
<emphasis box="[1064,1121,1852,1874]" italics="true" pageId="8" pageNumber="820">et al.</emphasis>
|
|
, 2013
|
|
</bibRefCitation>
|
|
). The olfactory region shows bulbous dorsolateral expansions expressed as broad depressions on the internal surfaces of the prefrontals with partial extension onto the lacrimals (
|
|
<figureCitation box="[172,338,258,280]" captionStart="Figure 5" captionStartId="10.[163,246,1678,1700]" captionTargetBox="[165,1446,305,1637]" captionTargetId="figure-80@10.[163,1446,303,1638]" captionTargetPageId="10" captionText="Figure 5. Internal rostral anatomy of metriorhynchids Cricosaurus araucanensis [MLP 72-IV-7-1 (A, B, G, H)], Thalattosuchus superciliosus [NHMUK PV R 11999 (C, D, I, J)] and TorƲoneustes coryphaeus [MJML K1863 (E, F, K, L)]. A, B, Cricosaurus araucanensis in dorsal (A) and left lateral view (B). C, D, Thalattosuchus in dorsal (C) and left lateral view (D). E, F, TorƲoneustes in dorsal (E) and left lateral view (F). G-L, transverse CT slices of nasal cavity olfactory region of Cricosaurus araucanensis (G, H), Thalattosuchus (I, J) and TorƲoneustes (K, L) showing the pre-orbital fenestra external opening (G, I, K) and dorsolateral expansions of olfactory region where salt glands are inferred (H, J, L). All skulls are rendered transparent. Internal reconstruction colours: nasal cavity, yellow; nasopharyngeal ducts, pale yellow; antorbital cavity/sinus, green; preobrital fenestra, light green. Scale bars equal 50 mm (A-F). and 10 mm (G-L)." figureDoi="http://doi.org/10.5281/zenodo.7695281" httpUri="https://zenodo.org/record/7695281/files/figure.png" pageId="10" pageNumber="822">Fig. 5A, B, G</figureCitation>
|
|
) and a shallow dorsolateral ridge separating the inferred location of salt glands from the nasal capsule (
|
|
<figureCitation box="[1060,1151,228,250]" captionStart="Figure 5" captionStartId="10.[163,246,1678,1700]" captionTargetBox="[165,1446,305,1637]" captionTargetId="figure-80@10.[163,1446,303,1638]" captionTargetPageId="10" captionText="Figure 5. Internal rostral anatomy of metriorhynchids Cricosaurus araucanensis [MLP 72-IV-7-1 (A, B, G, H)], Thalattosuchus superciliosus [NHMUK PV R 11999 (C, D, I, J)] and TorƲoneustes coryphaeus [MJML K1863 (E, F, K, L)]. A, B, Cricosaurus araucanensis in dorsal (A) and left lateral view (B). C, D, Thalattosuchus in dorsal (C) and left lateral view (D). E, F, TorƲoneustes in dorsal (E) and left lateral view (F). G-L, transverse CT slices of nasal cavity olfactory region of Cricosaurus araucanensis (G, H), Thalattosuchus (I, J) and TorƲoneustes (K, L) showing the pre-orbital fenestra external opening (G, I, K) and dorsolateral expansions of olfactory region where salt glands are inferred (H, J, L). All skulls are rendered transparent. Internal reconstruction colours: nasal cavity, yellow; nasopharyngeal ducts, pale yellow; antorbital cavity/sinus, green; preobrital fenestra, light green. Scale bars equal 50 mm (A-F). and 10 mm (G-L)." figureDoi="http://doi.org/10.5281/zenodo.7695281" httpUri="https://zenodo.org/record/7695281/files/figure.png" pageId="10" pageNumber="822">Fig. 5H</figureCitation>
|
|
). Identical nasal cavity morphologies are present in both
|
|
<taxonomicName family="Metriorhynchidae" genus="Thalattosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" lastPageId="11" lastPageNumber="823" pageId="10" pageNumber="822" phylum="Chordata" rank="species" species="superciliosus">
|
|
<emphasis italics="true" lastPageId="11" lastPageNumber="823" pageId="10" pageNumber="822">Thalattosuchus superciliosus</emphasis>
|
|
</taxonomicName>
|
|
(NHMUK PV R11999) (
|
|
