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<document ID-DOI="10.1093/zoolinnean/zlaa061" ID-ISSN="0024-4082" ID-Zenodo-Dep="5300243" approvalRequired="62" approvalRequired_for_taxonomicNames="11" approvalRequired_for_textStreams="45" approvalRequired_for_treatments="6" checkinTime="1630095814227" checkinUser="felipe" docAuthor="Norman, David B" docDate="2021" docId="B66BDD2A082BFF9EE36774C5FEB6E16B" docLanguage="en" docName="zlaa061.pdf" docOrigin="Zoological Journal of the Linnean Society 191 (1)" docSource="https://academic.oup.com/zoolinnean/article/191/1/1/5893854" docStyle="DocumentStyle:36B3BD6A90C22AB4F7F465C853188CC8.5:ZoolJLinnSoc.2017-.journal_article" docStyleId="36B3BD6A90C22AB4F7F465C853188CC8" docStyleName="ZoolJLinnSoc.2017-.journal_article" docStyleVersion="5" docTitle="Scelidosaurus" docType="treatment" docVersion="3" lastPageNumber="9" masterDocId="4A52A552082FFF96E03F7400FFA5E61A" masterDocTitle="Scelidosaurus harrisonii (Dinosauria: Ornithischia) from the Early Jurassic of Dorset, England: biology and phylogenetic relationships" masterLastPageNumber="86" masterPageNumber="1" pageNumber="5" updateTime="1631151263965" updateUser="ExternalLinkService">
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<mods:title>Scelidosaurus harrisonii (Dinosauria: Ornithischia) from the Early Jurassic of Dorset, England: biology and phylogenetic relationships</mods:title>
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<mods:namePart>Norman, David B</mods:namePart>
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<mods:title>Zoological Journal of the Linnean Society</mods:title>
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<mods:date>2021</mods:date>
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<mods:number>2021-01-01</mods:number>
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<mods:number>191</mods:number>
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<subSubSection pageId="4" pageNumber="5" type="nomenclature">
<paragraph blockId="4.[856,1377,197,221]" box="[856,1377,197,221]" pageId="4" pageNumber="5">
<heading box="[856,1377,197,221]" centered="true" fontSize="9" level="2" pageId="4" pageNumber="5" reason="2">
<emphasis box="[856,1106,197,220]" italics="true" pageId="4" pageNumber="5">
<taxonomicName baseAuthorityName="NHMUK R" baseAuthorityYear="1111" box="[856,1099,197,220]" class="Reptilia" family="Scelidosauridae" genus="Scelidosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">SCELIDOSAURUS</taxonomicName>
:
</emphasis>
CRANIAL BIOLOGY
</heading>
</paragraph>
<paragraph blockId="4.[996,1237,242,267]" box="[996,1237,242,267]" pageId="4" pageNumber="5">EXTERNAL FEATURES</paragraph>
</subSubSection>
<subSubSection lastPageId="8" lastPageNumber="9" pageId="4" pageNumber="5" type="description">
<paragraph blockId="4.[809,1425,283,795]" pageId="4" pageNumber="5">
The surfaces of the skull and mandible (
<figureCitation box="[1301,1369,283,305]" captionStart="Figure 4" captionStartId="5.[163,243,1729,1751]" captionTargetBox="[174,1434,211,1677]" captionTargetId="figure-23@5.[163,1443,195,1689]" captionTargetPageId="5" captionText="Figure 4. Scelidosaurus harrisonii Owen, 1861. Cranial anatomy reconstructed in: A, lateral; B, dorsal views (modified from Norman, 2020a: figs 8, 9)." figureDoi="http://doi.org/10.5281/zenodo.5300253" httpUri="https://zenodo.org/record/5300253/files/figure.png" pageId="4" pageNumber="5">Fig. 4</figureCitation>
) are coated by widespread exostoses and a small number of discrete osteoderms. This suggests that the scelidosaur skull was encased by an array of keratinous epidermal scutes or plates.
</paragraph>
<paragraph blockId="4.[809,1425,283,795]" pageId="4" pageNumber="5">
For comparison, an osteological preparation of the skull of a similarly sized subadult green turtle [
<taxonomicName authority="(Linnaeus, 1758)" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[817,1195,497,519]" class="Reptilia" family="Cheloniidae" genus="Chelonia" kingdom="Animalia" order="Testudines" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="mydas">
<emphasis box="[817,999,497,518]" italics="true" pageId="4" pageNumber="5">Chelonia mydas</emphasis>
(
<bibRefCitation author="Linnaeus C" box="[1014,1187,497,519]" pageId="4" pageNumber="5" refId="ref61651" refString="Linnaeus C. 1758. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus differentiis, synonymis, locis, 10 th edn. Stockholm: L. Salvius." type="book" year="1758">Linnaeus, 1758</bibRefCitation>
)
</taxonomicName>
Norman, personal collection
<figureCitation box="[942,1054,527,550]" captionStart="Figure 5" captionStartId="6.[145,225,1070,1092]" captionTargetBox="[153,1419,205,1025]" captionTargetId="figure-382@6.[145,1428,195,1030]" captionTargetPageId="6" captionText="Figure 5. Chelonia mydas (Linnaeus, 1758) the Green Turtle. External skull anatomy. A, lateral and B, dorsal views, illustrating the superficial cranial osteology. C, lateral and D, dorsal views showing the distribution of keratinous scutes. Norman, pers.colln (ex-University of Cambridge Zoology Department teachingcollection).Scale bar indicated.Abbreviations: au, auditory recess; au.sc, auditory recess scute covering; tom, the chelonian tomium (= rhamphotheca of Scelidosaurus)." figureDoi="http://doi.org/10.5281/zenodo.5300255" httpUri="https://zenodo.org/record/5300255/files/figure.png" pageId="4" pageNumber="5">Fig. 5A, B</figureCitation>
] a taxon belonging to the clade Testudinata with putative (sister-taxon) archosaur affinities (
<bibRefCitation author="Crawford NG &amp; Faircloth BC &amp; McCormack JE &amp; Brumfield RT &amp; Winker K &amp; Glenn TC" box="[933,1184,589,611]" pageId="4" pageNumber="5" pagination="783 - 786" refId="ref59251" refString="Crawford NG, Faircloth BC, McCormack JE, Brumfield RT, Winker K, Glenn TC. 2012. More than 1000 ultraconserved elements provide evidence that turtles are the sister group of archosaurs. Biology Letters 8: 783 - 786." type="journal article" year="2012">
Crawford
<emphasis box="[1053,1112,589,611]" italics="true" pageId="4" pageNumber="5">et al.</emphasis>
, 2012
</bibRefCitation>
) reveals similarly heavily textured skull roofing bones that, in life, are known to anchor a tessellate covering of keratinous scutes (
<figureCitation box="[904,1028,681,703]" captionStart="Figure 5" captionStartId="6.[145,225,1070,1092]" captionTargetBox="[153,1419,205,1025]" captionTargetId="figure-382@6.[145,1428,195,1030]" captionTargetPageId="6" captionText="Figure 5. Chelonia mydas (Linnaeus, 1758) the Green Turtle. External skull anatomy. A, lateral and B, dorsal views, illustrating the superficial cranial osteology. C, lateral and D, dorsal views showing the distribution of keratinous scutes. Norman, pers.colln (ex-University of Cambridge Zoology Department teachingcollection).Scale bar indicated.Abbreviations: au, auditory recess; au.sc, auditory recess scute covering; tom, the chelonian tomium (= rhamphotheca of Scelidosaurus)." figureDoi="http://doi.org/10.5281/zenodo.5300255" httpUri="https://zenodo.org/record/5300255/files/figure.png" pageId="4" pageNumber="5">Fig. 5C, D</figureCitation>
). Unfortunately, this particular specimen does not show the pattern of smooth, shallow grooves that mark the edges of some of the principal cranial scutes (Penkalski, pers. comm.
