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<mods:title id="BBFBA2F712822F8C946546A92330DEE6">Limb histology of the Triassic stem turtles Proterochersis porebensis Szczygielski &amp; Sulej, 2016 and Proganochelys quenstedtii Baur, 1887 with insights into growth patterns of early turtles</mods:title>
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All sectioned bones of
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<emphasis id="B903EACFFF16FFD0C9491FACFACCF906" box="[1081,1345,1746,1772]" italics="true" pageId="7" pageNumber="640">Proterochersis porebensis</emphasis>
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present a well-developed, compact cortex and medullary area filled with cancellous bone (
<figureCitation id="134C2A58FF16FFD0C9551E6CFBE8F8C6" box="[1061,1125,1810,1836]" 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="7" pageNumber="640">Fig. 2</figureCitation>
A-I). Thickness of the cortices (without erosion cavities and remodeling) is
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, and
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for the middle-sized and small humerus specimen and
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and
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for the large and small femora, respectively. The matrix of the cortex is composed predominantly of parallel-fibered bone (
<figureCitation id="134C2A58FF16FFD0C9801ECCFABEF826" box="[1264,1331,1970,1996]" 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="7" pageNumber="640">Figs 2</figureCitation>
A-I; 3A-D; 4A-F; 5; 6) that often shows high organization (approaching but not exactly the same as the lamellar bone in the same sections) and is even locally grading into lamellar bone in some samples, and in the middle-sized humerus ZPAL V. 39/433 (
<figureCitation id="134C2A58FF18FFDECDFD1849FF57FEB8" box="[141,218,311,338]" captionStart="FIG" captionStartId="12.[132,143,1849,1866]" captionTargetBox="[161,1427,215,1802]" captionTargetId="figure-220@12.[702,1246,774,1336]" captionTargetPageId="12" captionText="FIG. 4. — Proterochersis porebensis Szczygielski &amp; Sulej, 2016, thin sections of middle-sized right humerus ZPAL V. 39/433: A, B, anterodorsal region showing predominantly parallel-fibered bone matrix externally grading into higher organized, nearly lamellar bone with primary vasculature (richer in the deeper cortex than externally, longitudinal to radial) and undulating LAGs; C, D, dorsal region with sparse primary longitudinal vasculature and trabeculae incorporating cortical parallel-fibered tissue; E, F, anteroventral region showing coarse collagen bundles in the parallel-fibered matrix, longitudinal to radial predominantly primary vasculature with sparse secondary osteons, and undulating LAGs; G, H, interior of the medullary region presenting thick trabeculae composed of endosteal lamellar bone. A, C, E, G in normal transmitted light, B, D, F, H in polarized light. Abbreviations and symbols: ELB, endosteal lamellar bone; IS, intertrabecular space; PFB, parallel-fibered bone; OPFB, higher organization of parallel-fibered bone; *, primary vascular canal; red +, primary osteon; black and white +, secondary osteon. Lines of arrested growth (LAGs) indicated with arrowheads. Scale bars: 500 µm." pageId="9" pageNumber="642">Fig. 4B</figureCitation>
) and large femur ZPAL V. 39/500 (
<figureCitation id="134C2A58FF18FFDECF3F1849FD2DFEB8" box="[591,672,311,338]" captionStart="FIG" captionStartId="14.[132,143,1872,1889]" captionTargetBox="[150,1438,215,1825]" captionTargetId="figure-195@14.[701,1255,782,1352]" captionTargetPageId="14" captionText="FIG. 6. — Proterochersis porebensis Szczygielski &amp; Sulej, 2016, thin sections of large right femur ZPAL V. 39/500: A, B, dorsal region showing predominantly parallel-fibered bone matrix with longitudinal primary canals in the external part,numerous large,meandering secondary osteons in the deeper cortex,and gradual gradation into the medullary region;C, D, anterior region with parallel-fibered bone locally grading into higher organized,nearly lamellar bone;E, F, posteroventral region locally showing slightly less intensive invasion of remodeling, preserving undulating LAGs.Note the scalloped external surface. A, C, E in normal transmitted light, B, D, F in polarized light.Abbreviations and symbols: ELB, endosteal lamellar bone;IS, intertrabecular space; PFB, parallel-fibered bone;OPFB, higher organization of parallel-fibered bone; *, primary vascular canal; red +, primary osteon; black and white +, secondary osteon. Lines of arrested growth (LAGs) indicated with arrowheads. Scale bars: 500 µm." pageId="9" pageNumber="642">Fig. 6D</figureCitation>
) it locally takes a highly organized appearance, approaching lamellar bone. The matrix shows annular (ring-like) lines of arrested growth (LAGs), which appear relatively even in the small femur ZPAL V. 39/499 (
<figureCitation id="134C2A58FF18FFDECCFF18C9FE52FE3B" box="[399,479,439,465]" 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="9" pageNumber="642">Figs 2F</figureCitation>
;
<figureCitation id="134C2A58FF18FFDECC9B18C9FD81FE3B" box="[491,524,439,466]" captionStart="FIG" captionStartId="13.[131,142,1637,1654]" captionTargetBox="[132,1455,214,1595]" captionTargetId="figure-328@13.[698,1267,766,1320]" captionTargetPageId="13" captionText="FIG. 5. — Proterochersis porebensis Szczygielski &amp; Sulej, 2016, thin sections of small right femur ZPAL V. 39/499: A, B, anterior region showing parallel-fibered matrix with longitudinal primary canals and straight LAGs; C, D, posterior region with thinner and less vascularized cortex (note that the external cortex is destroyed posteroventrally) and indistinct growth marks.Despite the small size of the specimen and primary nature of the cortical tissue,the boundary between the cortex and medullary region reveals active remodeling evidenced by two to three generations of secondary lamellar bone (marked with red rectangles). A and C in normal transmitted light, B and D in polarized light. Abbreviations and symbols: ELB, endosteal lamellar bone; IS, intertrabecular space; PFB, parallel-fibered bone; *, primary vascular canal. Lines of arrested growth (LAGs) indicated with arrowheads. Scale bars: 500 µm." pageId="9" pageNumber="642">5A</figureCitation>
) and in the deeper part of the cortex of the small humerus ZPAL V. 39/439 (
<figureCitation id="134C2A58FF18FFDECFC018A9FC8FFE1B" box="[688,770,471,497]" 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="9" pageNumber="642">Figs 2A</figureCitation>
;
<figureCitation id="134C2A58FF18FFDECDF41889FF4BFDFB" box="[132,198,503,529]" captionStart="FIG" captionStartId="10.[132,143,1827,1844]" captionTargetBox="[132,1455,215,1667]" captionTargetId="figure-26@10.[698,1267,759,1305]" captionTargetPageId="10" captionText="FIG. 3. — Proterochersis porebensis Szczygielski &amp; Sulej, 2016, thin sections of small right humerus ZPAL V. 39/439: A, anterodorsal region showing relatively thin, nearly avascular cortex composed of parallel-fibered bone with gently undulating collagen fibers and thin trabeculae composed mostly of endosteal lamellar bone; B, dorsal region showing undulating collagen fiber layout parallel to bone surface and sparse collagen fibers oblique to the bone surface in the cortex, slightly increased longitudinal primary vasculature, and trabeculae incorporating cortical parallel-fibered bone; C, D, ventral region showing more developed vasculature (mostly primary canals with sporadic primary osteons, predominantly longitudinal), growth marks, and parallel-fibered matrix grading externally into more ordered, nearly lamellar bone; E, border between the cortex and the medullary region in the ventral part of the bone, adjacent (slightly more proximally) to A-C showing a large chunk of cortical parallel-fibered bone incorporated into the medullary region. A, B, D in polarized light with quarter lambda plate, C and E in normal transmitted light.Abbreviations and symbols: ELB, endosteal lamellar bone;IS, intertrabecular space; PFB, parallel-fibered bone;OPFB, higher organization of parallel-fibered bone; *, primary vascular canal; red +, primary osteon. Lines of arrested growth (LAGs) indicated with arrowheads. Scale bars: 500 µm." pageId="9" pageNumber="642">3C, E</figureCitation>
), but show more pronounced waviness in the external part of the cortex of ZPAL V. 39/439 (
<figureCitation id="134C2A58FF18FFDECF641B69FDE8FDDB" box="[532,613,535,561]" 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="9" pageNumber="642">Figs 2A</figureCitation>
;