<figureCitation box="[583,752,197,219]" captionStart="Figure 5" captionStartId="10.[163,246,1678,1700]" captionTargetBox="[165,1446,305,1637]" captionTargetId="figure-80@10.[163,1446,303,1638]" captionTargetPageId="10" captionText="Figure 5. Internal rostral anatomy of metriorhynchids Cricosaurus araucanensis [MLP 72-IV-7-1 (A, B, G, H)], Thalattosuchus superciliosus [NHMUK PV R 11999 (C, D, I, J)] and TorƲoneustes coryphaeus [MJML K1863 (E, F, K, L)]. A, B, Cricosaurus araucanensis in dorsal (A) and left lateral view (B). C, D, Thalattosuchus in dorsal (C) and left lateral view (D). E, F, TorƲoneustes in dorsal (E) and left lateral view (F). G-L, transverse CT slices of nasal cavity olfactory region of Cricosaurus araucanensis (G, H), Thalattosuchus (I, J) and TorƲoneustes (K, L) showing the pre-orbital fenestra external opening (G, I, K) and dorsolateral expansions of olfactory region where salt glands are inferred (H, J, L). All skulls are rendered transparent. Internal reconstruction colours: nasal cavity, yellow; nasopharyngeal ducts, pale yellow; antorbital cavity/sinus, green; preobrital fenestra, light green. Scale bars equal 50 mm (A-F). and 10 mm (G-L)." figureDoi="http://doi.org/10.5281/zenodo.7695281" httpUri="https://zenodo.org/record/7695281/files/figure.png" pageId="11" pageNumber="823">Fig. 5C, D, I, J</figureCitation>
|
|
) and
|
|
<taxonomicName authority="(MJML K 1863)" baseAuthorityName="MJML K" baseAuthorityYear="1863" box="[200,696,228,250]" class="Reptilia" family="Metriorhynchidae" genus="Torvoneustes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="11" pageNumber="823" phylum="Chordata" rank="species" species="coryphaeus">
|
|
<emphasis box="[200,501,228,249]" italics="true" pageId="11" pageNumber="823">TorƲoneustes coryphaeus</emphasis>
|
|
(MJML K1863)
|
|
</taxonomicName>
|
|
(
|
|
<figureCitation captionStart="Figure 5" captionStartId="10.[163,246,1678,1700]" captionTargetBox="[165,1446,305,1637]" captionTargetId="figure-80@10.[163,1446,303,1638]" captionTargetPageId="10" captionText="Figure 5. Internal rostral anatomy of metriorhynchids Cricosaurus araucanensis [MLP 72-IV-7-1 (A, B, G, H)], Thalattosuchus superciliosus [NHMUK PV R 11999 (C, D, I, J)] and TorƲoneustes coryphaeus [MJML K1863 (E, F, K, L)]. A, B, Cricosaurus araucanensis in dorsal (A) and left lateral view (B). C, D, Thalattosuchus in dorsal (C) and left lateral view (D). E, F, TorƲoneustes in dorsal (E) and left lateral view (F). G-L, transverse CT slices of nasal cavity olfactory region of Cricosaurus araucanensis (G, H), Thalattosuchus (I, J) and TorƲoneustes (K, L) showing the pre-orbital fenestra external opening (G, I, K) and dorsolateral expansions of olfactory region where salt glands are inferred (H, J, L). All skulls are rendered transparent. Internal reconstruction colours: nasal cavity, yellow; nasopharyngeal ducts, pale yellow; antorbital cavity/sinus, green; preobrital fenestra, light green. Scale bars equal 50 mm (A-F). and 10 mm (G-L)." figureDoi="http://doi.org/10.5281/zenodo.7695281" httpUri="https://zenodo.org/record/7695281/files/figure.png" pageId="11" pageNumber="823">Fig. 5E, F, K, L</figureCitation>
|
|
). However, in
|
|
<taxonomicName box="[433,698,258,280]" class="Reptilia" family="Metriorhynchidae" genus="Cricosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="11" pageNumber="823" phylum="Chordata" rank="species" species="schroederi">
|
|
<emphasis box="[433,698,258,280]" italics="true" pageId="11" pageNumber="823">Cricosaurus schroederi</emphasis>
|
|
</taxonomicName>
|
|
(MM unnumbered) no bulbous expansions of the olfactory region are present. The olfactory region itself is dorsoventrally narrow and the nasopharyngeal ducts and antorbital cavities located ventral to this region are larger compared to the other metriorhynchids (
|
|
<figureCitation box="[683,747,412,434]" captionStart="Figure 6" captionStartId="11.[146,226,1787,1809]" captionTargetBox="[150,1426,1096,1746]" captionTargetId="figure-454@11.[145,1429,1090,1747]" captionTargetPageId="11" captionText="Figure 6. Internal rostral anatomy of Cricosaurus schroederi (MM unnumbered). A, B, dorsal (A) and left lateral views (B) with the skull rendered transparent. C, D, transverse CT slices of the nasal cavity olfactory region showing salt ducts (C) and small lateral olfactory region expansions where salt glands are inferred (D). Internal reconstruction colours: nasal cavity, yellow; nasopharyngeal ducts, pale yellow; antorbital cavity/sinus, green; preobrital fenestra, light green. Scale bars equal 50 mm (A, B) and 10 mm (C, D)." figureDoi="http://doi.org/10.5281/zenodo.7695283" httpUri="https://zenodo.org/record/7695283/files/figure.png" pageId="11" pageNumber="823">Fig. 6</figureCitation>
