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<mods:titleInfo id="C4444849331632AEF4DF7ECFD9F0054A">
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<mods:title id="3DB043A802034F55439D6EDF2B761F78">A new dentition-based phylogeny of Litopterna (Mammalia: Placentalia) and ‘ archaic’ South American ungulates</mods:title>
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<mods:namePart id="780B31BD1A885A9E8D3FC014E702BBF2">Püschel, Hans P.</mods:namePart>
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<mods:affiliation id="965BAEE2CBFE86E5F0320052CEE55B7B">Millenium Nucleus Early Evolutionary Transitions of Mammals (EVOTEM), Red Palentológica U-Chile, Santiago, 7800003, Chile & School of GeoSciences, University of Edinburgh, Grant Institute, James Hutton Road, Edinburgh, EH 9 3 FE, United Kingdom</mods:affiliation>
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<mods:nameIdentifier id="1C5FA41D8A84500A5810D6A8B5B39547" type="email">hpuschelr@gmail.com</mods:nameIdentifier>
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<mods:namePart id="3A3D3F7B0E97452AF8D2F2EF334E83BD">Shelley, Sarah L.</mods:namePart>
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<mods:affiliation id="06345BAB95F3298983D445DC62A6CF49">School of GeoSciences, University of Edinburgh, Grant Institute, James Hutton Road, Edinburgh, EH 9 3 FE, United Kingdom</mods:affiliation>
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<mods:namePart id="75DB5082C8B967B325212D46F754B33B">Williamson, Thomas E.</mods:namePart>
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<mods:affiliation id="FFC2B8B720D15B27481860E2D58DE66B">New Mexico Museum of Natural History and Science, Albuquerque, NM 87104, United States</mods:affiliation>
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<mods:namePart id="A9183B2F8B22A19F8DFFACCAB8995A01">Perini, Fernando A.</mods:namePart>
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<mods:affiliation id="75DAB92D587FC39A042D70C4A0279408">Instituto de Ciência Biológicas, UFMG, BH 31270 - 901, Brazil</mods:affiliation>
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<mods:namePart id="A509DC2BE60B180910E52D1665713B74">Wible, John R.</mods:namePart>
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<mods:affiliation id="1501E5930B764C8CD879B78B4FC6C561">Section of Mammals, Carnegie Museum of Natural History, 5800 Baum Boulevard, Pittsburgh, PA 15206, United States</mods:affiliation>
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<mods:namePart id="6B723F1FA18F1A7EF1542137C754AB6C">Brusatte, Stephen L.</mods:namePart>
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<mods:affiliation id="607CC022013F819C8C98E35C7D47D83D">School of GeoSciences, University of Edinburgh, Grant Institute, James Hutton Road, Edinburgh, EH 9 3 FE, United Kingdom</mods:affiliation>
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<mods:typeOfResource id="464C8F4AA043CD67BF1291724AC88ECA">text</mods:typeOfResource>
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<mods:title id="9C959D74835D2E21DD33C1A67DB4DF42">Zoological Journal of the Linnean Society</mods:title>
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<mods:date id="2A7D23EAF86CB8EF84B0A18ADCBBE59C">2024</mods:date>
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<mods:number id="AAE93538A9219059B47BCAF0B8D1F954">2024-09-07</mods:number>
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<mods:url id="AB4B3A4943B8921E39C1E32DFFD0F391">http://dx.doi.org/10.1093/zoolinnean/zlae095</mods:url>
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<treatment id="03B9B735FFE90D10C7E1FF2E88DCF99D" LSID="urn:lsid:plazi:treatment:03B9B735FFE90D10C7E1FF2E88DCF99D" httpUri="http://treatment.plazi.org/id/03B9B735FFE90D10C7E1FF2E88DCF99D" lastPageNumber="16" pageId="15" pageNumber="16">
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<subSubSection id="C30A55A8FFE90D10C7E1FF2E8BBAFEB3" box="[227,648,248,274]" pageId="15" pageNumber="16" type="nomenclature">
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<paragraph id="8BAF0623FFE90D10C7E1FF2E8BBAFEB3" blockId="15.[227,648,248,274]" box="[227,648,248,274]" pageId="15" pageNumber="16">
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<emphasis id="B964DA31FFE90D10C7E1FF2E8BBAFEB3" box="[227,648,248,274]" italics="true" pageId="15" pageNumber="16">
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<taxonomicName id="4C107DA0FFE90D10C7E1FF2E88AAFEB3" box="[227,408,248,274]" class="Mammalia" family="Amilnedwardsiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="family">Amilnedwardsiidae</taxonomicName>
|
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and the order
|
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<taxonomicName id="4C107DA0FFE90D10C519FF2F8BBAFEB3" authorityName="Soria" authorityYear="1989" box="[539,648,249,274]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="order">Notopterna</taxonomicName>
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</emphasis>
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</paragraph>
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<subSubSection id="C30A55A8FFE90D10C773FEC988DCF99D" pageId="15" pageNumber="16" type="multiple">
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<paragraph id="8BAF0623FFE90D10C773FEC988A1FBE7" blockId="15.[112,764,287,1596]" pageId="15" pageNumber="16">
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<taxonomicName id="4C107DA0FFE90D10C773FEC98805FE96" box="[113,311,287,311]" class="Mammalia" family="Amilnedwardsiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="family">Amilnedwardsiidae</taxonomicName>
|
||||
was proposed preliminarily by Soria (1984b) and better justified in a later article (Soria 1989a). It is based on two genera (i.e.,
|
||||
<emphasis id="B964DA31FFE90D10C687FE888B25FED7" box="[389,535,350,374]" italics="true" pageId="15" pageNumber="16">Amilnedwarsia</emphasis>
|
||||
<bibRefCitation id="EF817BD2FFE90D10C522FE888BFEFED7" author="Ameghino" box="[544,716,350,374]" firstAuthor="Ameghino" pageId="15" pageNumber="16" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation>
|
||||
and
|
||||
<taxonomicName id="4C107DA0FFE90D10C773FEA88890FE34" authority="Ameghino, 1901" authorityName="Ameghino" authorityYear="1901" box="[113,418,381,405]" class="Mammalia" family="Amilnedwardsiidae" genus="Rutimeyeria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="genus">
|
||||
<emphasis id="B964DA31FFE90D10C773FEA889DAFE34" box="[113,232,382,405]" italics="true" pageId="15" pageNumber="16">Rutimeyeria</emphasis>
|
||||
<bibRefCitation id="EF817BD2FFE90D10C7F1FEAB8890FE34" author="Ameghino" box="[243,418,381,405]" firstAuthor="Ameghino" pageId="15" pageNumber="16" pagination="349 - 426" refId="ref39297" refString="Ameghino F. Notices preliminaires sur des ongules nouveaux des terrains cretaces de Patagonie. Boletin de la Academia de Ciencias de Cordoba 1901; 16: 349 - 426." type="journal article" year="1901">Ameghino, 1901</bibRefCitation>
|
||||
</taxonomicName>
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||||
) and three species named after small bunodont isolated upper molars whose positions were assigned tentatively by Soria (1989a) by comparing them with the indaleciid
|
||||
<taxonomicName id="4C107DA0FFE90D10C606FE0D88FAFE52" authorityName="Bond & Vucetich" authorityYear="1983" box="[260,456,475,499]" class="Mammalia" family="Adianthidae" genus="Indalecia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="grandensis">
|
||||
<emphasis id="B964DA31FFE90D10C606FE0D88FAFE52" box="[260,456,475,499]" italics="true" pageId="15" pageNumber="16">Indalecia grandensis</emphasis>
|
||||
</taxonomicName>
|
||||
.
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||||
<taxonomicName id="4C107DA0FFE90D10C6D6FE0D8BA8FE52" box="[468,666,475,499]" class="Mammalia" family="Amilnedwardsiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="family">Amilnedwardsiidae</taxonomicName>
|
||||
was considered to be the most basal family of the order
|
||||
<taxonomicName id="4C107DA0FFE90D10C580FE2D89D9FD93" authority="Soria, 1989" authorityName="Soria" authorityYear="1989" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="order">Notopterna Soria, 1989</taxonomicName>
|
||||
, which also includes the families
|
||||
<taxonomicName id="4C107DA0FFE90D10C54DFDCC8BF9FD93" baseAuthorityName="Bond & Vucetich" baseAuthorityYear="1983" box="[591,715,538,562]" class="Mammalia" family="Indaleciidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="16" phylum="Chordata" rank="family">Indaleciidae</taxonomicName>
|
||||
and
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||||
<taxonomicName id="4C107DA0FFE90D10C773FDEF888BFDF0" authority="(Soria 1989 b)" baseAuthorityName="Soria" baseAuthorityYear="1989" box="[113,441,569,593]" class="Mammalia" family="Notonychopidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="16" phylum="Chordata" rank="family">Notonychopidae (Soria 1989b)</taxonomicName>
|
||||
.
|
||||
<taxonomicName id="4C107DA0FFE90D10C6C4FDEF8B0CFDF1" authorityName="Soria" authorityYear="1989" box="[454,574,569,592]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="order">Notopterna</taxonomicName>
|
||||
is diagnosed by a well-developed and buccally projected parastyle and the absence of a mesostyle in the upper molars, bicrescentic and selenodont lower molars, and anteriorly acuminate petrosals, among other anatomical features that distinguish them from other SANUs (Soria 1989a). Soria (1989a, b) argued that, considering their dental similarities, ‘amilnedwarsids’ could be the structural ancestor of both indaleciids and notonychopids. However, a phylogenetic analysis failed to recover the members of
|
||||
<taxonomicName id="4C107DA0FFE90D10C558FCE28BE0FCEA" authorityName="Soria" authorityYear="1989" box="[602,722,820,843]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="order">Notopterna</taxonomicName>
|
||||
as a monophyletic group (
|
||||
<bibRefCitation id="EF817BD2FFE90D10C656FC858B4FFCCA" author="Bonaparte and Morales" box="[340,637,851,875]" firstAuthor="Bonaparte" pageId="15" pageNumber="16" pagination="263 - 74" refId="ref40092" refString="Bonaparte JF, Morales J. Un primitivo Notonychopidae (Litopterna) del Paleoceno inferior de Punta Peligro, Chubut, Argentina. Estudios Geologicos 1997; 53: 263 - 74." type="journal article" year="1997">Bonaparte and Morales 1997</bibRefCitation>
|
||||
), which has led to some authors to include its members tentatively within
|
||||
<taxonomicName id="4C107DA0FFE90D10C773FC4489ECFC0B" box="[113,222,914,938]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="order">Litopterna</taxonomicName>
|
||||
(e.g.,
|
||||
<bibRefCitation id="EF817BD2FFE90D10C622FC4488F8FC0B" author="Gelfo" box="[288,458,914,938]" etAl="et al." firstAuthor="Gelfo" pageId="15" pageNumber="16" pagination="285 - 91" refId="ref42550" refString="Gelfo JN, Lopez GM, Lorente M. Los ungulados arcaicos de America del Sur: ' Condylarthra' y Litopterna. Contribuciones Cientificas Del Museo Argentino de Ciencias Naturales ' Bernardino Rivadavia' 2016; 6: 285 - 91." type="journal article" year="2016">
|
||||
Gelfo
|
||||
<emphasis id="B964DA31FFE90D10C65DFC4488A2FC0B" box="[351,400,914,938]" italics="true" pageId="15" pageNumber="16">et al.</emphasis>
|
||||
2016
|
||||
</bibRefCitation>
|
||||
, Croft
|
||||
<emphasis id="B964DA31FFE90D10C514FC448B75FC0B" box="[534,583,914,938]" italics="true" pageId="15" pageNumber="16">et al.</emphasis>
|
||||
2020,
|
||||
<bibRefCitation id="EF817BD2FFE90D10C58CFC448997FC68" author="Goin" etAl="et al." firstAuthor="Goin" pageId="15" pageNumber="16" pagination="1029 - 53" refId="ref42920" refString="Goin FJ, Gelfo JN, Ortiz-Jaureguizar E, et al. Mamiferos del Banco Negro Inferior, Formacion Salamanca, Cuenca del Golfo de San Jorge. In: Giacosa RE (ed.), Geologia y Recursos Naturales de la Provincia de Chubut. Puerto Madryn, Relatorio del XXI Congreso Geologico Argentino, 2022, 1029 - 53." type="book chapter" year="2022">
|
||||
Goin
|
||||
<emphasis id="B964DA31FFE90D10C5C8FC448BC9FC0B" box="[714,763,914,938]" italics="true" pageId="15" pageNumber="16">et al.</emphasis>
|
||||
2022
|
||||
</bibRefCitation>
|
||||
). However, this is a problematic assumption as
|
||||
<bibRefCitation id="EF817BD2FFE90D10C593FC678873FC49" author="Bonaparte and Morales" firstAuthor="Bonaparte" pageId="15" pageNumber="16" pagination="263 - 74" refId="ref40092" refString="Bonaparte JF, Morales J. Un primitivo Notonychopidae (Litopterna) del Paleoceno inferior de Punta Peligro, Chubut, Argentina. Estudios Geologicos 1997; 53: 263 - 74." type="journal article" year="1997">Bonaparte and Morales (1997)</bibRefCitation>
|
||||
used a matrix that includes only litopterns, so the broader relationships of notopterns have not yet been tested in an analysis that includes other SANUs such as notoungulates and astrapotheres.
