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<mods:title id="0E7F0B551D27D8F354F12498B2566819">The first Pan-Podocnemididae turtle egg from the Presidente Prudente Formation (Late Cretaceous, Bauru Group), Brazil</mods:title>
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<mods:namePart id="5BC0DA4FD28421A06370EF9CD1934516">Marsola, Júlio C. De A.</mods:namePart>
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<mods:namePart id="5D7A01A887A5DDF222EAB9D982C28608">Grellet-Tinner, Gerald</mods:namePart>
<mods:affiliation id="76BD81E15CFD6562995AEF596CF18940">Orcas Island Museum, PO Box 134, 181 North Beach Road, Eastsound, WA 98245; Investigador Correspondiente at Departamento de Geociencias, CRILAR, CONICET, Argentina. E-mail: locarnolugano @ gmail. com</mods:affiliation>
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<mods:namePart id="6AC4F1DB05607BB7C1B6D83F07ACB7A5">Montefeltro, Felipe C.</mods:namePart>
<mods:affiliation id="8D1DA623B9DF4D91DF990A99CB7434C1">Departamento de Zoologia, Universidade Estadual Paulista, Avenida 24 A 1515, Rio Claro, Brazil, felipecmontefeltro @ gmail. com Corresponding author. E-mail: juliomarsola @ gmail. com. Tel.: + 55 16 36023844</mods:affiliation>
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<taxonomicName id="4C634D308814FF8977B3FBDAE65D3AF0" authority="Joyce, Praham &amp; Gauthier, 2004" box="[151,877,898,923]" kingdom="Animalia" pageId="2" pageNumber="189" rank="family">
<emphasis id="B917EAA18814FF8977B3FBDAE65D3AF0" bold="true" box="[151,877,898,923]" pageId="2" pageNumber="189">PAN-PODOCNEMIDIDAE Joyce, Praham &amp; Gauthier 2004</emphasis>
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<emphasis id="B917EAA18814FF8977B3FB92E46F3A88" bold="true" box="[151,351,970,995]" pageId="2" pageNumber="189">Gen. et sp. indet.</emphasis>
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<heading id="D09481DF8814FF8977B3FC54E4F83D48" bold="true" box="[151,456,1036,1061]" fontSize="10" level="2" pageId="2" pageNumber="189" reason="2">
<emphasis id="B917EAA18814FF8977B3FC54E4F83D48" box="[151,456,1036,1061]" inLineHeading="true" pageId="2" pageNumber="189" reason="1">
<emphasis id="B917EAA18814FF8977B3FC54E4F03D4E" bold="true" box="[151,448,1036,1061]" pageId="2" pageNumber="189">Comparative description</emphasis>
.
</emphasis>
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<paragraph id="8BDC36B38814FF8977B3FC77E19B3CBE" blockId="2.[151,1437,1036,2033]" pageId="2" pageNumber="189">
LPRP-USP 0052 (
<figureCitation id="13582A368814FF897642FC77E4EE3D23" box="[358,478,1071,1096]" captionStart="FIGURE 2" captionStartId="3.[151,250,1799,1821]" captionTargetBox="[302,1252,193,1780]" captionTargetId="figure-175@3.[302,1252,193,1780]" captionTargetPageId="3" captionText="FIGURE 2. A, LPRP-USP 0052 in two views showing the missing pole and the folded eggshell. B, MO of a thin section of LPRP-USP 0052 eggshell. C, interpretative drawing of the thin section. Gray color in represents the additional cuticle layer above the units. D, E, F and G, SEM images of LPRP-USP 0052. D, eggshell in radial section showing the additional cuticle layer. Black arrow indicates the boundary between the biomineralized eggshell and the cuticle. E, eggshell in radial section showing two loosely abutted and triangular basic units and the “cavern” between them (translucent triangle). Black arrows point to the large primary spherites, from which the acicular aragonitic crystals radiate. F, outer surface of the eggshell showing two pore apertures (black arrows). Note that the shell basic units outlines are not easily seen. G, Enlargment of the rounded pore aperture (black arrow)." figureDoi="http://doi.org/10.5281/zenodo.4948050" httpUri="https://zenodo.org/record/4948050/files/figure.png" pageId="2" pageNumber="189">figure 2 A</figureCitation>
) is elliptic (0,5 width/length ratio) with a missing pole. Its main axis is 5,1 cm long, whereas the minor axis range from 2,9 to 2,2 cm due to compression. Although taphonomicaly distorted, the elongation of LPRP-USP 0052 still matches that of chelids such as
<taxonomicName id="4C634D308814FF897487FC2FE1283DE4" box="[931,1048,1143,1168]" class="Reptilia" family="Chelidae" genus="Elseya" kingdom="Animalia" order="Testudines" pageId="2" pageNumber="189" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B917EAA18814FF897487FC2FE6DE3DFB" box="[931,1006,1143,1168]" italics="true" pageId="2" pageNumber="189">Elseya</emphasis>
sp.
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,
<taxonomicName id="4C634D308814FF897303FC2FE1F53DE4" box="[1063,1221,1143,1168]" class="Reptilia" family="Chelidae" genus="Chelodina" kingdom="Animalia" order="Testudines" pageId="2" pageNumber="189" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B917EAA18814FF897303FC2FE1AC3DFB" box="[1063,1180,1143,1168]" italics="true" pageId="2" pageNumber="189">Chelodina</emphasis>
sp.