|
|
). The lack of any dorsoventral compression in the rest of the skull demonstrates this was not a deformation artefact. However, two small lateral expansions of the olfactory region are present medial to the prefrontals in
|
|
<taxonomicName box="[176,444,565,586]" class="Reptilia" family="Metriorhynchidae" genus="Cricosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="11" pageNumber="823" phylum="Chordata" rank="species" species="schroederi">
|
|
<emphasis box="[176,444,565,586]" italics="true" pageId="11" pageNumber="823">Cricosaurus schroederi</emphasis>
|
|
</taxonomicName>
|
|
and correspond to the salt gland locations in natural casts.
|
|
</paragraph>
|
|
</subSubSection>
|
|
<caption ID-DOI="http://doi.org/10.5281/zenodo.7695277" ID-Zenodo-Dep="7695277" httpUri="https://zenodo.org/record/7695277/files/figure.png" pageId="9" pageNumber="821" startId="9.[145,225,1611,1633]" targetBox="[305,1266,199,1570]" targetPageId="9">
|
|
<paragraph blockId="9.[145,1425,1611,1868]" pageId="9" pageNumber="821">
|
|
<emphasis bold="true" box="[145,252,1611,1633]" pageId="9" pageNumber="821">Figure 4.</emphasis>
|
|
Internal rostral anatomy of basal metriorhynchoids
|
|
<emphasis box="[832,1040,1611,1633]" italics="true" pageId="9" pageNumber="821">
|
|
<taxonomicName baseAuthorityName="Witmer" baseAuthorityYear="1997" box="[832,975,1611,1633]" class="Reptilia" family="Metriorhynchidae" genus="Pelagosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="9" pageNumber="821" phylum="Chordata" rank="genus">Pelagosaurus</taxonomicName>
|
|
typus
|
|
</emphasis>
|
|
[NHMUK PV OR 32599 (A-E)] and
|
|
<taxonomicName box="[145,337,1641,1663]" class="Reptilia" genus="Eoneustes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="9" pageNumber="821" phylum="Chordata" rank="species" species="gaudryi">
|
|
<emphasis box="[145,337,1641,1663]" italics="true" pageId="9" pageNumber="821">Eoneustes gaudryi</emphasis>
|
|
</taxonomicName>
|
|
[NHMUK PV R 3263 (F-H)]. A-C,
|
|
<taxonomicName baseAuthorityName="Witmer" baseAuthorityYear="1997" box="[698,837,1641,1663]" class="Reptilia" family="Metriorhynchidae" genus="Pelagosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="9" pageNumber="821" phylum="Chordata" rank="genus">
|
|
<emphasis box="[698,837,1641,1663]" italics="true" pageId="9" pageNumber="821">Pelagosaurus</emphasis>
|
|
</taxonomicName>
|
|
in dorsal (A), anterodorsal (B) and left lateral views (C) with the skull rendered transparent. D, E, transverse CT slices of nasal cavity olfactory region showing dorsal expansions where salt glands are inferred, the dorsolateral groove separating the inferred positions of salt glands and the nasal capsule (D) and the nasal ridge creating a V-shaped depression of the olfactory region ventral to the nasals (E). F-G,
|
|
<taxonomicName authorityName="Young, Brusatte, Ruta & Andrade" authorityYear="2010" box="[1292,1396,1729,1750]" class="Reptilia" genus="Eoneustes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="9" pageNumber="821" phylum="Chordata" rank="genus">
|
|
<emphasis box="[1292,1396,1729,1750]" italics="true" pageId="9" pageNumber="821">Eoneustes</emphasis>
|
|
</taxonomicName>
|
|
in dorsal (F) and left lateral views (G) with the skull rendered transparent. H, transverse CT slice of nasal cavity olfactory region showing the left salt duct, larger dorsolateral expansions where salt glands are inferred and ridge on the ventral surface of the nasals. Internal reconstruction colours: nasal cavity, yellow; nasopharyngeal ducts, pale yellow; antorbital fenestra, green; preobrital fenestra, light green. Scale bars equal 50 mm (A-C, F, G) and 10 mm (D, E, H).
