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<taxonomicName id="4C774D5EFF1AFFDCCCEA1D2BFD5AFB87" authority="Baur, 1887" authorityName="Baur" authorityYear="1887" box="[410,727,1109,1134]" class="Reptilia" family="Proganochelyidae" genus="Proganochelys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="11" pageNumber="665" phylum="Chordata" rank="species" species="quenstedtii">
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Most of the humeral cross section of SMF 09-F2 (about 77% of the lateromedial diameter) is formed by a spongious inner medullary region (
<figureCitation id="134C2A58FF1AFFDCCC3D1DCAFE34FB24" box="[333,441,1204,1231]" captionStart="FIG" captionStartId="8.[132,143,1940,1957]" captionTargetBox="[173,1414,216,1879]" captionTargetId="figure-173@8.[745,1320,770,1326]" captionTargetPageId="8" captionText="FIG. 2. — Proterochersis porebensis Szczygielski &amp; Sulej, 2016 (A-I) and Proganochelys quenstedtii Baur, 1887 (J, K), thin sections in normal transmitted (A, D, F, H, J), polarized (B, E, G, I, K), and polarized with quarter lambda plate I (C) light: A-E, right humeri:A-C, small ZPAL V. 39/439;D, E, middle-sized ZPAL V. 39/433; C adjacent (slightly more distal) to A and B; F-I, right femora:F, G, small ZPAL V. 39/499; H, I, large ZPAL V. 39/500; J, K, left humerus, large SMF 09-F2. Dorsal surface towards the top of the page, anterior surface to the left. Scale bar: 1 cm." pageId="11" pageNumber="644">Fig. 2J, K</figureCitation>
), which is typical for all turtles and independent of their respective life style (e.g.
<bibRefCitation id="EFE64B2CFF1AFFDCCFEE1DAAFF77FAE4" author="NAKAJIMA Y. &amp; HIRAYAMA R. &amp; ENDO H." pageId="11" pageNumber="644" pagination="719 - 734" refId="ref20289" refString="NAKAJIMA Y., HIRAYAMA R. &amp; ENDO H. 2014. - Turtle humeral microanatomy and its relationship to lifestyle. Biological Journal of the Linnean Society 112 (4): 719 - 734. https: // doi. org / 10.1111 / bij. 12336" type="journal article" year="2014">
Nakajima
<emphasis id="B903EACFFF1AFFDCCDF41D8BFF34FAE4" box="[132,185,1268,1294]" italics="true" pageId="11" pageNumber="644">et al.</emphasis>
2014
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). The cortical thickness varies between approximately
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and
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. The “pure” cortex (i.e., without any erosion or remodeling) is difficult to measure due to scattered erosion cavities and secondary osteons (see below). The medullary region consists of endosteal bone and irregularly formed medium-sized and small erosion cavities (
<figureCitation id="134C2A58FF1AFFDCCFE71CEDFC8EFA44" box="[663,771,1427,1454]" captionStart="FIG" captionStartId="8.[132,143,1940,1957]" captionTargetBox="[173,1414,216,1879]" captionTargetId="figure-173@8.[745,1320,770,1326]" captionTargetPageId="8" captionText="FIG. 2. — Proterochersis porebensis Szczygielski &amp; Sulej, 2016 (A-I) and Proganochelys quenstedtii Baur, 1887 (J, K), thin sections in normal transmitted (A, D, F, H, J), polarized (B, E, G, I, K), and polarized with quarter lambda plate I (C) light: A-E, right humeri:A-C, small ZPAL V. 39/439;D, E, middle-sized ZPAL V. 39/433; C adjacent (slightly more distal) to A and B; F-I, right femora:F, G, small ZPAL V. 39/499; H, I, large ZPAL V. 39/500; J, K, left humerus, large SMF 09-F2. Dorsal surface towards the top of the page, anterior surface to the left. Scale bar: 1 cm." pageId="11" pageNumber="644">Figs 2J, K</figureCitation>
;