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and
<taxonomicName id="4C634D308814FF897224FC2FE4D73DDE" authority="(Winkler 2006)" baseAuthorityName="Winkler" baseAuthorityYear="2006" class="Reptilia" family="Chelidae" genus="Hydromedusa" kingdom="Animalia" order="Testudines" pageId="2" pageNumber="189" phylum="Chordata" rank="species" species="maximilliani">
<emphasis id="B917EAA18814FF897224FC2FE4143DDE" italics="true" pageId="2" pageNumber="189">Hydromedusa maximilliani</emphasis>
(Winkler 2006)
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. This differs from the rounded shape of almost all described turtle fossil eggs (
<bibRefCitation id="EFF24B428814FF8977BAFCE7E4B53DB3" author="Azevedo, S. A. &amp; Gallo, V. &amp; Ferigolo, J." box="[158,389,1215,1240]" pageId="2" pageNumber="189" pagination="187 - 193" refId="ref3026" refString="Azevedo, S. A., Gallo, V. &amp; Ferigolo, J. (2000) A possible chelonian egg from the Brazilian Late Cretaceous. Anais da Academia Brasileira de Ciencias, 72, 187 - 193. http: // dx. doi. org / 10.1590 / s 0001 - 37652000000200007" type="journal article" year="2000">
Azevedo
<emphasis id="B917EAA18814FF89762FFC99E40B3DB3" box="[267,315,1215,1240]" italics="true" pageId="2" pageNumber="189">et al</emphasis>
. 2000
</bibRefCitation>
;
<bibRefCitation id="EFF24B428814FF8976B4FCE7E7593DB3" author="Jackson, F. &amp; Jin, X. &amp; Varricchio, D. J. &amp; Azuma, Y. &amp; Jiang, Y." box="[400,617,1215,1240]" pageId="2" pageNumber="189" pagination="319 - 323" refId="ref4412" refString="Jackson, F., Jin, X., Varricchio, D. J., Azuma, Y. &amp; Jiang, Y. (2008) The first in situ turtle clutch from the Cretaceous Tiantai Basin, Zhejiang Province, China. Journal of Vertebrate Paleontology, 28, 319 - 323. http: // dx. doi. org / 10.1671 / 0272 - 4634 (2008) 28 [319: tfistc] 2.0. co; 2" type="journal article" year="2008">
Jackson
<emphasis id="B917EAA18814FF8976D4FC99E7103DB3" box="[496,544,1215,1240]" italics="true" pageId="2" pageNumber="189">et al</emphasis>
. 2008
</bibRefCitation>
, table 1;
<bibRefCitation id="EFF24B428814FF8975EAFCE7E6BE3DB3" author="Knell, M. J. &amp; Jackson, F. &amp; Titus, A. L. &amp; Albright III, L. B." box="[718,910,1215,1240]" pageId="2" pageNumber="189" pagination="57 - 62" refId="ref4645" refString="Knell, M. J., Jackson, F., Titus, A. L. &amp; Albright III, L. B. (2011) A gravid fossil turtle from the Upper Cretaceous (Campanian) Kaiparowits Formation, southern Utah. Historical Biology, 23, 57 - 62. http: // dx. doi. org / 10.1080 / 08912963.2010.499167" type="journal article" year="2011">
Knell
<emphasis id="B917EAA18814FF897430FC99E6753DB3" box="[788,837,1215,1240]" italics="true" pageId="2" pageNumber="189">et al</emphasis>
. 2011
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), except for those from the Jurassic of
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(
<bibRefCitation id="EFF24B428814FF8977BAFCBCE41D3D96" author="Hirsch, K. F." box="[158,301,1252,1277]" pageId="2" pageNumber="189" pagination="752 - 762" refId="ref4371" refString="Hirsch, K. F. (1996) Parataxonomic classification of fossil chelonian and gecko eggs. Journal of Vertebrate Paleontology, 16, 752 - 762. http: // dx. doi. org / 10.1080 / 02724634.1996.10011363" type="journal article" year="1996">Hirsch 1996</bibRefCitation>
;
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),
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(
<bibRefCitation id="EFF24B428814FF8975F8FCBCE6F43D96" author="Bray, E. S. &amp; Hirsch, K. F." box="[732,964,1252,1277]" pageId="2" pageNumber="189" pagination="219 - 240" refId="ref3435" refString="Bray, E. S. &amp; Hirsch, K. F. (1998) Eggshells from the Upper Jurassic Morrison Formation. Modern Geology, 23, 219 - 240." type="journal article" year="1998">Bray &amp; Hirsch 1998</bibRefCitation>
) and
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(
<bibRefCitation id="EFF24B428814FF897372FCBCE02C3D96" author="Wang, Q. &amp; Wang, X. &amp; Zhao, Z. &amp; Zhang, J. &amp; Jiang, S." box="[1110,1308,1252,1277]" pageId="2" pageNumber="189" pagination="103 - 111" refId="ref5377" refString="Wang, Q., Wang, X., Zhao, Z., Zhang, J. &amp; Jiang, S. (2013) New turtle egg fossil from the Upper Cretaceous of the Laiyang Baisn, Shandong Province, China. Anais da Academia Brasileira de Ciencias, 85 (1), 103 - 111." type="journal article" year="2013">
Wang
<emphasis id="B917EAA18814FF8973BBFCBDE1E13D96" box="[1183,1233,1252,1277]" italics="true" pageId="2" pageNumber="189">et al</emphasis>
. 2013
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). The outer surface of the egg shows folded areas that suggest that the eggshell was flexible before its fossilization (
<figureCitation id="13582A368814FF89723CFD5FE0A03C4B" box="[1304,1424,1287,1312]" captionStart="FIGURE 2" captionStartId="3.[151,250,1799,1821]" captionTargetBox="[302,1252,193,1780]" captionTargetId="figure-175@3.[302,1252,193,1780]" captionTargetPageId="3" captionText="FIGURE 2. A, LPRP-USP 0052 in two views showing the missing pole and the folded eggshell. B, MO of a thin section of LPRP-USP 0052 eggshell. C, interpretative drawing of the thin section. Gray color in represents the additional cuticle layer above the units. D, E, F and G, SEM images of LPRP-USP 0052. D, eggshell in radial section showing the additional cuticle layer. Black arrow indicates the boundary between the biomineralized eggshell and the cuticle. E, eggshell in radial section showing two loosely abutted and triangular basic units and the “cavern” between them (translucent triangle). Black arrows point to the large primary spherites, from which the acicular aragonitic crystals radiate. F, outer surface of the eggshell showing two pore apertures (black arrows). Note that the shell basic units outlines are not easily seen. G, Enlargment of the rounded pore aperture (black arrow)." figureDoi="http://doi.org/10.5281/zenodo.4948050" httpUri="https://zenodo.org/record/4948050/files/figure.png" pageId="2" pageNumber="189">figure 2 A</figureCitation>
). A different
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of sediment fills the egg from its damaged portion compared to that surrounding the egg, suggesting that it was buried and fossilized without the pole, thus helping keeping its biological shape intact. The missing pole may also indicate that the egg had hatched, which is independently supported by CT analysis not revealing any embryonic remains. The WDS analysis of the eggshell indicate calcium as the main component of the eggshell crystalline structure, suggesting that diagenetic modifications had been minimal.