|
|
</paragraph>
|
|
</caption>
|
|
<caption ID-DOI="http://doi.org/10.5281/zenodo.7695281" ID-Zenodo-Dep="7695281" httpUri="https://zenodo.org/record/7695281/files/figure.png" pageId="10" pageNumber="822" startId="10.[163,246,1678,1700]" targetBox="[165,1446,305,1637]" targetPageId="10">
|
|
<paragraph blockId="10.[163,1443,1678,1906]" pageId="10" pageNumber="822">
|
|
<emphasis bold="true" box="[163,276,1678,1700]" pageId="10" pageNumber="822">Figure 5.</emphasis>
|
|
Internal rostral anatomy of metriorhynchids
|
|
<taxonomicName baseAuthorityName="Fernandez & Herrera" baseAuthorityYear="2009" box="[809,1101,1679,1701]" class="Reptilia" family="Metriorhynchidae" genus="Cricosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="10" pageNumber="822" phylum="Chordata" rank="species" species="araucanensis">
|
|
<emphasis box="[809,1101,1679,1701]" italics="true" pageId="10" pageNumber="822">Cricosaurus araucanensis</emphasis>
|
|
</taxonomicName>
|
|
[MLP 72-IV-7-1 (A, B, G, H)],
|
|
<taxonomicName box="[163,465,1708,1730]" family="Metriorhynchidae" genus="Thalattosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="822" phylum="Chordata" rank="species" species="superciliosus">
|
|
<emphasis box="[163,465,1708,1730]" italics="true" pageId="10" pageNumber="822">Thalattosuchus superciliosus</emphasis>
|
|
</taxonomicName>
|
|
[NHMUK PV R 11999 (C, D, I, J)] and
|
|
<taxonomicName baseAuthorityName="MJML K" baseAuthorityYear="1863" box="[871,1128,1708,1730]" class="Reptilia" family="Metriorhynchidae" genus="Torvoneustes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="10" pageNumber="822" phylum="Chordata" rank="species" species="coryphaeus">
|
|
<emphasis box="[871,1128,1708,1730]" italics="true" pageId="10" pageNumber="822">TorƲoneustes coryphaeus</emphasis>
|
|
</taxonomicName>
|
|
[MJML K1863 (E, F, K, L)]. A, B,
|
|
<taxonomicName baseAuthorityName="Fernandez & Herrera" baseAuthorityYear="2009" box="[190,462,1738,1759]" class="Reptilia" family="Metriorhynchidae" genus="Cricosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="10" pageNumber="822" phylum="Chordata" rank="species" species="araucanensis">
|
|
<emphasis box="[190,462,1738,1759]" italics="true" pageId="10" pageNumber="822">Cricosaurus araucanensis</emphasis>
|
|
</taxonomicName>
|
|
in dorsal (A) and left lateral view (B). C, D,
|
|
<taxonomicName box="[926,1087,1737,1758]" family="Metriorhynchidae" genus="Thalattosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="822" phylum="Chordata" rank="genus">
|
|
<emphasis box="[926,1087,1737,1758]" italics="true" pageId="10" pageNumber="822">Thalattosuchus</emphasis>
|
|
</taxonomicName>
|
|
in dorsal (C) and left lateral view (D). E, F,
|
|
<taxonomicName baseAuthorityName="MJML K" baseAuthorityYear="1863" box="[263,400,1767,1789]" class="Reptilia" family="Metriorhynchidae" genus="Torvoneustes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="10" pageNumber="822" phylum="Chordata" rank="genus">
|
|
<emphasis box="[263,400,1767,1789]" italics="true" pageId="10" pageNumber="822">TorƲoneustes</emphasis>
|
|
</taxonomicName>
|
|
in dorsal (E) and left lateral view (F). G-L, transverse CT slices of nasal cavity olfactory region of
|
|
<taxonomicName baseAuthorityName="Fernandez & Herrera" baseAuthorityYear="2009" box="[163,437,1796,1818]" class="Reptilia" family="Metriorhynchidae" genus="Cricosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="10" pageNumber="822" phylum="Chordata" rank="species" species="araucanensis">
|
|