<figureCitation id="134C2A58FF18FFDECF1E1B69FD1DFDDB" box="[622,656,535,561]" captionStart="FIG" captionStartId="10.[132,143,1827,1844]" captionTargetBox="[132,1455,215,1667]" captionTargetId="figure-26@10.[698,1267,759,1305]" captionTargetPageId="10" captionText="FIG. 3. — Proterochersis porebensis Szczygielski &amp; Sulej, 2016, thin sections of small right humerus ZPAL V. 39/439: A, anterodorsal region showing relatively thin, nearly avascular cortex composed of parallel-fibered bone with gently undulating collagen fibers and thin trabeculae composed mostly of endosteal lamellar bone; B, dorsal region showing undulating collagen fiber layout parallel to bone surface and sparse collagen fibers oblique to the bone surface in the cortex, slightly increased longitudinal primary vasculature, and trabeculae incorporating cortical parallel-fibered bone; C, D, ventral region showing more developed vasculature (mostly primary canals with sporadic primary osteons, predominantly longitudinal), growth marks, and parallel-fibered matrix grading externally into more ordered, nearly lamellar bone; E, border between the cortex and the medullary region in the ventral part of the bone, adjacent (slightly more proximally) to A-C showing a large chunk of cortical parallel-fibered bone incorporated into the medullary region. A, B, D in polarized light with quarter lambda plate, C and E in normal transmitted light.Abbreviations and symbols: ELB, endosteal lamellar bone;IS, intertrabecular space; PFB, parallel-fibered bone;OPFB, higher organization of parallel-fibered bone; *, primary vascular canal; red +, primary osteon. Lines of arrested growth (LAGs) indicated with arrowheads. Scale bars: 500 µm." pageId="9" pageNumber="642">3C</figureCitation>
) and in the larger ZPAL V. 39/433 (
<figureCitation id="134C2A58FF18FFDECCF21B48FE55FDBB" box="[386,472,566,593]" 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="9" pageNumber="642">Figs 2D</figureCitation>
;
<figureCitation id="134C2A58FF18FFDECC911B49FD90FDBB" box="[481,541,567,593]" captionStart="FIG" captionStartId="12.[132,143,1849,1866]" captionTargetBox="[161,1427,215,1802]" captionTargetId="figure-220@12.[702,1246,774,1336]" captionTargetPageId="12" captionText="FIG. 4. — Proterochersis porebensis Szczygielski &amp; Sulej, 2016, thin sections of middle-sized right humerus ZPAL V. 39/433: A, B, anterodorsal region showing predominantly parallel-fibered bone matrix externally grading into higher organized, nearly lamellar bone with primary vasculature (richer in the deeper cortex than externally, longitudinal to radial) and undulating LAGs; C, D, dorsal region with sparse primary longitudinal vasculature and trabeculae incorporating cortical parallel-fibered tissue; E, F, anteroventral region showing coarse collagen bundles in the parallel-fibered matrix, longitudinal to radial predominantly primary vasculature with sparse secondary osteons, and undulating LAGs; G, H, interior of the medullary region presenting thick trabeculae composed of endosteal lamellar bone. A, C, E, G in normal transmitted light, B, D, F, H in polarized light. Abbreviations and symbols: ELB, endosteal lamellar bone; IS, intertrabecular space; PFB, parallel-fibered bone; OPFB, higher organization of parallel-fibered bone; *, primary vascular canal; red +, primary osteon; black and white +, secondary osteon. Lines of arrested growth (LAGs) indicated with arrowheads. Scale bars: 500 µm." pageId="9" pageNumber="642">4A, E</figureCitation>
) and ZPAL V. 39/500 (
<figureCitation id="134C2A58FF18FFDECDFD1B28FF6AFD9B" box="[141,231,598,625]" 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="9" pageNumber="642">Figs 2H</figureCitation>
;
<figureCitation id="134C2A58FF18FFDECD831B28FE98FD9B" box="[243,277,598,625]" captionStart="FIG" captionStartId="14.[132,143,1872,1889]" captionTargetBox="[150,1438,215,1825]" captionTargetId="figure-195@14.[701,1255,782,1352]" captionTargetPageId="14" captionText="FIG. 6. — Proterochersis porebensis Szczygielski &amp; Sulej, 2016, thin sections of large right femur ZPAL V. 39/500: A, B, dorsal region showing predominantly parallel-fibered bone matrix with longitudinal primary canals in the external part,numerous large,meandering secondary osteons in the deeper cortex,and gradual gradation into the medullary region;C, D, anterior region with parallel-fibered bone locally grading into higher organized,nearly lamellar bone;E, F, posteroventral region locally showing slightly less intensive invasion of remodeling, preserving undulating LAGs.Note the scalloped external surface. A, C, E in normal transmitted light, B, D, F in polarized light.Abbreviations and symbols: ELB, endosteal lamellar bone;IS, intertrabecular space; PFB, parallel-fibered bone;OPFB, higher organization of parallel-fibered bone; *, primary vascular canal; red +, primary osteon; black and white +, secondary osteon. Lines of arrested growth (LAGs) indicated with arrowheads. Scale bars: 500 µm." pageId="9" pageNumber="642">6E</figureCitation>
). The vascularization in the cortex is generally poor, especially in the external layers, although some local variability is visible along the perimeter of the bones (e.g. the cortex of the humeri is clearly more vascularized ventrally than dorsally;
<figureCitation id="134C2A58FF18FFDECC511BA8FEECFD1A" box="[289,353,726,752]" 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="9" pageNumber="642">Figs 2</figureCitation>
A-E; 3A-D; 4A-F). The vascularization pattern is sub-longitudinal, arranged in circular patterns, with slight radial inclination visible in all specimens except ZPAL V. 39/499 (small femur). The vasculature in most sectioned bones is predominantly primary (primary canals without primary osteons and, locally, osteons). The incipient secondary remodeling (sparsely scattered secondary osteons,
<figureCitation id="134C2A58FF18FFDECFDF1AEBFD72FC45" box="[687,767,917,943]" captionStart="FIG" captionStartId="12.[132,143,1849,1866]" captionTargetBox="[161,1427,215,1802]" captionTargetId="figure-220@12.[702,1246,774,1336]" captionTargetPageId="12" captionText="FIG. 4. — Proterochersis porebensis Szczygielski &amp; Sulej, 2016, thin sections of middle-sized right humerus ZPAL V. 39/433: A, B, anterodorsal region showing predominantly parallel-fibered bone matrix externally grading into higher organized, nearly lamellar bone with primary vasculature (richer in the deeper cortex than externally, longitudinal to radial) and undulating LAGs; C, D, dorsal region with sparse primary longitudinal vasculature and trabeculae incorporating cortical parallel-fibered tissue; E, F, anteroventral region showing coarse collagen bundles in the parallel-fibered matrix, longitudinal to radial predominantly primary vasculature with sparse secondary osteons, and undulating LAGs; G, H, interior of the medullary region presenting thick trabeculae composed of endosteal lamellar bone. A, C, E, G in normal transmitted light, B, D, F, H in polarized light. Abbreviations and symbols: ELB, endosteal lamellar bone; IS, intertrabecular space; PFB, parallel-fibered bone; OPFB, higher organization of parallel-fibered bone; *, primary vascular canal; red +, primary osteon; black and white +, secondary osteon. Lines of arrested growth (LAGs) indicated with arrowheads. Scale bars: 500 µm." pageId="9" pageNumber="642">Fig. 4E</figureCitation>
) is present in the middle-sized humerus ZPAL V. 39/433. Only in the large femur ZPAL V. 39/500 is the secondary remodeling significant and, in some places, nearly reaches the external surface, although much of the primary tissue and some primary vascular canals are still locally present (
<figureCitation id="134C2A58FF18FFDECFCA1D4BFD7AFBA5" box="[698,759,1077,1103]" captionStart="FIG" captionStartId="14.[132,143,1872,1889]" captionTargetBox="[150,1438,215,1825]" captionTargetId="figure-195@14.[701,1255,782,1352]" captionTargetPageId="14" captionText="FIG. 6. — Proterochersis porebensis Szczygielski &amp; Sulej, 2016, thin sections of large right femur ZPAL V. 39/500: A, B, dorsal region showing predominantly parallel-fibered bone matrix with longitudinal primary canals in the external part,numerous large,meandering secondary osteons in the deeper cortex,and gradual gradation into the medullary region;C, D, anterior region with parallel-fibered bone locally grading into higher organized,nearly lamellar bone;E, F, posteroventral region locally showing slightly less intensive invasion of remodeling, preserving undulating LAGs.Note the scalloped external surface. A, C, E in normal transmitted light, B, D, F in polarized light.Abbreviations and symbols: ELB, endosteal lamellar bone;IS, intertrabecular space; PFB, parallel-fibered bone;OPFB, higher organization of parallel-fibered bone; *, primary vascular canal; red +, primary osteon; black and white +, secondary osteon. Lines of arrested growth (LAGs) indicated with arrowheads. Scale bars: 500 µm." pageId="9" pageNumber="642">Fig. 6</figureCitation>
). The small femur ZPAL V. 39/499 retains mostly non-remodeled cortex, but evidence of active remodeling is visible at the border between the cortex and the medullary region, where two to three overlapping generations of lamellar bone are present (
<figureCitation id="134C2A58FF18FFDECC771DAAFEC9FB05" box="[263,324,1236,1263]" captionStart="FIG" captionStartId="13.[131,142,1637,1654]" captionTargetBox="[132,1455,214,1595]" captionTargetId="figure-328@13.[698,1267,766,1320]" captionTargetPageId="13" captionText="FIG. 5. — Proterochersis porebensis Szczygielski &amp; Sulej, 2016, thin sections of small right femur ZPAL V. 39/499: A, B, anterior region showing parallel-fibered matrix with longitudinal primary canals and straight LAGs; C, D, posterior region with thinner and less vascularized cortex (note that the external cortex is destroyed posteroventrally) and indistinct growth marks.Despite the small size of the specimen and primary nature of the cortical tissue,the boundary between the cortex and medullary region reveals active remodeling evidenced by two to three generations of secondary lamellar bone (marked with red rectangles). A and C in normal transmitted light, B and D in polarized light. Abbreviations and symbols: ELB, endosteal lamellar bone; IS, intertrabecular space; PFB, parallel-fibered bone; *, primary vascular canal. Lines of arrested growth (LAGs) indicated with arrowheads. Scale bars: 500 µm." pageId="9" pageNumber="642">Fig. 5</figureCitation>