</paragraph>
</subSubSection>
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The egg outer surface is smooth, differing from the undulated and rough surfaces of turtle eggs from the Jurassic of the
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(
<bibRefCitation id="EFF24B428814FF8976DDFE5CE7DD3F76" author="Bray, E. S. &amp; Hirsch, K. F." box="[505,749,1540,1565]" pageId="2" pageNumber="189" pagination="219 - 240" refId="ref3435" refString="Bray, E. S. &amp; Hirsch, K. F. (1998) Eggshells from the Upper Jurassic Morrison Formation. Modern Geology, 23, 219 - 240." type="journal article" year="1998">Bray &amp; Hirsch 1998</bibRefCitation>
) and the Cretaceous of
<collectingCountry id="F37476238814FF897328FE5CE1613F76" box="[1036,1105,1540,1565]" name="Brazil" pageId="2" pageNumber="189">Brazil</collectingCountry>
(
<bibRefCitation id="EFF24B428814FF897340FE5CE0693F76" author="Azevedo, S. A. &amp; Gallo, V. &amp; Ferigolo, J." box="[1124,1369,1540,1565]" pageId="2" pageNumber="189" pagination="187 - 193" refId="ref3026" refString="Azevedo, S. A., Gallo, V. &amp; Ferigolo, J. (2000) A possible chelonian egg from the Brazilian Late Cretaceous. Anais da Academia Brasileira de Ciencias, 72, 187 - 193. http: // dx. doi. org / 10.1590 / s 0001 - 37652000000200007" type="journal article" year="2000">
Azevedo
<emphasis id="B917EAA18814FF8973F1FE5DE03A3F76" box="[1237,1290,1540,1565]" italics="true" pageId="2" pageNumber="189">et al</emphasis>
. 2000
</bibRefCitation>
). The eggshell is relatively thin (
<quantity id="4C9B9B568814FF8976E3FE7FE76A3F2B" box="[455,602,1575,1600]" metricMagnitude="-1" metricUnit="m" metricValue="1.525" metricValueMax="1.6" metricValueMin="1.45" pageId="2" pageNumber="189" unit="mm" value="152.5" valueMax="160.0" valueMin="145.0">145160 µm</quantity>
thick), including the biomineralized layer and an additional cuticle layer of about
<quantity id="4C9B9B568814FF8977D8FE14E4083F0F" box="[252,312,1612,1636]" metricMagnitude="-3" metricUnit="m" metricValue="7.0" pageId="2" pageNumber="189" unit="mm" value="7.0">7 µm</quantity>
thick (
<figureCitation id="13582A368814FF8976A3FE14E77E3F0E" box="[391,590,1612,1637]" captionStart="FIGURE 2" captionStartId="3.[151,250,1799,1821]" captionTargetBox="[302,1252,193,1780]" captionTargetId="figure-175@3.[302,1252,193,1780]" captionTargetPageId="3" captionText="FIGURE 2. A, LPRP-USP 0052 in two views showing the missing pole and the folded eggshell. B, MO of a thin section of LPRP-USP 0052 eggshell. C, interpretative drawing of the thin section. Gray color in represents the additional cuticle layer above the units. D, E, F and G, SEM images of LPRP-USP 0052. D, eggshell in radial section showing the additional cuticle layer. Black arrow indicates the boundary between the biomineralized eggshell and the cuticle. E, eggshell in radial section showing two loosely abutted and triangular basic units and the “cavern” between them (translucent triangle). Black arrows point to the large primary spherites, from which the acicular aragonitic crystals radiate. F, outer surface of the eggshell showing two pore apertures (black arrows). Note that the shell basic units outlines are not easily seen. G, Enlargment of the rounded pore aperture (black arrow)." figureDoi="http://doi.org/10.5281/zenodo.4948050" httpUri="https://zenodo.org/record/4948050/files/figure.png" pageId="2" pageNumber="189">figures 2 B, DE</figureCitation>
). Turtle fossil eggshells with a similar thickness are inferred to have either flexible or rigid eggs, e.g.
<emphasis id="B917EAA18814FF8976F7FE37E6733FE3" box="[467,835,1647,1672]" italics="true" pageId="2" pageNumber="189">Testudooflexoolithus bathonicae</emphasis>
(Hirsh 1996;
<bibRefCitation id="EFF24B428814FF8974C8FE28E1D13FE3" author="Bray, E. S. &amp; Hirsch, K. F." box="[1004,1249,1647,1672]" pageId="2" pageNumber="189" pagination="219 - 240" refId="ref3435" refString="Bray, E. S. &amp; Hirsch, K. F. (1998) Eggshells from the Upper Jurassic Morrison Formation. Modern Geology, 23, 219 - 240." type="journal article" year="1998">Bray &amp; Hirsch 1998</bibRefCitation>
),
<taxonomicName id="4C634D308814FF8973DCFE37E4AD3FC6" authority="(Kohring 1999)" baseAuthorityName="Kohring" baseAuthorityYear="1999" class="Reptilia" family="Testudoolithidae" genus="Testudoolithus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="2" pageNumber="189" phylum="Chordata" rank="species" species="hirschi">
<emphasis id="B917EAA18814FF8973DCFE37E5D53FC6" italics="true" pageId="2" pageNumber="189">Testudoolithus hirschi</emphasis>
(
<bibRefCitation id="EFF24B428814FF8977D7FECCE4A53FC6" author="Kohring, R." box="[243,405,1684,1709]" pageId="2" pageNumber="189" pagination="1 - 307" refId="ref4712" refString="Kohring, R. (1999) Structure, biostratinomy and systematic and phylogenetic relevance of egg shells of vertebrate amniotes. Courier Forschungsinstitut Senckenberg, 210, 1 - 307, A 1 - A 6." type="journal article" year="1999">Kohring 1999</bibRefCitation>
)
</taxonomicName>
, eggs from the Jurassic of Colorado (
<bibRefCitation id="EFF24B428814FF89741AFECCE1173FC6" author="Bray, E. S. &amp; Hirsch, K. F." box="[830,1063,1684,1709]" pageId="2" pageNumber="189" pagination="219 - 240" refId="ref3435" refString="Bray, E. S. &amp; Hirsch, K. F. (1998) Eggshells from the Upper Jurassic Morrison Formation. Modern Geology, 23, 219 - 240." type="journal article" year="1998">Bray &amp; Hirsch 1998</bibRefCitation>
),
<taxonomicName id="4C634D308814FF89731FFECCE42B3FBB" authority="(Schleich et al. 1988)" baseAuthorityName="Schleich" baseAuthorityYear="1988" class="Reptilia" genus="Haininchelys" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="2" pageNumber="189" phylum="Chordata" rank="species" species="curiosa">
<emphasis id="B917EAA18814FF89731FFECCE01B3FC7" box="[1083,1323,1684,1709]" italics="true" pageId="2" pageNumber="189">Haininchelys curiosa</emphasis>
(Schleich
<emphasis id="B917EAA18814FF8977B3FEE1E5FA3FBB" box="[151,202,1719,1744]" italics="true" pageId="2" pageNumber="189">et al</emphasis>
. 1988)
</taxonomicName>
and
<emphasis id="B917EAA18814FF897671FEEFE7053FBB" box="[341,565,1719,1744]" italics="true" pageId="2" pageNumber="189">Testudinarum ovum</emphasis>
(Schleich &amp; Kästle 1988; Schleich
<emphasis id="B917EAA18814FF8974F7FEE1E1353FBB" box="[979,1029,1719,1744]" italics="true" pageId="2" pageNumber="189">et al</emphasis>
. 1988).