|
||||
</paragraph>
|
||||
<paragraph id="8BAF0623FFE90D10C78FFB988B46FA3E" blockId="15.[112,764,287,1596]" pageId="15" pageNumber="16">
|
||||
<emphasis id="B964DA31FFE90D10C78FFB988811FBC7" box="[141,291,1102,1126]" italics="true" pageId="15" pageNumber="16">Amilnedwardia</emphasis>
|
||||
and
|
||||
<taxonomicName id="4C107DA0FFE90D10C655FB9888FCFBC4" authorityName="Ameghino" authorityYear="1901" box="[343,462,1102,1125]" class="Mammalia" family="Amilnedwardsiidae" genus="Rutimeyeria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="genus">
|
||||
<emphasis id="B964DA31FFE90D10C655FB9888FCFBC4" box="[343,462,1102,1125]" italics="true" pageId="15" pageNumber="16">Rutimeyeria</emphasis>
|
||||
</taxonomicName>
|
||||
were previously referred tentatively to the family
|
||||
<taxonomicName id="4C107DA0FFE90D10C64EFBBB88CEFB24" box="[332,508,1133,1157]" class="Mammalia" family="Macraucheniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="family">Macraucheniidae</taxonomicName>
|
||||
and regarded as close to
|
||||
<taxonomicName id="4C107DA0FFE90D10C771FB5A8837FB05" box="[115,261,1164,1188]" class="Mammalia" family="Proterotheriidae" genus="Victorlemoinea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="genus">
|
||||
<emphasis id="B964DA31FFE90D10C771FB5A8837FB05" box="[115,261,1164,1188]" italics="true" pageId="15" pageNumber="16">Victorlemoinea</emphasis>
|
||||
</taxonomicName>
|
||||
(Simpson 1945, 1948), although the latter now is now considered to be part of the family
|
||||
<taxonomicName id="4C107DA0FFE90D10C511FB7A8BC5FB65" box="[531,759,1196,1220]" class="Mammalia" family="Sparnotheriodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Notoungulata" pageId="15" pageNumber="16" phylum="Chordata" rank="family">Sparnotheriodontidae</taxonomicName>
|
||||
. Cifelli (1983a)considered referred specimens of these taxa to be too fragmentary to determine a clear affinity,but suggested the referred specimens represent deciduous teeth of a henricosborniid notoungulate based on the presence of a metaconule-derived crest connecting with the ectoloph wall. Indeed, probably due to their limited fossil record, thus far no members of this family have ever been included in a phylogenetic analysis.
|
||||
</paragraph>
|
||||
<paragraph id="8BAF0623FFE90D10C78FFA7088DCF99D" blockId="15.[112,764,287,1596]" pageId="15" pageNumber="16">
|
||||
If we consider
|
||||
<taxonomicName id="4C107DA0FFE90D10C633FA7088C5FA1F" box="[305,503,1446,1470]" class="Mammalia" family="Amilnedwardsiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Litopterna" pageId="15" pageNumber="16" phylum="Chordata" rank="family">Amilnedwardsiidae</taxonomicName>
|
||||
to be a valid family, as its three species all come from Patagonian fossil localities of a similar age that correlate with the Vacan SALMA, the temporal interval of this family is ~45–42 Mya (
|
||||
<figureCitation id="132B1AA6FFE90D10C6F6F9D28B09F9BD" box="[500,571,1540,1564]" captionStart="Figure 2" captionStartId="7.[113,178,1849,1873]" captionTargetBox="[174,1395,148,1822]" captionTargetId="graphics-9@7.[268,1359,583,1798]" captionTargetPageId="7" captionText="Figure 2. Diversity of litoptern and associated early SANU families expressed as the number of genera.A, pie chart expressing the relative diversity of each family. B, diversity over time of the different families of interest with colours corresponding to the families listed in (A). C, same as (B) but using a smoothing function that averages the different occurrences aiding the eye in seeing patterns. In (A) the number of genera for each family is indicated next to the name between brackets.In (C) the function geom_smooth from the R package ggplot v.3.4, method = loess and span = 0.5 was used.In (B) and (C) apart from the standard Periods, Epochs, and Ages below, the SALMAs are indicated in grey boxes crossing the plots. The Tiupampan and Peligran SALMAs and the Bonaerian and Lujanian SALMAs are taken as only two temporal units instead of four for the plots. The data on taxa and occurrences were taken from Supporting information, Table S1, and the temporal information on SALMAs were taken from Supporting information, File S6. Note that the two genera of ‘amilnedwarsids’ were not included in this plot.Note that in the Discussion section, we proposed to redefine the families Anisolambdidae and Sparnotheriodontidae as the subfamilies Anisolambdinae and Sparnotheriodontinae of the family Anisolambdidae. Abbreviations: Ba, Barrancan; Bo/Lu, Bonaerian and Lujanian; Cc, Colloncuran; Cd, Carodnia Zone; Ch, Chasicoan; Co, Colhuehuapian; Cp, Chapadmalalan; De, Deseadean; En, Ensenadan; Hu, Huayquerian; It, Itaboraian; La, Laventan; Ma, Marplatan; Mo, Montehermosan; Mu, Mustersan; Ri, Riochican; Sa, Sapoan; San, Santacrucian; Tg, Tinguirirican; Ti/Pe, Tiupampan and Peligran; Va, Vacan." pageId="15" pageNumber="16">Fig. 2B</figureCitation>
|
||||
; Supporting information,
|
||||
<tableCitation id="C6923398FFE90D10C7C7F9F2882DF99D" box="[197,287,1572,1596]" captionStart="Table 1" captionStartId="2.[110,165,500,525]" captionText="Table 1. Summary of the main taxonomic proposals for early SANUs (‘condylarths’), Litopterna, and Notopterna. Abbreviations: SF, Superfamily; SO, Suborder" pageId="15" pageNumber="16">Table S1</tableCitation>
|
||||
;
|
||||
<bibRefCitation id="EF817BD2FFE90D10C628F9F288EFF99D" author="Krause" box="[298,477,1572,1596]" etAl="et al." firstAuthor="Krause" pageId="15" pageNumber="16" pagination="886 - 903" refId="ref43716" refString="Krause JM, Clyde WC, Ibanez-Mejia M et al. New age constraints for Early Paleogene strata of central Patagonia, Argentina: implications for the timing of South American Land Mammal Ages. Geological Society of America Bulletin 2017; 129: 886 - 903. https: // doi. org / 10.1130 / b 31561.1" type="journal article" year="2017">
|
||||
Krause
|
||||
<emphasis id="B964DA31FFE90D10C674F9F28897F99D" box="[374,421,1572,1596]" italics="true" pageId="15" pageNumber="16">et al.</emphasis>
|
||||
2017
|
||||
</bibRefCitation>
|
||||
).
|
||||
</paragraph>
|
||||
</subSubSection>
|
||||
</treatment>
|
||||
</document>
|
||||
1126
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Genus
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|
||||
<emphasis id="D2B9EA99FFCDA777FE94FCDFFE6DFCEA" bold="true" box="[365,447,841,867]" italics="true" pageId="6" pageNumber="7">Spongia</emphasis>
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||||
<bibRefCitation id="845C4B7AFFCDA777FE3DFCDFFDB5FCEA" author="Linnaeus" box="[452,615,841,867]" firstAuthor="Linnaeus" pageId="6" pageNumber="7" pagination="825 - 1384" refId="ref17828" refString="Linnaeus C. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus II, Editio decima, reformata, Holmiae. (L. Salvii) 1759; 825 - 1384. https: // www. biodiversitylibrary. org / page / 587267" type="book chapter" year="1759">Linnaeus, 1759</bibRefCitation>
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</taxonomicName>
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||||
</emphasis>
|
||||
</heading>
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||||
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||||
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<paragraph id="E072368BFFCDA777FF78FC1CFD8AFC28" blockId="6.[129,600,905,929]" box="[129,600,905,929]" pageId="6" pageNumber="7">
|
||||
<emphasis id="D2B9EA99FFCDA777FF78FC1CFF2EFC28" box="[129,252,906,929]" italics="true" pageId="6" pageNumber="7">Type species:</emphasis>
|
||||
<taxonomicName id="27CD4D08FFCDA777FEF8FC1CFD86FC28" authority="Linnaeus, 1759" authorityName="Linnaeus" authorityYear="1759" box="[257,596,905,929]" class="Demospongiae" family="Spongiidae" genus="Spongia" kingdom="Animalia" order="Dictyoceratida" pageId="6" pageNumber="7" phylum="Porifera" rank="species" species="officinalis">
|
||||
<emphasis id="D2B9EA99FFCDA777FEF8FC1CFE62FC28" box="[257,432,905,929]" italics="true" pageId="6" pageNumber="7">Spongia officinalis</emphasis>
|
||||
<bibRefCitation id="845C4B7AFFCDA777FE4CFC1FFD86FC28" author="Linnaeus" box="[437,596,905,929]" firstAuthor="Linnaeus" pageId="6" pageNumber="7" pagination="825 - 1384" refId="ref17828" refString="Linnaeus C. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus II, Editio decima, reformata, Holmiae. (L. Salvii) 1759; 825 - 1384. https: // www. biodiversitylibrary. org / page / 587267" type="book chapter" year="1759">Linnaeus, 1759</bibRefCitation>
|
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</taxonomicName>
|
||||
.
|
||||
</paragraph>
|
||||
</subSubSection>
|
||||
<subSubSection id="A8D76500FFCDA777FF78FC55FD66FC73" pageId="6" pageNumber="7" type="description">
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<paragraph id="E072368BFFCDA777FF78FC55FD66FC73" blockId="6.[129,777,963,1018]" pageId="6" pageNumber="7">
|
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<emphasis id="D2B9EA99FFCDA777FF78FC55FF3BFC52" box="[129,233,963,987]" italics="true" pageId="6" pageNumber="7">Definition:</emphasis>
|
||||
Unarmoured spongiids, not heavily lacunose, with simple primary fibres (from Cook and Bergquist 2002).