). The large femur ZPAL V. 39/500 differs from the other sampled bones in having a clearly scalloped external surface due to a network of vascular grooves, which is visible also macroscopically (
<figureCitation id="134C2A58FF18FFDECCA01C4AFDC2FAA4" box="[464,591,1332,1358]" captionStart="FIG" captionStartId="6.[134,143,1238,1255]" captionTargetBox="[132,1455,214,1192]" captionTargetId="figure-598@6.[698,1267,213,749]" captionTargetPageId="6" captionText="FIG. 1. — Proterochersis porebensis Szczygielski &amp; Sulej, 2016, studied humeri (A-J) and femora (K-P), and the studied humerus of Proganochelys quenstedtii Baur, 1887 (Q, R) in dorsal (A, C, E, G, I, K, M, O, Q) and ventral (B, D, F, H, J, L, N, P, R) view: A, B, distal end of small right humerus ZPAL V. 39/439; C, D, small left humerus ZPAL V. 39/446; E, F, small right humerus ZPAL V. 39/156; G, H, distal end of middle-sized right humerus ZPAL V. 39/433; I, J, large right humerus ZPAL V. 39/50; K, L, distal end of small right femur ZPAL V. 39/499; M, N, distal end of large right femur ZPAL V. 39/500; O, P, large right femur ZPAL V. 39/432; Q, R, large left humerus SMF 09-F2. Sectioning planes for histology indicated in red. Scale bar: 1 cm." pageId="9" pageNumber="642">Figs 1M, N</figureCitation>
;
<figureCitation id="134C2A58FF18FFDECF2B1C4AFD14FAA4" box="[603,665,1332,1358]" 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="9" pageNumber="642">2H, I</figureCitation>
;
<figureCitation id="134C2A58FF18FFDECFD51C4AFD3EFAA4" box="[677,691,1332,1358]" captionStart="FIG" captionStartId="14.[132,143,1872,1889]" captionTargetBox="[150,1438,215,1825]" captionTargetId="figure-195@14.[701,1255,782,1352]" captionTargetPageId="14" captionText="FIG. 6. — Proterochersis porebensis Szczygielski &amp; Sulej, 2016, thin sections of large right femur ZPAL V. 39/500: A, B, dorsal region showing predominantly parallel-fibered bone matrix with longitudinal primary canals in the external part,numerous large,meandering secondary osteons in the deeper cortex,and gradual gradation into the medullary region;C, D, anterior region with parallel-fibered bone locally grading into higher organized,nearly lamellar bone;E, F, posteroventral region locally showing slightly less intensive invasion of remodeling, preserving undulating LAGs.Note the scalloped external surface. A, C, E in normal transmitted light, B, D, F in polarized light.Abbreviations and symbols: ELB, endosteal lamellar bone;IS, intertrabecular space; PFB, parallel-fibered bone;OPFB, higher organization of parallel-fibered bone; *, primary vascular canal; red +, primary osteon; black and white +, secondary osteon. Lines of arrested growth (LAGs) indicated with arrowheads. Scale bars: 500 µm." pageId="9" pageNumber="642">6</figureCitation>
). These grooves seem to be mostly imprints of superficial vasculature, but numerous vascular openings are also present on the dorsoposterior surface of the distal expansion of that specimen.
</paragraph>
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<paragraph id="8BC836DDFF19FFDFCDF41EEAFCAEF803" blockId="8.[132,1457,1940,2025]" pageId="8" pageNumber="641">
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. 2. —
<taxonomicName id="4C774D5EFF19FFDFCDA31EEAFD0BF84F" authority="Szczygielski &amp; Sulej, 2016" authorityName="Szczygielski &amp; Sulej" authorityYear="2016" box="[211,646,1940,1957]" class="Reptilia" family="Proterochersidae" genus="Proterochersis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="8" pageNumber="641" phylum="Chordata" rank="species" species="porebensis">
<emphasis id="B903EACFFF19FFDFCDA31EEAFE21F84F" box="[211,428,1940,1957]" italics="true" pageId="8" pageNumber="641">Proterochersis porebensis</emphasis>
<bibRefCitation id="EFE64B2CFF19FFDFCCDF1EEAFD0BF84F" author="SZCZYGIELSKI T. &amp; SULEJ T." box="[431,646,1940,1957]" pageId="8" pageNumber="641" pagination="395 - 427" refId="ref22723" refString="SZCZYGIELSKI T. &amp; SULEJ T. 2016. - Revision of the Triassic European turtles Proterochersis and Murrhardtia (Reptilia, Testudinata, Proterochersidae), with the description of new taxa from Poland and Germany. Zoological Journal of the Linnean Society 177 (2): 395 - 427. https: // doi. org / 10.1111 / zoj. 12374" type="journal article" year="2016">Szczygielski &amp; Sulej, 2016</bibRefCitation>
</taxonomicName>
(
<emphasis id="B903EACFFF19FFDFCFE01EEAFD27F84F" bold="true" box="[656,682,1940,1957]" pageId="8" pageNumber="641">A -I</emphasis>
) and
<taxonomicName id="4C774D5EFF19FFDFCFA61EEAFB9CF84F" authority="Baur, 1887" authorityName="Baur" authorityYear="1887" box="[726,1041,1940,1957]" class="Reptilia" family="Proganochelyidae" genus="Proganochelys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="8" pageNumber="641" phylum="Chordata" rank="species" species="quenstedtii">
<emphasis id="B903EACFFF19FFDFCFA61EEAFC3FF84F" box="[726,946,1940,1957]" italics="true" pageId="8" pageNumber="641">Proganochelys quenstedtii</emphasis>
<bibRefCitation id="EFE64B2CFF19FFDFCEC61EEAFB9CF84F" author="BAUR G." box="[950,1041,1940,1957]" pageId="8" pageNumber="641" pagination="17 - 20" refId="ref15932" refString="BAUR G. 1887. - Ueber den ursprung der extremitaten der Ichthyopterygia. Berichte uber de Versammlungen des Oberrheinischen Vereines 20: 17 - 20." type="journal article" year="1887">Baur, 1887</bibRefCitation>
</taxonomicName>
(
<emphasis id="B903EACFFF19FFDFC96A1EEAFBA9F84F" bold="true" box="[1050,1060,1940,1957]" pageId="8" pageNumber="641">J</emphasis>
,
<emphasis id="B903EACFFF19FFDFC95E1EEAFBB6F84F" bold="true" box="[1070,1083,1940,1957]" pageId="8" pageNumber="641">K</emphasis>
), thin sections in normal transmitted (
<emphasis id="B903EACFFF19FFDFC8011EEAFAF3F84F" bold="true" box="[1393,1406,1940,1957]" pageId="8" pageNumber="641">A</emphasis>
,
<emphasis id="B903EACFFF19FFDFC8F71EEAFA18F84F" bold="true" box="[1415,1429,1940,1957]" pageId="8" pageNumber="641">D</emphasis>
,
<emphasis id="B903EACFFF19FFDFC8EF1EEAFA27F84F" bold="true" box="[1439,1450,1940,1957]" pageId="8" pageNumber="641">F</emphasis>
,
<emphasis id="B903EACFFF19FFDFCDF41ED5FF1FF856" bold="true" box="[132,146,1963,1980]" pageId="8" pageNumber="641">H</emphasis>
,
<emphasis id="B903EACFFF19FFDFCDEB1ED5FF28F856" bold="true" box="[155,165,1963,1980]" pageId="8" pageNumber="641">J</emphasis>
), polarized (
<emphasis id="B903EACFFF19FFDFCC791ED5FE9BF856" bold="true" box="[265,278,1963,1980]" pageId="8" pageNumber="641">B</emphasis>
,
<emphasis id="B903EACFFF19FFDFCC501ED5FEA1F856" bold="true" box="[288,300,1963,1980]" pageId="8" pageNumber="641">E</emphasis>
,
<emphasis id="B903EACFFF19FFDFCC451ED5FECEF856" bold="true" box="[309,323,1963,1980]" pageId="8" pageNumber="641">G</emphasis>
,
<emphasis id="B903EACFFF19FFDFCC3D1ED5FEDEF856" bold="true" box="[333,339,1963,1980]" pageId="8" pageNumber="641">I</emphasis>
,
<emphasis id="B903EACFFF19FFDFCC2B1ED5FEE5F856" bold="true" box="[347,360,1963,1980]" pageId="8" pageNumber="641">K</emphasis>
), and polarized with quarter lambda plate I (
<emphasis id="B903EACFFF19FFDFCFBF1ED5FD50F856" bold="true" box="[719,733,1963,1980]" pageId="8" pageNumber="641">C</emphasis>
) light:
<emphasis id="B903EACFFF19FFDFCE631ED5FCBEF856" bold="true" box="[787,819,1963,1980]" pageId="8" pageNumber="641">A -E</emphasis>
, right humeri:
<emphasis id="B903EACFFF19FFDFCED81ED5FC47F856" bold="true" box="[936,970,1963,1980]" pageId="8" pageNumber="641">A -C</emphasis>
, small ZPAL V. 39/439;
<emphasis id="B903EACFFF19FFDFC9FD1ED5FB16F856" bold="true" box="[1165,1179,1963,1980]" pageId="8" pageNumber="641">D</emphasis>
,
<emphasis id="B903EACFFF19FFDFC9D41ED5FB3DF856" bold="true" box="[1188,1200,1963,1980]" pageId="8" pageNumber="641">E</emphasis>
, middle-sized ZPAL V. 39/433;
<emphasis id="B903EACFFF19FFDFCDF41EBCFF1FF839" bold="true" box="[132,146,1986,2003]" pageId="8" pageNumber="641">C</emphasis>
adjacent (slightly more distal) to
<emphasis id="B903EACFFF19FFDFCCD21EBCFE22F839" bold="true" box="[418,431,1986,2003]" pageId="8" pageNumber="641">A</emphasis>
and
<emphasis id="B903EACFFF19FFDFCCA71EBCFE69F839" bold="true" box="[471,484,1986,2003]" pageId="8" pageNumber="641">B</emphasis>
;
<emphasis id="B903EACFFF19FFDFCC9E1EBCFD8AF839" bold="true" box="[494,519,1986,2003]" pageId="8" pageNumber="641">F -I</emphasis>
, right femora:
<emphasis id="B903EACFFF19FFDFCF0E1EBCFD04F839" bold="true" box="[638,649,1986,2003]" pageId="8" pageNumber="641">F</emphasis>
,
<emphasis id="B903EACFFF19FFDFCFE31EBCFD2CF839" bold="true" box="[659,673,1986,2003]" pageId="8" pageNumber="641">G</emphasis>
, small ZPAL V. 39/499;
<emphasis id="B903EACFFF19FFDFCE181EBCFCFBF839" bold="true" box="[872,886,1986,2003]" pageId="8" pageNumber="641">H</emphasis>
,
<emphasis id="B903EACFFF19FFDFCE0F1EBCFC08F839" bold="true" box="[895,901,1986,2003]" pageId="8" pageNumber="641">I</emphasis>
, large ZPAL V. 39/500;
<emphasis id="B903EACFFF19FFDFC9391EBCFBDEF839" bold="true" box="[1097,1107,1986,2003]" pageId="8" pageNumber="641">J</emphasis>
,
<emphasis id="B903EACFFF19FFDFC92D1EBCFBE7F839" bold="true" box="[1117,1130,1986,2003]" pageId="8" pageNumber="641">K</emphasis>
, left humerus, large SMF 09-F2. Dorsal surface towards the top of the page, anterior surface to the left. Scale bar: 1 cm.