<bibRefCitation id="EFF24B428814FF897343FEEFE0363FBB" author="Hirsch, K. F." box="[1127,1286,1719,1744]" pageId="2" pageNumber="189" pagination="382 - 397" refId="ref4336" refString="Hirsch, K. F. (1983) Contemporary and fossil chelonian eggshells. Copeia, 1983, 382 - 397. http: // dx. doi. org / 10.2307 / 1444381" type="journal article" year="1983">Hirsch (1983)</bibRefCitation>
noticed that, along with to the degree of rigidity, eggshell thickness may be ecological and biological indicators. The sea turtle
<taxonomicName id="4C634D308814FF8977B3FEA7E44E3E73" authorityName="Engelmann Et Al" authorityYear="1993" box="[151,382,1791,1816]" class="Reptilia" family="Cheloniidae" genus="Lepidochelys" kingdom="Animalia" order="Testudines" pageId="2" pageNumber="189" phylum="Chordata" rank="species" species="kempi">
<emphasis id="B917EAA18814FF8977B3FEA7E44E3E73" box="[151,382,1791,1816]" italics="true" pageId="2" pageNumber="189">Lepidochelys kempi</emphasis>
</taxonomicName>
lays highly pliable eggs, with eggshells about
<quantity id="4C9B9B568814FF8974F4FEA7E1113E73" box="[976,1057,1791,1816]" metricMagnitude="-2" metricUnit="m" metricValue="4.0" pageId="2" pageNumber="189" unit="mm" value="40.0">40 µm</quantity>
thick; the tortoise
<taxonomicName id="4C634D308814FF897231FEA7E41D3E56" authorityName="Ernst &amp; Barbour" authorityYear="1989" class="Reptilia" family="Testudinidae" genus="Geochelone" kingdom="Animalia" order="Testudines" pageId="2" pageNumber="189" phylum="Chordata" rank="species" species="elephantopus">
<emphasis id="B917EAA18814FF897231FEA7E41D3E56" italics="true" pageId="2" pageNumber="189">Geochelone elephantopus</emphasis>
</taxonomicName>
lays rigid-shelled eggs, with eggshells about
<quantity id="4C9B9B568814FF89744CFF7CE6F83E57" box="[872,968,1828,1853]" metricMagnitude="-1" metricUnit="m" metricValue="4.0" pageId="2" pageNumber="189" unit="mm" value="400.0">400 µm</quantity>
thick; the fresh-water turtle
<taxonomicName id="4C634D308814FF897217FF7CE43E3E34" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" class="Reptilia" family="Chelydridae" genus="Chelydra" kingdom="Animalia" order="Testudines" pageId="2" pageNumber="189" phylum="Chordata" rank="species" species="serpentina">
<emphasis id="B917EAA18814FF897217FF7CE43E3E34" italics="true" pageId="2" pageNumber="189">Chelydra serpentina</emphasis>
</taxonomicName>
has moderately flexible eggs, with
<quantity id="4C9B9B568814FF8975BBFF1FE7C53E0B" box="[671,757,1863,1888]" metricMagnitude="-1" metricUnit="m" metricValue="1.1" pageId="2" pageNumber="189" unit="mm" value="110.0">110 µm</quantity>
thick eggshells, although other fresh-water turtles also have rigid eggshells. The
<taxonomicName id="4C634D308814FF89765AFF34E7563EEF" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[382,614,1900,1925]" class="Reptilia" family="Chelydridae" genus="Chelydra" kingdom="Animalia" order="Testudines" pageId="2" pageNumber="189" phylum="Chordata" rank="species" species="serpentina">
<emphasis id="B917EAA18814FF89765AFF34E7563EEF" box="[382,614,1900,1925]" italics="true" pageId="2" pageNumber="189">Chelydra serpentina</emphasis>
</taxonomicName>
condition best compares to that of LPRP-USP 0052, which is congruent with the host freshwater deposits. In addition, the thickness of the additional cuticle layer of LPRP-USP 0052 resembles that of the other freshwater taxon
<taxonomicName id="4C634D308814FF8975BEFFECE1FA3EA6" authority="(Winkler 2006)" baseAuthorityName="Winkler" baseAuthorityYear="2006" box="[666,1226,1972,1997]" class="Reptilia" family="Podocnemididae" genus="Erymnochelys" kingdom="Animalia" order="Testudines" pageId="2" pageNumber="189" phylum="Chordata" rank="species" species="madagascariensis">
<emphasis id="B917EAA18814FF8975BEFFECE13F3EA6" box="[666,1039,1972,1997]" italics="true" pageId="2" pageNumber="189">Erymnochelys madagascariensis</emphasis>
(Winkler 2006)
</taxonomicName>
. The functionality of the cuticle layer in turtles is rarely mentioned, however, analogous cuticle structures in bird eggs are directly related to nesting in wet conditions, as described in
<taxonomicName id="4C634D308812FF8F75C9F8CFE6A039DB" authorityName="Lesson" authorityYear="1831" box="[749,912,151,176]" class="Aves" family="Megapodiidae" kingdom="Animalia" order="Craciformes" pageId="4" pageNumber="191" phylum="Chordata" rank="family">Megapodiidae</taxonomicName>
,
<taxonomicName id="4C634D308812FF8F74B9F8CFE10C39DB" box="[925,1084,151,176]" class="Aves" family="Podicipedidae" kingdom="Animalia" order="Podicipediformes" pageId="4" pageNumber="191" phylum="Chordata" rank="family">Podicipedidae</taxonomicName>
and
<taxonomicName id="4C634D308812FF8F7352F8CFE00B39DB" box="[1142,1339,151,176]" class="Aves" family="Phoenicopteridae" kingdom="Animalia" order="Phoenicopteriformes" pageId="4" pageNumber="191" phylum="Chordata" rank="family">Phoenicopteridae</taxonomicName>
, and are thought to preclude the blocking of pores apertures by foreign material to faciliate gas exchange and limit chemical erosion from microorganisms in the soil (
<bibRefCitation id="EFF24B428812FF8F75A8F887E62B3993" author="Board, R. G." box="[652,795,223,248]" pageId="4" pageNumber="191" pagination="132 - 136" refId="ref3203" refString="Board, R. G. (1981) The microstructure of avian eggshells, adaptive significance and practical implications in aviculture. Wildfowl, 32, 132 - 136." type="journal article" year="1981">Board 1981</bibRefCitation>
;
<bibRefCitation id="EFF24B428812FF8F7409F887E1363993" author="Board, R. G. &amp; Perrott, H. R. &amp; Love, G. &amp; Seymour, R. S." box="[813,1030,223,248]" pageId="4" pageNumber="191" pagination="131 - 134" refId="ref3234" refString="Board, R. G., Perrott, H. R., Love, G. &amp; Seymour, R. S. (1982) A novel pore system in the eggshells of the Mallee fowl, Leipoa ocellata. Journal of Experimental Zoology, 220, 131 - 134." type="journal article" year="1982">
Board
<emphasis id="B917EAA18812FF8F745BF8B9E6873993" box="[895,951,223,248]" italics="true" pageId="4" pageNumber="191">et al</emphasis>
. 1982
</bibRefCitation>
;
<bibRefCitation id="EFF24B428812FF8F733CF887E0123993" author="Board, R. G. &amp; Sparks, N. H. C." box="[1048,1314,223,248]" pageId="4" pageNumber="191" pagination="53 - 70" refId="ref3288" refString="Board, R. G. &amp; Sparks, N. H. C. (1991) Shell structure and formation in avian eggs, In: Deeming, D. C. &amp; Ferguson, M. W. J. (Eds.), Egg incubation: its effects on embryonic development in birds and reptiles. Cambridge University Press, Cambridge, pp. 53 - 70." type="book chapter" year="1991">Board &amp; Sparks 1991</bibRefCitation>
;
<bibRefCitation id="EFF24B428812FF8F7210F887E4613876" author="Booth, D. T. &amp; Thompson, M. B." pageId="4" pageNumber="191" pagination="325 - 344" refId="ref3361" refString="Booth, D. T. &amp; Thompson, M. B. (1991) A comparison of reptilian eggs with those of megapode birds, In: Deeming, D. C. &amp; Ferguson, M. W. J. (Eds.), Egg incubation: its effects on embryonic development in birds and reptiles. Cambridge University Press, Cambridge, pp. 325 - 344." type="book chapter" year="1991">Booth &amp; Thompson 1991</bibRefCitation>
).
</paragraph>
<caption id="DF1C663B8815FF8877B3FF5FE4B03E9F" ID-DOI="http://doi.org/10.5281/zenodo.4948050" ID-Zenodo-Dep="4948050" httpUri="https://zenodo.org/record/4948050/files/figure.png" pageId="3" pageNumber="190" startId="3.[151,250,1799,1821]" targetBox="[302,1252,193,1780]" targetPageId="3">
<paragraph id="8BDC36B38815FF8877B3FF5FE4B03E9F" blockId="3.[151,1436,1799,2036]" pageId="3" pageNumber="190">
<emphasis id="B917EAA18815FF8877B3FF5FE41D3E77" bold="true" box="[151,301,1799,1821]" pageId="3" pageNumber="190">FIGURE 2. A</emphasis>
, LPRP-USP 0052 in two views showing the missing pole and the folded eggshell.
