269:(i) Neural 1, (ii) Neural 2, (iii) Neural 3, (iv) Neural 4, (v) Neural 5, (vi) Neural 6, (vii) Neural 7, (viii) Neural 8, (ix) extra neural, divided, (x) suprapygal, (xi) nuchal, (xii) right peripheral 1, (xiii) right peripheral 2, (xiv) right peripheral 3, (xv) right peripheral 4, (xvi) right peripheral 5, (xvii) right peripheral 6, (xviii) right peripheral 7, (xix) right peripheral 8, (xx) right peripheral 9, (xxi) right peripheral 10, (xxii) right peripheral 11, (xxiii) pygal, (xxiv) left peripheral 11, (xxv) left peripheral 10, (xxvi) left peripheral 9, (xxvii) left peripheral 8, (xxviii) left peripheral 7, (xxix) left peripheral 6, xxx left peripheral 5, xxxi left peripheral 4, (xxxii) left peripheral 3, (xxxiii) left peripheral 2, (xxxiv) left peripheral 1, (xxxv) right 1st rib, (xxxvi) right pleural 1, (xxxvii) right pleural 2, (xxxviii) right pleural 3, (xxxix) right pleural 4, (xl) right pleural 5, (xli) right pleural 6, (xlii) right pleural 7, (xliii) right pleural 8, (xliv) right 10th rib, (xlv) left 1st rib, (xlvi) left pleural 1, (xlvii) left pleural 2, (xlviii) left pleural 3, (xlix) left pleural 4, (l) left pleural 5, (li) left pleural 6, (lii) left pleural 7, (liii) left pleural 8, (liv) left 10th rib, (9-18) centrums.
560:
31:
544:
681:
378:
1204:
1002:, fused first to each other and then to the ribs beneath them. The theory persisted into the 21st century, when Olivier Rieppel proposed a hypothetical turtle precursor, its back covered by bony armour plates in the dermis, which he called the "Polka Dot Ancestor". Michael Lee proposed that the transformation of the carapace began with an unarmoured parareptile and then an armoured pareiasaur, and ended with modern turtles with a fully developed carapace and a relocated rib cage. The theory accounted for the evolution of fossil pareisaurs from
1220:
915:
1023:
395:
416:, what one would call the belly or ventral surface of the shell. It also includes within its structure the anterior and posterior bridge struts and the bridge of the shell. The plastron is made up of nine bones and the two epiplastra at the anterior border of the plastron are homologous to the clavicles of other tetrapods. The rest of the plastral bones are homologous to the
59:
44:
254:
1045:, some 260 million years ago, had a short broad trunk, and a body-case of broadened and somewhat overlapping ribs, suggesting an early stage in the acquisition of a shell. The fossil has been called "a diapsid reptile in the process of becoming secondarily anapsid". Olivier Rieppel summarizes the phylogenetic origins of the ancestral turtles: "
507:. There are six laterally symmetric pairs of scutes on the plastron: gular, humeral, pectoral, abdominal, femoral, and anal (going from the head to the tail down the seam); the abdominal and gular scute seams are approximately the same length, and the femoral and pectoral seams are approximately the same length.
440:
changes in the pathways of rib development often result in malformation or loss of the plastron. This phenomenon occurs in turtle development, but instead of experiencing complete loss of the sternum the turtle body plan repurposes the bone into the form of the plastron, although other analyses find that the
199:
240:
and are a lot like horn or nail tissue. In the center of the carapace are five vertebral scutes and out from these are four pairs of costal scutes. Around the edge of the shell are 12 pairs of marginal scutes. All these scutes are aligned so that for the most part the sutures between the bones are in
224:
Between each of the pleurals are a series of neural bones, which although always present are not always visible, in many species of
Pleurodire they are submerged below the pleurals. Beneath the neural bone is the neural arch which forms the upper half of the encasement for the spinal cord. Below this
925:
The carapacial ridge plays an essential role in the development of the turtle shell. Embryological analyses show that the carapacial ridge initiates the formation of the turtle shell. It causes axial arrest which causes the ribs to be dorsalized, the shoulder girdle to be rearranged and encapsulated
439:
The evolution of the plastron has remained more mysterious, though
Georges Cuvier, a French naturalist and zoologist in the 19th century, wrote that the plastron developed primarily from the sternum of the turtle. This fits well with the knowledge obtained through embryological studies, showing that
244:
On the plastron there are two gular scutes at the front, followed by a pair of pectorals, then abdominals, femorals and lastly anals. A particular variation is the
Pleurodiran turtles have an intergular scute between the gulars at the front, giving them a total of 13 plastral scutes. Compared to the
170:
process, which caused many microstructures to appear to aid survival and motion. Shell shape allows the animal to escape predatory situations. Microstructures can include the scutes mentioned prior or the ribs found internally of the shell. Many ribs can be found within the shell and throughout the
444:
sternum is absent and replaced by the exoskeletal plastron. The ventral ribs are effectively not present, replaced by the plastron, unless the gastralia from which the plastron evolved were once floating ventral ribs. During turtle evolution, there was probably a division of labor between the ribs,
1120:
in southwest China has a partial shell, consisting of a complete bony plastron and an incomplete carapace. The fossil showed that the plastron evolved before the carapace. Like crown turtles, it lacked intercostal muscles, so rib mobility was limited. The ribs were laterally expanded and broadened
232:
The skeletal elements of the plastron are also largely in pairs. Anteriorly there are two epiplastra, with the hyoplastra behind them. These enclose the singular entoplastron. These make up the front half of the plastron and the hyoplastron contains the anterior bridge strut. The posterior half is
194:
The bones of the shell are named for standard vertebrate elements. As such the carapace is made up of eight pleurals on each side, these are a combination of the ribs and fused dermal bone. Outside of this at the anterior of the shell is the single nuchal bone, a series of twelve paired periphals
162:
layer. This layer is important to the strength of the shell surrounding it. In an international study, the layer can be as thick as two to four cells. Even with such a small thickness, the epidermis allows the deformation the shell can experience and provides the shell more support. The epidermis
99:
The turtle's shell is an important study, not just because of the apparent protection it provides for the animal but also as an identification tool, in particular with fossils, as the shell is one of the likely parts of a turtle to survive fossilization. Hence understanding the shell structure in
476:
is different from its ancestor because the gastralia show signs of having once been fused, as indicated by the fossil specimens which show forked ends. This evidence shows a gradual change from paired gastralia, to paired and fused gastralia, and finally to the modern plastron across these three
713:
The appearance of scutes correlates to the transition from aquatic to terrestrial mode of life in tetrapods during the
Carboniferous period (340 Ma). In the evolution from amphibians to terrestrial amniotes, transition in a wide variety of skin structures occurred. Ancestors of turtles likely
709:
with carapace usually being a darker patterning than the plastron, though there are exceptions. Moustakas-Verho and
Cherepanov's embryological study reveals that the patterning of the plastral scutes appear independent from the patterning of carapacial scutes, suggesting that the carapace and
307:, a ridge that runs from front to the back of the animal is present in some species, these may be single, paired or even three rows of them. In most turtles the shell is relatively uniform in structure, species variation in general shape and color being the main differences. However, the
91:
Testudines), completely enclosing all the vital organs of the turtle and in some cases even the head. It is constructed of modified bony elements such as the ribs, parts of the pelvis and other bones found in most reptiles. The bone of the shell consists of both skeletal and
932:
presents evidence of axial arrest that is observed in embryos but lacks fan-shaped ribs and a carapace. This suggests that the primitive carapacial ridge functioned differently and must have gained the function of mediating the ribs and carapace development later. The
696:. The individual scutes as shown above have specific names and are generally consistent across the various species of turtles. Terrestrial tortoises do not shed their scutes. New scutes grow by the addition of keratin layers to the base of each scute. Aquatic
163:
layer is apparent in both sections of the shell, carapace, and plastron, and is thicker in critical areas. A thicker epidermis allows a higher stress force to be experienced without permanent deformation or critical failure of the shell.