</paragraph>
</caption>
<paragraph id="8BC836DDFF18FFDECDEB1CCDFC8AF806" blockId="9.[130,777,216,2028]" pageId="9" pageNumber="642">
The extent of the cancellous area appears to be smaller in the humeri than in the femora and more clearly defined in the small than in the middle-sized and large specimens (
<figureCitation id="134C2A58FF18FFDECFEA1C8DFD58F9E7" box="[666,725,1523,1549]" 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="9" pageNumber="642">Fig. 2</figureCitation>
A-I). The trabeculae partly retain primary tissue in interstitial spaces, which are lined by endosteal lamellar bone, especially close to the cortex, and even in the smallest sampled specimens clearly incorporate periosteal tissue (
<figureCitation id="134C2A58FF18FFDECC8C1F0DFDB4F967" box="[508,569,1651,1677]" 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="9" pageNumber="642">Figs 2</figureCitation>
A-I; 3; 4C, D;5; 6). Especially in the large femur ZPAL V. 39/500, the boundary between the endosteal cancellous and periosteal cortical region is gradual, with large areas of the periosteal cortex remodeled into either intertrabecular spaces or irregularly meandering secondary osteons (
<figureCitation id="134C2A58FF18FFDECC241E6CFE4DF8C7" box="[340,448,1810,1837]" 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="9" pageNumber="642">Figs 2H, I</figureCitation>
;
<figureCitation id="134C2A58FF18FFDECCBA1E6CFE55F8C6" box="[458,472,1810,1836]" captionStart="FIG" captionStartId="14.[132,143,1872,1889]" captionTargetBox="[150,1438,215,1825]" captionTargetId="figure-195@14.[701,1255,782,1352]" captionTargetPageId="14" captionText="FIG. 6. — Proterochersis porebensis Szczygielski &amp; Sulej, 2016, thin sections of large right femur ZPAL V. 39/500: A, B, dorsal region showing predominantly parallel-fibered bone matrix with longitudinal primary canals in the external part,numerous large,meandering secondary osteons in the deeper cortex,and gradual gradation into the medullary region;C, D, anterior region with parallel-fibered bone locally grading into higher organized,nearly lamellar bone;E, F, posteroventral region locally showing slightly less intensive invasion of remodeling, preserving undulating LAGs.Note the scalloped external surface. A, C, E in normal transmitted light, B, D, F in polarized light.Abbreviations and symbols: ELB, endosteal lamellar bone;IS, intertrabecular space; PFB, parallel-fibered bone;OPFB, higher organization of parallel-fibered bone; *, primary vascular canal; red +, primary osteon; black and white +, secondary osteon. Lines of arrested growth (LAGs) indicated with arrowheads. Scale bars: 500 µm." pageId="9" pageNumber="642">6</figureCitation>
). In the same specimen, trabeculae deeper inside the bone are comparatively very thin. The thickness and length of the trabeculae is, nonetheless, varied; they appear more slender and longer in the central and dorsal part of the medullary area (
<figureCitation id="134C2A58FF18FFDECC921EECFD90F846" box="[482,541,1938,1964]" 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="9" pageNumber="642">Fig. 2</figureCitation>
A-I), but some regions of increased thickness are locally evident, especially in the middle-sized humerus ZPAL V. 39/433 (
<figureCitation id="134C2A58FF18FFDECF451EAFFD25F806" box="[565,680,2001,2028]" 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="9" pageNumber="642">Figs 2D, E</figureCitation>
;
<figureCitation id="134C2A58FF18FFDECFDF1EACFD74F806" box="[687,761,2002,2028]" captionStart="FIG" captionStartId="12.[132,143,1849,1866]" captionTargetBox="[161,1427,215,1802]" captionTargetId="figure-220@12.[702,1246,774,1336]" captionTargetPageId="12" captionText="FIG. 4. — Proterochersis porebensis Szczygielski &amp; Sulej, 2016, thin sections of middle-sized right humerus ZPAL V. 39/433: A, B, anterodorsal region showing predominantly parallel-fibered bone matrix externally grading into higher organized, nearly lamellar bone with primary vasculature (richer in the deeper cortex than externally, longitudinal to radial) and undulating LAGs; C, D, dorsal region with sparse primary longitudinal vasculature and trabeculae incorporating cortical parallel-fibered tissue; E, F, anteroventral region showing coarse collagen bundles in the parallel-fibered matrix, longitudinal to radial predominantly primary vasculature with sparse secondary osteons, and undulating LAGs; G, H, interior of the medullary region presenting thick trabeculae composed of endosteal lamellar bone. A, C, E, G in normal transmitted light, B, D, F, H in polarized light. Abbreviations and symbols: ELB, endosteal lamellar bone; IS, intertrabecular space; PFB, parallel-fibered bone; OPFB, higher organization of parallel-fibered bone; *, primary vascular canal; red +, primary osteon; black and white +, secondary osteon. Lines of arrested growth (LAGs) indicated with arrowheads. Scale bars: 500 µm." pageId="9" pageNumber="642">4G, H</figureCitation>
).
</paragraph>
<paragraph id="8BC836DDFF18FFDECE3319A6FA3CFD1A" blockId="9.[811,1458,216,2028]" pageId="9" pageNumber="642">
The CT data (
<figureCitation id="134C2A58FF18FFDECEA319A6FB83FF18" box="[979,1038,216,242]" 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="9" pageNumber="642">Fig. 7</figureCitation>
) illustrate a gradual ontogenetic progression of the trends observed in the histological sections and reveal the microstructural patterns along the bone length. Both the absolute and relative thickness of the cortices appears to increase with increasing bone size. In that regard, ZPAL V. 39/446 (cortex
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) and ZPAL V. 39/156 (cortex
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), which are intermediate in size between the sectioned ZPAL V. 39/439 and ZPAL V. 39/433, are also intermediate structurally their cortices are thicker than in ZPAL V. 39/439 (
<quantity id="4C8F9B38FF18FFDECE431889FC30FDFB" box="[819,957,503,529]" metricMagnitude="-3" metricUnit="m" metricValue="1.45" metricValueMax="1.7" metricValueMin="1.2" pageId="9" pageNumber="642" unit="mm" value="1.45" valueMax="1.7" valueMin="1.2">1.2-1.7 mm</quantity>
), but thinner than in ZPAL V. 39/433 (
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; note that the cortical thickness in sectioned specimens is undervalued due to the slightly more distal plane of sectioning). The large humerus (ZPAL V. 39/50, cortex
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) and femur (ZPAL V. 39/432, cortex
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, compared to
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in ZPAL V.39/499 and
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in ZPAL V. 39.500) generally follow the trends of cortical thickening with increasing size suggested by the smaller specimens.
</paragraph>
<paragraph id="8BC836DDFF18FFDECE331B88FA2BF9E7" blockId="9.[811,1458,216,2028]" pageId="9" pageNumber="642">
As indicated by previous studies, the centers of ossification are located slightly more proximally than the midshaft and are associated with the largest thickness of the cortices (
<figureCitation id="134C2A58FF18FFDECE451A2BFC6CFC9A" box="[821,993,853,880]" 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="9" pageNumber="642">Fig. 7A, C, E, G</figureCitation>
). In small humeri, the cortices in that region are relatively compact (in ZPAL V. 39/446 more so than in ZPAL V. 39/156) and the trabecular bone in the bone center has a relatively uniform structure (
<figureCitation id="134C2A58FF18FFDEC9EA1ACBFB5AFC3A" box="[1178,1239,949,976]" 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="9" pageNumber="642">Fig. 7</figureCitation>
A-D) probably indicating low remodeling (although potentially also dependent on stress distribution patterns and load). In ZPAL V. 39/50, however, the cortex is somewhat more vascularized (although still relatively poorly), but the interior trabecular region is much less defined and denser than in smaller specimens, indicating more pronounced remodeling (
<figureCitation id="134C2A58FF18FFDEC9811D0AFAD2FB65" box="[1265,1375,1140,1167]" 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="9" pageNumber="642">Fig. 7E, F</figureCitation>
). There is no central marrow cavity in the studied humeri. Conversely, in the large femur ZPAL V. 39/432, the internal trabecular region is still distinct (even though it is denser than in the areas further away from the ossification center) and there is a small, irregular region devoid of trabeculae located around the level of the ossification center (
<figureCitation id="134C2A58FF18FFDEC9EC1C4AFA9EFAA4" box="[1180,1299,1332,1358]" 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="9" pageNumber="642">Fig. 7G, H</figureCitation>
). This cavity is connected with the bone exterior via two well-defined nutrient canals extending: 1) dorsoproximoanteriorly and eventually opening as a nutrient foramen at the anterodorsal base of the trochanter major; and 2) ventrodistally and opening as a nutrient foramen in the popliteal fossa, between the tibial and fibular condyle (see Appendices 1-16; and Morphosource).