<emphasis id="B917EAA18815FF8873ADFF50E1AA3E76" bold="true" box="[1161,1178,1800,1821]" pageId="3" pageNumber="190">B</emphasis>
, MO of a thin section of LPRP-USP 0052 eggshell.
<emphasis id="B917EAA18815FF887694FF7EE4F23E57" bold="true" box="[432,450,1830,1852]" pageId="3" pageNumber="190">C</emphasis>
, interpretative drawing of the thin section. Gray color in represents the additional cuticle layer above the units.
<emphasis id="B917EAA18815FF887664FF1DE4623E31" bold="true" box="[320,338,1861,1882]" pageId="3" pageNumber="190">D</emphasis>
,
<emphasis id="B917EAA18815FF887644FF1DE4413E31" bold="true" box="[352,369,1861,1882]" pageId="3" pageNumber="190">E</emphasis>
,
<emphasis id="B917EAA18815FF88765AFF1DE4BD3E31" bold="true" box="[382,397,1861,1882]" pageId="3" pageNumber="190">F</emphasis>
and
<emphasis id="B917EAA18815FF8876E5FF1DE4E23E30" bold="true" box="[449,466,1861,1883]" pageId="3" pageNumber="190">G</emphasis>
, SEM images of LPRP-USP 0052.
<emphasis id="B917EAA18815FF887465FF1DE6633E31" bold="true" box="[833,851,1861,1882]" pageId="3" pageNumber="190">D</emphasis>
, eggshell in radial section showing the additional cuticle layer. Black arrow indicates the boundary between the biomineralized eggshell and the cuticle.
<emphasis id="B917EAA18815FF88735DFF3CE1BA3E12" bold="true" box="[1145,1162,1892,1913]" pageId="3" pageNumber="190">E</emphasis>
, eggshell in radial section showing two loosely abutted and triangular basic units and the “cavern” between them (translucent triangle). Black arrows point to the large primary spherites, from which the acicular aragonitic crystals radiate.
<emphasis id="B917EAA18815FF88732DFFF9E1263EDD" bold="true" box="[1033,1046,1953,1974]" pageId="3" pageNumber="190">F</emphasis>
, outer surface of the eggshell showing two pore apertures (black arrows). Note that the shell basic units outlines are not easily seen.
<emphasis id="B917EAA18815FF88731AFFE7E17F3EBE" bold="true" box="[1086,1103,1983,2005]" pageId="3" pageNumber="190">G</emphasis>
, Enlargment of the rounded pore aperture (black arrow).
</paragraph>
</caption>
<paragraph id="8BDC36B38812FF8F77E3F97FE5EF3D7B" blockId="4.[151,1436,151,1040]" pageId="4" pageNumber="191">
The outlines of the shell basic units are not easily seen on the outer surface of LPRP-USP 0052 (
<figureCitation id="13582A368812FF8F723DF97FE0BF382B" box="[1305,1423,295,320]" captionStart="FIGURE 2" captionStartId="3.[151,250,1799,1821]" captionTargetBox="[302,1252,193,1780]" captionTargetId="figure-175@3.[302,1252,193,1780]" captionTargetPageId="3" captionText="FIGURE 2. A, LPRP-USP 0052 in two views showing the missing pole and the folded eggshell. B, MO of a thin section of LPRP-USP 0052 eggshell. C, interpretative drawing of the thin section. Gray color in represents the additional cuticle layer above the units. D, E, F and G, SEM images of LPRP-USP 0052. D, eggshell in radial section showing the additional cuticle layer. Black arrow indicates the boundary between the biomineralized eggshell and the cuticle. E, eggshell in radial section showing two loosely abutted and triangular basic units and the “cavern” between them (translucent triangle). Black arrows point to the large primary spherites, from which the acicular aragonitic crystals radiate. F, outer surface of the eggshell showing two pore apertures (black arrows). Note that the shell basic units outlines are not easily seen. G, Enlargment of the rounded pore aperture (black arrow)." figureDoi="http://doi.org/10.5281/zenodo.4948050" httpUri="https://zenodo.org/record/4948050/files/figure.png" pageId="4" pageNumber="191">figure 2 F</figureCitation>
). Pore openings are very sparse (
<figureCitation id="13582A368812FF8F7533F914E7A8380E" box="[535,664,332,357]" captionStart="FIGURE 2" captionStartId="3.[151,250,1799,1821]" captionTargetBox="[302,1252,193,1780]" captionTargetId="figure-175@3.[302,1252,193,1780]" captionTargetPageId="3" captionText="FIGURE 2. A, LPRP-USP 0052 in two views showing the missing pole and the folded eggshell. B, MO of a thin section of LPRP-USP 0052 eggshell. C, interpretative drawing of the thin section. Gray color in represents the additional cuticle layer above the units. D, E, F and G, SEM images of LPRP-USP 0052. D, eggshell in radial section showing the additional cuticle layer. Black arrow indicates the boundary between the biomineralized eggshell and the cuticle. E, eggshell in radial section showing two loosely abutted and triangular basic units and the “cavern” between them (translucent triangle). Black arrows point to the large primary spherites, from which the acicular aragonitic crystals radiate. F, outer surface of the eggshell showing two pore apertures (black arrows). Note that the shell basic units outlines are not easily seen. G, Enlargment of the rounded pore aperture (black arrow)." figureDoi="http://doi.org/10.5281/zenodo.4948050" httpUri="https://zenodo.org/record/4948050/files/figure.png" pageId="4" pageNumber="191">figure 2 F</figureCitation>
), a condition also present in the eggs of the extant pleurodires
<taxonomicName id="4C634D308812FF8F77B3F937E67138E3" authority="(Winkler &amp; Sanchez-Villagra 2006)" baseAuthorityName="Winkler &amp; Sanchez-Villagra" baseAuthorityYear="2006" box="[151,833,367,392]" class="Reptilia" family="Podocnemididae" genus="Podocnemis" kingdom="Animalia" order="Testudines" pageId="4" pageNumber="191" phylum="Chordata" rank="species" species="unifilis">
<emphasis id="B917EAA18812FF8F77B3F937E44F38E3" box="[151,383,367,392]" italics="true" pageId="4" pageNumber="191">Podocnemis unifilis</emphasis>
(
<bibRefCitation id="EFF24B428812FF8F76B3F937E60938E3" author="Winkler, J. D. &amp; Sanchez-Villagra, M. R." box="[407,825,367,392]" pageId="4" pageNumber="191" pagination="641 - 646" refId="ref5521" refString="Winkler, J. D. &amp; Sanchez-Villagra, M. R. (2006) A nesting site and egg morphology of a Miocene turtle from Urumaco, Venezuela: Evidence of marine adaptations in Pelomedusoides. Palaeontology, 49, 641 - 646. http: // dx. doi. org / 10.1111 / j. 1475 - 4983.2006.00557. x" type="journal article" year="2006">Winkler &amp; Sánchez-Villagra 2006</bibRefCitation>
)
</taxonomicName>
,