|
||||
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|
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85
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Bone histology and internal structure of vertebrae of
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|
||||
The CT examination of vertebrae was performed to nondestructively investigate the internal structure of each specimen and to assess the thickness of compact and cancellous bone in the different regions of the vertebrae. Within the anteroposteriorly terminal portions of all scanned ‘
|
||||
<emphasis id="D0E0BC47066DFF9054D4FD7761D26574" box="[470,484,748,771]" italics="true" pageId="5" pageNumber="6">P</emphasis>
|
||||
’.
|
||||
<taxonomicName id="25941BD6066DFF9054F4FD77627C6574" baseAuthorityName="Huene" baseAuthorityYear="1905" box="[502,586,748,771]" family="Tanystropheidae" genus="Protanystropheus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="antiquus">
|
||||
<emphasis id="D0E0BC47066DFF9054F4FD77627C6574" box="[502,586,748,771]" italics="true" pageId="5" pageNumber="6">antiquus</emphasis>
|
||||
</taxonomicName>
|
||||
centra, the core was filled with dense cancellous bone (identified in CT images so of uncertain tissue composition) and the cortex was relatively thin (
|
||||
<figureCitation id="7AAF7CD0066DFF9055F3FCD2617A6516" box="[241,332,841,865]" captionStart-0="Figure 2" captionStart-1="Figure 3" captionStart-2="Figure 4" captionStartId-0="3.[113,178,975,999]" captionStartId-1="6.[129,194,1669,1693]" captionStartId-2="7.[113,178,1841,1865]" captionTargetBox-0="[307,1266,157,946]" captionTargetBox-1="[305,1297,144,1642]" captionTargetBox-2="[175,1398,149,1813]" captionTargetId-0="figure-579@3.[305,1268,144,948]" captionTargetId-1="figure-103@6.[305,1297,144,1642]" captionTargetId-2="figure-6@7.[175,1398,149,1813]" captionTargetPageId-0="3" captionTargetPageId-1="6" captionTargetPageId-2="7" captionText-0="Figure 2. GIUS-7-3674, cervical of ‘P’.antiquus, surface model in left lateral view (A) and a longitudinal CT cross section (B). Notice the thin, but generally continuous neural canal floor.Anatomical abbreviations: ic—internal cavity; la—lamella that separates the neural canal from the internal cavity; nc—neural canal; ns—neural spine; poz—postzygapophysis; prz—prezygapophysis." captionText-1="Figure 3. CT transverse cross sections (from anterior to posterior) of ‘P’.antiquus cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum." captionText-2="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." pageId="5" pageNumber="6">Figs 2–4</figureCitation>
|
||||
), whereas in the middle portion, where the internal cavity was revealed, the cortical bone was distinctly thicker. The transverse sections revealed the changes of the cortical bone thickness and the shape of internal cavities, with the mid-centrum being roughly cylindrical. The cortex was thinner near the anteroposteriorly terminal portions of the vertebrae and its thickness gradually increased up to approximately four times the original thickness (about 10% of bone diameter), near the midpoint (
|
||||
<figureCitation id="7AAF7CD0066DFF90540AFBDF6176622B" box="[264,320,1092,1116]" captionStart="Figure 3" captionStartId="6.[129,194,1669,1693]" captionTargetBox="[305,1297,144,1642]" captionTargetId="figure-103@6.[305,1297,144,1642]" captionTargetPageId="6" captionText="Figure 3. CT transverse cross sections (from anterior to posterior) of ‘P’.antiquus cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum." pageId="5" pageNumber="6">Fig. 3</figureCitation>
|
||||
). The density of the cancellous bone of the anteroposteriorly terminal portions of the centrum decreased medially, with larger cavities being present closer to the midpoint. The transition to the empty internal cavity was relatively sharp. The anterior and posterior ends of the internal cavity were roughly hemispherical and surrounded circumferentially by marginalized trabeculae (
|
||||
<figureCitation id="7AAF7CD0066DFF905478FA9B6183636F" box="[378,437,1280,1304]" captionStart="Figure 3" captionStartId="6.[129,194,1669,1693]" captionTargetBox="[305,1297,144,1642]" captionTargetId="figure-103@6.[305,1297,144,1642]" captionTargetPageId="6" captionText="Figure 3. CT transverse cross sections (from anterior to posterior) of ‘P’.antiquus cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum." pageId="5" pageNumber="6">Fig. 3</figureCitation>
|
||||
). Long intermittent trabeculae composed of parallel-fibred bone moderately remodelled with endosteal secondary lamellar bone (
|
||||
<figureCitation id="7AAF7CD0066DFF9054DFFAA462106320" box="[477,550,1343,1367]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." pageId="5" pageNumber="6">Fig. 4D</figureCitation>
|
||||
) crossed the internal cavity in varied directions, mainly diagonally. In some vertebrae they divided the internal cavity into several smaller terminus-like pockets (
|
||||
<figureCitation id="7AAF7CD0066DFF9055D1FA06611763C2" box="[211,289,1437,1461]" captionStart="Figure 2" captionStartId="3.[113,178,975,999]" captionTargetBox="[307,1266,157,946]" captionTargetId="figure-579@3.[305,1268,144,948]" captionTargetPageId="3" captionText="Figure 2. GIUS-7-3674, cervical of ‘P’.antiquus, surface model in left lateral view (A) and a longitudinal CT cross section (B). Notice the thin, but generally continuous neural canal floor.Anatomical abbreviations: ic—internal cavity; la—lamella that separates the neural canal from the internal cavity; nc—neural canal; ns—neural spine; poz—postzygapophysis; prz—prezygapophysis." pageId="5" pageNumber="6">Figs 2B</figureCitation>
|
||||
,
|
||||
<figureCitation id="7AAF7CD0066DFF90542DFA06610A63C2" box="[303,316,1437,1461]" captionStart="Figure 3" captionStartId="6.[129,194,1669,1693]" captionTargetBox="[305,1297,144,1642]" captionTargetId="figure-103@6.[305,1297,144,1642]" captionTargetPageId="6" captionText="Figure 3. CT transverse cross sections (from anterior to posterior) of ‘P’.antiquus cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum." pageId="5" pageNumber="6">3</figureCitation>
|
||||
). The neural canal was separated ventrally from the internal cavity by a horizontal, plate-like lamella consisting of endosteal lamellar bone (
|
||||
<figureCitation id="7AAF7CD0066DFF9054D0FA40622A6384" box="[466,540,1499,1523]" captionStart="Figure 2" captionStartId="3.[113,178,975,999]" captionTargetBox="[307,1266,157,946]" captionTargetId="figure-579@3.[305,1268,144,948]" captionTargetPageId="3" captionText="Figure 2. GIUS-7-3674, cervical of ‘P’.antiquus, surface model in left lateral view (A) and a longitudinal CT cross section (B). Notice the thin, but generally continuous neural canal floor.Anatomical abbreviations: ic—internal cavity; la—lamella that separates the neural canal from the internal cavity; nc—neural canal; ns—neural spine; poz—postzygapophysis; prz—prezygapophysis." pageId="5" pageNumber="6">Figs 2B</figureCitation>
|
||||
,
|
||||
<figureCitation id="7AAF7CD0066DFF905725FA4062026384" box="[551,564,1499,1523]" captionStart="Figure 3" captionStartId="6.[129,194,1669,1693]" captionTargetBox="[305,1297,144,1642]" captionTargetId="figure-103@6.[305,1297,144,1642]" captionTargetPageId="6" captionText="Figure 3. CT transverse cross sections (from anterior to posterior) of ‘P’.antiquus cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum." pageId="5" pageNumber="6">3</figureCitation>
|
||||
,
|
||||
<figureCitation id="7AAF7CD0066DFF90573CFA40624A6384" box="[574,636,1499,1523]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." pageId="5" pageNumber="6">4A–C</figureCitation>
|
||||
), which was generally thinner than the internal trabeculae composed of the same tissue. This separation was not continuous, the neural canal floor was perforated by large openings in some specimens (
|
||||
<figureCitation id="7AAF7CD0066DFF9057D2F9A260B86006" captionStart="Figure 2" captionStartId="3.[113,178,975,999]" captionTargetBox="[307,1266,157,946]" captionTargetId="figure-579@3.[305,1268,144,948]" captionTargetPageId="3" captionText="Figure 2. GIUS-7-3674, cervical of ‘P’.antiquus, surface model in left lateral view (A) and a longitudinal CT cross section (B). Notice the thin, but generally continuous neural canal floor.Anatomical abbreviations: ic—internal cavity; la—lamella that separates the neural canal from the internal cavity; nc—neural canal; ns—neural spine; poz—postzygapophysis; prz—prezygapophysis." pageId="5" pageNumber="6">Figs 2B</figureCitation>
|
||||
and
|
||||
<figureCitation id="7AAF7CD0066DFF9055C1F9C261326006" box="[195,260,1625,1649]" captionStart="Figure 3" captionStartId="6.[129,194,1669,1693]" captionTargetBox="[305,1297,144,1642]" captionTargetId="figure-103@6.[305,1297,144,1642]" captionTargetPageId="6" captionText="Figure 3. CT transverse cross sections (from anterior to posterior) of ‘P’.antiquus cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum." pageId="5" pageNumber="6">3C–G</figureCitation>
|
||||
) and seemed to disappear completely in others. For example, in MGUWr 3889s at least two large openings were present in the lamella separating the neural canal from the internal cavity. In MGUWr 3902 and GPIH 5194c there appeared to be no bony separation between the neural canal and the internal cavity for most of the length of the centrum. Because no fragments of broken bone were present anywhere inside of the neural canal or the internal cavity of the centrum, and this condition was present in several specimens approximately in the same area, it appears not to be an artifact of preservation. The cortex was predominately composed of parallel-fibred matrix (
|
||||
<figureCitation id="7AAF7CD0066DFF9057B2F80962DF61DD" box="[688,745,1938,1962]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." pageId="5" pageNumber="6">Fig. 4</figureCitation>
|
||||
), with locally (especially in the dorsolateral part) highly organized arrays of mineralized collagen fibres. No rest lines were present. The vascularization was moderate, and its pattern was radial, which was especially evident in the ventral part of the vertebrae (
|
||||
<figureCitation id="7AAF7CD0066DFF905637FE9663E56752" box="[821,979,269,293]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." pageId="5" pageNumber="6">Fig. 4A–C, H–I</figureCitation>
|
||||
). Inside the neural arch, mostly dorsally to the neural canal (inside of the base of the neural spine) and in some specimens laterally to the neural canal there was a region of occurrence of secondary trabecular bone (
|
||||
<figureCitation id="7AAF7CD0066DFF9051C3FEF0653767F4" box="[1217,1281,363,387]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." pageId="5" pageNumber="6">Fig. 4J</figureCitation>
|
||||
). Both the cavity and the neural canal were clear cut and lined with a thin layer of endosteal lamellar bone (
|
||||
<figureCitation id="7AAF7CD0066DFF905137FE3164A067B5" box="[1077,1174,426,450]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." pageId="5" pageNumber="6">Fig. 4E–J</figureCitation>
|
||||
). Sharpey’s fibres could be seen extending throughout the cortex, especially in the dorsolateral and ventrolateral regions of the centrum (
|
||||
<figureCitation id="7AAF7CD0066DFF905017FE7265A56476" box="[1301,1427,489,513]" captionStart="Figure 4" captionStartId="7.[113,178,1841,1865]" captionTargetBox="[175,1398,149,1813]" captionTargetId="figure-6@7.[175,1398,149,1813]" captionTargetPageId="7" captionText="Figure 4. Thin sections of ‘P’.antiquus cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light.Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone.Scale bars for panels A–C equal 5 mm and 0.5 mm for D–J." pageId="5" pageNumber="6">Fig. 4C, G–I</figureCitation>
|
||||
).