<figureCitation id="134C2A58FF1AFFDCCDF41CCAFF5EFA24" box="[132,211,1460,1486]" captionStart="FIG" captionStartId="16.[132,143,1849,1866]" captionTargetBox="[199,1389,215,1802]" captionTargetId="figure-230@16.[747,1299,774,1336]" captionTargetPageId="16" captionText="FIG. 8. — Proganochelys quenstedtii Baur, 1887, thin sections of large left humerus SMF 09-F2: A, anterodorsal region showing predominantly parallel-fibered bone matrix with predominantly longitudinal primary canals and primary osteons and pathological exostosis; B, closeup of the pathology, showing osteocyte lacunae aggregations; C, D, posterodorsal region with parallel-fibered bone locally grading into higher organized, nearly lamellar bone, secondary remodeling and gradual gradation into the medullary region; E, F, ventral region locally showing more intensive remodeling and Sharpey’s fibers; G, posteroventral region showing intensive remodeling of the deeper cortex and numerous radially inclined primary and secondary osteons, H, interior of the medullary region presenting thick trabeculae composed of endosteal lamellar bone. A, D, F, G in polarized light, B, C, E in normal transmitted light, H in polarized light with 530 nm λ plate. Abbreviations and symbols: ELB, endosteal lamellar bone; IS, intertrabecular space; OPFB, higher organization of parallel-fibered bone; PAT, pathology; PFB, parallel-fibered bone; SF, Sharpey’s fibers; *, primary vascular canal; red +, primary osteon; black and white +, secondary osteon. Scale bars: 500 µm." pageId="11" pageNumber="644">8G, H</figureCitation>
). The medullary region grades into a perimedullary region (
<figureCitation id="134C2A58FF1AFFDCCC601CADFE09FA04" box="[272,388,1491,1518]" captionStart="FIG" captionStartId="8.[132,143,1940,1957]" captionTargetBox="[173,1414,216,1879]" captionTargetId="figure-173@8.[745,1320,770,1326]" captionTargetPageId="8" captionText="FIG. 2. — Proterochersis porebensis Szczygielski &amp; Sulej, 2016 (A-I) and Proganochelys quenstedtii Baur, 1887 (J, K), thin sections in normal transmitted (A, D, F, H, J), polarized (B, E, G, I, K), and polarized with quarter lambda plate I (C) light: A-E, right humeri:A-C, small ZPAL V. 39/439;D, E, middle-sized ZPAL V. 39/433; C adjacent (slightly more distal) to A and B; F-I, right femora:F, G, small ZPAL V. 39/499; H, I, large ZPAL V. 39/500; J, K, left humerus, large SMF 09-F2. Dorsal surface towards the top of the page, anterior surface to the left. Scale bar: 1 cm." pageId="11" pageNumber="644">Figs 2J, K</figureCitation>
;
<figureCitation id="134C2A58FF1AFFDCCCE41CADFE2DFA07" box="[404,416,1491,1517]" captionStart="FIG" captionStartId="16.[132,143,1849,1866]" captionTargetBox="[199,1389,215,1802]" captionTargetId="figure-230@16.[747,1299,774,1336]" captionTargetPageId="16" captionText="FIG. 8. — Proganochelys quenstedtii Baur, 1887, thin sections of large left humerus SMF 09-F2: A, anterodorsal region showing predominantly parallel-fibered bone matrix with predominantly longitudinal primary canals and primary osteons and pathological exostosis; B, closeup of the pathology, showing osteocyte lacunae aggregations; C, D, posterodorsal region with parallel-fibered bone locally grading into higher organized, nearly lamellar bone, secondary remodeling and gradual gradation into the medullary region; E, F, ventral region locally showing more intensive remodeling and Sharpey’s fibers; G, posteroventral region showing intensive remodeling of the deeper cortex and numerous radially inclined primary and secondary osteons, H, interior of the medullary region presenting thick trabeculae composed of endosteal lamellar bone. A, D, F, G in polarized light, B, C, E in normal transmitted light, H in polarized light with 530 nm λ plate. Abbreviations and symbols: ELB, endosteal lamellar bone; IS, intertrabecular space; OPFB, higher organization of parallel-fibered bone; PAT, pathology; PFB, parallel-fibered bone; SF, Sharpey’s fibers; *, primary vascular canal; red +, primary osteon; black and white +, secondary osteon. Scale bars: 500 µm." pageId="11" pageNumber="644">8</figureCitation>