<taxonomicName id="4C634D308812FF8F7472F937E1B938E3" baseAuthorityName="Mikan" baseAuthorityYear="1820" box="[854,1161,367,392]" class="Reptilia" family="Chelidae" genus="Hydromedusa" kingdom="Animalia" order="Testudines" pageId="4" pageNumber="191" phylum="Chordata" rank="species" species="maximiliani">
<emphasis id="B917EAA18812FF8F7472F937E1B938E3" box="[854,1161,367,392]" italics="true" pageId="4" pageNumber="191">Hydromedusa maximiliani</emphasis>
</taxonomicName>
,
<taxonomicName id="4C634D308812FF8F7384F937E05238E3" baseAuthorityName="Dumeril &amp; Bibron" baseAuthorityYear="1835" box="[1184,1378,367,392]" class="Reptilia" family="Chelidae" genus="Phrynops" kingdom="Animalia" order="Testudines" pageId="4" pageNumber="191" phylum="Chordata" rank="species" species="hilarii">
<emphasis id="B917EAA18812FF8F7384F937E05238E3" box="[1184,1378,367,392]" italics="true" pageId="4" pageNumber="191">Phrynops hilarii</emphasis>
</taxonomicName>
and
<taxonomicName id="4C634D308812FF8F77B3F9CCE70638C6" authority="(Winkler 2006)" baseAuthorityName="Winkler" baseAuthorityYear="2006" box="[151,566,404,429]" class="Reptilia" family="Chelidae" genus="Acanthochelys" kingdom="Animalia" order="Testudines" pageId="4" pageNumber="191" phylum="Chordata" rank="species" species="spixii">
<emphasis id="B917EAA18812FF8F77B3F9CCE44E38C7" box="[151,382,404,429]" italics="true" pageId="4" pageNumber="191">Acanthochelys spixii</emphasis>
(Winkler 2006)
</taxonomicName>
. The pores apertures are typical of those seen in podocnemidid eggs, but also of some chelids (Winkler 2006). The diameters range from
<quantity id="4C9B9B568812FF8F746FF9EFE6EC38BB" box="[843,988,439,464]" metricMagnitude="-2" metricUnit="m" metricValue="8.55" metricValueMax="9.5" metricValueMin="7.6" pageId="4" pageNumber="191" unit="mm" value="85.5" valueMax="95.0" valueMin="76.0">76 to 95 µm</quantity>
(
<figureCitation id="13582A368812FF8F74C9F9EFE18738BB" box="[1005,1207,439,464]" captionStart="FIGURE 2" captionStartId="3.[151,250,1799,1821]" captionTargetBox="[302,1252,193,1780]" captionTargetId="figure-175@3.[302,1252,193,1780]" captionTargetPageId="3" captionText="FIGURE 2. A, LPRP-USP 0052 in two views showing the missing pole and the folded eggshell. B, MO of a thin section of LPRP-USP 0052 eggshell. C, interpretative drawing of the thin section. Gray color in represents the additional cuticle layer above the units. D, E, F and G, SEM images of LPRP-USP 0052. D, eggshell in radial section showing the additional cuticle layer. Black arrow indicates the boundary between the biomineralized eggshell and the cuticle. E, eggshell in radial section showing two loosely abutted and triangular basic units and the “cavern” between them (translucent triangle). Black arrows point to the large primary spherites, from which the acicular aragonitic crystals radiate. F, outer surface of the eggshell showing two pore apertures (black arrows). Note that the shell basic units outlines are not easily seen. G, Enlargment of the rounded pore aperture (black arrow)." figureDoi="http://doi.org/10.5281/zenodo.4948050" httpUri="https://zenodo.org/record/4948050/files/figure.png" pageId="4" pageNumber="191">figure 2 F and G</figureCitation>
), differing from the podocnemidids
<taxonomicName id="4C634D308812FF8F766CF984E716389E" baseAuthorityName="Winkler &amp; Sanchez-Villagra" baseAuthorityYear="2006" box="[328,550,476,501]" class="Reptilia" family="Podocnemididae" genus="Podocnemis" kingdom="Animalia" order="Testudines" pageId="4" pageNumber="191" phylum="Chordata" rank="species" species="unifilis">
<emphasis id="B917EAA18812FF8F766CF984E716389E" box="[328,550,476,501]" italics="true" pageId="4" pageNumber="191">Podocnemis unifilis</emphasis>
</taxonomicName>
(pore openings about 27,7 µm wide) and
<taxonomicName id="4C634D308812FF8F74DCF984E15F389E" authorityName="Gaffney &amp; Wood" authorityYear="2002" box="[1016,1135,476,501]" class="Reptilia" family="Podocnemididae" genus="Bairdemys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="4" pageNumber="191" phylum="Chordata" rank="genus">
<emphasis id="B917EAA18812FF8F74DCF984E15F389E" box="[1016,1135,476,501]" italics="true" pageId="4" pageNumber="191">Bairdemys</emphasis>
</taxonomicName>
(pores openings about
<quantity id="4C9B9B568812FF8F7256F984E4253B73" metricMagnitude="-1" metricUnit="m" metricValue="1.85" metricValueMax="2.0" metricValueMin="1.7" pageId="4" pageNumber="191" unit="mm" value="185.0" valueMax="200.0" valueMin="170.0">170 to 200 µm</quantity>
wide,
<bibRefCitation id="EFF24B428812FF8F764EF9A7E7CC3B73" author="Winkler, J. D. &amp; Sanchez-Villagra, M. R." box="[362,764,511,536]" pageId="4" pageNumber="191" pagination="641 - 646" refId="ref5521" refString="Winkler, J. D. &amp; Sanchez-Villagra, M. R. (2006) A nesting site and egg morphology of a Miocene turtle from Urumaco, Venezuela: Evidence of marine adaptations in Pelomedusoides. Palaeontology, 49, 641 - 646. http: // dx. doi. org / 10.1111 / j. 1475 - 4983.2006.00557. x" type="journal article" year="2006">Winkler &amp; Sánchez-Villagra 2006</bibRefCitation>
). In radial view, the shell units display the characteristic acicular crystallographic pattern of aragonitic calcium carbonate crystals (
<figureCitation id="13582A368812FF8F7324FA7CE1B03B56" box="[1024,1152,548,573]" captionStart="FIGURE 2" captionStartId="3.[151,250,1799,1821]" captionTargetBox="[302,1252,193,1780]" captionTargetId="figure-175@3.[302,1252,193,1780]" captionTargetPageId="3" captionText="FIGURE 2. A, LPRP-USP 0052 in two views showing the missing pole and the folded eggshell. B, MO of a thin section of LPRP-USP 0052 eggshell. C, interpretative drawing of the thin section. Gray color in represents the additional cuticle layer above the units. D, E, F and G, SEM images of LPRP-USP 0052. D, eggshell in radial section showing the additional cuticle layer. Black arrow indicates the boundary between the biomineralized eggshell and the cuticle. E, eggshell in radial section showing two loosely abutted and triangular basic units and the “cavern” between them (translucent triangle). Black arrows point to the large primary spherites, from which the acicular aragonitic crystals radiate. F, outer surface of the eggshell showing two pore apertures (black arrows). Note that the shell basic units outlines are not easily seen. G, Enlargment of the rounded pore aperture (black arrow)." figureDoi="http://doi.org/10.5281/zenodo.4948050" httpUri="https://zenodo.org/record/4948050/files/figure.png" pageId="4" pageNumber="191">figure 2 E</figureCitation>
