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<mods:title id="E8A1180E049D4C57913ECC672EF311EF">Cranial skeletogenesis of one of the largest amphibians, Andrias japonicus, provides insight into ontogenetic adaptations for feeding in salamanders</mods:title>
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<paragraph id="1C0B36EEFF85E258FF48FA8BD4AD7375" blockId="6.[145,486,1283,1307]" box="[145,486,1283,1307]" pageId="6" pageNumber="305">
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<taxonomicName id="DBB44D6DFF85E258FF48FA8BD42D7375" ID-CoL="4KBDC" authorityName="Michahelles" authorityYear="1830" box="[145,358,1283,1307]" class="Amphibia" family="Salamandridae" genus="Pleurodeles" kingdom="Animalia" order="Caudata" pageId="6" pageNumber="305" phylum="Chordata" rank="species" species="waltl">Pleurodeles waltl</taxonomicName>
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(
<figureCitation id="848F2A6BFF85E258FEAFFA8CD4957375" box="[374,478,1283,1307]" captionStart-0="Figure 3" captionStart-1="Figure 4" captionStart-2="Figure 5" captionStartId-0="9.[163,246,1759,1781]" captionStartId-1="9.[163,243,1206,1228]" captionStartId-2="10.[145,225,1055,1077]" captionTargetBox-1="[166,1443,198,1162]" captionTargetBox-2="[148,1429,198,1012]" captionTargetId-0="figure-24@8.[189,1380,195,1923]" captionTargetId-1="figure-213@9.[163,1449,195,1166]" captionTargetId-2="figure-397@10.[145,1432,195,1015]" captionTargetPageId-0="8" captionTargetPageId-1="9" captionTargetPageId-2="10" captionText-0="Figure 3. Cranial skeletogenesis (dorsal view) in Andrias japonicus, Hynobius nebulosus, Pleurodeles waltl and Ambystoma mexicanum. Pink elements are bone. Blue elements are cartilage. Scale bars are 1 mm, except those for phase XIII An. japonicus and Am. mexicanum, which are 10 mm. Abbreviations: cl, columella; exo, exoccipital; fr, frontal; ma, maxilla; na, nasal; oc, otic capsule; opi, opisthotic, osph, orbitosphenoid; pa, parietal; pfr, prefrontal; pma, premaxilla; po, prootic; psph, parasphenoid; pt, pterygoid; q, quadrate; seth, sphenothmoid; sq, squamosal; vo, vomer." captionText-1="Figure 4. Select phases of cranial skeletogenesis (ventral view) in Andrias japonicus, Hynobius nebulosus, Pleurodeles waltl and Ambystoma mexicanum. Pink elements are bone. Blue elements are cartilage. Scale bars are 1 mm, except for the 10 mm scale bar for phase XIII An. japonicus. Abbreviations: cl, columella; exo, exoccipital; fr, frontal; ma, maxilla; na, nasal; oc, otic capsule; opi, opisthotic, osph, orbitosphenoid; pa, parietal; pfr, prefrontal; pma, premaxilla; po, prootic; psph, parasphenoid; pt, pterygoid; q, quadrate; seth, sphenothmoid; sq, squamosal; vo, vomer." captionText-2="Figure 5. Osteogenesis of the lower jaw and hyobranchial skeletons (ventral view) at larval (phases I or II) and postmetamorphosis (phases XII or XIII) stages of Andrias japonicus, Hynobius nebulosus, Pleurodeles waltl and Ambystoma mexicanum. Pink elements are bone. Blue elements are cartilage. Scale bars: 1 mm.Abbreviations: ang, angular; ar, anterior radial; art, articular; bb, basibranchial; cb, ceratobranchial; ce, ceratohyal; den, dentary; hb, hypobranchial; hc, hypohyal; pr, posterior radial." figureDoi-0="http://doi.org/10.5281/zenodo.6535618" figureDoi-1="http://doi.org/10.5281/zenodo.6530552" figureDoi-2="http://doi.org/10.5281/zenodo.6530554" httpUri-0="https://zenodo.org/record/6535618/files/figure.png" httpUri-1="https://zenodo.org/record/6530552/files/figure.png" httpUri-2="https://zenodo.org/record/6530554/files/figure.png" pageId="6" pageNumber="305">Figs 35</figureCitation>
)
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<bibRefCitation id="78254B1FFF85E258FF48FAA4D4CB732C" author="Smirnov SV &amp; Merkulova KM &amp; Vassilieva AB" box="[145,384,1324,1346]" pageId="6" pageNumber="305" pagination="543 - 555" refId="ref10331" refString="Smirnov SV, Merkulova KM, Vassilieva AB. 2020. Skull development in the Iberian newt, Pleurodeles waltl (Salamandridae: Caudata: Amphibia): timing, sequence, variations, and thyroid hormone mediation of bone appearance. Journal of Anatomy 237: 543 - 555." type="journal article" year="2020">
Smirnov
<emphasis id="2EC0EAFCFF85E258FF20FAA4D479732F" box="[249,306,1324,1345]" italics="true" pageId="6" pageNumber="305">et al.</emphasis>
(2020)
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described cranial skeletogenesis of
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<emphasis id="2EC0EAFCFF85E258FF77FAC3D44D7331" box="[174,262,1354,1376]" italics="true" pageId="6" pageNumber="305">P. waltl</emphasis>
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and reported that ossification began soon after hatching. The results of
<bibRefCitation id="78254B1FFF85E258FE04FAE1D78C7311" author="Smirnov SV &amp; Merkulova KM &amp; Vassilieva AB" box="[477,711,1385,1407]" pageId="6" pageNumber="305" pagination="543 - 555" refId="ref10331" refString="Smirnov SV, Merkulova KM, Vassilieva AB. 2020. Skull development in the Iberian newt, Pleurodeles waltl (Salamandridae: Caudata: Amphibia): timing, sequence, variations, and thyroid hormone mediation of bone appearance. Journal of Anatomy 237: 543 - 555." type="journal article" year="2020">