<emphasis box="[163,437,1796,1818]" italics="true" pageId="10" pageNumber="822">Cricosaurus araucanensis</emphasis>
|
|
</taxonomicName>
|
|
(G, H),
|
|
<taxonomicName box="[521,684,1796,1818]" family="Metriorhynchidae" genus="Thalattosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="822" phylum="Chordata" rank="genus">
|
|
<emphasis box="[521,684,1796,1818]" italics="true" pageId="10" pageNumber="822">Thalattosuchus</emphasis>
|
|
</taxonomicName>
|
|
(I, J) and
|
|
<taxonomicName baseAuthorityName="MJML K" baseAuthorityYear="1863" box="[794,930,1796,1818]" class="Reptilia" family="Metriorhynchidae" genus="Torvoneustes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="10" pageNumber="822" phylum="Chordata" rank="genus">
|
|
<emphasis box="[794,930,1796,1818]" italics="true" pageId="10" pageNumber="822">TorƲoneustes</emphasis>
|
|
</taxonomicName>
|
|
(K, L) showing the pre-orbital fenestra external opening (G, I, K) and dorsolateral expansions of olfactory region where salt glands are inferred (H, J, L). All skulls are rendered transparent. Internal reconstruction colours: nasal cavity, yellow; nasopharyngeal ducts, pale yellow; antorbital cavity/sinus, green; preobrital fenestra, light green. Scale bars equal 50 mm (A-F). and 10 mm (G-L).
|
|
</paragraph>
|
|
</caption>
|
|
<subSubSection pageId="11" pageNumber="823" type="description">
|
|
<paragraph blockId="11.[145,762,197,1047]" lastBlockId="11.[809,1426,197,556]" pageId="11" pageNumber="823">
|
|
Unlike
|
|
<taxonomicName baseAuthorityName="Witmer" baseAuthorityYear="1997" box="[264,428,626,647]" class="Reptilia" family="Metriorhynchidae" genus="Pelagosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="11" pageNumber="823" phylum="Chordata" rank="genus">
|
|
<emphasis box="[264,428,626,647]" italics="true" pageId="11" pageNumber="823">Pelagosaurus</emphasis>
|
|
</taxonomicName>
|
|
and
|
|
<taxonomicName authorityName="Young, Brusatte, Ruta & Andrade" authorityYear="2010" box="[496,618,627,648]" class="Reptilia" genus="Eoneustes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="11" pageNumber="823" phylum="Chordata" rank="genus">
|
|
<emphasis box="[496,618,627,648]" italics="true" pageId="11" pageNumber="823">Eoneustes</emphasis>
|
|
</taxonomicName>
|
|
, no medial V-shaped depression of the nasal cavity occurs in the metriorhynchids, and the olfactory region expansions are instead separated by a smaller ridge on the ventral surface of the nasals creating a shallower depression. The depth and breadth of this depression varies among the sample; in
|
|
<taxonomicName baseAuthorityName="Fernandez & Herrera" baseAuthorityYear="2009" class="Reptilia" family="Metriorhynchidae" genus="Cricosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="11" pageNumber="823" phylum="Chordata" rank="species" species="araucanensis">
|
|
<emphasis italics="true" pageId="11" pageNumber="823">Cricosaurus araucanensis</emphasis>
|
|
</taxonomicName>
|
|
and
|
|
<taxonomicName box="[373,562,841,862]" family="Metriorhynchidae" genus="Thalattosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="11" pageNumber="823" phylum="Chordata" rank="genus">
|
|
<emphasis box="[373,562,841,862]" italics="true" pageId="11" pageNumber="823">Thalattosuchus</emphasis>
|
|
</taxonomicName>
|
|
this depression extends across the medial third of the dorsal surface of the olfactory region, but in
|
|
<taxonomicName box="[494,761,902,924]" class="Reptilia" family="Metriorhynchidae" genus="Cricosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="11" pageNumber="823" phylum="Chordata" rank="species" species="schroederi">