|
||||
</paragraph>
|
||||
</subSubSection>
|
||||
</treatment>
|
||||
</document>
|
||||
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<document id="F432FB799579E2FB39BD36A8C6FE82B9" ID-DOI="10.1093/zoolinnean/zlae126" ID-ISSN="0024-4082" IM.bibliography_approvedBy="jonas" IM.illustrations_approvedBy="jonas" IM.materialsCitations_approvedBy="jonas" IM.metadata_approvedBy="jonas" IM.taxonomicNames_approvedBy="jonas" IM.treatments_approvedBy="jonas" checkinTime="1732850508520" checkinUser="plazi" docAuthor="Rytel, Adam, Surmik, Dawid, Szczygielski, Tomasz, Spiekman, Stephan N. F., Kamp, Thomas van de, Zuber, Marcus & Scheyer, Torsten M." docDate="2024" docId="6A3DD143066DFF9F5659FDBB64A963F0" docLanguage="en" docName="zlae126.pdf" docOrigin="Zoological Journal of the Linnean Society 202 (3)" docSource="http://dx.doi.org/10.1093/zoolinnean/zlae126" docStyle="DocumentStyle:4F230B9370E98E256D973D6DFB57F36C.9:ZoolJLinnSoc.2023-.journal_article" docStyleId="4F230B9370E98E256D973D6DFB57F36C" docStyleName="ZoolJLinnSoc.2023-.journal_article" docStyleVersion="9" docTitle="Tanystropheus Meyer 1852" docType="treatment" docVersion="1" lastPageNumber="11" masterDocId="9604A93B0668FF955502FF9B60366677" masterDocTitle="Unique internal anatomy of vertebrae as a key factor for neck elongation in Triassic archosauromorphs" masterLastPageNumber="21" masterPageNumber="1" pageNumber="6" updateTime="1733770130157" updateUser="jonas">
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<mods:title id="55D904495AFD91E8CA7A072BCC06B489">Unique internal anatomy of vertebrae as a key factor for neck elongation in Triassic archosauromorphs</mods:title>
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<mods:namePart id="56CFBFCAF87A551F3A500120B66154D9">Rytel, Adam</mods:namePart>
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<mods:affiliation id="3BF3BA990BFA6838E1E2A660C51B397C">Institute of Paleobiology, Polish Academy of Sciences, Twarda 51 / 55, 00 - 818 Warsaw, Poland & Department of Paleontology, University of Zurich, Karl Schmid-Strasse 4, 8006 Zurich, Switzerland</mods:affiliation>
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<mods:namePart id="3582F54FD8BBF179D2E10C486B697652">Surmik, Dawid</mods:namePart>
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<mods:affiliation id="8033441A6ACCA03CB02CED969587F97B">Institute of Earth Sciences, Faculty of Natural Sciences, University of Silesia in Katowice, Bedzinska 60, 41 - 200 Sosnowiec, Poland</mods:affiliation>
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<mods:nameIdentifier id="C354CBA16E7C9F6D2FE985117B7E59A4" type="email">adam.rytel@twarda.pan.pl</mods:nameIdentifier>
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<mods:namePart id="224654880F8AC92EB17F9B07DD660DA4">Szczygielski, Tomasz</mods:namePart>
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<mods:affiliation id="4439AEC49F1053E4EACB3D65963E7710">Institute of Paleobiology, Polish Academy of Sciences, Twarda 51 / 55, 00 - 818 Warsaw, Poland</mods:affiliation>
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<mods:namePart id="B6173128E8E1271B47019658F5F323F0">Spiekman, Stephan N. F.</mods:namePart>
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<mods:affiliation id="EF9847D22673D85DD2FCC32DF202107E">Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, 70191 Stuttgart, Germany</mods:affiliation>
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<mods:namePart id="7F54C8F6CFE78D0ADF52892BC6B677A2">Kamp, Thomas van de</mods:namePart>
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<mods:affiliation id="DCC5A2B3BE76B4C011956D671286C640">Institute for Photon Science and Synchrotron Radiation (IPS), Karlsruhe Institute of Technology (KIT), Hermann-von-Helmholtz-Platz 1, 76344 Eggenstein-Leopoldshafen, Germany & Laboratory for Applications of Synchrotron Radiation (LAS), Karlsruhe Institute of Technology (KIT), Kaiserstrasse 12, 76131 Karlsruhe, Germany</mods:affiliation>
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<mods:namePart id="9A0539229FF64D0AC2206921D742FC3D">Zuber, Marcus</mods:namePart>
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<mods:affiliation id="B986845BF3A1947D3AEE18FFF0502287">Institute for Photon Science and Synchrotron Radiation (IPS), Karlsruhe Institute of Technology (KIT), Hermann-von-Helmholtz-Platz 1, 76344 Eggenstein-Leopoldshafen, Germany</mods:affiliation>
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<mods:namePart id="BA75F830DC7959299FF9131C29B27465">Scheyer, Torsten M.</mods:namePart>
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<mods:affiliation id="6E4AAD8C7BAB36FABF8F46AA5F90437B">Department of Paleontology, University of Zurich, Karl Schmid-Strasse 4, 8006 Zurich, Switzerland</mods:affiliation>
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<mods:title id="CE206F9964933AA6B66E0E2502D03E05">Zoological Journal of the Linnean Society</mods:title>
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<mods:date id="80DA6D0A824D49806C8810422822F87A">2024</mods:date>
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<mods:number id="499ED76663CDE25D6F885D1944DE75BE">202</mods:number>
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<mods:number id="0BB8F9DEE5C5A32AF03E81210B73E81A">3</mods:number>
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<mods:url id="389162C0128F93933CCB8AA5386CF43B">http://dx.doi.org/10.1093/zoolinnean/zlae126</mods:url>
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<mods:identifier id="5A286777B2052DBD3EFEABB7AE45A00C" type="DOI">10.1093/zoolinnean/zlae126</mods:identifier>
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<mods:identifier id="708E4364E656B19A89F71BC8A2FC8840" type="ISSN">0024-4082</mods:identifier>
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<treatment id="6A3DD143066DFF9F5659FDBB64A963F0" LSID="urn:lsid:plazi:treatment:6A3DD143066DFF9F5659FDBB64A963F0" httpUri="http://treatment.plazi.org/id/6A3DD143066DFF9F5659FDBB64A963F0" lastPageId="10" lastPageNumber="11" pageId="5" pageNumber="6">
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<subSubSection id="AA8E33DE066DFF905659FDBB64F9642D" pageId="5" pageNumber="6" type="nomenclature">
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<emphasis id="D0E0BC47066DFF905659FDBB64F9642D" bold="true" pageId="5" pageNumber="6">
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Bone histology and internal structure of vertebrae of
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<taxonomicName id="25941BD6066DFF90510CFDA464F9642D" authority="Meyer, 1852" authorityName="Meyer" authorityYear="1852" box="[1038,1231,575,602]" class="Reptilia" family="Tanystropheidae" genus="Tanystropheus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Protorosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
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<emphasis id="D0E0BC47066DFF90510CFDA464A9642D" bold="true" box="[1038,1183,575,602]" italics="true" pageId="5" pageNumber="6">Tanystropheus</emphasis>
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spp.
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</emphasis>
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</paragraph>
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<emphasis id="D0E0BC47066DFF9056DEFDF1653464F4" box="[988,1282,617,644]" italics="true" pageId="5" pageNumber="6">Anatomy of the middle cervicals</emphasis>
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</paragraph>
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<paragraph id="E22B6055066DFF9D5628FD0B62E063A9" blockId="5.[809,1463,617,1965]" lastBlockId="8.[128,779,1322,1753]" lastPageId="8" lastPageNumber="9" pageId="5" pageNumber="6">
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Within all of the studied vertebrae, a large cavity could be identified, surrounded by relatively thick cortex composed of alternating layers of well-organized parallel-fibred to lamellar bone (
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<figureCitation id="7AAF7CD0066DFF905637FD7563BD6571" box="[821,907,750,774]" captionStart-0="Figure 5" captionStart-1="Figure 6" captionStart-2="Figure 7" captionStart-3="Figure 8" captionStartId-0="8.[130,195,1138,1162]" captionStartId-1="9.[113,178,1191,1215]" captionStartId-2="10.[129,194,695,719]" captionStartId-3="11.[113,178,1841,1865]" captionTargetBox-0="[289,1313,144,1110]" captionTargetBox-1="[146,1426,144,1163]" captionTargetBox-2="[289,1313,144,667]" captionTargetBox-3="[224,1348,150,1812]" captionTargetId-0="figure-299@8.[289,1313,144,1110]" captionTargetId-1="figure-396@9.[146,1426,144,1163]" captionTargetId-2="figure-733@10.[289,1313,144,667]" captionTargetId-3="figure-6@11.[223,1349,149,1813]" captionTargetPageId-0="8" captionTargetPageId-1="9" captionTargetPageId-2="10" captionTargetPageId-3="11" captionText-0="Figure 5.Middle cervicals of Tanystropheus sp.from Miedary: A, B, D—ZPAL V.36/101; C, E, F—ZPAL V. 36/102; A is a surface model in left lateral view.B, C, longitudinal CT cross sections of an anterior (B) and a posterior portion (C) of the vertebra. D, E, F, transverse CT cross sections. Sectioning planes and walls of the neural canal and internal cavity have been outlined with white dashed lines, with the lines signifing the sectioning planes being thicker than the internal cavity outline. Anatomical abbreviations: ic—internal cavity; nc—neural canal; ns— neural spine; poz—postzygapophysis; prz—prezygapophysis." captionText-1="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." captionText-2="Figure 7. Internal cavity volume in the cervical vertebrae of ‘P’.antiquus MGUWr 3895s (A) and Tanystropheus sp. from Miedary ZPAL V. 36/181 (B). Anatomical abbreviation:ic—internal cavity. Images obtained using ‘xray’ shader in MeshLab v.2020.12 (Cignoni et al.2008)." captionText-3="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="5" pageNumber="6">Figs 5–8</figureCitation>
|
||||
). In the anterior and posterior terminal portions of the cervicals, discs of secondary cancellous bone of endosteal origin formed the articular regions of the centrum (
|
||||
<figureCitation id="7AAF7CD0066DFF9051F1FCB6656D6533" box="[1267,1371,812,837]" captionStart="Figure 5" captionStartId="8.[130,195,1138,1162]" captionTargetBox="[289,1313,144,1110]" captionTargetId="figure-299@8.[289,1313,144,1110]" captionTargetPageId="8" captionText="Figure 5.Middle cervicals of Tanystropheus sp.from Miedary: A, B, D—ZPAL V.36/101; C, E, F—ZPAL V. 36/102; A is a surface model in left lateral view.B, C, longitudinal CT cross sections of an anterior (B) and a posterior portion (C) of the vertebra. D, E, F, transverse CT cross sections. Sectioning planes and walls of the neural canal and internal cavity have been outlined with white dashed lines, with the lines signifing the sectioning planes being thicker than the internal cavity outline. Anatomical abbreviations: ic—internal cavity; nc—neural canal; ns— neural spine; poz—postzygapophysis; prz—prezygapophysis." pageId="5" pageNumber="6">Fig. 5B–D</figureCitation>