<date box="[1248,1388,773,795]" pageId="4" pageNumber="5" value="2020-03">March 2020</date>
).
</paragraph>
<paragraph blockId="4.[809,1085,851,875]" box="[809,1085,851,875]" pageId="4" pageNumber="5">
<emphasis box="[809,1085,851,875]" italics="true" pageId="4" pageNumber="5">The premaxillary beak</emphasis>
</paragraph>
<paragraph blockId="4.[809,1425,892,1834]" pageId="4" pageNumber="5">
In
<taxonomicName baseAuthorityName="NHMUK R" baseAuthorityYear="1111" box="[842,1004,892,913]" class="Reptilia" family="Scelidosauridae" genus="Scelidosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis box="[842,1004,892,913]" italics="true" pageId="4" pageNumber="5">Scelidosaurus</emphasis>
</taxonomicName>
a rhamphotheca was, without much doubt, an externally smooth casque-like structure referred to as a tomium in
<taxonomicName box="[1111,1213,953,974]" class="Reptilia" family="Cheloniidae" genus="Chelonia" kingdom="Animalia" order="Testudines" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis box="[1111,1213,953,974]" italics="true" pageId="4" pageNumber="5">Chelonia</emphasis>
</taxonomicName>
(
<figureCitation box="[1227,1306,953,975]" captionStart="Figure 5" captionStartId="6.[145,225,1070,1092]" captionTargetBox="[153,1419,205,1025]" captionTargetId="figure-382@6.[145,1428,195,1030]" captionTargetPageId="6" captionText="Figure 5. Chelonia mydas (Linnaeus, 1758) the Green Turtle. External skull anatomy. A, lateral and B, dorsal views, illustrating the superficial cranial osteology. C, lateral and D, dorsal views showing the distribution of keratinous scutes. Norman, pers.colln (ex-University of Cambridge Zoology Department teachingcollection).Scale bar indicated.Abbreviations: au, auditory recess; au.sc, auditory recess scute covering; tom, the chelonian tomium (= rhamphotheca of Scelidosaurus)." figureDoi="http://doi.org/10.5281/zenodo.5300255" httpUri="https://zenodo.org/record/5300255/files/figure.png" pageId="4" pageNumber="5">Fig. 5C</figureCitation>
, tom) that enveloped the premaxillae (
<figureCitation box="[1137,1203,984,1006]" captionStart="Figure 6" captionStartId="7.[163,241,1637,1659]" captionTargetBox="[175,1431,208,1582]" captionTargetId="figure-23@7.[163,1443,195,1597]" captionTargetPageId="7" captionText="Figure 6. Scelidosaurus. An interpretation of the cranial keratinous scute pattern on the skull and mandible in (A) lateral and (B) dorsal views. Abbreviations: asc, angular scute; fsc, frontal scute; dsc, dentary scute or sheath; hsc, occipital horn scute; itsc, infratemporal scutes; jsc, jugal arch scutes; lsc, lacrimal scute; msc, maxillary scute (there may have been several); nmsc, nasal median scute; nsc, nasal scutes restored as overlapping plates; pdsc, predentary scute (rhamphotheca); posc, postorbital crater-like scute; qjsc, quadratojugalquadrate scute; rsc, rhamphothecal scute of the premaxilla; sosc, supraorbital (brow-ridge) scute (there may have been several); stsc, supratemporal scutes; tym, tympanic membrane;?, area on the surface of the lacrimal lamina that is devoid of exostoses and may mark the location of a diverticulum of a cranial sinus." figureDoi="http://doi.org/10.5281/zenodo.5300257" httpUri="https://zenodo.org/record/5300257/files/figure.png" pageId="4" pageNumber="5">Fig. 6</figureCitation>
, rsc). The external surface of the premaxilla (not including the narial fossa) is slightly rugose and pitted with small foramina that in combination would have supported, anchored and indicate the presence of a vascular supply that provided nutrients for the growth of an overlying rhamphotheca; closely comparable osteological features are seen underlying the tomium of the turtle. The scelidosaur rhamphotheca formed a short cuttingedge along the edentulous margin of the premaxilla (
<figureCitation box="[817,885,1291,1313]" captionStart="Figure 4" captionStartId="5.[163,243,1729,1751]" captionTargetBox="[174,1434,211,1677]" captionTargetId="figure-23@5.[163,1443,195,1689]" captionTargetPageId="5" captionText="Figure 4. Scelidosaurus harrisonii Owen, 1861. Cranial anatomy reconstructed in: A, lateral; B, dorsal views (modified from Norman, 2020a: figs 8, 9)." figureDoi="http://doi.org/10.5281/zenodo.5300253" httpUri="https://zenodo.org/record/5300253/files/figure.png" pageId="4" pageNumber="5">Figs 4</figureCitation>
,
<figureCitation box="[897,911,1290,1312]" captionStart="Figure 6" captionStartId="7.[163,241,1637,1659]" captionTargetBox="[175,1431,208,1582]" captionTargetId="figure-23@7.[163,1443,195,1597]" captionTargetPageId="7" captionText="Figure 6. Scelidosaurus. An interpretation of the cranial keratinous scute pattern on the skull and mandible in (A) lateral and (B) dorsal views. Abbreviations: asc, angular scute; fsc, frontal scute; dsc, dentary scute or sheath; hsc, occipital horn scute; itsc, infratemporal scutes; jsc, jugal arch scutes; lsc, lacrimal scute; msc, maxillary scute (there may have been several); nmsc, nasal median scute; nsc, nasal scutes restored as overlapping plates; pdsc, predentary scute (rhamphotheca); posc, postorbital crater-like scute; qjsc, quadratojugalquadrate scute; rsc, rhamphothecal scute of the premaxilla; sosc, supraorbital (brow-ridge) scute (there may have been several); stsc, supratemporal scutes; tym, tympanic membrane;?, area on the surface of the lacrimal lamina that is devoid of exostoses and may mark the location of a diverticulum of a cranial sinus." figureDoi="http://doi.org/10.5281/zenodo.5300257" httpUri="https://zenodo.org/record/5300257/files/figure.png" pageId="4" pageNumber="5">6</figureCitation>
). The slightly rugose and vascularized lateral wall to the dentulous portion of the premaxilla is likely to have been similarly encased by a posterior extension of the rhamphotheca, in conformity with the structure seen in
<emphasis box="[898,1008,1413,1434]" italics="true" pageId="4" pageNumber="5">
<taxonomicName box="[898,1004,1413,1434]" class="Reptilia" family="Cheloniidae" genus="Chelonia" kingdom="Animalia" order="Testudines" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">Chelonia</taxonomicName>
.