1072:, with strong shoulders and forelimbs, and increased muscle attachment structures such as their tubercle on the posterior coracoid and their large and wide terminal phalanges creating shovel-like "hands". Fossoriality may have helped
1911:
Hirasawa, Tatsuya; Pascual-Anaya, Juan; Kamezaki, Naoki; Taniguchi, Mari; Mine, Kanako; Kuratani, Shigeru (2015-05-01). "The evolutionary origin of the turtle shell and its dependence on the axial arrest of the embryonic rib cage".
1203:
241:
the middle of the scutes above. At the anterior of the shell there may be a cervical scute (sometimes incorrectly called a nuchal scute) however the presence or absence of this scute is highly variable, even within species.
704:
over the underlying bony structures; there is a very thin layer of subcutaneous tissue between the scute and the skeleton. The scutes can be brightly colored in some species, and turtle shells often follow
1455:
Wei Zhang, Chengwei Wu, Chenzhao Zhang, Zhen Chen, “Microstructure and mechanical property of turtle shell.” Theoretical and
Applied Mechanics Letters, Volume 2, Issue 1, (2012): 014009, ISSN 2095-0349.
921:
of the shell: seen in the egg at stage 16/17, the carapace is developing. In section, the ribs are growing sideways not downwards, into the carapacial ridge, seen here as a bud, to support the carapace.
150:
which have a floating pelvis. The anterior bridge strut and posterior bridge strut are part of the plastron. On the carapace are the sutures into which they insert, known as the Bridge carapace suture.
1613:
Nagashima, H.; Sugahara, F.; Takechi, M.; Ericsson, R.; Kawashima-Ohya, Y.; Narita, Y.; Kuratani, S. (2009). "Evolution of the turtle body plan by the folding and creation of new muscle connections".
1855:
MacCord, Kate; Caniglia, Guido; Moustakas-Verho, Jacqueline E.; Burke, Ann C. (2015-05-01). "The dawn of chelonian research: Turtles between comparative anatomy and embryology in the 19th century".
225:
the rest of the vertebral column. Some species of turtles have some extra bones called mesoplastra, which are located between the carapace and plastron in the bridge area. They are present in most
586:
The plastral formula is used to compare the sizes of the individual plastral scutes (measured along the midseam). The following plastral scutes are often distinguished (with their abbreviation):
445:
which specialized to stabilize the trunk, and the abdominal muscles, which specialized for respiration, and these changes took place 50 million years before the shell was fully ossified.
334:, are not linked directly to the vertebral column or rib cage allowing the ribs to move freely with the surrounding intercostal muscle. However, analysis of the transitional fossil,
2537:
Kuratani, S (2011). "Evolutionary developmental perspective for the origin of turtles: the folding theory for the shell based on the developmental nature of the carapacial ridge".
998:
Zoologists have sought to explain the evolutionary origin of the turtles, and in particular of their unique carapace. In 1914, J. Versluys proposed that bony plates in the dermis,
1504:
2064:
1107:, is a much larger animal, up to 1.8 metres (5.9 ft) long, with a long tail, and broadened but not overlapping ribs; like the earlier fossils, it has small teeth.
2256:
2201:
1388:
1955:
Lyson, Tyler R.; Schachner, Emma R.; Botha-Brink, Jennifer; Scheyer, Torsten M.; Lambertz, Markus; Bever, G. S.; Rubidge, Bruce S.; de
Queiroz, Kevin (2014-11-07).
1068:
origin for the turtles. During the
Permian, the broadened ribs may have provided great stability in burrowing, giving a body shape resembling the extant fossorial
503:
The plastral scutes join along a central seam down the middle of the plastron. The relative lengths of the seam segments can be used to help identify a species of
2700:
326:
The evolution of the turtle's shell is unique because of how the carapace represents transformed vertebrae and ribs. While other tetrapods have their scapula, or
1130:
of
Germany and Thailand. It lacked the ability to pull its head into its shell, and had a long neck and a long, spiked tail ending in a club, somewhat like an
684:
Pair of pond slider turtles with one on the left having a normal shell (somewhat muddy) and the other on the right, exhibiting scute shedding of shell segments.
3388:
1035:
Recent stem-turtle fossil discoveries provide a "comprehensive scenario" of the evolution of the turtle's shell. A fossil that may be a stem-turtle from the
1352:
107:, among other species, has been used as a material for a wide range of small decorative and practical items since antiquity, but is normally referred to as
1181:
shape underlying each scute. Factors which may contribute to pyramiding include inadequate water supply; the consumption of excessive animal or vegetable
1076:
survive the global mass extinction at the end of the
Permian period, and could have played an essential role in the early evolution of shelled turtles.
195:
then extend along each side. At the posterior of the shell is the pygal bone and in front of this nested behind the eighth pleurals is the suprapygal.
1219:
142:. These are joined by an area called the bridge. The actual suture between the bridge and the plastron is called the anterior bridge strut. In
1665:
The draft genomes of soft-shell turtle and green sea turtle yield insights into the development and evolution of the turtle-specific body plan
2381:
2118:"Variable plastron coloration of the eastern painted turtles Chrysemys picta picta in a single locality of South-Eastern Massachusetts, USA"
52:(left) and skeletal components (right) of a turtle's carapace. Scutes between the costal (= pleural) and marginals are called supramarginal.
3479:
500:, female. This is because of the mating position; the male's concave plastron allows it to more easily mount the female during copulation.
34:
A preserved turtle skeleton showing how the carapace and plastron connect with the rest of the skeleton to form a shell enclosing the body
120:
2400:
Cebra-Thomas, Judith; Tan, Fraser; Sistla, Seeta; Estes, Eileen; Bender, Gunes; Kim, Christine; Riccio, Paul; Gilbert, Scott F. (2005).
526:, while others have a single undivided gular scute. The gular scutes may be referred to as a gular projection if they stick out like a
2833:
2765:
Bever, G. S.; Lyson, Tyler R.; Field, Daniel J.; Bhullar, Bhart-Anjan S. (September 2015). "Evolutionary origin of the turtle skull".
2510:
1240:
171:
shell. The rib structures provide extra structural support but allows the shells to deform elastically depending on the situation the
3206:
Asher, J. Lichtig; Spencer G., Lucas; Klein, Hendrik; Lovelace, David M. (2018). "Triassic turtle tracks and the origin of turtles".
2676:
2343:
2149:
Moustakas-Verho, J. E., R. Zimm, J. Cebra-Thomas, N. K. Lempiainen, A. Kallonen, K. L. Mitchell, K. Hamalainen; et al. (2014).