|
||||
). Above them, the openings of the neural canal were present. In some of the middle cervicals the neural canal floor extended only slightly from the openings towards the middle of the vertebra, forming a shelf that partially separated the internal cavity from the neural canal (
|
||||
<figureCitation id="7AAF7CD0066DFF905673FC5263ED6596" box="[881,987,969,993]" captionStart="Figure 5" captionStartId="8.[130,195,1138,1162]" captionTargetBox="[289,1313,144,1110]" captionTargetId="figure-299@8.[289,1313,144,1110]" captionTargetPageId="8" captionText="Figure 5.Middle cervicals of Tanystropheus sp.from Miedary: A, B, D—ZPAL V.36/101; C, E, F—ZPAL V. 36/102; A is a surface model in left lateral view.B, C, longitudinal CT cross sections of an anterior (B) and a posterior portion (C) of the vertebra. D, E, F, transverse CT cross sections. Sectioning planes and walls of the neural canal and internal cavity have been outlined with white dashed lines, with the lines signifing the sectioning planes being thicker than the internal cavity outline. Anatomical abbreviations: ic—internal cavity; nc—neural canal; ns— neural spine; poz—postzygapophysis; prz—prezygapophysis." pageId="5" pageNumber="6">Figs 5B–E</figureCitation>
|
||||
,
|
||||
<figureCitation id="7AAF7CD0066DFF9056EAFC5264146596" box="[1000,1058,969,993]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="5" pageNumber="6">6C, F</figureCitation>
|
||||
), although in some other of these vertebrae the neural canal transitioned smoothly into the internal cavity (compare
|
||||
<figureCitation id="7AAF7CD0066DFF9056DBFB9364156257" box="[985,1059,1032,1056]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="5" pageNumber="6">Fig. 6B</figureCitation>
|
||||
to
|
||||
<figureCitation id="7AAF7CD0066DFF90514BFB9364A26257" box="[1097,1172,1032,1056]" captionStart="Figure 5" captionStartId="8.[130,195,1138,1162]" captionTargetBox="[289,1313,144,1110]" captionTargetId="figure-299@8.[289,1313,144,1110]" captionTargetPageId="8" captionText="Figure 5.Middle cervicals of Tanystropheus sp.from Miedary: A, B, D—ZPAL V.36/101; C, E, F—ZPAL V. 36/102; A is a surface model in left lateral view.B, C, longitudinal CT cross sections of an anterior (B) and a posterior portion (C) of the vertebra. D, E, F, transverse CT cross sections. Sectioning planes and walls of the neural canal and internal cavity have been outlined with white dashed lines, with the lines signifing the sectioning planes being thicker than the internal cavity outline. Anatomical abbreviations: ic—internal cavity; nc—neural canal; ns— neural spine; poz—postzygapophysis; prz—prezygapophysis." pageId="5" pageNumber="6">Fig. 5B</figureCitation>
|
||||
;
|
||||
<figureCitation id="7AAF7CD0066DFF9051A1FB9364DA6268" box="[1187,1260,1032,1056]" captionStart="Figure 7" captionStartId="10.[129,194,695,719]" captionTargetBox="[289,1313,144,667]" captionTargetId="figure-733@10.[289,1313,144,667]" captionTargetPageId="10" captionText="Figure 7. Internal cavity volume in the cervical vertebrae of ‘P’.antiquus MGUWr 3895s (A) and Tanystropheus sp. from Miedary ZPAL V. 36/181 (B). Anatomical abbreviation:ic—internal cavity. Images obtained using ‘xray’ shader in MeshLab v.2020.12 (Cignoni et al.2008)." pageId="5" pageNumber="6">Fig. 7B</figureCitation>
|
||||
). In the transverse cross section, the inner surface of the bone wall of the internal cavity was generally parallel to the circumferential outline of the vertebra (
|
||||
<figureCitation id="7AAF7CD0066DFF9056B4FBFD642D6209" box="[950,1051,1126,1150]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="5" pageNumber="6">Fig. 8A–E</figureCitation>
|
||||
). In most of the studied specimens, the dorsal part of the internal cavity seemed to preserve a semicircular shape in the transverse cross section, whereas tissue layout of its ventrolateral portion was usually much less uniform, with an irregular resorption front cutting through multiple annuli (
|
||||
<figureCitation id="7AAF7CD0066DFF905637FA986381636D" box="[821,951,1283,1307]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="5" pageNumber="6">Fig. 8A, C, E</figureCitation>
|
||||
). This was especially evident in larger specimens, whereas in the smallest cervical, the regions of resorption were much less developed, and the outline of the internal cavity in the transverse cross section was more regularly oval (
|
||||
<figureCitation id="7AAF7CD0066DFF905029FAFA6542630E" box="[1323,1396,1377,1401]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="5" pageNumber="6">Fig. 8B</figureCitation>
|
||||
). The neural spine extended over two deep fossae, located on both of its anteroposteriorly terminal portions. They were triangular in longitudinal cross section and ventrally delimited by thin bone trabeculae, that also constituted the roof of the neural canal. The posterior cavity was larger, forming the ‘postzygapophyseal trough’ (
|
||||
<bibRefCitation id="86051DA4066DFF905687F98664396042" author="Rieppel O" box="[901,1039,1565,1589]" pageId="5" pageNumber="6" pagination="271 - 87" refId="ref17778" refString="Rieppel O. A new species of Tanystropheus (Reptilia: Protorosauria) from the Middle Triassic of Makhtesh Ramon, Israel. Neues Jahrbuch fur Geologie und Palaontologie - Abhandlungen 2001; 221: 271 - 87. https: // doi. org / 10.1127 / njgpa / 221 / 2001 / 271" type="journal article" year="2001">Rieppel 2001</bibRefCitation>
|
||||
). Contrary to what was observed for ‘
|
||||
<emphasis id="D0E0BC47066DFF90509FF986659D6043" box="[1437,1451,1565,1588]" italics="true" pageId="5" pageNumber="6">P</emphasis>
|
||||
’.
|
||||
<taxonomicName id="25941BD6066DFF905628F9A663486023" baseAuthorityName="Huene" baseAuthorityYear="1905" box="[810,894,1597,1620]" family="Tanystropheidae" genus="Protanystropheus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="antiquus">
|
||||
<emphasis id="D0E0BC47066DFF905628F9A663486023" box="[810,894,1597,1620]" italics="true" pageId="5" pageNumber="6">antiquus</emphasis>
|
||||
</taxonomicName>
|
||||
, in none of the studied sections of the
|
||||
<taxonomicName id="25941BD6066DFF905025F9A763626004" class="Reptilia" family="Tanystropheidae" genus="Tanystropheus" kingdom="Animalia" order="Protorosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="undetermined">
|
||||
<emphasis id="D0E0BC47066DFF905025F9A765856023" box="[1319,1459,1596,1620]" italics="true" pageId="5" pageNumber="6">Tanystropheus</emphasis>
|
||||
spp.
|
||||
</taxonomicName>
|
||||
middle cervicals did the internal cavity contain bony trabeculae extending internally from the surface of the bone wall. Some semi-symmetrical, slanted trabeculae of endosteal origin were present only incidentally in the anteroposteriorly terminal sections of the vertebrae (see
|
||||
<bibRefCitation id="86051DA4066DFF905161F94264E36087" author="Broili F" box="[1123,1237,1752,1777]" pageId="5" pageNumber="6" pagination="51 - 62" refId="ref15553" refString="Broili F. Beobachtungen an Tanystropheus conspicuus H. v. Meyer. Neues Jahrbuch fur Mineralogie Geologie und Palaontologie 1915; 2: 51 - 62." type="journal article" year="1915">Broili 1915</bibRefCitation>
|
||||
: plate 3, fig. 4c). The roof of the neural canal was generally straight and parallel to the longitudinal axis of the vertebra. The ventral delimitation of the neural canal was seemingly absent for most of its length within a vertebra. Intriguingly, in one of the transverse polished sections from the anterior part of U-MO BT 738.00 (
|
||||
<figureCitation id="7AAF7CD0066DFF90500AF8EE656761FA" box="[1288,1361,1909,1933]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="5" pageNumber="6">Fig. 8F</figureCitation>
|
||||
), minute bony projections appeared symmetrically on each lateral side of the internal cavity. The colour of the limestone infill of the vertebra changed drastically at the same horizontal level, at which these projections were situated. Similar characteristics could be noted for the transverse thin section of ZPAL V. 36/166 (
|
||||
<figureCitation id="7AAF7CD0066EFF9357E1F8E460AD61C0" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="6" pageNumber="7">Fig. 8E</figureCitation>
|
||||
), in which corresponding projections composed of periosteal bone were also present in the dorsolateral regions of the internal cavity. Between them, there was a sharp transition in sediment fraction, with the more coarse-grained infill occupying the ventral-more space. Presence of this feature possibly shows the differences in taphonomical microenvironments of the ventral and dorsal portions of the internal cavity, which may have been caused by the original existence of a no longer preserved barrier between them. Near the anteroposterior midpoint of the middle cervicals, two symmetrically placed foramina were located on the ventral side of the centrum. They entered the internal cavity as straight canals (
|
||||
<figureCitation id="7AAF7CD00660FF9D5439FA5C61AD63A9" box="[315,411,1479,1503]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="8" pageNumber="9">Fig. 8B, E</figureCitation>
|
||||
), perforating the dense cortex.
|
||||
</paragraph>
|
||||
<caption id="B6EB30DD066EFF935583F91E618B6086" pageId="6" pageNumber="7" startId="6.[129,194,1669,1693]" targetBox="[305,1297,144,1642]" targetPageId="6" targetType="figure">
|
||||
<paragraph id="E22B6055066EFF935583F91E618B6086" blockId="6.[129,1449,1669,1777]" pageId="6" pageNumber="7">
|
||||
<emphasis id="D0E0BC47066EFF935583F91E60EE60EA" bold="true" box="[129,216,1669,1693]" pageId="6" pageNumber="7">Figure 3.</emphasis>
|
||||
CT transverse cross sections (from anterior to posterior) of ‘
|
||||
<emphasis id="D0E0BC47066EFF935616F91D631760EA" box="[788,801,1670,1693]" italics="true" pageId="6" pageNumber="7">P</emphasis>
|
||||
’.
|
||||
<taxonomicName id="25941BD6066EFF93562FF91D634D60EA" baseAuthorityName="Huene" baseAuthorityYear="1905" box="[813,891,1670,1693]" family="Tanystropheidae" genus="Protanystropheus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="antiquus">
|
||||
<emphasis id="D0E0BC47066EFF93562FF91D634D60EA" box="[813,891,1670,1693]" italics="true" pageId="6" pageNumber="7">antiquus</emphasis>
|
||||
</taxonomicName>
|
||||
cervical MGUWr 3889s (surface model in left lateral view) showing internal structure of the vertebra. Sectioning planes are marked with lines. Note the presence of a large internal cavity (ic), thicker cortex layer (tc), and slanted trabeculae (st) spanning across the internal chamber. Trabecular bone (tb) is present in the anteroposteriorly terminal portions of the centrum.