</paragraph>
<paragraph id="8BC836DDFF18FFDECE331F6DFA91F8C6" blockId="9.[811,1458,216,2028]" pageId="9" pageNumber="642">
Proximally and distally from the centers of ossification, the thickness of the cortex steadily decreases so in the areas of the proximal and distal articular surfaces the trabeculae nearly reach the bone exterior (
<figureCitation id="134C2A58FF18FFDEC9061F0DFAA8F967" box="[1142,1317,1651,1677]" 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="9" pageNumber="642">Fig. 7A, C, E, G</figureCitation>
). The cortex, however, retains a relatively constant thickness across the intertubercular fossa of the humeri (
<figureCitation id="134C2A58FF18FFDEC9B21FCCFAB7F927" box="[1218,1338,1714,1741]" 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="9" pageNumber="642">Fig. 7A, C</figureCitation>
; Appendices 1-16; and Morphosource). In ZPAL V. 39/
<quantity id="4C8F9B38FF18FFDEC8411FACFAF6F906" box="[1329,1403,1746,1772]" metricMagnitude="1" metricUnit="m" metricValue="1.1328399999999998" pageId="9" pageNumber="642" unit="in" value="446.0">446 in</quantity>
that area it is perforated by numerous vascular canals roughly parallel to the long axis of the bone (
<figureCitation id="134C2A58FF18FFDEC9CC1E6CFA9DF8C6" box="[1212,1296,1810,1837]" 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="9" pageNumber="642">Fig. 7A</figureCitation>
).
</paragraph>
<paragraph id="8BC836DDFF18FFDCCE331E4CFD1EFC25" blockId="9.[811,1458,216,2028]" lastBlockId="11.[131,776,758,1071]" lastPageId="11" lastPageNumber="644" pageId="9" pageNumber="642">
Trabeculae are relatively long and thick in the small humerus ZPAL V. 39/446 (
<figureCitation id="134C2A58FF18FFDEC9121E2CFB35F886" box="[1122,1208,1874,1901]" 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="9" pageNumber="642">Fig. 7A</figureCitation>
). In the slightly larger ZPAL V. 39/156 they retain a similar morphology close to the cortex (especially ventrally) and around the level of the center of ossification, but become finer and more densely packed in the middle of the more proximal and distal portions of the bone (
<figureCitation id="134C2A58FF1AFFDCCC7D1B88FEECFCFA" box="[269,353,758,784]" 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. 7C</figureCitation>
). In the large humerus ZPAL V. 39/50, they are generally more irregular, with larger and smaller intertrabecular spaces and thinner and thicker trabeculae scattered without a clear pattern (
<figureCitation id="134C2A58FF1AFFDCCC971A2BFDBAFC9A" box="[487,567,853,880]" 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. 7E</figureCitation>
). In the large femur ZPAL V. 39/432, the pattern of longer trabeculae closer to the cortex and finer trabeculae more interiorly is also noticeable, but less clear than in the humeri (
<figureCitation id="134C2A58FF1AFFDCCF401ACBFD0BFC25" box="[560,646,949,976]" 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. 7G</figureCitation>
).
</paragraph>
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<paragraph id="8BC836DDFF1BFFDDCDF41E5DFA22F803" blockId="10.[132,1457,1827,2025]" pageId="10" pageNumber="643">
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. 3. —
<taxonomicName id="4C774D5EFF1BFFDDCDA51E5DFD1EF8DE" authority="Szczygielski &amp; Sulej, 2016" authorityName="Szczygielski &amp; Sulej" authorityYear="2016" box="[213,659,1827,1844]" class="Reptilia" family="Proterochersidae" genus="Proterochersis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="10" pageNumber="643" phylum="Chordata" rank="species" species="porebensis">
<emphasis id="B903EACFFF1BFFDDCDA51E5DFE3DF8DE" box="[213,432,1827,1844]" italics="true" pageId="10" pageNumber="643">Proterochersis porebensis</emphasis>
<bibRefCitation id="EFE64B2CFF1BFFDDCCC61E5DFD1EF8DE" author="SZCZYGIELSKI T. &amp; SULEJ T." box="[438,659,1827,1844]" pageId="10" pageNumber="643" pagination="395 - 427" refId="ref22723" refString="SZCZYGIELSKI T. &amp; SULEJ T. 2016. - Revision of the Triassic European turtles Proterochersis and Murrhardtia (Reptilia, Testudinata, Proterochersidae), with the description of new taxa from Poland and Germany. Zoological Journal of the Linnean Society 177 (2): 395 - 427. https: // doi. org / 10.1111 / zoj. 12374" type="journal article" year="2016">Szczygielski &amp; Sulej, 2016</bibRefCitation>
</taxonomicName>
, thin sections of small right humerus ZPAL V. 39/439:
<emphasis id="B903EACFFF1BFFDDC92A1E5DFBEAF8DE" bold="true" box="[1114,1127,1827,1844]" pageId="10" pageNumber="643">A</emphasis>
, anterodorsal region showing relatively thin, nearly avascular cortex composed of parallel-fibered bone with gently undulating collagen fibers and thin trabeculae composed mostly of endosteal lamellar bone;
<emphasis id="B903EACFFF1BFFDDCDA21E2EFF52F88B" bold="true" box="[210,223,1872,1889]" pageId="10" pageNumber="643">B</emphasis>
, dorsal region showing undulating collagen fiber layout parallel to bone surface and sparse collagen fibers oblique to the bone surface in the cortex, slightly increased longitudinal primary vasculature, and trabeculae incorporating cortical parallel-fibered bone;
<emphasis id="B903EACFFF1BFFDDC95D1E19FBB6F892" bold="true" box="[1069,1083,1895,1912]" pageId="10" pageNumber="643">C</emphasis>
,
<emphasis id="B903EACFFF1BFFDDC9361E19FBD9F892" bold="true" box="[1094,1108,1895,1912]" pageId="10" pageNumber="643">D</emphasis>
, ventral region showing more developed vasculature (mostly primary canals with sporadic primary osteons, predominantly longitudinal), growth marks, and parallel-fibered matrix grading externally into more ordered, nearly lamellar bone;
<emphasis id="B903EACFFF1BFFDDCCC21EEAFE33F84F" bold="true" box="[434,446,1940,1957]" pageId="10" pageNumber="643">E</emphasis>
, border between the cortex and the medullary region in the ventral part of the bone, adjacent (slightly more proximally) to
<emphasis id="B903EACFFF1BFFDDCDF41ED5FF2BF856" bold="true" box="[132,166,1963,1980]" pageId="10" pageNumber="643">A -C</emphasis>
showing a large chunk of cortical parallel-fibered bone incorporated into the medullary region.
<emphasis id="B903EACFFF1BFFDDCEC31ED5FC4DF856" bold="true" box="[947,960,1963,1980]" pageId="10" pageNumber="643">A</emphasis>
,
<emphasis id="B903EACFFF1BFFDDCEB91ED5FC5BF856" bold="true" box="[969,982,1963,1980]" pageId="10" pageNumber="643">B</emphasis>
,
<emphasis id="B903EACFFF1BFFDDCE901ED5FC63F856" bold="true" box="[992,1006,1963,1980]" pageId="10" pageNumber="643">D</emphasis>
in polarized light with quarter lambda plate,
<emphasis id="B903EACFFF1BFFDDC82A1ED5FAE5F856" bold="true" box="[1370,1384,1963,1980]" pageId="10" pageNumber="643">C</emphasis>
and
<emphasis id="B903EACFFF1BFFDDC8E01ED5FA11F856" bold="true" box="[1424,1436,1963,1980]" pageId="10" pageNumber="643">E</emphasis>
in normal transmitted light.Abbreviations and symbols:
<emphasis id="B903EACFFF1BFFDDCF4B1EBCFDD2F839" bold="true" box="[571,607,1986,2003]" pageId="10" pageNumber="643">ELB</emphasis>
, endosteal lamellar bone;
<emphasis id="B903EACFFF1BFFDDCE461EBCFCC5F839" bold="true" box="[822,840,1986,2003]" pageId="10" pageNumber="643">IS</emphasis>
, intertrabecular space;
<emphasis id="B903EACFFF1BFFDDC97A1EBCFBA2F839" bold="true" box="[1034,1071,1986,2003]" pageId="10" pageNumber="643">PFB</emphasis>
, parallel-fibered bone;
<emphasis id="B903EACFFF1BFFDDC99C1EBCFA92F839" bold="true" box="[1260,1311,1986,2003]" pageId="10" pageNumber="643">OPFB</emphasis>
, higher organization of parallel-fibered bone;
<emphasis id="B903EACFFF1BFFDDCC051EA6FEF0F803" bold="true" box="[373,381,2008,2025]" pageId="10" pageNumber="643">*</emphasis>
, primary vascular canal;
<emphasis id="B903EACFFF1BFFDDCF3D1EA6FDF7F803" bold="true" box="[589,634,2008,2025]" pageId="10" pageNumber="643">red +</emphasis>
, primary osteon. Lines of arrested growth (LAGs) indicated with
<emphasis id="B903EACFFF1BFFDDC9E31EA6FB70F803" bold="true" box="[1171,1277,2008,2025]" pageId="10" pageNumber="643">arrowheads</emphasis>
. Scale bars: 500 µm.