680:
30:
3381:
2068:
131:
The turtle shell is made up of numerous bony elements, generally named after similar bones in other vertebrates, and a series of
1465:
Pritchard, P.C.H. (1988). "A survey of neural bone variation among recent chelonian species, with functional interpretations".
3358:
966:
further provide an understanding in key gene expression that could potentially be responsible for changing turtle morphology.
5512:
4595:
3268:
5639:
Turtles of the World 2017 Update: Annotated Checklist and Atlas of Taxonomy, Synonymy, Distribution, and Conservation Status
330:, found outside of the ribcage, the scapula for turtles is found inside the ribcage. The shells of other tetrapods, such as
2909:
Chen, Z.-Q.; Benton, M. J. (2012). "The timing and pattern of biotic recovery following the end-Permian mass extinction".
559:
543:
3332:
2458:
Ruckes, Herbert (December 1929). "STUDIES IN CHELONIAN OSTEOLOGY, Part I: TRUSS AND ARCH ANALOGIES IN CHELONIAN PELVES".
5489:
2944:
Schoch, Rainer R.; Sues, Hans-Dieter (2015). "A Middle Triassic stem-turtle and the evolution of the turtle body plan".
2012:
Schoch, Rainer R.; Sues, Hans-Dieter (2015). "A Middle Triassic stem-turtle and the evolution of the turtle body plan".
5637:
5669:
5648:
4171:
3555:
3374:
2116:
Surasinghe, T; Christen, R; Dewey, A; Gouthro, A; Tocchio, K; Sheehan, B; McCulley, T; Dobeib, Y (25 September 2019).
958:
expression in the ventral sclerotome and thus plays a key role in carapacial rib development. Genetic observations of
377:
214:
at pleural IV showing a narrow midline neural bone, lateral pleurals (P) and underlying vertebral neural arch. and C.
3904:
3416:
1026:
Diagram of origins of turtle body plan through the Triassic: isolated bony plates evolved to form a complete shell.
5679:
3282:
1371:
1112:
928:
441:
385:
360:
1149:
Septicemic cutaneous ulcerative disease (SCUD) or "shell rot" causes ulceration of the shell. This is caused by
5674:
4550:
3472:
978:
96:, showing that the complete enclosure of the shell likely evolved by including dermal armor into the rib cage.
2270:
Cherepanov, G. O. (2015). "Scute's polymorphism as a source of evolutionary development of the turtle shell".
5406:
2872:(14). B. S. Rubidge, T. M. Scheyer, K. De Queiroz, E. R. Schachner, R. M. Smith, J. Botha-Brink: 1887–1894.
312:
233:
made up of two hypoplastra (containing the posterior bridge strut) and the rear is a pair of xiphiplastra.
5378:
4178:
4136:
3776:
1165:. The disease progresses to a septicemic infection causing the degradation of the liver and other organs.
566:
336:
316:
175:
is in (i.e., predatory escape). Nonstructural mechanisms have also been in the turtle shell that aids the
108:
4122:
3624:
2580:
Moustakas-Verho, Jacqueline; Thomas, C. T.; Gilbert, Scott F. (2017). "Patterning of the turtle shell".
2250:
2195:
1382:
522:
is the most anterior part of the plastron, the underside of the shell. Some tortoises have paired gular
210:
1403:
1210:
653:
Comparison of the plastral formulas provides distinction between the two species. For example, for the
17:
2359:
Catalogue of the chelonians, rhynchocephalians, and crocodiles in the British Museum (Natural History)
2090:
319:
have lost the scutes and reduced the ossification of the shell. This leaves the shell covered only by
5364:
4030:
3654:
3138:
3069:
3011:
2953:
2918:
2774:
2723:
2279:
2021:
1968:
1801:
1622:
1094:, has more distinctly broadened ribs, T-shaped in cross-section. They vary in shape along the spine.
2858:
220:
at pleural IV showing location of a rudimentary neural bone underneath medially contiguous pleurals.
5349:
3708:
3465:
340:
shows that early ancestors of turtles lost that intercostal muscle usually found between the ribs.
159:
4756:
4721:
4588:
4386:
4327:
4297:
3223:
3162:
3101:
3035:
2977:
2891:
2806:
2747:
2694:
2562:
2483:
2440:
2337:
2295:
2183:
2150:
2045:
1767:
1663:
Wang, Z., J. Pascual-Anaya, A. Zadissa, W. Q. Li, Y. Niimura, Z. Y. Huang, C. Y. Li et al. 2013.
1646:
1527:
1178:
1158:
1117:
1010:
654:
216:
4305:
4059:
914:
412:
The plastron (plural: plastrons or plastra) is the nearly flat part of the shell structure of a
146:
the posterior pelvis is also part of the carapace, fully fused with it. This is not the case in
1737:"Morphogenesis of the turtle shell: The development of a novel structure in tetrapod evolution"
5299:
4282:
4234:
4225:
4108:
3685:
3264:
3188:
3154:
3093:
3085:
3027:
2969:
2883:
2839:
2829:
2798:
2790:
2739:
2682:
2672:
2649:
2597:
2554:
2516:
2506:
2475:
2432:
2424:
2377:
2238:
2175:
2037:
1994:
1986:
1937:
1929:
1890:
1882:
1837:
1819:
1759:
1717:
1638:
1595:
1282:
714:
diverged from amphibians to develop a horny cover in their early terrestrial ancestral forms.
204:
183:. A mucus film covers parts of the shell, allowing some physical protection and also reducing
180:
5091:
4479:
4454:
4415:
4363:
4094:
3639:
3311:
3215:
3146:
3077:
3019:
2961:
2926:
2873:
2782:
2731:
2639:
2589:
2546:
2467:
2416:
2287:
2228:
2165:
2129:
2029:
1976:
1921:
1872:
1864:
1827:
1809:
1751:
1707:
1630:
1585:
1519:
1330:
Zangerl, R. 1969. The turtle shell. In: Gans, C., Bellairs, D.d'A. and Parsons, T.A. (Eds).
1272:
1213:
with ongoing shell rot (circled in red) and scars from previous shell rot (circled in black)
1162:
1022:
288:
280:
259:
104:
5539:
5187:
4916:
4884:
4521:
4430:
4206:
3792:
3537:
1226:
1103:
1069:
424:, like osteoderms of other reptilians; but unlike osteoderms, the plastron also possesses
402:
366:
208:
showing the suture between the wide neural bone (N) and the vertebral neural arch (V). B.
88:
3352:
1755:
3142:
3073:
3015:
2995:
Li, Chun; Wu, Xiao-Chun; Rieppel, Olivier; Wang, Li-Ting; Zhao, Li-Jun (November 2008).
2957:
2922:
2778:
2727:
2714:
Lee, Michael S. Y. (February 1996). "Correlated progression and the origin of turtles".
2283:
2025:
1972:
1805:
1626:
5434:
5424:
5321:
5171:
5011:
4713:
4658:
4400:
4335:
3875:
3866:
3835:
3743:
3632:
3546:
2471:
1832:
1789:
941:
706:
550:
327:
188:
3123:
3054:
2996:
2401:
1736:
464:
the latter of which possesses a fully formed plastron. In place of a modern plastron,
5663:
5614:
5497:
5397:
5386:
5357:
5271:
5155:
5139:
5099:
5033:
4943:
4908:
4852:
4836:
4820:
4794:
4782:
4766:
4744:
4736:
4543:
4254:
4087:
3948:
3895:
3843:
3446:
3426:
3411:
3227:
3053:
Li, Chun; Wu, Xiao-Chun; Rieppel, Olivier; Wang, Li-Ting; Zhao, Li-Jun (2008-11-27).