), which is considered a
<taxonomicName id="4C634D308812FF8F77B3FA1FE43E3B0B" authorityName="Batsch" authorityYear="1788" box="[151,270,583,608]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="4" pageNumber="191" phylum="Chordata" rank="order">Testudines</taxonomicName>
apomorphy (
<bibRefCitation id="EFF24B428812FF8F7687FA1FE7013B0B" author="Young, J. D." box="[419,561,583,608]" pageId="4" pageNumber="191" pagination="455 - 469" refId="ref5585" refString="Young, J. D. (1950) The structure and physical properties of the testudinian eggshell. Proceedings of the Zoological Society of London, 120, 455 - 469. http: // dx. doi. org / 10.1111 / j. 1096 - 3642.1950. tb 00656. x" type="journal article" year="1950">Young 1950</bibRefCitation>
;
<bibRefCitation id="EFF24B428812FF8F7518FA1FE7F93B0B" author="Hirsch, K. F." box="[572,713,583,608]" pageId="4" pageNumber="191" pagination="382 - 397" refId="ref4336" refString="Hirsch, K. F. (1983) Contemporary and fossil chelonian eggshells. Copeia, 1983, 382 - 397. http: // dx. doi. org / 10.2307 / 1444381" type="journal article" year="1983">Hirsch 1983</bibRefCitation>
,
<bibRefCitation id="EFF24B428812FF8F75F0FA1FE63F3B0B" author="Hirsch, K. F." box="[724,783,583,608]" pageId="4" pageNumber="191" pagination="752 - 762" refId="ref4371" refString="Hirsch, K. F. (1996) Parataxonomic classification of fossil chelonian and gecko eggs. Journal of Vertebrate Paleontology, 16, 752 - 762. http: // dx. doi. org / 10.1080 / 02724634.1996.10011363" type="journal article" year="1996">1996</bibRefCitation>
;
<bibRefCitation id="EFF24B428812FF8F743FFA1FE1053B0B" author="Packard, G. C. &amp; Packard, M. J." box="[795,1077,583,608]" pageId="4" pageNumber="191" pagination="387 - 438" refId="ref4860" refString="Packard, G. C. &amp; Packard, M. J. (1988) The physiological ecology of reptilian eggs and embryos. In: Gans, C. &amp; Huey, R. B. (Eds), Biology of the Reptilia: Ecology (B) - Defense and life history. Vol. 16. Branta Books, New York, pp. 387 - 438." type="book chapter" year="1988">Packard &amp; Packard 1988</bibRefCitation>
; Winkler 2006). The metastable calcium carbonate crystals project radially from the large primary spherites (
<figureCitation id="13582A368812FF8F7315FA34E1813BEE" box="[1073,1201,620,645]" captionStart="FIGURE 2" captionStartId="3.[151,250,1799,1821]" captionTargetBox="[302,1252,193,1780]" captionTargetId="figure-175@3.[302,1252,193,1780]" captionTargetPageId="3" captionText="FIGURE 2. A, LPRP-USP 0052 in two views showing the missing pole and the folded eggshell. B, MO of a thin section of LPRP-USP 0052 eggshell. C, interpretative drawing of the thin section. Gray color in represents the additional cuticle layer above the units. D, E, F and G, SEM images of LPRP-USP 0052. D, eggshell in radial section showing the additional cuticle layer. Black arrow indicates the boundary between the biomineralized eggshell and the cuticle. E, eggshell in radial section showing two loosely abutted and triangular basic units and the “cavern” between them (translucent triangle). Black arrows point to the large primary spherites, from which the acicular aragonitic crystals radiate. F, outer surface of the eggshell showing two pore apertures (black arrows). Note that the shell basic units outlines are not easily seen. G, Enlargment of the rounded pore aperture (black arrow)." figureDoi="http://doi.org/10.5281/zenodo.4948050" httpUri="https://zenodo.org/record/4948050/files/figure.png" pageId="4" pageNumber="191">figure 2 E</figureCitation>
) differing from the condition in chelids
<taxonomicName id="4C634D308812FF8F76A8FAD7E7333BCC" box="[396,515,655,680]" class="Reptilia" family="Chelidae" genus="Elseya" kingdom="Animalia" order="Testudines" pageId="4" pageNumber="191" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B917EAA18812FF8F76A8FAD7E4E73BC3" box="[396,471,655,680]" italics="true" pageId="4" pageNumber="191">Elseya</emphasis>
sp.
</taxonomicName>
and
<taxonomicName id="4C634D308812FF8F7560FAD7E6223BC3" baseAuthorityName="Schneider" baseAuthorityYear="1783" box="[580,786,655,680]" class="Reptilia" family="Chelidae" genus="Chelus" kingdom="Animalia" order="Testudines" pageId="4" pageNumber="191" phylum="Chordata" rank="species" species="fimbriatus">
<emphasis id="B917EAA18812FF8F7560FAD7E6223BC3" box="[580,786,655,680]" italics="true" pageId="4" pageNumber="191">Chelus fimbriatus</emphasis>
</taxonomicName>
and the podocnemidids
<taxonomicName id="4C634D308812FF8F731EFAD7E0563BC3" box="[1082,1382,655,680]" class="Reptilia" family="Podocnemididae" genus="Peltocephalus" kingdom="Animalia" order="Testudines" pageId="4" pageNumber="191" phylum="Chordata" rank="species" species="dumerliana">
<emphasis id="B917EAA18812FF8F731EFAD7E0563BC3" box="[1082,1382,655,680]" italics="true" pageId="4" pageNumber="191">Peltocephalus dumerliana</emphasis>
</taxonomicName>
and
<taxonomicName id="4C634D308812FF8F77B3FAECE7393BA6" baseAuthorityName="Winkler" baseAuthorityYear="2006" box="[151,521,692,717]" class="Reptilia" family="Podocnemididae" genus="Erymnochelys" kingdom="Animalia" order="Testudines" pageId="4" pageNumber="191" phylum="Chordata" rank="species" species="madagascariensis">
<emphasis id="B917EAA18812FF8F77B3FAECE7393BA6" box="[151,521,692,717]" italics="true" pageId="4" pageNumber="191">Erymnochelys madagascariensis</emphasis>
</taxonomicName>
, with no visible spherites (Winkler 2006). Most shell units of LPRP-USP 0052 are roughly triangular, without marked borders, but more columnar unities are also present. This diversity of shapes results in relatively loosely abutting shell units (