|
||||
</paragraph>
|
||||
</caption>
|
||||
<caption id="B6EB30DD066FFF9D5573F8AA634161CD" lastPageId="8" lastPageNumber="9" pageId="7" pageNumber="8" startId="7.[113,178,1841,1865]" subCaptionStartIDs="8.[809,941,1899,1923]" subCaptionStarts="abbr" targetBox="[175,1398,149,1813]" targetPageId="7" targetType="figure">
|
||||
<paragraph id="E22B6055066FFF9D5573F8AA634161CD" blockId="7.[113,1457,1841,1978]" lastBlockId="8.[129,1454,1843,1979]" lastPageId="8" lastPageNumber="9" pageId="7" pageNumber="8">
|
||||
<emphasis id="D0E0BC47066FFF925573F8AA60FF613E" bold="true" box="[113,201,1841,1865]" pageId="7" pageNumber="8">Figure 4.</emphasis>
|
||||
Thin sections of ‘
|
||||
<emphasis id="D0E0BC47066FFF92546CF8A9614D613E" box="[366,379,1842,1865]" italics="true" pageId="7" pageNumber="8">P</emphasis>
|
||||
’.
|
||||
<taxonomicName id="25941BD6066FFF925485F8A961E3613E" baseAuthorityName="Huene" baseAuthorityYear="1905" box="[391,469,1842,1865]" family="Tanystropheidae" genus="Protanystropheus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="antiquus">
|
||||
<emphasis id="D0E0BC47066FFF925485F8A961E3613E" box="[391,469,1842,1865]" italics="true" pageId="7" pageNumber="8">antiquus</emphasis>
|
||||
</taxonomicName>
|
||||
cervical vertebrae SUT-MG/F/Tvert/2 (A, E, F, H, I) and GIUS-7-3674 (B–D, G, J). A–C, thin sections in normal transmitted light (A, B) and in polarized light (C). The slight lateral asymmetry results from minor antero-posterior skewing of the plane of sectioning. D, close-up of the internal supporting trabeculae in polarized light, presenting heavy remodelling. E–F, the lateral cortex of the centrum in polarized light (E) and polarized light with lambda compensator (F) showing lack of distinctive zonation and loosely ordered bone structure. G, cortex of the dorsolateral region of the centrum displaying more pronounced zonation and dorsal internal slanted trabeculae in polarized light. H–I, cortex of the ventrolateral region of centrum showing radial vascularization and a well-developed bundle of Sharpey’s fibres in polarized light (H) and polarized light with lambda compensator (I). J, the trabecular bone of the neural arch in polarized light. Arrows indicate a perforation of the neural canal floor. Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; pfb—parallel-fibred bone; pvc—primary vascular canals; sf—Sharpey’s fibres; st—slanted trabeculae; tb—trabecular bone. Scale bars for panels A–C equal
|
||||
<quantity id="256CCDB00660FF9D577BF838628661CD" box="[633,688,1955,1979]" metricMagnitude="-3" metricUnit="m" metricValue="5.0" pageId="8" pageNumber="9" unit="mm" value="5.0">5 mm</quantity>
|
||||
and
|
||||
<quantity id="256CCDB00660FF9D57DFF838631361CD" box="[733,805,1955,1979]" metricMagnitude="-4" metricUnit="m" metricValue="5.0" pageId="8" pageNumber="9" unit="mm" value="0.5">0.5 mm</quantity>
|
||||
for D–J.
|
||||
</paragraph>
|
||||
</caption>
|
||||
<caption id="B6EB30DD0660FF9D5580FBE962FF628D" pageId="8" pageNumber="9" startId="8.[130,195,1138,1162]" targetBox="[289,1313,144,1110]" targetPageId="8" targetType="figure">
|
||||
<paragraph id="E22B60550660FF9D5580FBE962FF628D" blockId="8.[129,1462,1138,1274]" pageId="8" pageNumber="9">
|
||||
<emphasis id="D0E0BC470660FF9D5580FBE960EC62FD" bold="true" box="[130,218,1138,1162]" pageId="8" pageNumber="9">Figure 5.</emphasis>
|
||||
Middle cervicals of
|
||||
<taxonomicName id="25941BD60660FF9D5494FBE9620162FD" box="[406,567,1138,1162]" class="Reptilia" family="Tanystropheidae" genus="Tanystropheus" kingdom="Animalia" order="Protorosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="undetermined">
|
||||
<emphasis id="D0E0BC470660FF9D5494FBE9622E62FD" box="[406,536,1138,1162]" italics="true" pageId="8" pageNumber="9">Tanystropheus</emphasis>
|
||||
sp.
|
||||
</taxonomicName>
|
||||
from Miedary: A, B, D—ZPAL V. 36/101; C, E, F—ZPAL V. 36/102; A is a surface model in left lateral view. B, C, longitudinal CT cross sections of an anterior (B) and a posterior portion (C) of the vertebra. D, E, F, transverse CT cross sections. Sectioning planes and walls of the neural canal and internal cavity have been outlined with white dashed lines, with the lines signifing the sectioning planes being thicker than the internal cavity outline. Anatomical abbreviations: ic—internal cavity; nc—neural canal; ns— neural spine; poz—postzygapophysis; prz—prezygapophysis.
|
||||
</paragraph>
|
||||
</caption>
|
||||
<paragraph id="E22B60550660FF9C559EFA7D620660C7" blockId="8.[128,779,1322,1753]" lastBlockId="9.[113,763,1375,1712]" lastPageId="9" lastPageNumber="10" pageId="8" pageNumber="9">
|
||||
While in ‘
|
||||
<emphasis id="D0E0BC470660FF9D5409FA7C612F6389" box="[267,281,1511,1534]" italics="true" pageId="8" pageNumber="9">P</emphasis>
|
||||
’.
|
||||
<taxonomicName id="25941BD60660FF9D542EFA7C61B66389" baseAuthorityName="Huene" baseAuthorityYear="1905" box="[300,384,1511,1534]" family="Tanystropheidae" genus="Protanystropheus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="antiquus">
|
||||
<emphasis id="D0E0BC470660FF9D542EFA7C61B66389" box="[300,384,1511,1534]" italics="true" pageId="8" pageNumber="9">antiquus</emphasis>
|
||||
</taxonomicName>
|
||||
the neural spine was easily identifiable, it was much less recognizable in the middle cervicals of
|
||||
<taxonomicName id="25941BD60660FF9D5583F9BE6109604B" box="[129,319,1573,1597]" class="Reptilia" family="Tanystropheidae" genus="Tanystropheus" kingdom="Animalia" order="Protorosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="undetermined">
|
||||
<emphasis id="D0E0BC470660FF9D5583F9BE613B604A" box="[129,269,1573,1597]" italics="true" pageId="8" pageNumber="9">Tanystropheus</emphasis>
|
||||
spp.
|
||||
</taxonomicName>
|
||||
It was impossible to demarcate the border between the neural arch and the vertebral centrum—they were completely fused, with no preserved remnants of the suture between them, even in the smallest specimens, and little to no trabeculae within the neural arch. Continuous layers of dense parallel-fibred tissue sheathed the internal cavity. In the middle section of the larger middle cervicals of
|
||||
<taxonomicName id="25941BD60660FF9D51DFFAB165AC6335" box="[1245,1434,1322,1346]" class="Reptilia" family="Tanystropheidae" genus="Tanystropheus" kingdom="Animalia" order="Protorosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="undetermined">
|
||||
<emphasis id="D0E0BC470660FF9D51DFFAB1655F6335" box="[1245,1385,1322,1346]" italics="true" pageId="8" pageNumber="9">Tanystropheus</emphasis>
|
||||
spp.
|
||||
</taxonomicName>
|
||||
the outermost of these zones formed complete rings, whereas the ones located innermost were discontinued only due to resorption of the region ventral into the internal cavity, as well as the presence of the neural spine (
|
||||
<figureCitation id="7AAF7CD00660FF9D5176FA3C64D663B7" box="[1140,1248,1447,1472]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="8" pageNumber="9">Fig. 8A, C</figureCitation>
|
||||
,
|
||||
<figureCitation id="7AAF7CD00660FF9D51ECFA3C650E63B7" box="[1262,1336,1447,1472]" captionStart="Figure 9" captionStartId="12.[130,195,1032,1056]" captionTargetBox="[243,1360,147,1002]" captionTargetId="figure-468@12.[240,1363,144,1005]" captionTargetPageId="12" captionText="Figure 9. Interpretative drawings of the internal anatomy of a cervical vertebra of ‘P’.antiquus (A) and a middle cervical of an adult individual of Tanystropheus spp.(B) shown in the transverse cross section near the midpoint of their anteroposterior length." pageId="8" pageNumber="9">Fig. 9B</figureCitation>
|
||||
). In some of the studied transverse thin sections, including the smallest specimens, the spinous process was developed only as a thin, vertically oriented plate. It was located directly above the internal cavity and emerged from the parallel-fibred tissue of the bone walls in the dorsalmost portion of the vertebra, projecting slightly from the smooth outline of the transverse cross section. In the smallest specimen from Miedary, ZPAL V. 36/181, the neural spine was never overlain by parallel-fibred bone (
|
||||
<figureCitation id="7AAF7CD00660FF9D504AF93965A460CD" box="[1352,1426,1698,1722]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="8" pageNumber="9">Fig. 8B</figureCitation>
|
||||
). In all of the larger specimens from Miedary, the anteroposteriorly middle part of the spinous process was obscured by thick layers of tissue that formed the walls of the internal cavity. Compared to what was noted for ‘
|
||||
<emphasis id="D0E0BC470661FF9C5464FA05614263C2" box="[358,372,1438,1461]" italics="true" pageId="9" pageNumber="10">P</emphasis>
|
||||
’.
|
||||
<taxonomicName id="25941BD60661FF9C5485FA0561ED63C2" baseAuthorityName="Huene" baseAuthorityYear="1905" box="[391,475,1438,1461]" family="Tanystropheidae" genus="Protanystropheus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="antiquus">
|
||||
<emphasis id="D0E0BC470661FF9C5485FA0561ED63C2" box="[391,475,1438,1461]" italics="true" pageId="9" pageNumber="10">antiquus</emphasis>
|
||||
</taxonomicName>
|
||||
, the dorsolateral regions of the
|
||||
<taxonomicName id="25941BD60661FF9C559AFA26616463A3" box="[152,338,1469,1493]" class="Reptilia" family="Tanystropheidae" genus="Tanystropheus" kingdom="Animalia" order="Protorosauria" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="undetermined">
|
||||
<emphasis id="D0E0BC470661FF9C559AFA26611263A2" box="[152,292,1469,1493]" italics="true" pageId="9" pageNumber="10">Tanystropheus</emphasis>
|
||||
spp.
|
||||
</taxonomicName>
|
||||
cervicals were thickened (see
|
||||
<figureCitation id="7AAF7CD00661FF9C5787FA26628B63A2" box="[645,701,1469,1493]" captionStart="Figure 9" captionStartId="12.[130,195,1032,1056]" captionTargetBox="[243,1360,147,1002]" captionTargetId="figure-468@12.[240,1363,144,1005]" captionTargetPageId="12" captionText="Figure 9. Interpretative drawings of the internal anatomy of a cervical vertebra of ‘P’.antiquus (A) and a middle cervical of an adult individual of Tanystropheus spp.(B) shown in the transverse cross section near the midpoint of their anteroposterior length." pageId="9" pageNumber="10">Fig. 9</figureCitation>
|
||||
). The neural spine was, thus, not relatively reduced in the latter, but rather embedded within the walls of the tubularly structured vertebra. As a result, the neural canal was located close to the middle of the height of the vertebra, as in ‘
|
||||
<emphasis id="D0E0BC470661FF9C571DF9A0621B6025" box="[543,557,1595,1618]" italics="true" pageId="9" pageNumber="10">P</emphasis>
|
||||
’.