2981:
2550:
2049:
1650:
1126:
918:
226:
2751:
2566:
2487:
2299:
1771:
973:, the ribs grow sideways into the carapacial ridge, unique to turtles, entering the
295:
beneath the skin to form a hard shell. Exterior to the skin the shell is covered by
5549:
5474:
5450:
4967:
4844:
4828:
4804:
4705:
4650:
4574:
4490:
4464:
4378:
4289:
4268:
4192:
4157:
3967:
3933:
3800:
3716:
3662:
3166:
3105:
3039:
2895:
2810:
2444:
2187:
1681:
1041:
665:
308:
292:
3219:
1788:
Rice, Ritva; Kallonen, Aki; Cebra-Thomas, Judith; Gilbert, Scott F. (2016-05-10).
3260:
1590:
1573:
1014:, but not for how the ribs could have become attached to the bony dermal plates.
657:, the plastral formula is: an > abd > gul > pect > hum >< fem.
5527:
5482:
5328:
5313:
5235:
5227:
5219:
5195:
5075:
5059:
5043:
4959:
4951:
4900:
4876:
4868:
4860:
4728:
4680:
4603:
4528:
4393:
4349:
4275:
4261:
4213:
4070:
3988:
3926:
3919:
3912:
3827:
3769:
3761:
3678:
3670:
3617:
3607:
1426:
Solomon, S. E., J. R. Hendrickson, and L. P. Hendrickson. "The structure of the
1090:
1004:
669:
421:
93:
119:
5606:
5519:
5505:
5466:
5243:
5163:
5131:
5115:
5083:
5021:
4994:
4975:
4892:
4697:
4617:
4610:
4558:
4535:
4445:
4312:
4044:
3851:
3784:
3701:
3693:
3596:
3587:
2878:
2843:
2593:
2520:
2402:"How the turtle forms its shell: a paracrine hypothesis of carapace formation"
2291:
1712:
1695:
1277:
1260:
1194:
1131:
661:
433:
425:
417:
147:
143:
69:
65:
3241:
Kaplan, H. M. (1957). "Septicemic, cutaneous ulcerative disease of turtles".
3192:
3089:
2794:
2743:
2686:
2653:
2479:
2428:
2409:
Journal of Experimental Zoology Part B: Molecular and Developmental Evolution
1990:
1933:
1914:
Journal of Experimental Zoology Part B: Molecular and Developmental Evolution
1886:
1857:
Journal of Experimental Zoology Part B: Molecular and Developmental Evolution
1823:
977:
of the back to support the carapace. The development is signalled locally by
5598:
5458:
5442:
5371:
5335:
5285:
5278:
5203:
5179:
5123:
5067:
4774:
4566:
4514:
4472:
4185:
4143:
4051:
4037:
4023:
4016:
4009:
4002:
3974:
3819:
3647:
1814:
1634:
1065:
1049:
is placed at the bottom of the stem section of the turtle tree, followed by
999:
485:
481:
394:
331:
167:
3158:
3097:
3031:
2973:
2887:
2802:
2601:
2558:
2436:
2242:
2179:
2041:
1998:
1941:
1894:
1841:
1790:"Development of the turtle plastron, the order-defining skeletal structure"
1763:
1721:
1642:
1599:
1286:
488:
scutes allowing the turtle to almost completely enclose itself. In certain
287:, consisting of the animal's ossified ribs fused with the dermal bone. The
1664:
950:) serve as key regulators during the development of the vertebral column.
493:
253:
5413:
5342:
5262:
5251:
5147:
4812:
4689:
4581:
4423:
4408:
4370:
4319:
4245:
4150:
4129:
4115:
4101:
3957:
3884:
3753:
3732:
3724:
3506:
3406:
2372:
Pritchard, Peter Charles Howard; Pritchard, Peter Charles Howard (1979).
2233:
2216:
1431:
1427:
1174:
1150:
1085:
276:
184:
3150:
3081:
3023:
2965:
2786:
2420:
2134:
2117:
2033:
1925:
1877:
1868:
1484:
Thomson, S. & Georges, A. (1996). "Neural bones in chelid turtles".
497:
348:"Plastron" redirects here. For the arthropod structural adaptation, see
58:
43:
5557:
5306:
5211:
5107:
5051:
4631:
4624:
4506:
4356:
4342:
4199:
4164:
4079:
3981:
3521:
3512:
2644:
2628:"The origin of the turtle body plan: evidence from fossils and embryos"
2627:
2170:
1981:
1956:
1531:
1186:
1182:
1098:
1036:
693:
489:
429:
300:
237:
236:
Overlying the boney elements are a series of scutes, which are made of
132:
1574:"Leatherback sea turtle shell: A tough and flexible biological design"
1404:"Turtle without shell: Can They Survive? What's Inside? #WhatTheShell"
1110:
Also in the Late Triassic, some 220 million years ago, the freshwater
5292:
4935:
3527:
3500:
3488:
3366:
2930:
2735:
1124:
The development of a shell reaches completion with the late Triassic
974:
970:
697:
527:
519:
504:
413:
296:
284:
176:
172:
155:
84:
64:
Scutes (left) and skeletal components (right) of a turtle's plastron
3283:"Effects of diet on the systematic utility of the tortoise carapace"
1523:
68:
have an extra scute known as the intergular. It is mostly absent in
2315:
The turtle shell. In: C. Gans (ed.), Biology of the Reptilia, vol 1
1562:
Bojanus, L. H. 1819. Anatome testudinis Europaeae. 178pp, 31 plates
1549:
Pritchard, P.C.H., and P. Trebbau. 1984. The Turtles of Venezuela.
198:
4499:
3566:
3124:"An ancestral turtle from the Late Triassic of southwestern China"
3055:"An ancestral turtle from the Late Triassic of southwestern China"
2997:"An ancestral turtle from the Late Triassic of southwestern China"
1154:
1021:
982:
954:
expression in the neural tube is essential for the maintenance of
913:
689:
679:
523:
252:
197:
135:
118:
49:
29:
3122:
Li, C.; Wu, X.-C.; Rieppel, O.; Wang, L.-T.; Zhao, L.-J. (2008).
1735:
Gilbert, S. F.; Loredo, G. A.; Brukman, A.; Burke, A. C. (2001).
138:
which are also uniquely named. The ventral surface is called the
3995:
2151:"The origin and loss of periodic patterning in the turtle shell"
935:
349:
320:
3461:
3370:
2067:. Netherlands Biodiversity Information Facility. Archived from
660:
Turtle plastrons were used by the ancient Chinese in a type of
456:
serves as an intermediate form between two early stem-turtles,
1505:"Neural bones in chelid turtles from Australia and New Guinea"
1190:
492:
the sex of a testudine can be told by whether the plastron is
3457:
3362:
article (including video) on how the turtle evolved its shell
2626:
Schoch, Rainer R.; Sues, Hans‐Dieter; Benson, Roger (2019).