C-F) where erosion cavities and endosteal bone are intermixed or grade into, respectively, large secondary osteons. Smaller scattered secondary osteons reach far into the outer cortex. Only the outer third of the cross section displays primary cortex (
<figureCitation id="134C2A58FF1AFFDCCF661F2DFD0FF987" box="[534,642,1619,1645]" captionStart="FIG" captionStartId="8.[132,143,1940,1957]" captionTargetBox="[173,1414,216,1879]" captionTargetId="figure-173@8.[745,1320,770,1326]" captionTargetPageId="8" captionText="FIG. 2. — Proterochersis porebensis Szczygielski &amp; Sulej, 2016 (A-I) and Proganochelys quenstedtii Baur, 1887 (J, K), thin sections in normal transmitted (A, D, F, H, J), polarized (B, E, G, I, K), and polarized with quarter lambda plate I (C) light: A-E, right humeri:A-C, small ZPAL V. 39/439;D, E, middle-sized ZPAL V. 39/433; C adjacent (slightly more distal) to A and B; F-I, right femora:F, G, small ZPAL V. 39/499; H, I, large ZPAL V. 39/500; J, K, left humerus, large SMF 09-F2. Dorsal surface towards the top of the page, anterior surface to the left. Scale bar: 1 cm." pageId="11" pageNumber="644">Figs 2J, K</figureCitation>
;
<figureCitation id="134C2A58FF1AFFDCCFFA1F2DFD15F987" box="[650,664,1619,1645]" captionStart="FIG" captionStartId="16.[132,143,1849,1866]" captionTargetBox="[199,1389,215,1802]" captionTargetId="figure-230@16.[747,1299,774,1336]" captionTargetPageId="16" captionText="FIG. 8. — Proganochelys quenstedtii Baur, 1887, thin sections of large left humerus SMF 09-F2: A, anterodorsal region showing predominantly parallel-fibered bone matrix with predominantly longitudinal primary canals and primary osteons and pathological exostosis; B, closeup of the pathology, showing osteocyte lacunae aggregations; C, D, posterodorsal region with parallel-fibered bone locally grading into higher organized, nearly lamellar bone, secondary remodeling and gradual gradation into the medullary region; E, F, ventral region locally showing more intensive remodeling and Sharpey’s fibers; G, posteroventral region showing intensive remodeling of the deeper cortex and numerous radially inclined primary and secondary osteons, H, interior of the medullary region presenting thick trabeculae composed of endosteal lamellar bone. A, D, F, G in polarized light, B, C, E in normal transmitted light, H in polarized light with 530 nm λ plate. Abbreviations and symbols: ELB, endosteal lamellar bone; IS, intertrabecular space; OPFB, higher organization of parallel-fibered bone; PAT, pathology; PFB, parallel-fibered bone; SF, Sharpey’s fibers; *, primary vascular canal; red +, primary osteon; black and white +, secondary osteon. Scale bars: 500 µm." pageId="11" pageNumber="644">8</figureCitation>
A-F). This periosteal tissue consists generally of in overall low vascularized parallel-fibered tissue, locally even grading into lamellar tissue. The inner preserved cortex is subsequently followed or partially intermixed with the perimedullary region. It shows less organized and less strongly vascularized parallel-fibered tissue. Small longitudinal and few reticular to radial simple vascular canals, as well as some longitudinal primary osteons occur, which are not ordered in a clear pattern. The number of osteocyte lacunae is low and they remain small and flattened throughout. At the dorsolateral bone side, the highly organized, poorly vascularized outer cortex is followed by local regions of less organized parallel-fibered tissue, indicating an area of temporary faster growth (