</emphasis>
It is probable that the premaxillary dentition was ensheathed (and supported) by this portion of the keratinous beak; the crowns of the teeth are envisioned projecting from behind the rhamphothecal parapet (
<figureCitation box="[1091,1171,1536,1558]" captionStart="Figure 6" captionStartId="7.[163,241,1637,1659]" captionTargetBox="[175,1431,208,1582]" captionTargetId="figure-23@7.[163,1443,195,1597]" captionTargetPageId="7" captionText="Figure 6. Scelidosaurus. An interpretation of the cranial keratinous scute pattern on the skull and mandible in (A) lateral and (B) dorsal views. Abbreviations: asc, angular scute; fsc, frontal scute; dsc, dentary scute or sheath; hsc, occipital horn scute; itsc, infratemporal scutes; jsc, jugal arch scutes; lsc, lacrimal scute; msc, maxillary scute (there may have been several); nmsc, nasal median scute; nsc, nasal scutes restored as overlapping plates; pdsc, predentary scute (rhamphotheca); posc, postorbital crater-like scute; qjsc, quadratojugalquadrate scute; rsc, rhamphothecal scute of the premaxilla; sosc, supraorbital (brow-ridge) scute (there may have been several); stsc, supratemporal scutes; tym, tympanic membrane;?, area on the surface of the lacrimal lamina that is devoid of exostoses and may mark the location of a diverticulum of a cranial sinus." figureDoi="http://doi.org/10.5281/zenodo.5300257" httpUri="https://zenodo.org/record/5300257/files/figure.png" pageId="4" pageNumber="5">Fig. 6A</figureCitation>
), even though there is no modern analogue for such a composite arrangement. The dorsal portion of the rhamphotheca coated the external surface of the premaxillae and would have extended as far dorsally as the base of the dorsomedian premaxillary process, but would have been cut back so that it skirted the ventrolateral portion of the external naris. Its posterodorsal edge would have merged with the rhamphothecal margin near the posterior end of the premaxilla on either side of the snout (
<figureCitation box="[1294,1375,1812,1834]" captionStart="Figure 6" captionStartId="7.[163,241,1637,1659]" captionTargetBox="[175,1431,208,1582]" captionTargetId="figure-23@7.[163,1443,195,1597]" captionTargetPageId="7" captionText="Figure 6. Scelidosaurus. An interpretation of the cranial keratinous scute pattern on the skull and mandible in (A) lateral and (B) dorsal views. Abbreviations: asc, angular scute; fsc, frontal scute; dsc, dentary scute or sheath; hsc, occipital horn scute; itsc, infratemporal scutes; jsc, jugal arch scutes; lsc, lacrimal scute; msc, maxillary scute (there may have been several); nmsc, nasal median scute; nsc, nasal scutes restored as overlapping plates; pdsc, predentary scute (rhamphotheca); posc, postorbital crater-like scute; qjsc, quadratojugalquadrate scute; rsc, rhamphothecal scute of the premaxilla; sosc, supraorbital (brow-ridge) scute (there may have been several); stsc, supratemporal scutes; tym, tympanic membrane;?, area on the surface of the lacrimal lamina that is devoid of exostoses and may mark the location of a diverticulum of a cranial sinus." figureDoi="http://doi.org/10.5281/zenodo.5300257" httpUri="https://zenodo.org/record/5300257/files/figure.png" pageId="4" pageNumber="5">Fig. 6A</figureCitation>
).
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.5300253" ID-Zenodo-Dep="5300253" httpUri="https://zenodo.org/record/5300253/files/figure.png" pageId="5" pageNumber="6" startId="5.[163,243,1729,1751]" targetBox="[174,1434,211,1677]" targetPageId="5">
<paragraph blockId="5.[163,1442,1729,1781]" pageId="5" pageNumber="6">
<emphasis bold="true" box="[163,269,1729,1751]" pageId="5" pageNumber="6">Figure 4.</emphasis>
<taxonomicName authority="Owen, 1861" authorityName="Owen" authorityYear="1861" box="[280,675,1729,1751]" class="Reptilia" family="Scelidosauridae" genus="Scelidosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="harrisonii">
<emphasis box="[280,544,1729,1751]" italics="true" pageId="5" pageNumber="6">Scelidosaurus harrisonii</emphasis>
<bibRefCitation author="Owen R" box="[550,675,1729,1751]" pageId="5" pageNumber="6" pagination="1 - 14" refId="ref63737" refString="Owen R. 1861. Monograph of the fossil Reptilia of the Liassic Formations. A monograph of a fossil dinosaur (Scelidosaurus harrisonii, Owen) of the Lower Lias. Palaeontographical Society Monographs XII: 1 - 14." type="journal article" year="1861">Owen, 1861</bibRefCitation>
</taxonomicName>
. Cranial anatomy reconstructed in: A, lateral; B, dorsal views (modified from
<bibRefCitation author="Norman DB" box="[218,379,1759,1781]" pageId="5" pageNumber="6" pagination="1 - 81" refId="ref63078" refString="Norman DB. 2020 a. Scelidosaurus harrisonii Owen, 1861 (Dinosauria: Ornithischia) from the Early Jurassic of Dorset, England: cranial anatomy. Zoological Journal of the Linnean Society 188: 1 - 81." type="journal article" year="2020">Norman, 2020a</bibRefCitation>
: figs 8, 9).