1057:
along the turtle stem and on to more crown-ward turtles".
1334:, Vol 1, Morphology A. London: Academic Press. pp. 311–340
1121:
without ossification, like the embryos of modern turtles.
420:
of other tetrapods. The plastron has been described as an
100:
living species provides comparable material with fossils.
452:
provides additional clues as to how the plastron formed.
127:. Pe=Peripheral, P1=Pleural 1, BCS=Bridge Carapace Suture
2395:
2393:
2317:. New York and London: Academic Press. pp. 311–339.
700:
shed individual scutes. The scute effectively forms the
1957:"Origin of the unique ventilatory apparatus of turtles"
3117:
3115:
2217:"Environmental causation of turtle scutes anomalies"
291:
and expanded ribs are fused through ossification to
5589:
5582:
5575:
5537:
5423:
5396:
5261:
5031:
5009:
5002:
4993:
4926:
4792:
4754:
4679:
4672:
4641:
4489:
4453:
4444:
4244:
4223:
4069:
3956:
3947:
3894:
3874:
3865:
3810:
3742:
3606:
3595:
3586:
3554:
3545:
3536:
2621:
2619:
2617:
2615:
2613:
2611:
2332:. Ithaca, New York: Comstock Publishing Associates.
1680:Hirasawa, T., H. Nagashima, and S. Kuratani. 2013.
1177:, in which the shell grows unevenly resulting in a
994:
Bony dermal plates theory: the "Polka Dot Ancestor"
303:that protect the shell from scrapes and bruises. A
202:Transverse sections through the first neural of A.
2826:Turtles as hopeful monsters: origins and evolution
2669:Turtles as hopeful monsters: origins and evolution
2503:Turtles as hopeful monsters: origins and evolution
27:Shield for the ventral and dorsal parts of turtles
3182:The comparative osteology of the Triassic turtle
480:In certain families there is a hinge between the
2532:
2530:
1351:Thomson, S., White, A. & Georges, A (1997).
283:(back), convex part of the shell structure of a
83:is a shield for the ventral and dorsal parts of
1794:Proceedings of the National Academy of Sciences
1503:Rhodin, A.G.J. & Mittermeier, R.A. (1977).
1682:The endoskeletal origin of the turtle carapace
1304:Romer, A.S. (1956) Osteology of the Reptiles.
926:in the rib cage, and the carapace to develop.
5636:Phylogenetic arrangement of turtles based on
3473:
3382:
3312:"BASIC HUSBANDRY AND NUTRITION OF CHELONIANS"
2582:Current Opinion in Genetics & Development
448:The discovery of an ancestral turtle fossil,
8:
2255:: CS1 maint: multiple names: authors list (
2200:: CS1 maint: multiple names: authors list (
2063:C.H. Ernst; R.G.M. Altenburg; R.W. Barbour.
1387:: CS1 maint: multiple names: authors list (
577:Gular anatomical formations in other species
1173:Pyramiding is a shell deformity of captive
718:Names of epidermal scutes in turtle shells
5586:
5579:
5006:
4999:
4676:
4450:
3953:
3871:
3603:
3592:
3560:
3551:
3542:
3480:
3466:
3458:
3389:
3375:
3367:
2699:: CS1 maint: location missing publisher (
2460:Annals of the New York Academy of Sciences
2361:. London: British Museum. pp. 311 pp.
1545:
1543:
1541:
468:has paired gastralia, like those found in
2877:
2643:
2505:. Indiana University Press. p. 146.
2232:
2169:
2133:
1980:
1876:
1831:
1813:
1711:
1589:
1276:
358:Comparison of plastrons of a Cryptodire (
2828:. Indiana University Press. p. 70.
1326:
1324:
1322:
1320:
1318:
1316:
1314:
1088:of Germany, some 240 million years ago,
1060:Tyler Lyson and colleagues suggest that
716:
2215:Zimm, R.; Wyneken, Bentley, J. (2017).
1346:
1344:
1342:
1340:
1300:
1298:
1296:
1251:
1199:
18:Septicemic cutaneous ulcerative disease
2859:"Fossorial Origin of the Turtle Shell"
2692:
2376:. Neptune, N.J.: T.F.H. Publications.
2335:
2248:
2193:
1696:"Palaeontology: Turtles in transition"
1380:
323:. These are all highly aquatic forms.
2671:. Bloomington, Indiana. p. 157.
1906:
1904:
1783:
1781:
1080:Triassic: evolution of complete shell
969:During the development of the turtle
7:
3355:, has more information on pyramiding
2097:. SeaWorld Parks & Entertainment
1756:10.1046/j.1525-142x.2001.003002047.x
1357:: Identification of a Living Fossil"
536:The gular scutes or gular projection
2221:Integrative and Comparative Biology
1584:. W. Yang, and M. A. Meyers: 2–12.
1551:SSAR Contributions to Herpetology 2
166:The shape of the shell is from its
2857:Lyson, T. R.; et al. (2016).
2472:10.1111/j.1749-6632.1929.tb55191.x
1486:Chelonian Conservation and Biology
1241:Evolutionary developmental biology
25:
1097:A Late Triassic stem-turtle from
688:The turtle's shell is covered in
299:, which are horny plates made of
257:Exploded view of the carapace of
2551:10.1111/j.1525-142x.2010.00451.x
1364:Memoirs of the Queensland Museum
1218:
1202:
565:The paired gular scutes of this
558:
542:
393:
376:
57:
42:
3436:
1084:A stem-turtle from the Middle
245:12 in all Cryptodiran turtles.
123:Internal anterior carapace of
1:
3431:
3421:
3220:10.1080/08912963.2017.1339037
2342:: CS1 maint: date and year (
710:plastron evolved separately.
569:are visible beneath its neck.
3261:Reptile Medicine and Surgery
1591:10.1016/j.actbio.2015.09.023
1370:(1): 327–336. Archived from
5649:List of Testudines families
3180:Gaffney, Eugene S. (1990).
2539:Evolution & Development
1744:Evolution & Development
1467:Acta Zoologica Cracoviensia
1031:Permian: first stem-turtles
5696:
2091:"Physical Characteristics"
1684:. Nature Communications 4.
1442:(the loggerhead turtle)."
1402:Lum, Steven (2021-12-21).
347:
5632:
3563:
3495:
3402:
3333:"Pyramiding in Tortoises"
2879:10.1016/j.cub.2016.05.020
2824:Rieppel, Olivier (2017).
2667:Rieppel, Olivier (2017).
2594:10.1016/j.gde.2017.03.016
2501:Rieppel, Olivier (2017).
2292:10.1134/S003103011514004X
1713:10.1016/j.cub.2013.05.011
1278:10.1016/j.cub.2016.12.040
1113:Odontochelys semitestacea
979:fibroblast growth factors
929:Odontochelys semitestacea
386:Chrysemys picta marginata
361:Chrysemys picta marginata
3353:African Tortoise Website
3337:www.reptilesmagazine.com
2357:Boulenger, G.A. (1889).
154:In the shell there is a
3243:Proc. Animal Care Panel
2374:Encyclopedia of Turtles
2272:Paleontological Journal
2122:Herpetological Bulletin
1815:10.1073/pnas.1600958113
1635:10.1126/science.1173826
1438:(the green turtle) and
1332:Biology of the Reptilia
1259:Cordero, G. A. (2017).
