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<document id="1537F736DF994DBD28BFE613B47F2B01" ID-CLB-Dataset="276646" ID-DOI="10.1111/j.1096-3642.2009.00533.x" ID-GBIF-Dataset="fdee393a-0668-4e46-9807-91b0e351b4f3" ID-ISSN="0024-4082" ID-Zenodo-Dep="10114794" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1630456506483" checkinUser="felipe" docAuthor="Maxwell, Erin E." docDate="2009" docId="7F4D879C757FFF9FF857FD5BD2C9FA86" docLanguage="en" docName="j.1096-3642.2009.00480.x.pdf" docOrigin="Zoological Journal of the Linnean Society 156 (1)" docSource="https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2009.00533.x" docStyle="DocumentStyle:0DD8C314D74634CE09062A86991413F8.2:ZoolJLinnSoc.2002-2009.journal_article" docStyleId="0DD8C314D74634CE09062A86991413F8" docStyleName="ZoolJLinnSoc.2002-2009.journal_article" docStyleVersion="2" docTitle="Rhea americana" docType="treatment" docVersion="7" lastPageNumber="191" masterDocId="8374FFE47578FF98F909FFF1D66FFF8B" masterDocTitle="Comparative ossification and development of the skull in palaeognathous birds (Aves: Palaeognathae)" masterLastPageNumber="200" masterPageNumber="184" pageNumber="191" updateTime="1700040445798" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0" zenodo-license-figures="CC-BY-4.0">
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<mods:title id="3E87B6B44861C9B0FAC2E54D51A39A8C">Comparative ossification and development of the skull in palaeognathous birds (Aves: Palaeognathae)</mods:title>
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<mods:namePart id="A1B43623AF859C477E1D8CF2E026DAD3">Maxwell, Erin E.</mods:namePart>
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<mods:date id="F9FD19FF6DAC66BA88BF6ADC5E522BB5">2009</mods:date>
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<treatment id="7F4D879C757FFF9FF857FD5BD2C9FA86" ID-DOI="http://doi.org/10.5281/zenodo.5492307" ID-GBIF-Taxon="217238449" ID-Zenodo-Dep="5492307" LSID="urn:lsid:plazi:treatment:7F4D879C757FFF9FF857FD5BD2C9FA86" httpUri="http://treatment.plazi.org/id/7F4D879C757FFF9FF857FD5BD2C9FA86" lastPageNumber="191" pageId="7" pageNumber="191">
<subSubSection id="BFFE6501757FFF9FF857FD5BD736FD6E" pageId="7" pageNumber="191" type="nomenclature">
<paragraph id="F75B368A757FFF9FF857FD5BD447FD36" blockId="7.[350,552,682,703]" box="[350,552,682,703]" pageId="7" pageNumber="191">
<heading id="AC1381E6757FFF9FF857FD5BD447FD36" box="[350,552,682,703]" centered="true" fontSize="9" level="2" pageId="7" pageNumber="191" reason="2">
<taxonomicName id="30E44D09757FFF9FF857FD5BD447FD36" ID-CoL="4S5KY" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[350,552,682,703]" class="Aves" family="Rheidae" genus="Rhea" kingdom="Animalia" order="Rheiformes" pageId="7" pageNumber="191" phylum="Chordata" rank="species" species="americana">
<emphasis id="C590EA98757FFF9FF857FD5BD447FD36" box="[350,552,682,703]" italics="true" pageId="7" pageNumber="191">
<emphasis id="C590EA98757FFF9FF857FD5BD7F2FD36" box="[350,413,682,703]" italics="true" pageId="7" pageNumber="191">RHEA</emphasis>
AMERICANA
</emphasis>
</taxonomicName>
</heading>
</paragraph>
<paragraph id="F75B368A757FFF9FF999FD21D736FD6E" blockId="7.[144,759,720,987]" box="[144,345,720,741]" pageId="7" pageNumber="191">
<emphasis id="C590EA98757FFF9FF999FD21D736FD6E" box="[144,345,720,741]" italics="true" pageId="7" pageNumber="191">
Stage 34 (