</paragraph>
</caption>
<caption ID-DOI="http://doi.org/10.5281/zenodo.5300255" ID-Zenodo-Dep="5300255" httpUri="https://zenodo.org/record/5300255/files/figure.png" pageId="6" pageNumber="7" startId="6.[145,225,1070,1092]" targetBox="[153,1419,205,1025]" targetPageId="6">
<paragraph blockId="6.[145,1426,1070,1180]" pageId="6" pageNumber="7">
<emphasis bold="true" box="[145,251,1070,1092]" pageId="6" pageNumber="7">Figure 5.</emphasis>
<taxonomicName authority="(Linnaeus, 1758)" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[263,621,1070,1092]" class="Reptilia" family="Cheloniidae" genus="Chelonia" kingdom="Animalia" order="Testudines" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="mydas">
<emphasis box="[263,434,1070,1092]" italics="true" pageId="6" pageNumber="7">Chelonia mydas</emphasis>
(
<bibRefCitation author="Linnaeus C" box="[449,613,1070,1092]" pageId="6" pageNumber="7" refId="ref61651" refString="Linnaeus C. 1758. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus differentiis, synonymis, locis, 10 th edn. Stockholm: L. Salvius." type="book" year="1758">Linnaeus, 1758</bibRefCitation>
)
</taxonomicName>
the Green Turtle. External skull anatomy. A, lateral and B, dorsal views, illustrating the superficial cranial osteology. C, lateral and D, dorsal views showing the distribution of keratinous scutes. Norman, pers. colln (ex-University of Cambridge Zoology Department teaching collection). Scale bar indicated.Abbreviations: au, auditory recess; au.sc, auditory recess scute covering; tom, the chelonian tomium (= rhamphotheca of
<taxonomicName baseAuthorityName="NHMUK R" baseAuthorityYear="1111" box="[1243,1389,1158,1180]" class="Reptilia" family="Scelidosauridae" genus="Scelidosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis box="[1243,1389,1158,1180]" italics="true" pageId="6" pageNumber="7">Scelidosaurus</emphasis>
</taxonomicName>
).
</paragraph>
</caption>
<paragraph blockId="6.[145,265,1236,1260]" box="[145,265,1236,1260]" pageId="6" pageNumber="7">
<emphasis box="[145,265,1236,1260]" italics="true" pageId="6" pageNumber="7">The snout</emphasis>
</paragraph>
<paragraph blockId="6.[145,761,1277,1881]" lastBlockId="6.[809,1426,1237,1596]" pageId="6" pageNumber="7">
Anteriorly, the dorsal surface of the nasals (
<figureCitation box="[668,752,1277,1299]" captionStart="Figure 4" captionStartId="5.[163,243,1729,1751]" captionTargetBox="[174,1434,211,1677]" captionTargetId="figure-23@5.[163,1443,195,1689]" captionTargetPageId="5" captionText="Figure 4. Scelidosaurus harrisonii Owen, 1861. Cranial anatomy reconstructed in: A, lateral; B, dorsal views (modified from Norman, 2020a: figs 8, 9)." figureDoi="http://doi.org/10.5281/zenodo.5300253" httpUri="https://zenodo.org/record/5300253/files/figure.png" pageId="6" pageNumber="7">Fig. 4B</figureCitation>
) is characterized by a radiating pattern of strands of bony tissue that may have anchored a midline scute (
<figureCitation box="[154,282,1369,1391]" captionStart="Figure 6" captionStartId="7.[163,241,1637,1659]" captionTargetBox="[175,1431,208,1582]" captionTargetId="figure-23@7.[163,1443,195,1597]" captionTargetPageId="7" captionText="Figure 6. Scelidosaurus. An interpretation of the cranial keratinous scute pattern on the skull and mandible in (A) lateral and (B) dorsal views. Abbreviations: asc, angular scute; fsc, frontal scute; dsc, dentary scute or sheath; hsc, occipital horn scute; itsc, infratemporal scutes; jsc, jugal arch scutes; lsc, lacrimal scute; msc, maxillary scute (there may have been several); nmsc, nasal median scute; nsc, nasal scutes restored as overlapping plates; pdsc, predentary scute (rhamphotheca); posc, postorbital crater-like scute; qjsc, quadratojugalquadrate scute; rsc, rhamphothecal scute of the premaxilla; sosc, supraorbital (brow-ridge) scute (there may have been several); stsc, supratemporal scutes; tym, tympanic membrane;?, area on the surface of the lacrimal lamina that is devoid of exostoses and may mark the location of a diverticulum of a cranial sinus." figureDoi="http://doi.org/10.5281/zenodo.5300257" httpUri="https://zenodo.org/record/5300257/files/figure.png" pageId="6" pageNumber="7">Fig. 6A, B</figureCitation>
, nmsc); this underlying bony pattern corresponds to that which supports similar midline scutes on the chelonian skull (
<figureCitation box="[521,643,1430,1452]" captionStart="Figure 5" captionStartId="6.[145,225,1070,1092]" captionTargetBox="[153,1419,205,1025]" captionTargetId="figure-382@6.[145,1428,195,1030]" captionTargetPageId="6" captionText="Figure 5. Chelonia mydas (Linnaeus, 1758) the Green Turtle. External skull anatomy. A, lateral and B, dorsal views, illustrating the superficial cranial osteology. C, lateral and D, dorsal views showing the distribution of keratinous scutes. Norman, pers.colln (ex-University of Cambridge Zoology Department teachingcollection).Scale bar indicated.Abbreviations: au, auditory recess; au.sc, auditory recess scute covering; tom, the chelonian tomium (= rhamphotheca of Scelidosaurus)." figureDoi="http://doi.org/10.5281/zenodo.5300255" httpUri="https://zenodo.org/record/5300255/files/figure.png" pageId="6" pageNumber="7">Fig. 5B, D</figureCitation>
). Farther posteriorly, the surface of these bones develops a thicker knobbly texture that is overprinted by a series of repeated curved ridges. The ridges are oriented (more or less) transversely across the roof of the snout and extend down the sides of the snout where it is walled by the maxilla and prefrontal. The repetition of the curved ridges is suggestive of attachment sites for successive (possibly overlapping) scutes that encased the snout above the buccal emargination and extended posteriorly as far as the nasofrontal suture (
<figureCitation box="[153,215,1767,1789]" captionStart="Figure 6" captionStartId="7.[163,241,1637,1659]" captionTargetBox="[175,1431,208,1582]" captionTargetId="figure-23@7.[163,1443,195,1597]" captionTargetPageId="7" captionText="Figure 6. Scelidosaurus. An interpretation of the cranial keratinous scute pattern on the skull and mandible in (A) lateral and (B) dorsal views. Abbreviations: asc, angular scute; fsc, frontal scute; dsc, dentary scute or sheath; hsc, occipital horn scute; itsc, infratemporal scutes; jsc, jugal arch scutes; lsc, lacrimal scute; msc, maxillary scute (there may have been several); nmsc, nasal median scute; nsc, nasal scutes restored as overlapping plates; pdsc, predentary scute (rhamphotheca); posc, postorbital crater-like scute; qjsc, quadratojugalquadrate scute; rsc, rhamphothecal scute of the premaxilla; sosc, supraorbital (brow-ridge) scute (there may have been several); stsc, supratemporal scutes; tym, tympanic membrane;?, area on the surface of the lacrimal lamina that is devoid of exostoses and may mark the location of a diverticulum of a cranial sinus." figureDoi="http://doi.org/10.5281/zenodo.5300257" httpUri="https://zenodo.org/record/5300257/files/figure.png" pageId="6" pageNumber="7">Fig. 6</figureCitation>