1694:Lee, M. S. Y. (2013).
1306:Univ. of Chicago Press
1229:with severe pyramiding
1209:Plastron of wild male
1027:
922:
685:
567:Aldabra giant tortoise
549:Gular projection on a
337:Eunotosaurus africanus
317:leatherback sea turtle
272:
221:
128:
35:
3263:, 2nd ed., Saunders,
1961:Nature Communications
1434:of juvenile turtles,
1025:
1018:Broadened ribs theory
917:
683:
350:Gill § Plastrons
256:
211:Chelodina longicollis
201:
122:
33:
2313:Zangerl, R. (1969).
2065:"Terrapene carolina"
1572:Chen, I. H. (2015).
1408:Journeying The Globe
1355:Emydura lavarackorum
1261:"The Turtle's Shell"
1157:entering through an
450:Pappochelys rosinae,
364:) and a Pleurodire (
3417:Fibropapillomatosis
3290:Island Biodiversity
3281:Gerlach, J (2004).
3151:10.1038/nature07533
3143:2008Natur.456..497L
3082:10.1038/nature07533
3074:2008Natur.456..497L
3024:10.1038/nature07533
3016:2008Natur.456..497L
2966:10.1038/nature14472
2958:2015Natur.523..584S
2923:2012NatGe...5..375C
2787:10.1038/nature14900
2779:2015Natur.525..239B
2728:1996Natur.379..812L
2421:10.1002/jez.b.21059
2330:Handbook of Turtles
2328:Carr, A.F. (1952).
2284:2015PalJ...49.1635C
2135:10.33256/hb150.2325
2034:10.1038/nature14472
2026:2015Natur.523..584S
1973:2014NatCo...5.5211L
1926:10.1002/jez.b.22579
1869:10.1002/jez.b.22587
1806:2016PNAS..113.5317R
1627:2009Sci...325..193N
989:Evolutionary origin
719:
5670:Turtle terminology
4757:Nanhsiungchelyidae
4722:Macrocephalochelys
3397:Turtle terminology
3339:. 23 January 2014.
3208:Historical Biology
2645:10.1111/pala.12460
2234:10.1093/icb/icx066
2171:10.1242/dev.109041
1982:10.1038/ncomms6211
1578:Acta Biomaterialia
1444:Journal of anatomy
1353:"Re-Evaluation of
1211:Hermann's tortoise
1197:; poor nutrition.
1028:
923:
717:
686:
655:eastern box turtle
273:
222:
217:Emydura subglobosa
129:
115:Shell nomenclature
36:
5657:
5656:
5628:
5627:
5624:
5623:
5571:
5570:
5567:
5566:
4989:
4988:
4985:
4984:
4668:
4667:
4440:
4439:
3943:
3942:
3861:
3860:
3582:
3581:
3578:
3577:
3574:
3573:
3455:
3454:
3259:Mader, D. (2006)
3137:(7221): 497–501.
3068:(7221): 497–501.
3010:(7221): 497–501.
2952:(7562): 584–587.
2911:Nature Geoscience
2773:(7568): 239–242.
2722:(6568): 812–815.
2383:978-0-87666-918-1
2278:(14): 1635–1644.
2164:(15): 3033–3039.
2020:(7562): 584–587.
1800:(19): 5317–5322.
1706:(12): R513–R515.
1621:(5937): 193–196.
1039:of South Africa,
904:
903:
692:that are made of
649:
648:
383:Plastral view of
205:Aspideretes hurum
103:The shell of the
16:(Redirected from
5687:
5680:Armour (zoology)
5587:
5580:
5092:Chupacabrachelys
5007:
5000:
4677:
4455:Carettochelyidae
4451:
3954:
3872:
3604:
3593:
3561:
3552:
3543:
3482:
3475:
3468:
3459:
3391:
3384:
3377:
3368:
3341:
3340:
3329:
3323:
3322:
3316:
3310:Innis, Charles.
3307:
3301:
3300:
3298:
3296:
3287:
3278:
3272:
3257:
3251:
3250:
3238:
3232:
3231:
3214:(8): 1112–1122.
3203:
3197:
3196:
3177:
3171:
3170:
3128:
3119:
3110:
3109:
3059:
3050:
3044:
3043:
3001:
2992:
2986:
2985:
2941:
2935:
2934:
2931:10.1038/ngeo1475
2906:
2900:
2899:
2881:
2863:
2854:
2848:
2847:
2821:
2815:
2814:
2762:
2756:
2755:
2736:10.1038/379812a0
2711:
2705:
2704:
2698:
2690:
2664:
2658:
2657:
2647:
2623:
2606:
2605:
2577:
2571:
2570:
2534:
2525:
2524:
2498:
2492:
2491:
2455:
2449:
2448:
2406:
2397:
2388:
2387:
2369:
2363:
2362:
2354:
2348:
2347:
2341:
2333:
2325:
2319:
2318:
2310:
2304:
2303:
2267:
2261:
2260:
2254:
2246:
2236:
2227:(6): 1303–1311.
2212:
2206:
2205:
2199:
2191:
2173:
2155:
2146:
2140:
2139:
2137:
2113:
2107:
2106:
2104:
2102:
2087:
2081:
2080:
2078:
2076:
2060:
2054:
2053:
2009:
2003:
2002:
1984:
1952:
1946:
1945:
1908:
1899:
1898:
1880:
1852:
1846:
1845:
1835:
1817:
1785:
1776:
1775:
1741:
1732:
1726:
1725:
1715:
1691:
1685:
1678:
1672:
1661:
1655:
1654:
1610:
1604:
1603:
1593:
1569:
1563:
1560:
1554:
1547:
1536:
1535:
1509:
1500:
1494:
1493:
1481:
1475:
1474:
1462:
1456:
1453:
1447:
1446:145 (1986): 123.
1424:
1418:
1417:
1415:
1414:
1399:
1393:
1392:
1386:
1378:
1376:
1361:
1348:
1335:
1328:
1309:
1302:
1291:
1290:
1280:
1271:(5): R168–R169.