<figureCitation id="134C2A58FF1AFFDCC90E1B88FB64FCFA" box="[1150,1257,758,784]" captionStart="FIG" captionStartId="16.[132,143,1849,1866]" captionTargetBox="[199,1389,215,1802]" captionTargetId="figure-230@16.[747,1299,774,1336]" captionTargetPageId="16" captionText="FIG. 8. — Proganochelys quenstedtii Baur, 1887, thin sections of large left humerus SMF 09-F2: A, anterodorsal region showing predominantly parallel-fibered bone matrix with predominantly longitudinal primary canals and primary osteons and pathological exostosis; B, closeup of the pathology, showing osteocyte lacunae aggregations; C, D, posterodorsal region with parallel-fibered bone locally grading into higher organized, nearly lamellar bone, secondary remodeling and gradual gradation into the medullary region; E, F, ventral region locally showing more intensive remodeling and Sharpey’s fibers; G, posteroventral region showing intensive remodeling of the deeper cortex and numerous radially inclined primary and secondary osteons, H, interior of the medullary region presenting thick trabeculae composed of endosteal lamellar bone. A, D, F, G in polarized light, B, C, E in normal transmitted light, H in polarized light with 530 nm λ plate. Abbreviations and symbols: ELB, endosteal lamellar bone; IS, intertrabecular space; OPFB, higher organization of parallel-fibered bone; PAT, pathology; PFB, parallel-fibered bone; SF, Sharpey’s fibers; *, primary vascular canal; red +, primary osteon; black and white +, secondary osteon. Scale bars: 500 µm." pageId="11" pageNumber="644">Fig. 8A, B</figureCitation>
). This area is barely recognizable as a slight rugosity on the bone surface. It likely represents a kind of localized pathology, potentially a small surface exostosis (
<bibRefCitation id="EFE64B2CFF1AFFDCCE9E1A2BFB6EFC9A" author="ROTHSCHILD B. M. &amp; SCHULTZE H. - P. &amp; PELLEGRINI R." box="[1006,1251,853,880]" pageId="11" pageNumber="644" refId="ref21429" refString="ROTHSCHILD B. M., SCHULTZE H. - P. &amp; PELLEGRINI R. 2012. - Herpetological osteopathology. Springer, New York, 450 p. https: // doi. org / 10.1007 / 978 - 1 - 4614 - 0824 - 6" type="book" year="2012">
Rothschild
<emphasis id="B903EACFFF1AFFDCC91C1A28FB2FFC85" box="[1132,1186,853,879]" italics="true" pageId="11" pageNumber="644">et al.</emphasis>
2012
</bibRefCitation>
). The outer cortex contains 6-7 growth marks, indicated by thin annuli in the inner cortex and LAGs (lines of arrested growth) or multiple closely spaced rest lines in the outermost cortex. This resembles, at least locally, an outer circumferential layer (OCL)/ external fundamental system (EFS;
<bibRefCitation id="EFE64B2CFF1AFFDCC9DC1A8BFAF9FBE5" author="PONTON F. &amp; ELZANOWSKI A. &amp; CASTANET J. &amp; CHINSAMY A. &amp; DE MARGERIE E. &amp; RICQLES A. J. DE &amp; CUBO J." box="[1196,1396,1013,1039]" pageId="11" pageNumber="644" pagination="137 - 140" refId="ref20791" refString="PONTON F., ELZANOWSKI A., CASTANET J., CHINSAMY A., DE MARGERIE E., RICQLES A. J. DE &amp; CUBO J. 2004. - Variation of the outer circumferential layer in the limb bones of birds. Acta Ornithologica 39 (2): 137 - 140. https: // doi. org / 10.3161 / 068.039.0210" type="journal article" year="2004">
Ponton