<figureCitation id="6FDF2A0F757FFF9FF80EFD21D73EFD6E" box="[263,337,720,741]" captionStart="Figure 1" captionStartId="7.[144,223,198,217]" captionText="Figure 1. Lateral view of the skull of palaeognath embryos. A–C, Dromaius novaehollandiae. A, stage 34 (day 22 of incubation, RM 8020); B, stage 38 (day 32 of incubation, RM 8030); C, stage 40+ (day 43 of incubation, RM 8039). D–E, Eudromia elegans. D, day 11 of incubation (YPM 112520); E, day 15 of incubation (YPM 112525). F–H, Struthio camelus. F, day 15 of incubation (YPM 112437); G, day 23 of incubation (YPM 112448); H, day 35 of incubation (YPM 112465). I–J, Rhea americana. I, stage 34 (day 14 of incubation, RM 72172); J, stage 40+ (day 26 of incubation, RM 7223). Grey shaded regions represent cartilage; black regions represent ossified tissue. The density of stippling reflects the relative degree of ossification. Scale bars = 5 mm. Abbreviations: ac, auditory capsule; ang, angular; d, dentary; f, frontal; ip, infrapolar process; j, jugal; lac, lacrimal; m, Meckel’s cartilage; me, mesethmoid; met, metotic cartilage; mx, maxilla; mx (pn), nasal process of the maxilla; n, nasal; n (pm), maxillary process of the nasal; oo, opisthotic; pa, parietal; pal, palatine; pao, planum antorbitale; pmx, premaxilla; pnp, prenasal process; poc, postorbital cartilage; pt, pterygoid; q, quadrate; qj; quadratojugal; sa, supra-angular; so, supraoccipital; soc, supraorbital cartilage; sq, squamosal." pageId="7" pageNumber="191">Fig. 1I</figureCitation>
)
</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="BFFE6501757FFF9FF999FD1ED2C9FA86" pageId="7" pageNumber="191" type="description">
<paragraph id="F75B368A757FFF9FF999FD1ED41DFC51" blockId="7.[144,759,720,987]" pageId="7" pageNumber="191">No ossification is present at this stage. The prenasal process is slightly swollen at its tip, and is only slightly longer than Meckel’s cartilage. The roof of the nasal tectum has grown posteriorly, dorsal to the orbits. The pars canaliculi of the auditory capsule is prominent, encompassing the entire lateral posterior margin of the skull. The contact between the two rami of Meckel’s cartilage is relatively narrow.</paragraph>
<paragraph id="F75B368A757FFF9FF999FC0BD69BFB84" blockId="7.[144,759,1018,1315]" box="[144,244,1018,1039]" pageId="7" pageNumber="191">
<emphasis id="C590EA98757FFF9FF999FC0BD69BFB84" box="[144,244,1018,1039]" italics="true" pageId="7" pageNumber="191">Stage 35</emphasis>
</paragraph>
<paragraph id="F75B368A757FFF9FF999FBE9D7A9FAA8" blockId="7.[144,759,1018,1315]" pageId="7" pageNumber="191">The contact between the two rami of Meckel’s cartilage becomes broader in this stage. The squamosal, palatine, vomer, pterygoid, jugal, and quadratojugal are ossifying. The lacrimal is ossifying, beginning from its orbital process. The frontal process of the premaxilla is ossifying, as is the jugal process of the maxilla. The supra-angular and angular are ossified in the lower jaw. This is followed by the ossification of the parasphenoid rostrum.</paragraph>
<paragraph id="F75B368A757FFF9FF999FAB2D70EFADC" blockId="7.[144,759,1346,1644]" box="[144,353,1346,1368]" pageId="7" pageNumber="191">
<emphasis id="C590EA98757FFF9FF999FAB2D70EFADC" box="[144,353,1346,1368]" italics="true" pageId="7" pageNumber="191">
Stage 37 (
<figureCitation id="6FDF2A0F757FFF9FF80EFAB2D735FADC" box="[263,346,1346,1368]" captionStart="Figure 2" captionStartId="8.[826,905,198,217]" captionTargetBox="[173,773,195,1791]" captionTargetId="figure-315@8.[168,776,193,1794]" captionTargetPageId="8" captionText="Figure 2. Palatal view of selected palaeognath embryos. A, B, Struthio camelus (modified from Parker, 1866). C, Rhea americana, stage 37 (day 17 of incubation, RM 7219). D, Rhea americana (modified from Müller, 1963). E, Dromaius novaehollandiae, stage 37 (day 28 of incubation, RM 8026). F, Eudromia elegans, day 14 of incubation (YPM 112524). Scale bars = 5 mm. Abbreviations: bo, basioccipital; exo, exoccipital; mx, maxilla; pal, palatine; pmx, premaxilla; psl, parasphenoid lamina; psr, parasphenoid rostrum; pt, pterygoid; q, quadrate; v, vomer." figureDoi="http://doi.org/10.5281/zenodo.10132793" httpUri="https://zenodo.org/record/10132793/files/figure.png" pageId="7" pageNumber="191">Fig. 2C</figureCitation>
)
</emphasis>
</paragraph>