, nsc). The lateral wall of the snout formed by the maxilla and premaxilla is reconstructed here covered by a large maxillary scute (
<figureCitation box="[463,528,1829,1851]" captionStart="Figure 6" captionStartId="7.[163,241,1637,1659]" captionTargetBox="[175,1431,208,1582]" captionTargetId="figure-23@7.[163,1443,195,1597]" captionTargetPageId="7" captionText="Figure 6. Scelidosaurus. An interpretation of the cranial keratinous scute pattern on the skull and mandible in (A) lateral and (B) dorsal views. Abbreviations: asc, angular scute; fsc, frontal scute; dsc, dentary scute or sheath; hsc, occipital horn scute; itsc, infratemporal scutes; jsc, jugal arch scutes; lsc, lacrimal scute; msc, maxillary scute (there may have been several); nmsc, nasal median scute; nsc, nasal scutes restored as overlapping plates; pdsc, predentary scute (rhamphotheca); posc, postorbital crater-like scute; qjsc, quadratojugalquadrate scute; rsc, rhamphothecal scute of the premaxilla; sosc, supraorbital (brow-ridge) scute (there may have been several); stsc, supratemporal scutes; tym, tympanic membrane;?, area on the surface of the lacrimal lamina that is devoid of exostoses and may mark the location of a diverticulum of a cranial sinus." figureDoi="http://doi.org/10.5281/zenodo.5300257" httpUri="https://zenodo.org/record/5300257/files/figure.png" pageId="6" pageNumber="7">Fig. 6</figureCitation>
, msc), but there are indications (
<figureCitation box="[285,366,1859,1881]" captionStart="Figure 4" captionStartId="5.[163,243,1729,1751]" captionTargetBox="[174,1434,211,1677]" captionTargetId="figure-23@5.[163,1443,195,1689]" captionTargetPageId="5" captionText="Figure 4. Scelidosaurus harrisonii Owen, 1861. Cranial anatomy reconstructed in: A, lateral; B, dorsal views (modified from Norman, 2020a: figs 8, 9)." figureDoi="http://doi.org/10.5281/zenodo.5300253" httpUri="https://zenodo.org/record/5300253/files/figure.png" pageId="6" pageNumber="7">Fig. 4A</figureCitation>
) of faint attachment ridges, so it is possible that a series of overlapping scutes were found here as well. Posterior to the reconstructed lateral maxillary scute there is a smooth patch of bone (
<figureCitation box="[1356,1419,1298,1320]" captionStart="Figure 6" captionStartId="7.[163,241,1637,1659]" captionTargetBox="[175,1431,208,1582]" captionTargetId="figure-23@7.[163,1443,195,1597]" captionTargetPageId="7" captionText="Figure 6. Scelidosaurus. An interpretation of the cranial keratinous scute pattern on the skull and mandible in (A) lateral and (B) dorsal views. Abbreviations: asc, angular scute; fsc, frontal scute; dsc, dentary scute or sheath; hsc, occipital horn scute; itsc, infratemporal scutes; jsc, jugal arch scutes; lsc, lacrimal scute; msc, maxillary scute (there may have been several); nmsc, nasal median scute; nsc, nasal scutes restored as overlapping plates; pdsc, predentary scute (rhamphotheca); posc, postorbital crater-like scute; qjsc, quadratojugalquadrate scute; rsc, rhamphothecal scute of the premaxilla; sosc, supraorbital (brow-ridge) scute (there may have been several); stsc, supratemporal scutes; tym, tympanic membrane;?, area on the surface of the lacrimal lamina that is devoid of exostoses and may mark the location of a diverticulum of a cranial sinus." figureDoi="http://doi.org/10.5281/zenodo.5300257" httpUri="https://zenodo.org/record/5300257/files/figure.png" pageId="6" pageNumber="7">Fig. 6</figureCitation>
,?) behind which the lacrimal bears irregular exostotic growth that would have supported an overlying scute (
<figureCitation box="[817,880,1390,1412]" captionStart="Figure 6" captionStartId="7.[163,241,1637,1659]" captionTargetBox="[175,1431,208,1582]" captionTargetId="figure-23@7.[163,1443,195,1597]" captionTargetPageId="7" captionText="Figure 6. Scelidosaurus. An interpretation of the cranial keratinous scute pattern on the skull and mandible in (A) lateral and (B) dorsal views. Abbreviations: asc, angular scute; fsc, frontal scute; dsc, dentary scute or sheath; hsc, occipital horn scute; itsc, infratemporal scutes; jsc, jugal arch scutes; lsc, lacrimal scute; msc, maxillary scute (there may have been several); nmsc, nasal median scute; nsc, nasal scutes restored as overlapping plates; pdsc, predentary scute (rhamphotheca); posc, postorbital crater-like scute; qjsc, quadratojugalquadrate scute; rsc, rhamphothecal scute of the premaxilla; sosc, supraorbital (brow-ridge) scute (there may have been several); stsc, supratemporal scutes; tym, tympanic membrane;?, area on the surface of the lacrimal lamina that is devoid of exostoses and may mark the location of a diverticulum of a cranial sinus." figureDoi="http://doi.org/10.5281/zenodo.5300257" httpUri="https://zenodo.org/record/5300257/files/figure.png" pageId="6" pageNumber="7">Fig. 6</figureCitation>
, lsc).
</paragraph>
<paragraph blockId="6.[809,1426,1237,1596]" pageId="6" pageNumber="7">
There is no equivalent patterning of successive curved exostotic ridges on the chelonian skulls that I have examined, so the scute pattern in the scelidosaur skull probably differs from the mosaic-like tessellate pattern of scutes seen in these living, albeit distantly related, diapsid taxa (
<figureCitation box="[1058,1121,1574,1596]" captionStart="Figure 5" captionStartId="6.[145,225,1070,1092]" captionTargetBox="[153,1419,205,1025]" captionTargetId="figure-382@6.[145,1428,195,1030]" captionTargetPageId="6" captionText="Figure 5. Chelonia mydas (Linnaeus, 1758) the Green Turtle. External skull anatomy. A, lateral and B, dorsal views, illustrating the superficial cranial osteology. C, lateral and D, dorsal views showing the distribution of keratinous scutes. Norman, pers.colln (ex-University of Cambridge Zoology Department teachingcollection).Scale bar indicated.Abbreviations: au, auditory recess; au.sc, auditory recess scute covering; tom, the chelonian tomium (= rhamphotheca of Scelidosaurus)." figureDoi="http://doi.org/10.5281/zenodo.5300255" httpUri="https://zenodo.org/record/5300255/files/figure.png" pageId="6" pageNumber="7">Fig. 5</figureCitation>
).