</paragraph>
</caption>
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<paragraph id="8BC836DDFF1AFFDCCDF319A2FBE3FEF0" blockId="11.[131,1455,220,282]" pageId="11" pageNumber="644">
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3. — Carapace, humerus, and femur measurements and length ratios in
<taxonomicName id="4C774D5EFF1AFFDCCE5619A2FBE3FF07" authority="Baur, 1887" authorityName="Baur" authorityYear="1887" box="[806,1134,220,237]" class="Reptilia" family="Proganochelyidae" genus="Proganochelys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="11" pageNumber="644" phylum="Chordata" rank="species" species="quenstedtii">
<emphasis id="B903EACFFF1AFFDCCE5619A2FB88FF07" box="[806,1029,220,237]" italics="true" pageId="11" pageNumber="644">Proganochelys quenstedtii</emphasis>
<bibRefCitation id="EFE64B2CFF1AFFDCC97D19A2FBE3FF07" author="BAUR G." box="[1037,1134,220,237]" pageId="11" pageNumber="644" pagination="17 - 20" refId="ref15932" refString="BAUR G. 1887. - Ueber den ursprung der extremitaten der Ichthyopterygia. Berichte uber de Versammlungen des Oberrheinischen Vereines 20: 17 - 20." type="journal article" year="1887">Baur, 1887</bibRefCitation>
</taxonomicName>
,
<taxonomicName id="4C774D5EFF1AFFDCC90519A2FADCFF07" authorityName="Szczygielski &amp; Sulej" authorityYear="2016" box="[1141,1361,220,237]" class="Reptilia" family="Proterochersidae" genus="Proterochersis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="11" pageNumber="644" phylum="Chordata" rank="species" species="porebensis">
<emphasis id="B903EACFFF1AFFDCC90519A2FADCFF07" box="[1141,1361,220,237]" italics="true" pageId="11" pageNumber="644">Proterochersis porebensis</emphasis>
</taxonomicName>
Szczygiel- ski &amp; Sulej, 2016, and
<taxonomicName id="4C774D5EFF1AFFDCCC31198CFCC4FEE9" authority="Rougier, Fuente &amp; Arcucci, 1995" authorityName="Rougier, Fuente &amp; Arcucci" authorityYear="1995" box="[321,841,242,259]" class="Reptilia" genus="Palaeochersis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="11" pageNumber="644" phylum="Chordata" rank="species" species="talampayensis">
<emphasis id="B903EACFFF1AFFDCCC31198CFDBDFEE9" box="[321,560,242,259]" italics="true" pageId="11" pageNumber="644">Palaeochersis talampayensis</emphasis>
<bibRefCitation id="EFE64B2CFF1AFFDCCF46198CFCC4FEE9" author="ROUGIER G. W. &amp; FUENTE M. S. DE LA &amp; ARCUCCI A. B." box="[566,841,242,259]" pageId="11" pageNumber="644" pagination="855 - 858" refId="ref21477" refString="ROUGIER G. W., FUENTE M. S. DE LA &amp; ARCUCCI A. B. 1995. - Late Triassic turtles from South America. Science 268 (5212): 855 - 858. https: // doi. org / 10.1126 / science. 268.5212.855" type="journal article" year="1995">Rougier, Fuente &amp; Arcucci, 1995</bibRefCitation>
</taxonomicName>
. Carapace length estimation for SMNS 17203 taken from
<bibRefCitation id="EFE64B2CFF1AFFDCC840198CFA24FEE9" author="GAFFNEY E. S." box="[1328,1449,242,259]" 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>
. Abbreviations:
<emphasis id="B903EACFFF1AFFDCCC711877FE83FEF0" bold="true" box="[257,270,265,282]" pageId="11" pageNumber="644">R</emphasis>
, right;
<emphasis id="B903EACFFF1AFFDCCC391877FED9FEF0" bold="true" box="[329,340,265,282]" pageId="11" pageNumber="644">L</emphasis>
, left;
<emphasis id="B903EACFFF1AFFDCCCF21877FE07FEF0" bold="true" box="[386,394,265,282]" pageId="11" pageNumber="644">*</emphasis>
, preservation (e.g. damage or deformation) possibly affecting length and resulting ratios.
</paragraph>
</caption>
<paragraph id="8BC836DDFF1AFFDCCDF61810FEFFFD5F" pageId="11" pageNumber="644">
<table id="F977C47DFF1A0028CDF6182AFA20FD5F" box="[134,1453,340,693]" gridcols="8" gridrows="11" pageId="11" pageNumber="644">
<tr id="3547349FFF1A0028CDF6182AFA20FE68" box="[134,1453,340,386]" gridrow="0" pageId="11" pageNumber="644">
<th id="76965DE3FF1A0028CDF6182AFE96FE68" box="[134,283,340,386]" gridcol="0" gridrow="0" pageId="11" pageNumber="644">
<emphasis id="B903EACFFF1AFFDCCDF61810FF5AFE68" bold="true" box="[134,215,366,386]" pageId="11" pageNumber="644">Species</emphasis>
</th>
<th id="76965DE3FF1A0028CC5C182AFE1AFE68" box="[300,407,340,386]" gridcol="1" gridrow="0" pageId="11" pageNumber="644">
<emphasis id="B903EACFFF1AFFDCCC5C1810FE1CFE68" bold="true" box="[300,401,366,386]" pageId="11" pageNumber="644">Specimen</emphasis>
</th>
<th id="76965DE3FF1A0028CCDC182AFDD7FE68" box="[428,602,340,386]" gridcol="2" gridrow="0" pageId="11" pageNumber="644">
<emphasis id="B903EACFFF1AFFDCCCDC182AFDD7FE82" bold="true" box="[428,602,340,360]" pageId="11" pageNumber="644">Midline carapace</emphasis>
<emphasis id="B903EACFFF1AFFDCCCBA1810FDB6FE68" bold="true" box="[458,571,366,386]" pageId="11" pageNumber="644">length (cm)</emphasis>
</th>
<th id="76965DE3FF1A0028CF00182AFD6FFE68" box="[624,738,340,386]" gridcol="3" gridrow="0" pageId="11" pageNumber="644">
<emphasis id="B903EACFFF1AFFDCCF0B182AFD5AFE82" bold="true" box="[635,727,340,360]" pageId="11" pageNumber="644">Humerus</emphasis>
<emphasis id="B903EACFFF1AFFDCCF001810FD6FFE68" bold="true" box="[624,738,366,386]" pageId="11" pageNumber="644">length (cm)</emphasis>
</th>
<th id="76965DE3FF1A0028CF89182AFC31FE68" box="[761,956,340,386]" gridcol="4" gridrow="0" pageId="11" pageNumber="644">
<emphasis id="B903EACFFF1AFFDCCF89182AFC31FE82" bold="true" box="[761,956,340,360]" pageId="11" pageNumber="644">Humerus/carapace</emphasis>
<emphasis id="B903EACFFF1AFFDCCE511810FC18FE68" bold="true" box="[801,917,366,386]" pageId="11" pageNumber="644">length ratio</emphasis>
</th>
<th id="76965DE3FF1A0028CEB9182AFBDDFE68" box="[969,1104,340,386]" gridcol="5" gridrow="0" pageId="11" pageNumber="644">
<emphasis id="B903EACFFF1AFFDCCEB9182AFBDDFE82" bold="true" box="[969,1104,340,360]" pageId="11" pageNumber="644">Femur length</emphasis>
<emphasis id="B903EACFFF1AFFDCCE861810FBAFFE68" bold="true" box="[1014,1058,366,386]" pageId="11" pageNumber="644">(cm)</emphasis>
</th>
<th id="76965DE3FF1A0028C92B182AFA8FFE68" box="[1115,1282,340,386]" gridcol="6" gridrow="0" pageId="11" pageNumber="644">
<emphasis id="B903EACFFF1AFFDCC92B182AFA8FFE82" bold="true" box="[1115,1282,340,360]" pageId="11" pageNumber="644">Femur/carapace</emphasis>
<emphasis id="B903EACFFF1AFFDCC9051810FB64FE68" bold="true" box="[1141,1257,366,386]" pageId="11" pageNumber="644">length ratio</emphasis>
</th>
<th id="76965DE3FF1A0028C87D182AFA20FE68" box="[1293,1453,340,386]" gridcol="7" gridrow="0" pageId="11" pageNumber="644">
<emphasis id="B903EACFFF1AFFDCC87D182AFA20FE82" bold="true" box="[1293,1453,340,360]" pageId="11" pageNumber="644">Humerus/femur</emphasis>
<emphasis id="B903EACFFF1AFFDCC8531810FA1AFE68" bold="true" box="[1315,1431,366,386]" pageId="11" pageNumber="644">length ratio</emphasis>
</th>
</tr>
<tr id="3547349FFF1A0028CDF618ECFA20FE4C" box="[134,1453,402,422]" gridrow="1" pageId="11" pageNumber="644">
<th id="76965DE3FF1A0028CDF618ECFE96FE55" box="[134,283,402,447]" gridcol="0" gridrow="1" pageId="11" pageNumber="644" rowspan="2" rowspanBelow="1">
<taxonomicName id="4C774D5EFF1AFFDCCDF618ECFE87FE4C" authority="PULR" authorityName="PULR" box="[134,266,402,422]" class="Reptilia" genus="Palaeochersis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="11" pageNumber="644" phylum="Chordata" rank="genus">