<paragraph id="F75B368A757FFF9FF999FA90D691F9E7" blockId="7.[144,759,1346,1644]" pageId="7" pageNumber="191">The nasal is ossifying, as are the splenial, prearticular, and dentary. The jugal process of the premaxilla is ossified, but is not fused with the frontal process. The maxilla is triradiate. There are two independent, parallel ossification centres posterior to the parasphenoid rostrum; these represent the initiation of ossification of the parasphenoid lamina. The squamosal forms a process that parallels the otic process of the quadrate.</paragraph>
<paragraph id="F75B368A757FFF9FF999F97AD757F92B" blockId="7.[144,758,1675,1758]" box="[144,312,1675,1696]" pageId="7" pageNumber="191">
<emphasis id="C590EA98757FFF9FF999F97AD757F92B" box="[144,312,1675,1696]" italics="true" pageId="7" pageNumber="191">Stage 38 (late)</emphasis>
</paragraph>
<paragraph id="F75B368A757FFF9FF999F958D446F955" blockId="7.[144,758,1675,1758]" pageId="7" pageNumber="191">The parietal and frontal are ossifying, as are the quadrate and the ceratobranchials.</paragraph>
<paragraph id="F75B368A757FFF9FF999F90CD76DF899" blockId="7.[144,759,1789,1903]" box="[144,258,1789,1810]" pageId="7" pageNumber="191">
<emphasis id="C590EA98757FFF9FF999F90CD76DF899" box="[144,258,1789,1810]" italics="true" pageId="7" pageNumber="191">Stage 40+</emphasis>
</paragraph>
<paragraph id="F75B368A757FFF9FF999F8EDD204FC91" blockId="7.[144,759,1789,1903]" lastBlockId="7.[806,1421,588,794]" pageId="7" pageNumber="191">
<emphasis id="C590EA98757FFF9FF999F8EDD686F8BA" box="[144,233,1820,1841]" italics="true" pageId="7" pageNumber="191">Day 22:</emphasis>
The supraoccipital is ossifying from a single centre. The parasphenoid alae are ossifying, as is the basisphenoid. The lacrimal is triradiate. Although the nasal lacks a descending process, the ascending process of the maxilla in
<taxonomicName id="30E44D09757FFF9FFD4FFD9AD2B0FD0B" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[1094,1247,619,640]" class="Aves" family="Rheidae" genus="Rhea" kingdom="Animalia" order="Rheiformes" pageId="7" pageNumber="191" phylum="Chordata" rank="species" species="americana">
<emphasis id="C590EA98757FFF9FFD4FFD9AD2B0FD0B" box="[1094,1247,619,640]" italics="true" pageId="7" pageNumber="191">R. americana</emphasis>
</taxonomicName>
is much better developed than in
<taxonomicName id="30E44D09757FFF9FFAF0FD7BD2BAFD15" baseAuthorityName="Latham" baseAuthorityYear="1790" box="[1017,1237,649,671]" class="Aves" family="Dromaiidae" genus="Dromaius" kingdom="Animalia" order="Casuariiformes" pageId="7" pageNumber="191" phylum="Chordata" rank="species" species="novaehollandiae">
<emphasis id="C590EA98757FFF9FFAF0FD7BD2BAFD15" box="[1017,1237,649,671]" italics="true" pageId="7" pageNumber="191">D. novaehollandiae</emphasis>
</taxonomicName>
. There is a hole located in the middle of the squamosal, perhaps because of osteological restructuring caused by muscle development, as hypothesized for
<taxonomicName id="30E44D09757FFF9FFC17FD14D208FC91" authority="(Maxwell, 2008 a)" baseAuthorityName="Maxwell" baseAuthorityYear="2008" class="Aves" family="Phasianidae" genus="Meleagris" kingdom="Animalia" order="Galliformes" pageId="7" pageNumber="191" phylum="Chordata" rank="species" species="gallopavo">
<emphasis id="C590EA98757FFF9FFC17FD14D5FCFC92" italics="true" pageId="7" pageNumber="191">Meleagris gallopavo</emphasis>
(
<bibRefCitation id="93754B7B757FFF9FFAACFCF5D20FFC91" author="Maxwell EE" box="[933,1120,772,794]" pageId="7" pageNumber="191" pagination="242 - 257" refId="ref10387" refString="Maxwell EE. 2008 a. Comparative embryonic development of the skeleton of the domestic turkey (Meleagris gallopavo) and other galliform birds. Zoology 111: 242 - 257." type="journal article" year="2008">Maxwell, 2008a</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
<paragraph id="F75B368A757FFF9FFA2FFCB4D5D3FBBB" blockId="7.[806,1421,836,1072]" pageId="7" pageNumber="191">