</paragraph>
<paragraph blockId="6.[809,1127,1633,1658]" box="[809,1127,1633,1658]" pageId="6" pageNumber="7">
<emphasis box="[809,1127,1633,1658]" italics="true" pageId="6" pageNumber="7">The skull roof and occiput</emphasis>
</paragraph>
<paragraph blockId="6.[809,1425,1674,1880]" pageId="6" pageNumber="7">
The frontals are dominated by a dense pattern of strand-like superficial bone that radiates from a midline groove (
<figureCitation box="[995,1078,1735,1757]" captionStart="Figure 4" captionStartId="5.[163,243,1729,1751]" captionTargetBox="[174,1434,211,1677]" captionTargetId="figure-23@5.[163,1443,195,1689]" captionTargetPageId="5" captionText="Figure 4. Scelidosaurus harrisonii Owen, 1861. Cranial anatomy reconstructed in: A, lateral; B, dorsal views (modified from Norman, 2020a: figs 8, 9)." figureDoi="http://doi.org/10.5281/zenodo.5300253" httpUri="https://zenodo.org/record/5300253/files/figure.png" pageId="6" pageNumber="7">Fig. 4B</figureCitation>
). As with the anterior portion of the nasal, this morphology is suggestive of the presence of a large, shield-like scute (
<figureCitation box="[1280,1367,1797,1819]" captionStart="Figure 6" captionStartId="7.[163,241,1637,1659]" captionTargetBox="[175,1431,208,1582]" captionTargetId="figure-23@7.[163,1443,195,1597]" captionTargetPageId="7" captionText="Figure 6. Scelidosaurus. An interpretation of the cranial keratinous scute pattern on the skull and mandible in (A) lateral and (B) dorsal views. Abbreviations: asc, angular scute; fsc, frontal scute; dsc, dentary scute or sheath; hsc, occipital horn scute; itsc, infratemporal scutes; jsc, jugal arch scutes; lsc, lacrimal scute; msc, maxillary scute (there may have been several); nmsc, nasal median scute; nsc, nasal scutes restored as overlapping plates; pdsc, predentary scute (rhamphotheca); posc, postorbital crater-like scute; qjsc, quadratojugalquadrate scute; rsc, rhamphothecal scute of the premaxilla; sosc, supraorbital (brow-ridge) scute (there may have been several); stsc, supratemporal scutes; tym, tympanic membrane;?, area on the surface of the lacrimal lamina that is devoid of exostoses and may mark the location of a diverticulum of a cranial sinus." figureDoi="http://doi.org/10.5281/zenodo.5300257" httpUri="https://zenodo.org/record/5300257/files/figure.png" pageId="6" pageNumber="7">Fig. 6B</figureCitation>
, fsc) that extended across to the adjacent surfaces of the prefrontal, middle supraorbital and postorbital.
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.5300257" ID-Zenodo-Dep="5300257" httpUri="https://zenodo.org/record/5300257/files/figure.png" pageId="7" pageNumber="8" startId="7.[163,241,1637,1659]" targetBox="[175,1431,208,1582]" targetPageId="7">
<paragraph blockId="7.[163,1443,1637,1864]" pageId="7" pageNumber="8">
<emphasis bold="true" box="[163,266,1637,1659]" pageId="7" pageNumber="8">Figure 6.</emphasis>
<taxonomicName baseAuthorityName="NHMUK R" baseAuthorityYear="1111" box="[277,423,1637,1659]" class="Reptilia" family="Scelidosauridae" genus="Scelidosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
<emphasis box="[277,423,1637,1659]" italics="true" pageId="7" pageNumber="8">Scelidosaurus</emphasis>
</taxonomicName>
. An interpretation of the cranial keratinous scute pattern on the skull and mandible in (A) lateral and (B) dorsal views. Abbreviations: asc, angular scute; fsc, frontal scute; dsc, dentary scute or sheath; hsc, occipital horn scute; itsc, infratemporal scutes; jsc, jugal arch scutes; lsc, lacrimal scute; msc, maxillary scute (there may have been several); nmsc, nasal median scute; nsc, nasal scutes restored as overlapping plates; pdsc, predentary scute (rhamphotheca); posc, postorbital crater-like scute; qjsc, quadratojugalquadrate scute; rsc, rhamphothecal scute of the premaxilla; sosc, supraorbital (brow-ridge) scute (there may have been several); stsc, supratemporal scutes; tym, tympanic membrane;?, area on the surface of the lacrimal lamina that is devoid of exostoses and may mark the location of a diverticulum of a cranial sinus.
</paragraph>
</caption>
<paragraph blockId="8.[145,761,197,1905]" pageId="8" pageNumber="9">
Lateral to the frontal plate, the palpebral (= anterior supraorbital) and posterior supraorbital osteoderms form a shallowly arched bar of bone (= brow ridge) that flanks the dorsal orbital margin of the skull roof. The rugose external surfaces of these bones are structurally distinct from the frontal plate, and are likely to have anchored their own substantial keratinous sheaths (
<figureCitation box="[154,226,412,434]" captionStart="Figure 6" captionStartId="7.[163,241,1637,1659]" captionTargetBox="[175,1431,208,1582]" captionTargetId="figure-23@7.[163,1443,195,1597]" captionTargetPageId="7" captionText="Figure 6. Scelidosaurus. An interpretation of the cranial keratinous scute pattern on the skull and mandible in (A) lateral and (B) dorsal views. Abbreviations: asc, angular scute; fsc, frontal scute; dsc, dentary scute or sheath; hsc, occipital horn scute; itsc, infratemporal scutes; jsc, jugal arch scutes; lsc, lacrimal scute; msc, maxillary scute (there may have been several); nmsc, nasal median scute; nsc, nasal scutes restored as overlapping plates; pdsc, predentary scute (rhamphotheca); posc, postorbital crater-like scute; qjsc, quadratojugalquadrate scute; rsc, rhamphothecal scute of the premaxilla; sosc, supraorbital (brow-ridge) scute (there may have been several); stsc, supratemporal scutes; tym, tympanic membrane;?, area on the surface of the lacrimal lamina that is devoid of exostoses and may mark the location of a diverticulum of a cranial sinus." figureDoi="http://doi.org/10.5281/zenodo.5300257" httpUri="https://zenodo.org/record/5300257/files/figure.png" pageId="8" pageNumber="9">Fig. 6</figureCitation>
, sosc) that served to shield the orbit and its associated soft tissues. It is possible that these keratinous scutes were subdivided into smaller units than illustrated here, and may have been superficially ornate for behaviourally related reasons; enlarged, sculpted and colourful circumorbital scutes are seen in many living squamates.