<emphasis id="B903EACFFF1AFFDCC8721A88FAB8FBE5" box="[1282,1333,1013,1039]" italics="true" pageId="11" pageNumber="644">et al.</emphasis>
2004
</bibRefCitation>
), but is not deposited all around the outermost cortex of the cross section. Growth marks in the inner cortex, which would represent earlier ontogenetic stages, are already lost due to remodeling. Extensive secondary remodeling is also revealed along the bone, including the area of the ossification center, by CT data (
<figureCitation id="134C2A58FF1AFFDCCEC21DCAFB9DFB24" box="[946,1040,1204,1231]" captionStart="FIG" captionStartId="15.[132,143,1390,1407]" captionTargetBox="[132,1455,215,1347]" captionTargetId="figure-518@15.[698,1267,780,1348]" captionTargetPageId="15" captionText="FIG. 7. — CT slices of humeri (A-F) and femur (G, H) of Proterochersis porebensis Szczygielski &amp; Sulej, 2016 and humerus of Proganochelys quenstedtii Baur, 1887 (I-J), sagittal (A, C, E, G, I), and transverse at the level of the ossification center or as close as possible allowed by specimen preservation (B, D, F, H, J): A, B, small left humerus ZPAL V.39/446;C, D, small right humerus ZPAL V.39/156;E, F, large right humerus ZPAL V.39/50;G, H, large right femur ZPAL V.39/432; I, J, large left humerus SMF 09-F2. Dorsal surface to the right for A, C, E, G, I and towards the top of the page for B, D, F, H, J. Scale bar: 1 cm." pageId="11" pageNumber="644">Fig. 7I, J</figureCitation>
). There is no medullary cavity proximal or distal to the thin-sectioning plane. Based on the increase in tissue organization and the nearly avascular condition and increase of the number of growth marks in the outer cortex, growth rate was clearly reduced, and the specimen likely represents an adult individual close to or already fully-grown. This late ontogenetic stage of SMF 09-F2 is also supported by humerus and carapace lengths that are in the upper range of other known individuals of
<taxonomicName id="4C774D5EFF1AFFDCC93C1CCDFAC6FA27" authorityName="Baur" authorityYear="1887" box="[1100,1355,1459,1485]" class="Reptilia" family="Proganochelyidae" genus="Proganochelys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="11" pageNumber="644" phylum="Chordata" rank="species" species="quenstedtii">
<emphasis id="B903EACFFF1AFFDCC93C1CCDFAC6FA27" box="[1100,1355,1459,1485]" italics="true" pageId="11" pageNumber="644">Proganochelys quenstedtii</emphasis>
</taxonomicName>
(
<bibRefCitation id="EFE64B2CFF1AFFDCC82A1CCDFCE8FA07" author="GAFFNEY E. S." pageId="11" pageNumber="644" pagination="1 - 263" refId="ref18361" refString="GAFFNEY E. S. 1990. - The comparative osteology of the Triassic turtle Proganochelys. Bulletin of the American Museum of Natural History 194: 1 - 263." type="journal article" year="1990">Gaffney 1990</bibRefCitation>
;
<bibRefCitation id="EFE64B2CFF1AFFDCCE031CADFBB2FA07" author="SCHEYER T. M. &amp; KLEIN N. &amp; EVERS S. W. &amp; MAUTNER A. - K. &amp; PABST B." box="[883,1087,1491,1517]" pageId="11" pageNumber="644" refId="ref21642" refString="SCHEYER T. M., KLEIN N., EVERS S. W., MAUTNER A. - K. &amp; PABST B. 2022. - First evidence of Proganochelys quenstedtii (Testudinata) from the Plateosaurus bonebeds (Norian, Late Triassic) of Frick, Canton Aargau, Switzerland. Swiss Journal of Palaeontology 141: 17." type="journal volume" year="2022">
Scheyer
<emphasis id="B903EACFFF1AFFDCCEBC1CAAFB8DFA07" box="[972,1024,1491,1517]" italics="true" pageId="11" pageNumber="644">et al.</emphasis>
2022
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).
</paragraph>
</subSubSection>
</treatment>
</document>
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