|
||||
<taxonomicName id="25941BD60661FF9C5738F9A062B86025" baseAuthorityName="Huene" baseAuthorityYear="1905" box="[570,654,1595,1618]" family="Tanystropheidae" genus="Protanystropheus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="antiquus">
|
||||
<emphasis id="D0E0BC470661FF9C5738F9A062B86025" box="[570,654,1595,1618]" italics="true" pageId="9" pageNumber="10">antiquus</emphasis>
|
||||
</taxonomicName>
|
||||
. However, in
|
||||
<taxonomicName id="25941BD60661FF9C558FF9C2617D6006" box="[141,331,1625,1649]" class="Reptilia" family="Tanystropheidae" genus="Tanystropheus" kingdom="Animalia" order="Protorosauria" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="undetermined">
|
||||
<emphasis id="D0E0BC470661FF9C558FF9C2612F6006" box="[141,281,1625,1649]" italics="true" pageId="9" pageNumber="10">Tanystropheus</emphasis>
|
||||
spp.
|
||||
</taxonomicName>
|
||||
there was more tissue dorsolaterally (see
|
||||
<figureCitation id="7AAF7CD00661FF9C5573F9E2609A60E6" box="[113,172,1657,1681]" captionStart="Figure 9" captionStartId="12.[130,195,1032,1056]" captionTargetBox="[243,1360,147,1002]" captionTargetId="figure-468@12.[240,1363,144,1005]" captionTargetPageId="12" captionText="Figure 9. Interpretative drawings of the internal anatomy of a cervical vertebra of ‘P’.antiquus (A) and a middle cervical of an adult individual of Tanystropheus spp.(B) shown in the transverse cross section near the midpoint of their anteroposterior length." pageId="9" pageNumber="10">Fig. 9</figureCitation>
|
||||
), due to the neural spine being hypertrophied along the middle section of its anteroposterior length.
|
||||
</paragraph>
|
||||
<caption id="B6EB30DD0661FF9C5573FB3C64746358" pageId="9" pageNumber="10" startId="9.[113,178,1191,1215]" targetBox="[146,1426,144,1163]" targetPageId="9" targetType="figure">
|
||||
<paragraph id="E22B60550661FF9C5573FB3C64746358" blockId="9.[113,1454,1191,1327]" pageId="9" pageNumber="10">
|
||||
<emphasis id="D0E0BC470661FF9C5573FB3C60FF62C8" bold="true" box="[113,201,1191,1215]" pageId="9" pageNumber="10">Figure 6.</emphasis>
|
||||
Vertebrae of
|
||||
<taxonomicName id="25941BD60661FF9C5444FB3C61D062C9" box="[326,486,1191,1215]" class="Reptilia" family="Tanystropheidae" genus="Tanystropheus" kingdom="Animalia" order="Protorosauria" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="undetermined">
|
||||
<emphasis id="D0E0BC470661FF9C5444FB3C61FE62C8" box="[326,456,1191,1215]" italics="true" pageId="9" pageNumber="10">Tanystropheus</emphasis>
|
||||
sp.
|
||||
</taxonomicName>
|
||||
from Miedary: A, E, I—ZPAL V. 36/106, 11th cervical; B, F, J—ZPAL V. 36/108, 12th cervical; C, G, K—ZPAL V. 36/110, 13th (last) cervical; D—ZPAL V. 36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections. White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations: ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels.
|
||||
</paragraph>
|
||||
</caption>
|
||||
<paragraph id="E22B60550661FF9C5418F94B6267609D" blockId="9.[113,765,1744,1963]" box="[282,593,1744,1770]" pageId="9" pageNumber="10">
|
||||
<emphasis id="D0E0BC470661FF9C5418F94B6267609D" box="[282,593,1744,1770]" italics="true" pageId="9" pageNumber="10">Anatomy of the posterior cervicals</emphasis>
|
||||
</paragraph>
|
||||
<paragraph id="E22B60550661FF9F5573F96C62506310" blockId="9.[113,765,1744,1963]" lastBlockId="10.[128,779,795,1728]" lastPageId="10" lastPageNumber="11" pageId="9" pageNumber="10">
|
||||
In
|
||||
<taxonomicName id="25941BD60661FF9C558CF96D610B6179" box="[142,317,1782,1806]" class="Reptilia" family="Tanystropheidae" genus="Tanystropheus" kingdom="Animalia" order="Protorosauria" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="undetermined">
|
||||
<emphasis id="D0E0BC470661FF9C558CF96D612C6179" box="[142,282,1782,1806]" italics="true" pageId="9" pageNumber="10">Tanystropheus</emphasis>
|
||||
sp.
|
||||
</taxonomicName>
|
||||
from Miedary, the external morphology of the cervicals changed near the posterior subregion of the neck. The 10th and 11th vertebrae were still elongate, but, contrary to the middle cervicals, exhibited a well exposed, relatively tall and continuous neural spine (
|
||||
<figureCitation id="7AAF7CD00661FF9C54B5F8E861C361FC" box="[439,501,1907,1931]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="9" pageNumber="10">Fig. 6</figureCitation>
|
||||
). The two last cervicals were relatively much shorter (
|
||||
<figureCitation id="7AAF7CD00661FF9C54AEF808622361DC" box="[428,533,1939,1963]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="9" pageNumber="10">Fig. 6B, C</figureCitation>
|
||||
), and more similar in proportions to the dorsals (
|
||||
<bibRefCitation id="86051DA40661FF9C513DFAC464CE6300" author="Rieppel O & Jiang DY & Fraser NC" box="[1087,1272,1375,1399]" pageId="9" pageNumber="10" pagination="1082 - 9" refId="ref17833" refString="Rieppel O, Jiang DY, Fraser NC et al. Tanystropheus cf. T. longobardicus from the early Late Triassic of Guizhou Province, Southwestern China. Journal of Vertebrate Paleontology 2010; 30: 1082 - 9. https: // doi. org / 10.1080 / 02724634.2010.483548" type="journal article" year="2010">
|
||||
Rieppel
|
||||
<emphasis id="D0E0BC470661FF9C5191FAC464F76300" box="[1171,1217,1375,1399]" italics="true" pageId="9" pageNumber="10">et al.</emphasis>
|
||||
2010
|
||||
</bibRefCitation>
|
||||
,
|
||||
<bibRefCitation id="86051DA40661FF9C5000FAC465946300" author="Rytel A & Bohmer C & Spiekman SNF" box="[1282,1442,1375,1399]" pageId="9" pageNumber="10" refId="ref17884" refString="Rytel A, Bohmer C, Spiekman SNF et al. Extreme neck elongation evolved despite strong developmental constraints in bizarre Triassic reptiles - implications for neck modularity in archosaurs. Royal Society Open Science 2024; 11: 240233. https: // doi. org / 10.1098 / rsos. 240233" type="journal volume" year="2024">
|
||||
Rytel
|
||||
<emphasis id="D0E0BC470661FF9C503FFAC4655D6300" box="[1341,1387,1375,1399]" italics="true" pageId="9" pageNumber="10">et al.</emphasis>
|
||||
2024
|
||||
</bibRefCitation>
|
||||
). This transition was paired with the modifications in the internal structure of these elements. The relative volume occupied by the trabeculae within the centra increased caudally, especially along the neck-torso transition. In the middle cervicals the centrum was nearly devoid of trabecular bone, but this changed within the more posterior vertebrae, with the last two cervicals and the dorsals exhibiting sparse trabeculae occupying the space between the articular discs. The 10th and 11th vertebrae were generally similar in their anatomy to the middle cervicals, with their internal cavities being surrounded by a dense cortex, forming a tube-like structure (
|
||||
<figureCitation id="7AAF7CD00661FF9C56F8F92C640B60B8" box="[1018,1085,1719,1743]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="9" pageNumber="10">Fig. 6I</figureCitation>
|
||||
). However, in the succeeding vertebrae the internal cavity was greatly reduced and exhibited much thicker walls, composed of two regions: a relatively thin cortex and porous, cancellous bone encompassed by it. Especially in the middle portion of the centrum of the last two cervicals, there were several large chambers, similar to those present in the anteroposteriorly terminal portions of the more anteriorly located vertebrae (
|
||||
<figureCitation id="7AAF7CD00661FF9C56F0F808647761DC" box="[1010,1089,1939,1963]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="9" pageNumber="10">Fig. 8D</figureCitation>
|
||||
). Some of them connected to the neural canal through its floor (
|
||||
<figureCitation id="7AAF7CD00662FF9F54C3FC80626A6544" box="[449,604,795,819]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="10" pageNumber="11">Fig. 6E, F, H, I</figureCitation>
|
||||
). The transverse cross sections of the posteriormost cervicals were less oval than in the preceding vertebrae, with the centrum being much more ventrally expanded, forming a distinct keel (
|
||||
<figureCitation id="7AAF7CD00662FF9F5762FCE262F165E6" box="[608,711,889,913]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="10" pageNumber="11">Fig. 6J, K</figureCitation>
|
||||
). The diameter of the neural canal was relatively constant along the length of each of the two last vertebrae. In the longitudinal cross section, it had a sagging appearance, with its middle portion being located more ventrally. In the transverse cross section, it was oval, with its dorsolateral regions being roughly semicircular and symmetrical in outline (
|
||||
<figureCitation id="7AAF7CD00662FF9F54A3FBAE61CB623A" box="[417,509,1077,1101]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="10" pageNumber="11">Fig. 6J, K</figureCitation>
|
||||
). A similar shape could be noted in the anteroposteriorly terminal sections of the rest of the postaxial cervicals, in which the neural canal floor was still present (e.g.
|
||||
<figureCitation id="7AAF7CD00662FF9F55E1FB08611862DC" box="[227,302,1171,1195]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="10" pageNumber="11">Fig. 8D</figureCitation>
|
||||
), and also in the dorsal vertebrae (
|
||||
<figureCitation id="7AAF7CD00662FF9F5794FB0862EB62DC" box="[662,733,1171,1195]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="10" pageNumber="11">Fig. 6L</figureCitation>
|
||||
). In each of the two posteriormost neck vertebrae of
|
||||
<taxonomicName id="25941BD60662FF9F577FFB2860AA629E" class="Reptilia" family="Tanystropheidae" genus="Tanystropheus" kingdom="Animalia" order="Protorosauria" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="undetermined">
|
||||
<emphasis id="D0E0BC470662FF9F577FFB28633F62BC" box="[637,777,1203,1227]" italics="true" pageId="10" pageNumber="11">Tanystropheus</emphasis>
|
||||
sp.
|
||||
</taxonomicName>
|
||||
from Miedary, up to two small subcentral foramina could be traced, with some specimens exhibiting only one subcentral foramen or none at all. In the studied posteriormost cervical ZPAL V. 36/110, a straight, non-branching canal connected the one present foramen with the neural canal (
|
||||
<figureCitation id="7AAF7CD00662FF9F5712FAD462636310" box="[528,597,1359,1383]" captionStart="Figure 10" captionStartId="13.[113,178,1140,1164]" captionTargetBox="[306,1266,144,1112]" captionTargetId="figure-458@13.[306,1266,144,1112]" captionTargetPageId="13" captionText="Figure 10. Longitudinal cross section of ZPAL V.36/110, 13th (last) cervical vertebra of Tanystropheus sp.from Miedary, showing the course of the arterial canal (locations of its end sections are indicated with arrows). The anterior end of the vertebra points towards the left. Anatomical abbreviations: nc—neural canal; ns—neural spine." pageId="10" pageNumber="11">Fig. 10</figureCitation>
|
||||
).