</paragraph>
<paragraph blockId="8.[145,761,197,1905]" pageId="8" pageNumber="9">
A rugose, double-ridged sagittal crest dominates the posterior part of the skull roof. The ridges are flanked by large ovoid supratemporal fenestrae and behind these latter is a divergent pair of prominent, curved, horn-like occipital osteoderms. The temporal arches and occipital margin show some evidence of irregular exostotic growth that may well reflect the attachment of overlying scutes (
<figureCitation box="[367,429,841,863]" captionStart="Figure 6" captionStartId="7.[163,241,1637,1659]" captionTargetBox="[175,1431,208,1582]" captionTargetId="figure-23@7.[163,1443,195,1597]" captionTargetPageId="7" captionText="Figure 6. Scelidosaurus. An interpretation of the cranial keratinous scute pattern on the skull and mandible in (A) lateral and (B) dorsal views. Abbreviations: asc, angular scute; fsc, frontal scute; dsc, dentary scute or sheath; hsc, occipital horn scute; itsc, infratemporal scutes; jsc, jugal arch scutes; lsc, lacrimal scute; msc, maxillary scute (there may have been several); nmsc, nasal median scute; nsc, nasal scutes restored as overlapping plates; pdsc, predentary scute (rhamphotheca); posc, postorbital crater-like scute; qjsc, quadratojugalquadrate scute; rsc, rhamphothecal scute of the premaxilla; sosc, supraorbital (brow-ridge) scute (there may have been several); stsc, supratemporal scutes; tym, tympanic membrane;?, area on the surface of the lacrimal lamina that is devoid of exostoses and may mark the location of a diverticulum of a cranial sinus." figureDoi="http://doi.org/10.5281/zenodo.5300257" httpUri="https://zenodo.org/record/5300257/files/figure.png" pageId="8" pageNumber="9">Fig. 6</figureCitation>
, stsc). It is, of course, possible that this posterior part of the skull table was less extensively scute-covered. The fenestrae themselves would have been spanned by skin that (although scaly) retained a degree of flexibility to allow movement of the underlying temporal musculature. The same consideration should also apply to the adjacent infratemporal fenestrae (itsc). However, there are areas of the skull of
<taxonomicName baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[385,573,1086,1107]" class="Reptilia" family="Cheloniidae" genus="Chelonia" kingdom="Animalia" order="Testudines" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="mydas">
<emphasis box="[385,573,1086,1107]" italics="true" pageId="8" pageNumber="9">Chelonia mydas</emphasis>
</taxonomicName>
where exostotic bone is absent and the bone surface is, instead, smooth-surfaced (
<figureCitation box="[352,435,1147,1170]" captionStart="Figure 5" captionStartId="6.[145,225,1070,1092]" captionTargetBox="[153,1419,205,1025]" captionTargetId="figure-382@6.[145,1428,195,1030]" captionTargetPageId="6" captionText="Figure 5. Chelonia mydas (Linnaeus, 1758) the Green Turtle. External skull anatomy. A, lateral and B, dorsal views, illustrating the superficial cranial osteology. C, lateral and D, dorsal views showing the distribution of keratinous scutes. Norman, pers.colln (ex-University of Cambridge Zoology Department teachingcollection).Scale bar indicated.Abbreviations: au, auditory recess; au.sc, auditory recess scute covering; tom, the chelonian tomium (= rhamphotheca of Scelidosaurus)." figureDoi="http://doi.org/10.5281/zenodo.5300255" httpUri="https://zenodo.org/record/5300255/files/figure.png" pageId="8" pageNumber="9">Fig. 5A</figureCitation>
, au); this area (the margins of which are dotted in
<figureCitation box="[420,506,1178,1200]" captionStart="Figure 5" captionStartId="6.[145,225,1070,1092]" captionTargetBox="[153,1419,205,1025]" captionTargetId="figure-382@6.[145,1428,195,1030]" captionTargetPageId="6" captionText="Figure 5. Chelonia mydas (Linnaeus, 1758) the Green Turtle. External skull anatomy. A, lateral and B, dorsal views, illustrating the superficial cranial osteology. C, lateral and D, dorsal views showing the distribution of keratinous scutes. Norman, pers.colln (ex-University of Cambridge Zoology Department teachingcollection).Scale bar indicated.Abbreviations: au, auditory recess; au.sc, auditory recess scute covering; tom, the chelonian tomium (= rhamphotheca of Scelidosaurus)." figureDoi="http://doi.org/10.5281/zenodo.5300255" httpUri="https://zenodo.org/record/5300255/files/figure.png" pageId="8" pageNumber="9">Fig. 5C</figureCitation>
) is covered in life by several scutes (
<figureCitation box="[323,405,1209,1231]" captionStart="Figure 5" captionStartId="6.[145,225,1070,1092]" captionTargetBox="[153,1419,205,1025]" captionTargetId="figure-382@6.[145,1428,195,1030]" captionTargetPageId="6" captionText="Figure 5. Chelonia mydas (Linnaeus, 1758) the Green Turtle. External skull anatomy. A, lateral and B, dorsal views, illustrating the superficial cranial osteology. C, lateral and D, dorsal views showing the distribution of keratinous scutes. Norman, pers.colln (ex-University of Cambridge Zoology Department teachingcollection).Scale bar indicated.Abbreviations: au, auditory recess; au.sc, auditory recess scute covering; tom, the chelonian tomium (= rhamphotheca of Scelidosaurus)." figureDoi="http://doi.org/10.5281/zenodo.5300255" httpUri="https://zenodo.org/record/5300255/files/figure.png" pageId="8" pageNumber="9">Fig. 5C</figureCitation>
, ausc). Therefore, it is possible that in
<taxonomicName baseAuthorityName="NHMUK R" baseAuthorityYear="1111" box="[230,389,1240,1261]" class="Reptilia" family="Scelidosauridae" genus="Scelidosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
<emphasis box="[230,389,1240,1261]" italics="true" pageId="8" pageNumber="9">Scelidosaurus</emphasis>
</taxonomicName>
tessellated scutes enveloped the posterior skull roof, as well as its lateral flanks. The restoration (
<figureCitation box="[286,397,1301,1323]" captionStart="Figure 6" captionStartId="7.[163,241,1637,1659]" captionTargetBox="[175,1431,208,1582]" captionTargetId="figure-23@7.[163,1443,195,1597]" captionTargetPageId="7" captionText="Figure 6. Scelidosaurus. An interpretation of the cranial keratinous scute pattern on the skull and mandible in (A) lateral and (B) dorsal views. Abbreviations: asc, angular scute; fsc, frontal scute; dsc, dentary scute or sheath; hsc, occipital horn scute; itsc, infratemporal scutes; jsc, jugal arch scutes; lsc, lacrimal scute; msc, maxillary scute (there may have been several); nmsc, nasal median scute; nsc, nasal scutes restored as overlapping plates; pdsc, predentary scute (rhamphotheca); posc, postorbital crater-like scute; qjsc, quadratojugalquadrate scute; rsc, rhamphothecal scute of the premaxilla; sosc, supraorbital (brow-ridge) scute (there may have been several); stsc, supratemporal scutes; tym, tympanic membrane;?, area on the surface of the lacrimal lamina that is devoid of exostoses and may mark the location of a diverticulum of a cranial sinus." figureDoi="http://doi.org/10.5281/zenodo.5300257" httpUri="https://zenodo.org/record/5300257/files/figure.png" pageId="8" pageNumber="9">Fig. 6A, B</figureCitation>
) includes an imaginary array of rather large scutes in these areas.