|
|
<emphasis box="[494,761,902,924]" italics="true" pageId="11" pageNumber="823">Cricosaurus schroederi</emphasis>
|
|
</taxonomicName>
|
|
and
|
|
<taxonomicName baseAuthorityName="MJML K" baseAuthorityYear="1863" box="[196,344,933,954]" class="Reptilia" family="Metriorhynchidae" genus="Torvoneustes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="11" pageNumber="823" phylum="Chordata" rank="genus">
|
|
<emphasis box="[196,344,933,954]" italics="true" pageId="11" pageNumber="823">TorƲoneustes</emphasis>
|
|
</taxonomicName>
|
|
it is limited to a small medial ridge marking the dorsal surface of the cartilaginous nasal septum. The metriorhynchids all have a fenestra in the classic antorbital fenestra position of archosaurs sharing a near identical morphology with each other and
|
|
<taxonomicName authorityName="Young, Brusatte, Ruta & Andrade" authorityYear="2010" box="[935,1053,228,249]" class="Reptilia" genus="Eoneustes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="11" pageNumber="823" phylum="Chordata" rank="genus">
|
|
<emphasis box="[935,1053,228,249]" italics="true" pageId="11" pageNumber="823">Eoneustes</emphasis>
|
|
</taxonomicName>
|
|
, including an oblique external antorbital fossa leading to the internal fenestra. The inferred antorbital cavities of all the metriorhynchids are internalized and do not connect to the fenestrae; at their closest point, the anterior of the antorbital cavity is ventromedial to the fenestra. In
|
|
<taxonomicName class="Reptilia" family="Metriorhynchidae" genus="Cricosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="11" pageNumber="823" phylum="Chordata" rank="species" species="schroederi">
|
|
<emphasis italics="true" pageId="11" pageNumber="823">Cricosaurus schroederi</emphasis>
|
|
</taxonomicName>
|
|
, short ducts located dorsal to the antorbital cavity connect the fenestra to the lateral expansions of the olfactory region (
|
|
<figureCitation box="[1045,1125,473,495]" captionStart="Figure 6" captionStartId="11.[146,226,1787,1809]" captionTargetBox="[150,1426,1096,1746]" captionTargetId="figure-454@11.[145,1429,1090,1747]" captionTargetPageId="11" captionText="Figure 6. Internal rostral anatomy of Cricosaurus schroederi (MM unnumbered). A, B, dorsal (A) and left lateral views (B) with the skull rendered transparent. C, D, transverse CT slices of the nasal cavity olfactory region showing salt ducts (C) and small lateral olfactory region expansions where salt glands are inferred (D). Internal reconstruction colours: nasal cavity, yellow; nasopharyngeal ducts, pale yellow; antorbital cavity/sinus, green; preobrital fenestra, light green. Scale bars equal 50 mm (A, B) and 10 mm (C, D)." figureDoi="http://doi.org/10.5281/zenodo.7695283" httpUri="https://zenodo.org/record/7695283/files/figure.png" pageId="11" pageNumber="823">Fig. 6C</figureCitation>
|
|
). Thus, as with
|
|
<taxonomicName authorityName="Young, Brusatte, Ruta & Andrade" authorityYear="2010" box="[1306,1419,474,495]" class="Reptilia" genus="Eoneustes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="11" pageNumber="823" phylum="Chordata" rank="genus">
|
|
<emphasis box="[1306,1419,474,495]" italics="true" pageId="11" pageNumber="823">Eoneustes</emphasis>
|
|
</taxonomicName>
|
|
, we consider the fenestra in these metriorhynchids to be a pre-orbital fenestra.
|
|
</paragraph>
|
|
</subSubSection>
|
|
</treatment>
|
|
</document> |