|
||||
</paragraph>
|
||||
<caption id="B6EB30DD0662FF9F5583FD2C65AF649B" pageId="10" pageNumber="11" startId="10.[129,194,695,719]" targetBox="[289,1313,144,667]" targetPageId="10" targetType="figure">
|
||||
<paragraph id="E22B60550662FF9F5583FD2C65AF649B" blockId="10.[129,1433,695,748]" pageId="10" pageNumber="11">
|
||||
<emphasis id="D0E0BC470662FF9F5583FD2C60EE64B8" bold="true" box="[129,216,695,719]" pageId="10" pageNumber="11">Figure 7.</emphasis>
|
||||
Internal cavity volume in the cervical vertebrae of ‘
|
||||
<emphasis id="D0E0BC470662FF9F57BAFD2362F364B8" box="[696,709,696,719]" italics="true" pageId="10" pageNumber="11">P</emphasis>
|
||||
’.
|
||||
<taxonomicName id="25941BD60662FF9F57D3FD23632964B8" baseAuthorityName="Huene" baseAuthorityYear="1905" box="[721,799,696,719]" family="Tanystropheidae" genus="Protanystropheus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="antiquus">
|
||||
<emphasis id="D0E0BC470662FF9F57D3FD23632964B8" box="[721,799,696,719]" italics="true" pageId="10" pageNumber="11">antiquus</emphasis>
|
||||
</taxonomicName>
|
||||
MGUWr 3895s (A) and
|
||||
<taxonomicName id="25941BD60662FF9F510CFD2C649864B8" box="[1038,1198,695,719]" class="Reptilia" family="Tanystropheidae" genus="Tanystropheus" kingdom="Animalia" order="Protorosauria" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="undetermined">
|
||||
<emphasis id="D0E0BC470662FF9F510CFD2C64A664B8" box="[1038,1168,695,719]" italics="true" pageId="10" pageNumber="11">Tanystropheus</emphasis>
|
||||
sp.
|
||||
</taxonomicName>
|
||||
from Miedary ZPAL V. 36/181 (B). Anatomical abbreviation: ic—internal cavity. Images obtained using ‘xray’ shader in MeshLab v.2020.12 (
|
||||
<bibRefCitation id="86051DA40662FF9F51DAFD4865BF649C" author="Cignoni P & Callieri M & Corsini M" box="[1240,1417,723,747]" pageId="10" pageNumber="11" pagination="29 - 36" refId="ref15718" refString="Cignoni P, Callieri M, Corsini M et al. MeshLab: an open-source mesh processing tool. In: 6 th Eurographics Italian Chapter Conference 2008 - Proceedings, Salerno, Italy, 2008. p. 29 - 36. Eindhoven, The Netherlands: Eurographics Association, 2008." type="book chapter" year="2008">
|
||||
Cignoni
|
||||
<emphasis id="D0E0BC470662FF9F5028FD4F6560649C" box="[1322,1366,723,747]" italics="true" pageId="10" pageNumber="11">et al.</emphasis>
|
||||
2008
|
||||
</bibRefCitation>
|
||||
).
|
||||
</paragraph>
|
||||
</caption>
|
||||
<paragraph id="E22B60550662FF9F559EFAF462AB60B7" blockId="10.[128,779,795,1728]" pageId="10" pageNumber="11">
|
||||
The dorsal vertebrae were generally similar to the posteriormost cervicals in their internal anatomy. They differed in a complete lack of subcentral foramina. Moreover, the neural canal of the dorsals extended anteroposteriorly straight, with its roof and floor remaining on a relatively constant horizontal level (
|
||||
<figureCitation id="7AAF7CD00662FF9F5589F99060E16054" box="[139,215,1547,1571]" captionStart="Figure 6" captionStartId="9.[113,178,1191,1215]" captionTargetBox="[146,1426,144,1163]" captionTargetId="figure-396@9.[146,1426,144,1163]" captionTargetPageId="9" captionText="Figure 6. Vertebrae of Tanystropheus sp. from Miedary:A, E, I—ZPAL V.36/106, 11th cervical; B, F, J—ZPAL V.36/108, 12th cervical; C, G, K—ZPAL V.36/110, 13th (last) cervical; D—ZPAL V.36/1036, dorsal; H, L—ZPAL V. 36/112, dorsal. A–D, surface models in left lateral view. E–H, longitudinal CT cross sections. I–L, transverse CT cross sections.White dashed lines mark the sectioning planes (yellow) and walls of the neural canal and internal cavity (white). Anatomical abbreviations:ic—internal cavity; nc—neural canal; ns—neural spine; poz— postzygapophysis; prz—prezygapophysis. Scale bars equal 2 cm for A–H, and 1 cm for the other panels." pageId="10" pageNumber="11">Fig. 6H</figureCitation>
|
||||
). Its transverse cross section was similar in shape to the posteriormost cervicals, but more regular. Congruently to what could be observed in the 12th and 13th cervical vertebrae, the internal portion of the centrum was composed of sparsely distributed trabeculae containing a large number of cavities, some of which opened to the neural canal through its floor.
|
||||
</paragraph>
|
||||
<paragraph id="E22B60550662FF9F546DF944622D608E" blockId="10.[367,539,1759,1785]" box="[367,539,1759,1785]" pageId="10" pageNumber="11">
|
||||
<emphasis id="D0E0BC470662FF9F546DF944622D608E" box="[367,539,1759,1785]" italics="true" pageId="10" pageNumber="11">Vertebral histology</emphasis>
|
||||
</paragraph>
|
||||
<paragraph id="E22B60550662FF9F5583F89D64A963F0" blockId="10.[129,780,1798,1979]" lastBlockId="10.[824,1476,795,1415]" pageId="10" pageNumber="11">
|
||||
The histological characteristics of the studied vertebrae of
|
||||
<taxonomicName id="25941BD60662FF9F5583F8BE610A614A" box="[129,316,1829,1853]" class="Reptilia" family="Tanystropheidae" genus="Tanystropheus" kingdom="Animalia" order="Protorosauria" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="undetermined">
|
||||
<emphasis id="D0E0BC470662FF9F5583F8BE613B614A" box="[129,269,1829,1853]" italics="true" pageId="10" pageNumber="11">Tanystropheus</emphasis>
|
||||
spp.
|
||||
</taxonomicName>
|
||||
were congruent with the results of the study performed by
|
||||
<bibRefCitation id="86051DA40662FF9F5417F8DE6210612A" author="Jaquier VP & Scheyer TM" box="[277,550,1861,1885]" pageId="10" pageNumber="11" refId="ref16697" refString="Jaquier VP, Scheyer TM. Bone histology of the Middle Triassic long-necked reptiles Tanystropheus and Macrocnemus (Archosauromorpha, Protorosauria). Journal of Vertebrate Paleontology 2017; 37: e 1296456. https: // doi. org / 10.1080 / 02724634.2017.1296456" type="journal volume" year="2017">Jaquier and Scheyer (2017</bibRefCitation>
|
||||
and the supplemental data provided therein). New data presented herein expand our knowledge on the anteroposterior and ontogenetic variation of their structure. The walls of the internal cavity were built of primary compact lamellar to parallel-fibred matrix of periosteal origin. Some intervals with decreased vascularization could be traced within these bone walls—they formed ellipsoidal annuli that alternated with more vascularized zones in parallel to the circumference of the vertebra (see
|
||||
<bibRefCitation id="86051DA40662FF9F51A4FC02658765C6" author="Jaquier VP & Scheyer TM" box="[1190,1457,921,945]" pageId="10" pageNumber="11" refId="ref16697" refString="Jaquier VP, Scheyer TM. Bone histology of the Middle Triassic long-necked reptiles Tanystropheus and Macrocnemus (Archosauromorpha, Protorosauria). Journal of Vertebrate Paleontology 2017; 37: e 1296456. https: // doi. org / 10.1080 / 02724634.2017.1296456" type="journal volume" year="2017">Jaquier and Scheyer 2017</bibRefCitation>
|
||||
). Closer to the terminal portions of the anteroposterior length of the cervicals, the tissue composition changed, with the internal cavity being dorsolaterally lined with secondary endosteal lamellar bone (
|
||||
<figureCitation id="7AAF7CD00662FF9F56C0FB8D64386259" box="[962,1038,1046,1070]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="10" pageNumber="11">Fig. 8H</figureCitation>
|
||||
) and the ventrolateral portions of the bone centrum not exhibiting the regular annuli, but rather being nearly completely composed of endosteal lamellar bone scattered with secondary osteons and erosion cavities of varying size, forming a net of sparse trabeculae (
|
||||
<figureCitation id="7AAF7CD00662FF9F5150FB0864F662DC" box="[1106,1216,1171,1195]" captionStart="Figure 5" captionStartId="8.[130,195,1138,1162]" captionTargetBox="[289,1313,144,1110]" captionTargetId="figure-299@8.[289,1313,144,1110]" captionTargetPageId="8" captionText="Figure 5.Middle cervicals of Tanystropheus sp.from Miedary: A, B, D—ZPAL V.36/101; C, E, F—ZPAL V. 36/102; A is a surface model in left lateral view.B, C, longitudinal CT cross sections of an anterior (B) and a posterior portion (C) of the vertebra. D, E, F, transverse CT cross sections. Sectioning planes and walls of the neural canal and internal cavity have been outlined with white dashed lines, with the lines signifing the sectioning planes being thicker than the internal cavity outline. Anatomical abbreviations: ic—internal cavity; nc—neural canal; ns— neural spine; poz—postzygapophysis; prz—prezygapophysis." pageId="10" pageNumber="11">Figs 5B–D</figureCitation>
|
||||
,
|
||||
<figureCitation id="7AAF7CD00662FF9F51CEFB08653D62DC" box="[1228,1291,1171,1195]" captionStart="Figure 8" captionStartId="11.[113,178,1841,1865]" captionTargetBox="[224,1348,150,1812]" captionTargetId="figure-6@11.[223,1349,149,1813]" captionTargetPageId="11" captionText="Figure 8. Transverse cross sections through the cervicals of Tanystropheus ‘conspicuus’ (F) and Tanystropheus sp. from Miedary (all other panels). A, ZPAL V. 36/150, middle cervical, middle portion; (B) ZPAL V. 36/181, middle cervical, middle portion; (C) ZPAL V. 36/193, middle cervical, middle portion; (D) ZPAL V. 36/1099, 10th cervical, posterior portion; (E) ZPAL V.36/166, 10th cervical, middle portion; (F) UMO BT 738.00, 11th cervical, anterior portion; (G) ZPAL V.36/1099, 10th cervical, posterior portion, close-up on the right ventrolateral portion of the vertebra with bundles of longitudinally oriented secondary osteons; (H) ZPAL V. 36/156, middle cervical, middle section, close-up on the base of neural spine with the endosteal bone lining.A, C, D, E, G, H, thin sections viewed in polarized light. B, µCT image.F, polished section.Anatomical abbreviations: elb—endosteal lamellar bone; ic—internal cavity; nc—neural canal; ns—neural spine; scf—subcentral foramen; so—secondary osteons; tb—trabecular bone.Scale bar equals 2 mm for B, G, H, and 5 mm for the other panels." pageId="10" pageNumber="11">8D, G</figureCitation>
|
||||
). The core of the neural spine in larger specimens was highly remodelled with longitudinally oriented secondary osteons. Simple primary vascular canals extended radially in the middle part of the vertebrae, but became more longitudinally oriented, larger, and less organized in the anteroposteriorly terminal portions. The number of vascular canals was also smaller in the more compact sections of the cortex within a single cross section.
|
||||
</paragraph>
|
||||
</subSubSection>
|
||||
</treatment>
|
||||
</document>
|
||||
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