<emphasis id="B903EACFFF1AFFDCCDF618ECFE87FE4C" box="[134,266,402,422]" italics="true" pageId="11" pageNumber="644">Palaeochersis</emphasis>
</taxonomicName>
<taxonomicName id="4C774D5EFF1AFFDCCDE518D5FE96FE55" authorityName="Rougier, Fuente &amp; Arcucci" authorityYear="1995" box="[149,283,427,447]" class="Reptilia" genus="Palaeochersis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="11" pageNumber="644" phylum="Chordata" rank="species" species="talampayensis">
<emphasis id="B903EACFFF1AFFDCCDE518D5FE96FE55" box="[149,283,427,447]" italics="true" pageId="11" pageNumber="644">talampayensis</emphasis>
</taxonomicName>
</th>
<td id="76965DE3FF1A0028CC5C18ECFE1AFE55" box="[300,407,402,447]" gridcol="1" gridrow="1" pageId="11" pageNumber="644" rowspan="2" rowspanBelow="1">PULR 068</td>
<td id="76965DE3FF1A0028CCDC18ECFDD7FE55" box="[428,602,402,447]" gridcol="2" gridrow="1" pageId="11" pageNumber="644" rowspan="2" rowspanBelow="1">45.5</td>
<td id="76965DE3FF1A0028CF0018ECFD6FFE4C" box="[624,738,402,422]" gridcol="3" gridrow="1" pageId="11" pageNumber="644">L: 12.7</td>
<td id="76965DE3FF1A0028CF8918ECFC31FE4C" box="[761,956,402,422]" gridcol="4" gridrow="1" pageId="11" pageNumber="644">L: 0.28</td>
<td id="76965DE3FF1A0028CEB918ECFBDDFE4C" box="[969,1104,402,422]" gridcol="5" gridrow="1" pageId="11" pageNumber="644">L: 11.9</td>
<td id="76965DE3FF1A0028C92B18ECFA8FFE4C" box="[1115,1282,402,422]" gridcol="6" gridrow="1" pageId="11" pageNumber="644">L: 0.26</td>
<td id="76965DE3FF1A0028C87D18ECFA20FE4C" box="[1293,1453,402,422]" gridcol="7" gridrow="1" pageId="11" pageNumber="644">L: 1.07</td>
</tr>
<tr id="3547349FFF1A0028CDF618D5FA20FE55" box="[134,1453,427,447]" gridrow="2" pageId="11" pageNumber="644">
<td id="76965DE3FF1A0028CF0018D5FD6FFE55" box="[624,738,427,447]" gridcol="3" gridrow="2" pageId="11" pageNumber="644">R: 12.0</td>
<td id="76965DE3FF1A0028CF8918D5FC31FE55" box="[761,956,427,447]" gridcol="4" gridrow="2" pageId="11" pageNumber="644">R: 0.26</td>
<td id="76965DE3FF1A0028CEB918D5FBDDFE55" box="[969,1104,427,447]" gridcol="5" gridrow="2" pageId="11" pageNumber="644">R: 11.6</td>
<td id="76965DE3FF1A0028C92B18D5FA8FFE55" box="[1115,1282,427,447]" gridcol="6" gridrow="2" pageId="11" pageNumber="644">R: 0.25</td>
<td id="76965DE3FF1A0028C87D18D5FA20FE55" box="[1293,1453,427,447]" gridcol="7" gridrow="2" pageId="11" pageNumber="644">R: 1.03</td>
</tr>
<tr id="3547349FFF1A0028CDF618B1FA20FE09" box="[134,1453,463,483]" gridrow="3" pageId="11" pageNumber="644">
<th id="76965DE3FF1A0028CDF618B1FE96FD92" box="[134,283,463,632]" gridcol="0" gridrow="3" pageId="11" pageNumber="644" rowspan="7" rowspanBelow="6">
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<emphasis id="B903EACFFF1AFFDCCDF618B1FE9EFE09" box="[134,275,463,483]" italics="true" pageId="11" pageNumber="644">Proganochelys</emphasis>
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<taxonomicName id="4C774D5EFF1AFFDCCDE51896FF73FE16" authorityName="Baur" authorityYear="1887" box="[149,254,488,508]" class="Reptilia" family="Proganochelyidae" genus="Proganochelys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="11" pageNumber="644" phylum="Chordata" rank="species" species="quenstedtii">
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</th>
<td id="76965DE3FF1A0028CC5C18B1FE1AFE16" box="[300,407,463,508]" gridcol="1" gridrow="3" pageId="11" pageNumber="644" rowspan="2" rowspanBelow="1">SMNS 16980</td>
<td id="76965DE3FF1A0028CCDC18B1FDD7FE16" box="[428,602,463,508]" gridcol="2" gridrow="3" pageId="11" pageNumber="644" rowspan="2" rowspanBelow="1">48.5</td>
<td id="76965DE3FF1A0028CF0018B1FD6FFE09" box="[624,738,463,483]" gridcol="3" gridrow="3" pageId="11" pageNumber="644">L: 13.4</td>
<td id="76965DE3FF1A0028CF8918B1FC31FE09" box="[761,956,463,483]" gridcol="4" gridrow="3" pageId="11" pageNumber="644">L: 0.28</td>
<td id="76965DE3FF1A0028CEB918B1FBDDFE09" box="[969,1104,463,483]" gridcol="5" gridrow="3" pageId="11" pageNumber="644">L: 15.3</td>
<td id="76965DE3FF1A0028C92B18B1FA8FFE09" box="[1115,1282,463,483]" gridcol="6" gridrow="3" pageId="11" pageNumber="644">L: 0.32</td>
<td id="76965DE3FF1A0028C87D18B1FA20FE09" box="[1293,1453,463,483]" gridcol="7" gridrow="3" pageId="11" pageNumber="644">L: 0.88</td>
</tr>
<tr id="3547349FFF1A0028CDF61896FA20FE16" box="[134,1453,488,508]" gridrow="4" pageId="11" pageNumber="644">
<td id="76965DE3FF1A0028CF001896FD6FFE16" box="[624,738,488,508]" gridcol="3" gridrow="4" pageId="11" pageNumber="644">R: 13.9</td>
<td id="76965DE3FF1A0028CF891896FC31FE16" box="[761,956,488,508]" gridcol="4" gridrow="4" pageId="11" pageNumber="644">R: 0.29</td>
<td id="76965DE3FF1A0028CEB91896FBDDFE16" box="[969,1104,488,508]" gridcol="5" gridrow="4" pageId="11" pageNumber="644">R: 15.7*</td>
<td id="76965DE3FF1A0028C92B1896FA8FFE16" box="[1115,1282,488,508]" gridcol="6" gridrow="4" pageId="11" pageNumber="644">R: 0.32*</td>
<td id="76965DE3FF1A0028C87D1896FA20FE16" box="[1293,1453,488,508]" gridcol="7" gridrow="4" pageId="11" pageNumber="644">R: 0.89*</td>
</tr>
<tr id="3547349FFF1A0028CDF61B7EFA20FDFE" box="[134,1453,512,532]" gridrow="5" pageId="11" pageNumber="644">
<td id="76965DE3FF1A0028CC5C1B7EFE1AFDC4" box="[300,407,512,558]" gridcol="1" gridrow="5" pageId="11" pageNumber="644" rowspan="2" rowspanBelow="1">SMNS 17203</td>
<td id="76965DE3FF1A0028CCDC1B7EFDD7FDC4" box="[428,602,512,558]" gridcol="2" gridrow="5" pageId="11" pageNumber="644" rowspan="2" rowspanBelow="1">40* (estimated)</td>
<td id="76965DE3FF1A0028CF001B7EFD6FFDFE" box="[624,738,512,532]" gridcol="3" gridrow="5" pageId="11" pageNumber="644">L: 9.8*</td>
<td id="76965DE3FF1A0028CF891B7EFC31FDFE" box="[761,956,512,532]" gridcol="4" gridrow="5" pageId="11" pageNumber="644">L: 0.25*</td>
<td id="76965DE3FF1A0028CEB91B7EFBDDFDFE" box="[969,1104,512,532]" gridcol="5" gridrow="5" pageId="11" pageNumber="644">L: 13.1</td>
<td id="76965DE3FF1A0028C92B1B7EFA8FFDFE" box="[1115,1282,512,532]" gridcol="6" gridrow="5" pageId="11" pageNumber="644">L: 0.32*</td>
<td id="76965DE3FF1A0028C87D1B7EFA20FDFE" box="[1293,1453,512,532]" gridcol="7" gridrow="5" pageId="11" pageNumber="644">L: 0.75*</td>
</tr>
<tr id="3547349FFF1A0028CDF61B64FA20FDC4" box="[134,1453,538,558]" gridrow="6" pageId="11" pageNumber="644">
<td id="76965DE3FF1A0028CF001B64FD6FFDC4" box="[624,738,538,558]" gridcol="3" gridrow="6" pageId="11" pageNumber="644"></td>
<td id="76965DE3FF1A0028CF891B64FC31FDC4" box="[761,956,538,558]" gridcol="4" gridrow="6" pageId="11" pageNumber="644"></td>
<td id="76965DE3FF1A0028CEB91B64FBDDFDC4" box="[969,1104,538,558]" gridcol="5" gridrow="6" pageId="11" pageNumber="644">R: 14.4*</td>
<td id="76965DE3FF1A0028C92B1B64FA8FFDC4" box="[1115,1282,538,558]" gridcol="6" gridrow="6" pageId="11" pageNumber="644">R: 0.36*</td>
<td id="76965DE3FF1A0028C87D1B64FA20FDC4" box="[1293,1453,538,558]" gridcol="7" gridrow="6" pageId="11" pageNumber="644"></td>
</tr>
<tr id="3547349FFF1A0028CDF61B4CFA20FDAC" box="[134,1453,562,582]" gridrow="7" pageId="11" pageNumber="644">
<td id="76965DE3FF1A0028CC5C1B4CFE1AFD8A" box="[300,407,562,608]" gridcol="1" gridrow="7" pageId="11" pageNumber="644" rowspan="2" rowspanBelow="1">SMNS 17204</td>