Smirnov
<emphasis id="2EC0EAFCFF85E258FD9DFAE2D7307310" box="[580,635,1385,1407]" italics="true" pageId="6" pageNumber="305">et al.</emphasis>
(2020)
</bibRefCitation>
and our results show a similar pattern of ossification. In the study by
<bibRefCitation id="78254B1FFF85E258FEFCFA2ED75873D3" author="Smirnov SV &amp; Merkulova KM &amp; Vassilieva AB" box="[293,531,1446,1469]" pageId="6" pageNumber="305" pagination="543 - 555" refId="ref10331" refString="Smirnov SV, Merkulova KM, Vassilieva AB. 2020. Skull development in the Iberian newt, Pleurodeles waltl (Salamandridae: Caudata: Amphibia): timing, sequence, variations, and thyroid hormone mediation of bone appearance. Journal of Anatomy 237: 543 - 555." type="journal article" year="2020">
Smirnov
<emphasis id="2EC0EAFCFF85E258FE54FA2FD48E73D5" box="[397,453,1446,1468]" italics="true" pageId="6" pageNumber="305">et al.</emphasis>
(2020)
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, specimen body size was
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at the hatching stage and
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at the stage when metamorphosis occurs. However, in our study, these measurements were
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and
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, respectively. Given that the growth rate of salamanders is influenced strongly by external factors, such as temperature and food, this difference in the timing of ossification onset could be attributable to the different rearing environments (
<bibRefCitation id="78254B1FFF85E258FDD8F914D7EB70DC" author="Stewart MM" box="[513,672,1692,1714]" pageId="6" pageNumber="305" pagination="47 - 56" refId="ref10461" refString="Stewart MM. 1956. The separate effects of food and temperature differences on development of marbled salamander larvae. Journal of the Mitchell Society 72: 47 - 56." type="journal article" year="1956">Stewart, 1956</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="54AE6565FF85E258FF48F943D0327283" pageId="6" pageNumber="305" type="description">
<paragraph id="1C0B36EEFF85E258FF48F943D11E7776" blockId="6.[145,761,1738,1883]" lastBlockId="6.[809,1426,197,1261]" pageId="6" pageNumber="305">In phases IV, ossification has not yet started. In the skull, the sphenethmoid, otic capsule and exoccipital are observed as cartilaginous elements. In the mandible and pharynx, the Meckels cartilage, the first and second basibranchials, ceratohyal, hypohyal, the first and second hypobranchials and the first to fourth ceratobranchials are detected as cartilaginous elements.</paragraph>
<paragraph id="1C0B36EEFF85E258FCF0FEA9D0D977FD" blockId="6.[809,1426,197,1261]" pageId="6" pageNumber="305">Ossification begins in phase VI. The premaxilla, vomer, prootic, squamosal, exoccipital, parasphenoid, dentary and angular begin to ossify. The palatine and pterygoid bones appear fused into a single bone.</paragraph>
<paragraph id="1C0B36EEFF85E258FCF0FE14D1D27460" blockId="6.[809,1426,197,1261]" pageId="6" pageNumber="305">In phase VII, ossification of the skull progresses rapidly, and all bones except the nasal and orbitosphenoid begin to ossify. In the mandible, the coronoid is ossified from the medial side.</paragraph>
<paragraph id="1C0B36EEFF85E258FCF0FD9FD6BC74E6" blockId="6.[809,1426,197,1261]" pageId="6" pageNumber="305">In phase VIII, the nasal begins to ossify, meaning that all bones of the skull are ossified. In the hyobranchial skeleton, the second basibranchial starts to ossify.</paragraph>
<paragraph id="1C0B36EEFF85E258FCF0FD19D12A754C" blockId="6.[809,1426,197,1261]" pageId="6" pageNumber="305">In phase IX, the posterolateral projection of the frontal and anteromedial projection of the squamosal are visible. The pterygoid is elongated in the anterior posterior direction, forming a bony bar in the ventral side of the skull.</paragraph>
<paragraph id="1C0B36EEFF85E258FCF0FCA3D69A7531" blockId="6.[809,1426,197,1261]" pageId="6" pageNumber="305">In phases X and XI, the third to fourth ceratobranchials degenerate.</paragraph>
<paragraph id="1C0B36EEFF85E258FCF0FCE0D0327283" blockId="6.[809,1426,197,1261]" pageId="6" pageNumber="305">
In phases XII and XIII, the tips of the posterolateral projection of the frontal and the anteromedial projection of the squamosal fuse to form the frontosquamosal arch, a characteristic skull feature of the family
<taxonomicName id="DBB44D6DFF85E258FB9CFC6AD1B17596" authorityName="Goldfuss" authorityYear="1820" box="[1093,1274,994,1016]" class="Amphibia" family="Salamandridae" kingdom="Animalia" order="Caudata" pageId="6" pageNumber="305" phylum="Chordata" rank="family">Salamandridae</taxonomicName>
. The maxilla is elongated posterolaterally and its posterior end is now bifurcated, possessing short lateral and medial projections. The bony bar formed by the formerly continuous anterior palatine and posterior pterygoid is now separated in the middle. In the mandible, the second basibranchial degenerates, and the previously cartilaginous ceratohyal, second ceratobranchial and second hypobranchial ossify.
</paragraph>
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