<emphasis id="C590EA98757FFF9FFA2FFCB4D59DFCD2" box="[806,1010,836,858]" italics="true" pageId="7" pageNumber="191">
Day 26 (
<figureCitation id="6FDF2A0F757FFF9FFA9AFCB5D58EFCD2" box="[915,993,836,857]" captionStart="Figure 1" captionStartId="7.[144,223,198,217]" captionText="Figure 1. Lateral view of the skull of palaeognath embryos. A–C, Dromaius novaehollandiae. A, stage 34 (day 22 of incubation, RM 8020); B, stage 38 (day 32 of incubation, RM 8030); C, stage 40+ (day 43 of incubation, RM 8039). D–E, Eudromia elegans. D, day 11 of incubation (YPM 112520); E, day 15 of incubation (YPM 112525). F–H, Struthio camelus. F, day 15 of incubation (YPM 112437); G, day 23 of incubation (YPM 112448); H, day 35 of incubation (YPM 112465). I–J, Rhea americana. I, stage 34 (day 14 of incubation, RM 72172); J, stage 40+ (day 26 of incubation, RM 7223). Grey shaded regions represent cartilage; black regions represent ossified tissue. The density of stippling reflects the relative degree of ossification. Scale bars = 5 mm. Abbreviations: ac, auditory capsule; ang, angular; d, dentary; f, frontal; ip, infrapolar process; j, jugal; lac, lacrimal; m, Meckel’s cartilage; me, mesethmoid; met, metotic cartilage; mx, maxilla; mx (pn), nasal process of the maxilla; n, nasal; n (pm), maxillary process of the nasal; oo, opisthotic; pa, parietal; pal, palatine; pao, planum antorbitale; pmx, premaxilla; pnp, prenasal process; poc, postorbital cartilage; pt, pterygoid; q, quadrate; qj; quadratojugal; sa, supra-angular; so, supraoccipital; soc, supraorbital cartilage; sq, squamosal." pageId="7" pageNumber="191">Fig. 1J</figureCitation>
):
</emphasis>
The basioccipital, exoccipital, laterosphenoid, prootic, opisthotic, epiotic, and mesethmoid are ossifying. The lamina dorsalis of the mesethmoid is ossifying from paired ossification centres, rather than from a single median centre. The prearticular has developed a second ossification centre, located along the posterior edge of the articular cartilage.
</paragraph>
<paragraph id="F75B368A757FFF9FFA2FFBADD356FB25" blockId="7.[806,1421,1115,1198]" pageId="7" pageNumber="191">
<emphasis id="C590EA98757FFF9FFA2FFBADD5ECFBFB" box="[806,899,1115,1137]" italics="true" pageId="7" pageNumber="191">Day 28:</emphasis>
The facial region of the skull has become extremely elongate. The articular is ossifying, and the dermal bones of the skull roof are in contact.
</paragraph>
<paragraph id="F75B368A757FFF9FFA2FFB28D2C9FA86" blockId="7.[806,1421,1241,1293]" pageId="7" pageNumber="191">
<emphasis id="C590EA98757FFF9FFA2FFB28D5EBFB65" box="[806,900,1241,1262]" italics="true" pageId="7" pageNumber="191">Day 30:</emphasis>
The opisthotic and epiotic have developed large lateral exposures (
<figureCitation id="6FDF2A0F757FFF9FFD4BFB06D2F5FA86" box="[1090,1178,1271,1293]" captionStart="Figure 2" captionStartId="8.[826,905,198,217]" captionTargetBox="[173,773,195,1791]" captionTargetId="figure-315@8.[168,776,193,1794]" captionTargetPageId="8" captionText="Figure 2. Palatal view of selected palaeognath embryos. A, B, Struthio camelus (modified from Parker, 1866). C, Rhea americana, stage 37 (day 17 of incubation, RM 7219). D, Rhea americana (modified from Müller, 1963). E, Dromaius novaehollandiae, stage 37 (day 28 of incubation, RM 8026). F, Eudromia elegans, day 14 of incubation (YPM 112524). Scale bars = 5 mm. Abbreviations: bo, basioccipital; exo, exoccipital; mx, maxilla; pal, palatine; pmx, premaxilla; psl, parasphenoid lamina; psr, parasphenoid rostrum; pt, pterygoid; q, quadrate; v, vomer." figureDoi="http://doi.org/10.5281/zenodo.10132793" httpUri="https://zenodo.org/record/10132793/files/figure.png" pageId="7" pageNumber="191">Fig. 2D</figureCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>
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