<td id="76965DE3FF1A0028CCDC1B4CFDD7FD8A" box="[428,602,562,608]" gridcol="2" gridrow="7" pageId="11" pageNumber="644" rowspan="2" rowspanBelow="1">≥ 50*</td>
<td id="76965DE3FF1A0028CF001B4CFD6FFDAC" box="[624,738,562,582]" gridcol="3" gridrow="7" pageId="11" pageNumber="644"></td>
<td id="76965DE3FF1A0028CF891B4CFC31FDAC" box="[761,956,562,582]" gridcol="4" gridrow="7" pageId="11" pageNumber="644"></td>
<td id="76965DE3FF1A0028CEB91B4CFBDDFDAC" box="[969,1104,562,582]" gridcol="5" gridrow="7" pageId="11" pageNumber="644">L: 15.8</td>
<td id="76965DE3FF1A0028C92B1B4CFA8FFDAC" box="[1115,1282,562,582]" gridcol="6" gridrow="7" pageId="11" pageNumber="644">L: 0.32</td>
<td id="76965DE3FF1A0028C87D1B4CFA20FDAC" box="[1293,1453,562,582]" gridcol="7" gridrow="7" pageId="11" pageNumber="644">&gt; 0.82*</td>
</tr>
<tr id="3547349FFF1A0028CDF61B32FA20FD8A" box="[134,1453,588,608]" gridrow="8" pageId="11" pageNumber="644">
<td id="76965DE3FF1A0028CF001B32FD6FFD8A" box="[624,738,588,608]" gridcol="3" gridrow="8" pageId="11" pageNumber="644">R:&gt; 13*</td>
<td id="76965DE3FF1A0028CF891B32FC31FD8A" box="[761,956,588,608]" gridcol="4" gridrow="8" pageId="11" pageNumber="644">R:&gt; 0.26*</td>
<td id="76965DE3FF1A0028CEB91B32FBDDFD8A" box="[969,1104,588,608]" gridcol="5" gridrow="8" pageId="11" pageNumber="644">R: 15.9</td>
<td id="76965DE3FF1A0028C92B1B32FA8FFD8A" box="[1115,1282,588,608]" gridcol="6" gridrow="8" pageId="11" pageNumber="644">R: 0.32</td>
<td id="76965DE3FF1A0028C87D1B32FA20FD8A" box="[1293,1453,588,608]" gridcol="7" gridrow="8" pageId="11" pageNumber="644"></td>
</tr>
<tr id="3547349FFF1A0028CDF61B1AFA20FD92" box="[134,1453,612,632]" gridrow="9" pageId="11" pageNumber="644">
<td id="76965DE3FF1A0028CC5C1B1AFE1AFD92" box="[300,407,612,632]" gridcol="1" gridrow="9" pageId="11" pageNumber="644">SMF 09-F2</td>
<td id="76965DE3FF1A0028CCDC1B1AFDD7FD92" box="[428,602,612,632]" gridcol="2" gridrow="9" pageId="11" pageNumber="644">55</td>
<td id="76965DE3FF1A0028CF001B1AFD6FFD92" box="[624,738,612,632]" gridcol="3" gridrow="9" pageId="11" pageNumber="644">L: 15.6</td>
<td id="76965DE3FF1A0028CF891B1AFC31FD92" box="[761,956,612,632]" gridcol="4" gridrow="9" pageId="11" pageNumber="644">L: 0.28</td>
<td id="76965DE3FF1A0028CEB91B1AFBDDFD92" box="[969,1104,612,632]" gridcol="5" gridrow="9" pageId="11" pageNumber="644"></td>
<td id="76965DE3FF1A0028C92B1B1AFA8FFD92" box="[1115,1282,612,632]" gridcol="6" gridrow="9" pageId="11" pageNumber="644"></td>
<td id="76965DE3FF1A0028C87D1B1AFA20FD92" box="[1293,1453,612,632]" gridcol="7" gridrow="9" pageId="11" pageNumber="644"></td>
</tr>
<tr id="3547349FFF1A0028CDF61BF9FAEFFD5F" box="[134,1378,647,693]" gridrow="10" pageId="11" pageNumber="644">
<th id="76965DE3FF1A0028CDF61BF9FE96FD5F" box="[134,283,647,693]" gridcol="0" gridrow="10" pageId="11" pageNumber="644">
<taxonomicName id="4C774D5EFF1AFFDCCDF61BF9FE9DFD71" authority="ZPAL V." authorityName="ZPAL V." box="[134,272,647,667]" class="Reptilia" family="Proterochersidae" genus="Proterochersis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="11" pageNumber="644" phylum="Chordata" rank="genus">
<emphasis id="B903EACFFF1AFFDCCDF61BF9FE9DFD71" box="[134,272,647,667]" italics="true" pageId="11" pageNumber="644">Proterochersis</emphasis>
</taxonomicName>
<taxonomicName id="4C774D5EFF1AFFDCCDE51BDFFF73FD5F" authorityName="Szczygielski &amp; Sulej" authorityYear="2016" box="[149,254,673,693]" class="Reptilia" family="Proterochersidae" genus="Proterochersis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="11" pageNumber="644" phylum="Chordata" rank="species" species="porebensis">
<emphasis id="B903EACFFF1AFFDCCDE51BDFFF73FD5F" box="[149,254,673,693]" italics="true" pageId="11" pageNumber="644">porebensis</emphasis>
</taxonomicName>
</th>
<td id="76965DE3FF1A0028CC5C1BF9FE1AFD5F" box="[300,407,647,693]" gridcol="1" gridrow="10" pageId="11" pageNumber="644">ZPAL V. 39/48</td>
<paragraph id="8BC836DDFF1AFFDCCCCC1BF9FAEFFD71" box="[444,1378,647,667]" pageId="11" pageNumber="644">
<td id="76965DE3FF1A0028CCDC1BF9FDD7FD5F" box="[428,602,647,693]" gridcol="2" gridrow="10" pageId="11" pageNumber="644">39.5</td>
<td id="76965DE3FF1A0028CF001BF9FD6FFD5F" box="[624,738,647,693]" gridcol="3" gridrow="10" pageId="11" pageNumber="644"></td>
<td id="76965DE3FF1A0028CF891BF9FC31FD5F" box="[761,956,647,693]" gridcol="4" gridrow="10" pageId="11" pageNumber="644"></td>
<td id="76965DE3FF1A0028CEB91BF9FBDDFD5F" box="[969,1104,647,693]" gridcol="5" gridrow="10" pageId="11" pageNumber="644">R: 12.7</td>
<td id="76965DE3FF1A0028C92B1BF9FA8FFD5F" box="[1115,1282,647,693]" gridcol="6" gridrow="10" pageId="11" pageNumber="644">R: 0.32</td>
<td id="76965DE3FF1A0028C87D1BF9FA20FD5F" box="[1293,1453,647,693]" gridcol="7" gridrow="10" pageId="11" pageNumber="644"></td>
</paragraph>
</tr>
</table>
</paragraph>
<paragraph id="8BC836DDFF1AFFDCCDEB1AABFEDCFBC5" blockId="11.[131,776,758,1071]" pageId="11" pageNumber="644">Overall, based on the histological sections and CT images, the humeri appear denser than the femora of comparable ontogenetic stages.</paragraph>
</subSubSection>
</treatment>
</document>

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<mods:identifier id="74F86C38B52B711C9848AF239396E9EE" type="DOI">10.5852/cr-palevol2023v22a32</mods:identifier>
<mods:identifier id="22471619AA7302C7F4F9B5054608F602" type="ISSN">1777-571X</mods:identifier>
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<paragraph id="8BC836DDFF1AFFDCCDF41D0BFBC3FA04" blockId="11.[130,777,1109,2028]" lastBlockId="11.[811,1457,758,1518]" pageId="11" pageNumber="644">
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.
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Nakajima
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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 Sharpeys 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, Sharpeys 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 Sharpeys 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, Sharpeys 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 Sharpeys 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, Sharpeys 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 Sharpeys 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, Sharpeys 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 (
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Rothschild
<emphasis id="B903EACFFF1AFFDCC91C1A28FB2FFC85" box="[1132,1186,853,879]" italics="true" pageId="11" pageNumber="644">et al.</emphasis>
2012
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). 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
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), 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
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<emphasis id="B903EACFFF1AFFDCC93C1CCDFAC6FA27" box="[1100,1355,1459,1485]" italics="true" pageId="11" pageNumber="644">Proganochelys quenstedtii</emphasis>
</taxonomicName>
(
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;
<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
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2022
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).
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