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.5300259" ID-Zenodo-Dep="5300259" httpUri="https://zenodo.org/record/5300259/files/figure.png" pageId="8" pageNumber="9" startId="8.[808,890,903,925]" targetBox="[820,1416,203,861]" targetPageId="8">
<paragraph blockId="8.[808,1425,903,1101]" pageId="8" pageNumber="9">
<emphasis bold="true" box="[808,918,903,925]" pageId="8" pageNumber="9">Figure 7.</emphasis>
<taxonomicName authority="Linnaeus, 1758" authorityName="Linnaeus" authorityYear="1758" box="[931,1219,903,925]" class="Mammalia" family="Bovidae" genus="Ovis" kingdom="Animalia" order="Artiodactyla" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="aries">
<emphasis box="[931,1042,903,925]" italics="true" pageId="8" pageNumber="9">Ovis aries</emphasis>
<bibRefCitation author="Linnaeus C" box="[1051,1219,903,925]" pageId="8" pageNumber="9" refId="ref61651" refString="Linnaeus C. 1758. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus differentiis, synonymis, locis, 10 th edn. Stockholm: L. Salvius." type="book" year="1758">Linnaeus, 1758</bibRefCitation>
</taxonomicName>
. A, lateral view of cranial osteology showing the horn core. B, the horn core with its keratinous horn superimposed, showing the lack of correspondence in shape between the horn core and its overlying keratinous casque (horn). Norman, pers. colln (Rough Fell sheep, Sedbergh, Cumbria). Abbreviations: hc, horn core; kh, keratinous horn. Scale bar indicated.
</paragraph>
</caption>
<paragraph blockId="8.[145,761,197,1905]" pageId="8" pageNumber="9">
The occipital osteoderm horns have comparatively smooth, finely grooved surfaces pock-marked by many tiny foramina; these features are interpreted as a combination of points for the connective tissue that tethered an overlying keratinous horn (
<figureCitation box="[624,692,1485,1507]" captionStart="Figure 6" captionStartId="7.[163,241,1637,1659]" captionTargetBox="[175,1431,208,1582]" captionTargetId="figure-23@7.[163,1443,195,1597]" captionTargetPageId="7" captionText="Figure 6. Scelidosaurus. An interpretation of the cranial keratinous scute pattern on the skull and mandible in (A) lateral and (B) dorsal views. Abbreviations: asc, angular scute; fsc, frontal scute; dsc, dentary scute or sheath; hsc, occipital horn scute; itsc, infratemporal scutes; jsc, jugal arch scutes; lsc, lacrimal scute; msc, maxillary scute (there may have been several); nmsc, nasal median scute; nsc, nasal scutes restored as overlapping plates; pdsc, predentary scute (rhamphotheca); posc, postorbital crater-like scute; qjsc, quadratojugalquadrate scute; rsc, rhamphothecal scute of the premaxilla; sosc, supraorbital (brow-ridge) scute (there may have been several); stsc, supratemporal scutes; tym, tympanic membrane;?, area on the surface of the lacrimal lamina that is devoid of exostoses and may mark the location of a diverticulum of a cranial sinus." figureDoi="http://doi.org/10.5281/zenodo.5300257" httpUri="https://zenodo.org/record/5300257/files/figure.png" pageId="8" pageNumber="9">Fig. 6</figureCitation>
, hsc), and the vascular supply for its continued growth. Similar textures are visible on the horn-cores of living bovid mammals (e.g.
<emphasis box="[459,591,1577,1599]" italics="true" pageId="8" pageNumber="9">
<taxonomicName authorityName="Linnaeus" authorityYear="1758" box="[459,573,1577,1599]" class="Mammalia" family="Bovidae" genus="Ovis" kingdom="Animalia" order="Artiodactyla" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="aries">Ovis aries</taxonomicName>
</emphasis>
Norman, pers. colln
<figureCitation box="[232,318,1608,1630]" captionStart="Figure 7" captionStartId="8.[808,890,903,925]" captionTargetBox="[820,1416,203,861]" captionTargetId="figure-708@8.[811,1422,195,863]" captionTargetPageId="8" captionText="Figure 7. Ovis aries Linnaeus, 1758. A, lateral view of cranial osteology showing the horn core. B, the horn core with its keratinous horn superimposed, showing the lack of correspondence in shape between the horn core and its overlying keratinous casque (horn). Norman, pers. colln (Rough Fell sheep, Sedbergh, Cumbria). Abbreviations: hc, horn core; kh, keratinous horn. Scale bar indicated." figureDoi="http://doi.org/10.5281/zenodo.5300259" httpUri="https://zenodo.org/record/5300259/files/figure.png" pageId="8" pageNumber="9">Fig. 7A</figureCitation>
, hc). As can be seen in this example, the shape of the horn core may not necessarily have a direct bearing on that of the overlying keratinous sheath (
<figureCitation box="[246,335,1700,1722]" captionStart="Figure 7" captionStartId="8.[808,890,903,925]" captionTargetBox="[820,1416,203,861]" captionTargetId="figure-708@8.[811,1422,195,863]" captionTargetPageId="8" captionText="Figure 7. Ovis aries Linnaeus, 1758. A, lateral view of cranial osteology showing the horn core. B, the horn core with its keratinous horn superimposed, showing the lack of correspondence in shape between the horn core and its overlying keratinous casque (horn). Norman, pers. colln (Rough Fell sheep, Sedbergh, Cumbria). Abbreviations: hc, horn core; kh, keratinous horn. Scale bar indicated." figureDoi="http://doi.org/10.5281/zenodo.5300259" httpUri="https://zenodo.org/record/5300259/files/figure.png" pageId="8" pageNumber="9">Fig. 7B</figureCitation>
, kh); the same may be true in the case of the scelidosaur, but in the absence of new discoveries of scelidosaur material in the Lias exhibiting exceptional preservation (e.g.
<bibRefCitation author="Arbour VM &amp; Evans DC" pageId="8" pageNumber="9" pagination="161086" refId="ref57461" refString="Arbour VM, Evans DC. 2017. A new ankylosaurine dinosaur from the Judith River Formation of Montana, USA, based on an exceptional skeleton with soft tissue preservation. Royal Society Open Science 4: 161086." type="journal article" year="2017">Arbour &amp; Evans, 2017</bibRefCitation>
;
<bibRefCitation author="Brown CM" box="[298,443,1822,1844]" pageId="8" pageNumber="9" pagination="4066" refId="ref58176" refString="Brown CM. 2017. An exceptionally preserved armored dinosaur reveals the morphology and allometry of osteoderms and their horny epidermal coverings. PeerJ 5: e 4066." type="journal article" year="2017">Brown, 2017</bibRefCitation>
), there is no way of judging on the matter, so a conservative restoration has been illustrated.
</paragraph>
</subSubSection>
</treatment>
</document>