1256:
1222:
1206:
1163:animal husbandry
720:
591:
590:
582:Plastral formula
562:
546:
516:gular projection
397:
380:
315:turtles and the
260:Emys orbicularis
105:hawksbill turtle
61:
46:
21:
5695:
5694:
5690:
5689:
5688:
5686:
5685:
5684:
5675:Reptile anatomy
5660:
5659:
5658:
5653:
5620:
5563:
5540:Sahonachelyidae
5533:
5419:
5392:
5257:
5188:Phosphatochelys
5027:
4981:
4922:
4917:Terlinguachelys
4885:Pneumatoarthrus
4788:
4750:
4664:
4637:
4485:
4436:
4240:
4219:
4207:Siebenrockiella
4065:
3939:
3890:
3857:
3806:
3793:Mesodermochelys
3738:
3599:
3570:
3532:
3491:
3486:
3456:
3451:
3398:
3395:
3349:
3344:
3331:
3330:
3326:
3314:
3309:
3308:
3304:
3294:
3292:
3285:
3280:
3279:
3275:
3258:
3254:
3240:
3239:
3235:
3205:
3204:
3200:
3179:
3178:
3174:
3126:
3121:
3120:
3113:
3057:
3052:
3051:
3047:
2999:
2994:
2993:
2989:
2943:
2942:
2938:
2908:
2907:
2903:
2866:Current Biology
2861:
2856:
2855:
2851:
2836:
2823:
2822:
2818:
2764:
2763:
2759:
2713:
2712:
2708:
2691:
2679:
2666:
2665:
2661:
2625:
2624:
2609:
2579:
2578:
2574:
2536:
2535:
2528:
2513:
2500:
2499:
2495:
2457:
2456:
2452:
2404:
2399:
2398:
2391:
2384:
2371:
2370:
2366:
2356:
2355:
2351:
2334:
2327:
2326:
2322:
2312:
2311:
2307:
2269:
2268:
2264:
2247:
2214:
2213:
2209:
2192:
2153:
2148:
2147:
2143:
2115:
2114:
2110:
2100:
2098:
2089:
2088:
2084:
2074:
2072:
2071:on 24 July 2011
2062:
2061:
2057:
2011:
2010:
2006:
1954:
1953:
1949:
1910:
1909:
1902:
1854:
1853:
1849:
1787:
1786:
1779:
1739:
1734:
1733:
1729:
1700:Current Biology
1693:
1692:
1688:
1679:
1675:
1669:Nature Genetics
1662:
1658:
1612:
1611:
1607:
1571:
1570:
1566:
1561:
1557:
1548:
1539:
1524:10.2307/1443917
1507:
1502:
1501:
1497:
1483:
1482:
1478:
1464:
1463:
1459:
1454:
1450:
1440:Caretta caretta
1425:
1421:
1412:
1410:
1401:
1400:
1396:
1379:
1374:
1359:
1350:
1349:
1338:
1329:
1312:
1303:
1294:
1265:Current Biology
1258:
1257:
1253:
1249:
1237:
1230:
1227:gopher tortoise
1223:
1214:
1207:
1171:
1145:
1140:
1104:Eorhynchochelys
1082:
1070:gopher tortoise
1033:
1020:
996:
991:
912:
906:
827:(extra-) gular
732:Pritchard 1979
729:Boulenger 1889
678:
584:
574:
573:
572:
571:
570:
563:
555:
554:
547:
538:
537:
410:
409:
408:
407:
406:
403:Chelodina canni
398:
390:
389:
381:
372:
371:
367:Chelodina canni
353:
346:
328:shoulder blades
271:
270:
267:
251:
117:
77:
76:
75:
74:
73:
62:
54:
53:
47:
28:
23:
22:
15:
12:
11:
5:
5693:
5691:
5683:
5682:
5677:
5672:
5662:
5661:
5655:
5654:
5652:
5651:
5644:
5643:
5642:. † = extinct.
5633:
5630:
5629:
5626:
5625:
5622:
5621:
5619:
5618:
5610:
5602:
5593:
5591:
5584:
5577:
5573:
5572:
5569:
5568:
5565:
5564:
5562:
5561:
5553:
5544:
5542:
5535:
5534:
5532:
5531:
5523:
5516:
5509:
5501:
5493:
5486:
5478:
5470:
5462:
5454:
5446:
5438:
5435:Albertwoodemys
5429:
5427:
5425:Podocnemididae
5421:
5420:
5418:
5417:
5410:
5402:
5400:
5394:
5393:
5391:
5390:
5382:
5375:
5368:
5361:
5353:
5346:
5339:
5332:
5325:
5322:Lomalatachelys
5317:
5310:
5303:
5296:
5289:
5282:
5275:
5267:
5265:
5259:
5258:
5256:
5255:
5247:
5239:
5231:
5223:
5215:
5207:
5199:
5191:
5183:
5175:
5172:Labrostochelys
5167:
5159:
5151:
5143:
5135:
5127:
5119:
5111:
5103:
5095:
5087:
5079:
5071:
5063:
5055:
5047:
5038:
5036:
5029:
5028:
5026:
5025:
5016:
5014:
5012:Araripemydidae
5004:
4997:
4991:
4990:
4987:
4986:
4983:
4982:
4980:
4979:
4971:
4963:
4955:
4947:
4939:
4930:
4928:
4924:
4923:
4921:
4920:
4912:
4904:
4896:
4888:
4880:
4872:
4864:
4856:
4848:
4840:
4832:
4824:
4816:
4808:
4799:
4797:
4790:
4789:
4787:
4786:
4778:
4770:
4761:
4759:
4752:
4751:
4749:
4748:
4740:
4732:
4725:
4717:
4714:Emarginachelys
4709:
4701:
4693:
4685:
4683:
4674:
4670:
4669:
4666:
4665:
4663:
4662:
4659:Sinaspideretes
4654:
4645:
4643:
4639:
4638:
4636:
4635:
4628:
4621:
4614:
4607:
4599:
4592:
4585:
4578:
4570:
4562:
4554:
4547:
4539:
4532:
4525:
4518:
4510:
4503:
4495:
4493:
4487:
4486:
4484:
4483:
4476:
4468:
4459:
4457:
4448:
4442:
4441:
4438:
4437:
4435:
4434:
4427:
4419:
4412:
4404:
4397:
4390:
4382:
4374:
4367:
4360:
4353:
4346:
4339:
4336:Hesperotestudo
4331:
4323:
4316:
4309:
4301:
4293:
4286:
4279:
4272:
4265:
4258:
4250:
4248:
4242:
4241:
4239:
4238:
4230:
4228:
4226:Platysternidae
4221:
4220:
4218:
4217:
4210:
4203:
4196:
4189:
4182:
4175:
4168:
4161:
4154:
4147:
4140:
4133:
4126:
4119:
4112:
4105:
4098:
4091:
4083:
4075:
4073:
4067:
4066:
4064:
4063:
4055:
4048:
4041:
4034:
4027:
4020:
4013:
4006:
3999:
3992:
3985:
3978:
3971:
3962:
3960:
3951:
3945:
3944:
3941:
3940:
3938:
3937:
3930:
3923:
3916:
3908:
3900:
3898:
3892:
3891:
3889:
3888:
3880:
3878:
3876:Dermatemydidae
3869:
3867:Kinosternoidea
3863:
3862:
3859:
3858:
3856:
3855:
3847:
3839:
3836:Procolpochelys
3831:
3823:
3814:
3812:
3808:
3807:
3805:
3804:
3796:
3788:
3780:
3773:
3765:
3757:
3748:
3746:
3744:Dermochelyidae
3740:
3739:
3737:
3736:
3728:
3720:
3712:
3705:
3697:
3689:
3682:
3674:
3666:
3658:
3651:
3643:
3636:
3633:Carolinochelys
3628:
3621:
3612:
3610:
3601:
3590:
3584:
3583:
3580:
3579:
3576:
3575:
3572:
3571:
3564:
3558:
3549:
3540:
3534:
3533:
3531:
3530:
3524:
3515:
3509:
3503:
3496:
3493:
3492:
3487:
3485:
3484:
3477:
3470:
3462:
3453:
3452:
3450:
3449:
3444:
3439:
3434:
3429:
3424:
3419:
3414:
3409:
3403:
3400:
3399:
3396:
3394:
3393:
3386:
3379:
3371:
3365:
3364:
3356:
3348:
3347:External links
3345:
3343:
3342:
3324:
3302:
3273:
3252:
3233:
3198:
3172:
3111:
3045:
2987:
2936:
2917:(6): 375–383.