<figureCitation id="13582A368812FF8F7599FAA4E6963A7E" box="[701,934,764,789]" captionStart="FIGURE 2" captionStartId="3.[151,250,1799,1821]" captionTargetBox="[302,1252,193,1780]" captionTargetId="figure-175@3.[302,1252,193,1780]" captionTargetPageId="3" captionText="FIGURE 2. A, LPRP-USP 0052 in two views showing the missing pole and the folded eggshell. B, MO of a thin section of LPRP-USP 0052 eggshell. C, interpretative drawing of the thin section. Gray color in represents the additional cuticle layer above the units. D, E, F and G, SEM images of LPRP-USP 0052. D, eggshell in radial section showing the additional cuticle layer. Black arrow indicates the boundary between the biomineralized eggshell and the cuticle. E, eggshell in radial section showing two loosely abutted and triangular basic units and the “cavern” between them (translucent triangle). Black arrows point to the large primary spherites, from which the acicular aragonitic crystals radiate. F, outer surface of the eggshell showing two pore apertures (black arrows). Note that the shell basic units outlines are not easily seen. G, Enlargment of the rounded pore aperture (black arrow)." figureDoi="http://doi.org/10.5281/zenodo.4948050" httpUri="https://zenodo.org/record/4948050/files/figure.png" pageId="4" pageNumber="191">figure 2 B, C and E</figureCitation>
). According to
<bibRefCitation id="EFF24B428812FF8F737DFAA4E5D43A53" author="Winkler, J. D. &amp; Sanchez-Villagra, M. R." pageId="4" pageNumber="191" pagination="641 - 646" refId="ref5521" refString="Winkler, J. D. &amp; Sanchez-Villagra, M. R. (2006) A nesting site and egg morphology of a Miocene turtle from Urumaco, Venezuela: Evidence of marine adaptations in Pelomedusoides. Palaeontology, 49, 641 - 646. http: // dx. doi. org / 10.1111 / j. 1475 - 4983.2006.00557. x" type="journal article" year="2006">Winkler &amp; Sánchez-Villagra (2006)</bibRefCitation>
, this allows “caverns” (large inter-units spaces) among shell units in some portions of the eggshell (
<figureCitation id="13582A368812FF8F7218FB47E0A73A53" box="[1340,1431,799,824]" captionStart="FIGURE 2" captionStartId="3.[151,250,1799,1821]" captionTargetBox="[302,1252,193,1780]" captionTargetId="figure-175@3.[302,1252,193,1780]" captionTargetPageId="3" captionText="FIGURE 2. A, LPRP-USP 0052 in two views showing the missing pole and the folded eggshell. B, MO of a thin section of LPRP-USP 0052 eggshell. C, interpretative drawing of the thin section. Gray color in represents the additional cuticle layer above the units. D, E, F and G, SEM images of LPRP-USP 0052. D, eggshell in radial section showing the additional cuticle layer. Black arrow indicates the boundary between the biomineralized eggshell and the cuticle. E, eggshell in radial section showing two loosely abutted and triangular basic units and the “cavern” between them (translucent triangle). Black arrows point to the large primary spherites, from which the acicular aragonitic crystals radiate. F, outer surface of the eggshell showing two pore apertures (black arrows). Note that the shell basic units outlines are not easily seen. G, Enlargment of the rounded pore aperture (black arrow)." figureDoi="http://doi.org/10.5281/zenodo.4948050" httpUri="https://zenodo.org/record/4948050/files/figure.png" pageId="4" pageNumber="191">figure 2</figureCitation>
, E), which are absent from rigid-shelled eggs, such as those of the podocnemidid
<taxonomicName id="4C634D308812FF8F7309FB1CE1943A36" authorityName="Gaffney &amp; Wood" authorityYear="2002" box="[1069,1188,836,861]" class="Reptilia" family="Podocnemididae" genus="Bairdemys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="4" pageNumber="191" phylum="Chordata" rank="genus">
<emphasis id="B917EAA18812FF8F7309FB1CE1943A36" box="[1069,1188,836,861]" italics="true" pageId="4" pageNumber="191">Bairdemys</emphasis>
</taxonomicName>
(Winkler &amp; Sánchez-Villagra 2006). On the contrary, the semi-flexible egg of
<taxonomicName id="4C634D308812FF8F743CFB3FE1A03AEB" authority="(Foote 1978)" baseAuthorityName="Foote" baseAuthorityYear="1978" box="[792,1168,871,896]" class="Reptilia" family="Podocnemididae" genus="Podocnemis" kingdom="Animalia" order="Testudines" pageId="4" pageNumber="191" phylum="Chordata" rank="species" species="unifilis">
<emphasis id="B917EAA18812FF8F743CFB3FE6C73AEB" box="[792,1015,871,896]" italics="true" pageId="4" pageNumber="191">Podocnemis unifilis</emphasis>
(
<bibRefCitation id="EFF24B428812FF8F7321FB3FE1B73AEB" author="Foote, R. W." box="[1029,1159,871,896]" pageId="4" pageNumber="191" pagination="333 - 339" refId="ref3939" refString="Foote, R. W. (1978) Nesting of Podocnemis unifilis (Testudines: Pelomedusidae) in the Colombian Amazon. Herpetologica, 34, 333 - 339." type="journal article" year="1978">Foote 1978</bibRefCitation>
)
</taxonomicName>
has a mix of areas with and without “caverns” (
<bibRefCitation id="EFF24B428812FF8F7685FBD4E6183ACE" author="Winkler, J. D. &amp; Sanchez-Villagra, M. R." box="[417,808,908,933]" pageId="4" pageNumber="191" pagination="641 - 646" refId="ref5521" refString="Winkler, J. D. &amp; Sanchez-Villagra, M. R. (2006) A nesting site and egg morphology of a Miocene turtle from Urumaco, Venezuela: Evidence of marine adaptations in Pelomedusoides. Palaeontology, 49, 641 - 646. http: // dx. doi. org / 10.1111 / j. 1475 - 4983.2006.00557. x" type="journal article" year="2006">Winkler &amp; Sánchez-Villagra 2006</bibRefCitation>
). Finally, shell units of LPRP-USP 0052 are, in average slightly higher than wide (high/width ratio of 1,11,2). This is also notable in flexible turtle fossil eggs, as such
<emphasis id="B917EAA18812FF8F77B3FB8CE7373A86" box="[151,519,980,1005]" italics="true" pageId="4" pageNumber="191">Testudooflexoolithus bathonicae</emphasis>
(Hirsh 1996;
<bibRefCitation id="EFF24B428812FF8F7596FB8CE6963A86" author="Bray, E. S. &amp; Hirsch, K. F." box="[690,934,980,1005]" pageId="4" pageNumber="191" pagination="219 - 240" refId="ref3435" refString="Bray, E. S. &amp; Hirsch, K. F. (1998) Eggshells from the Upper Jurassic Morrison Formation. Modern Geology, 23, 219 - 240." type="journal article" year="1998">Bray &amp; Hirsch 1998</bibRefCitation>
) and
<emphasis id="B917EAA18812FF8F74CBFB8CE00C3A87" box="[1007,1340,980,1005]" italics="true" pageId="4" pageNumber="191">Testudooflexoolithus agassizi</emphasis>
(
<bibRefCitation id="EFF24B428812FF8F726BFB8CE5E33D7B" author="Hirsch, K. F." pageId="4" pageNumber="191" pagination="752 - 762" refId="ref4371" refString="Hirsch, K. F. (1996) Parataxonomic classification of fossil chelonian and gecko eggs. Journal of Vertebrate Paleontology, 16, 752 - 762. http: // dx. doi. org / 10.1080 / 02724634.1996.10011363" type="journal article" year="1996">Hirsch 1996</bibRefCitation>
).
</paragraph>
</subSubSection>
</subSection>
</treatment>
</document>