2901:
2849:
2835:978-0253024756
2834:
2816:
2757:
2706:
2677:
2659:
2638:(3): 375–393.
2607:
2572:
2526:
2512:978-0253024756
2511:
2493:
2450:
2415:(6): 558–569.
2389:
2382:
2364:
2349:
2320:
2305:
2262:
2207:
2141:
2108:
2082:
2055:
2004:
1947:
1920:(3): 194–207.
1900:
1863:(3): 169–180.
1847:
1777:
1727:
1686:
1673:
1656:
1605:
1564:
1555:
1537:
1518:(2): 370–372.
1495:
1476:
1473:(26): 625–686.
1457:
1448:
1436:Chelonia mydas
1419:
1394:
1377:on 2015-06-09.
1336:
1310:
1292:
1250:
1248:
1245:
1244:
1243:
1236:
1233:
1232:
1231:
1224:
1217:
1215:
1208:
1201:
1170:
1167:
1144:
1141:
1139:
1136:
1081:
1078:
1064:might imply a
1032:
1029:
1019:
1016:
995:
992:
990:
987:
942:Sonic hedgehog
911:
908:
902:
901:
898:
895:
892:
888:
887:
884:
881:
878:
874:
873:
870:
867:
864:
860:
859:
856:
853:
850:
846:
845:
842:
839:
836:
832:
831:
828:
825:
822:
818:
817:
814:
811:
808:
804:
803:
800:
797:
794:
793:12th marginal
790:
789:
786:
783:
780:
776:
775:
772:
769:
766:
762:
761:
758:
755:
752:
748:
747:
744:
741:
738:
734:
733:
730:
727:
724:
677:
674:
651:
650:
647:
646:
643:
639:
638:
635:
631:
630:
627:
623:
622:
619:
615:
614:
611:
607:
606:
603:
599:
598:
595:
583:
580:
579:
578:
564:
557:
556:
551:Texas tortoise
548:
541:
540:
539:
535:
534:
533:
532:
400:Plastral view
399:
392:
391:
382:
375:
374:
373:
357:
356:
355:
354:
345:
342:
268:
265:
264:
250:
247:
125:Elseya dentata
116:
113:
63:
56:
55:
48:
41:
40:
39:
38:
37:
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
5692:
5681:
5678:
5676:
5673:
5671:
5668:
5667:
5665:
5650:
5646:
5645:
5641:
5640:
5635:
5634:
5631:
5617:
5616:
5615:Tacuarembemys
5611:
5609:
5608:
5603:
5601:
5600:
5595:
5594:
5592:
5588:
5585:
5581:
5578:
5574:
5560:
5559:
5554:
5552:
5551:
5550:Sahonachelys
5546:
5545:
5543:
5541:
5536:
5530:
5529:
5524:
5522:
5521:
5517:
5515:
5514:
5513:Peltocephalus
5510:
5508:
5507:
5502:
5500:
5499:
5498:Lapparentemys
5494:
5492:
5491:
5487:
5485:
5484:
5479:
5477:
5476:
5471:
5469:
5468:
5463:
5461:
5460:
5455:
5453:
5452:
5447:
5445:
5444:
5439:
5437:
5436:
5431:
5430:
5428:
5426:
5422:
5416:
5415:
5411:
5409:
5408:
5404:
5403:
5401:
5399:
5398:Pelomedusidae
5395:
5389:
5388:
5387:Yaminuechelys
5383:
5381:
5380:
5376:
5374:
5373:
5369:
5367:
5366:
5362:
5360:
5359:
5358:Prochelidella
5354:
5352:
5351:
5347:
5345:
5344:
5340:
5338:
5337:
5333:
5331:
5330:
5326:
5324:
5323:
5318:
5316:
5315:
5311:
5309:
5308:
5304:
5302:
5301:
5297:
5295:
5294:
5290:
5288:
5287:
5283:
5281:
5280:
5276:
5274:
5273:
5272:Acanthochelys
5269:
5268:
5266:
5264:
5260:
5254:
5253:
5248:
5246:
5245:
5240:
5238:
5237:
5232:
5230:
5229:
5224:
5222:
5221:
5216:
5214:
5213:
5208:
5206:
5205:
5200:
5198:
5197:
5192:
5190:
5189:
5184:
5182:
5181:
5176:
5174:
5173:
5168:
5166:
5165:
5160:
5158:
5157:
5156:Kinkonychelys
5152:
5150:
5149:
5144:
5142:
5141:
5140:Itapecuruemys
5136:
5134:
5133:
5128:
5126:
5125:
5120:
5118:
5117:
5112:
5110:
5109:
5104:
5102:
5101:
5100:Eotaphrosphys
5096:
5094:
5093:
5088:
5086:
5085:
5080:
5078:
5077:
5072:
5070:
5069:
5064:
5062:
5061:
5056:
5054:
5053:
5048:
5046:
5045:
5040:
5039:
5037:
5035:
5034:Bothremydidae
5030:
5024:
5023:
5018:
5017:
5015:
5013:
5008:
5005:
5001:
4998:
4996:
4992:
4978:
4977:
4972:
4970:
4969:
4964:
4962:
4961:
4956:
4954:
4953:
4948:
4946:
4945:
4944:Argillochelys
4940:
4938:
4937:
4932:
4931:
4929:
4925:
4919:
4918:
4913:
4911:
4910:
4909:Santanachelys
4905:
4903:
4902:
4897:
4895:
4894:
4889:
4887:
4886:
4881:
4879:
4878:
4873:
4871:
4870:
4865:
4863:
4862:
4857:
4855:
4854:
4853:Desmatochelys
4849:
4847:
4846:
4841:
4839:
4838:
4837:Calcarichelys
4833:
4831:
4830:
4825:
4823:
4822:
4821:Atlantochelys
4817:
4815:
4814:
4809:
4807:
4806:
4801:
4800:
4798:
4796:
4795:Protostegidae
4791:
4785:
4784:
4783:Jiangxichelys
4779:
4777:
4776:
4771:
4769:
4768:
4767:Anomalochelys
4763:
4762:
4760:
4758:
4753:
4747:
4746:
4745:Protochelydra
4741:
4739:
4738:
4737:Planiplastron
4733:
4731:
4730:
4726:
4724:
4723:
4718:
4716:
4715:
4710:
4708:
4707:
4702:
4700:
4699:
4694:
4692:
4691:
4687:
4686:
4684:
4682:
4678:
4675:
4671:
4661:
4660:
4655:
4653:
4652:
4647:
4646:
4644:
4640:
4634:
4633:
4629:
4627:
4626:
4622:
4620:
4619:
4615:
4613:
4612:
4608:
4606:
4605:
4600:
4598:
4597:
4593:
4591:
4590:
4586:
4584:
4583:
4579:
4577:
4576:
4571:
4569:
4568:
4563:
4561:
4560:
4555:
4553:
4552:
4548:
4546:
4545:
4544:Drazinderetes
4540:
4538:
4537:
4533:
4531:
4530:
4526:
4524:
4523:
4519:
4517:
4516:
4511:
4509:
4508:
4504:
4502:
4501:
4497:
4496:
4494:
4492:
4488:
4482:
4481:
4480:Carettochelys
4477:
4475:
4474:
4469:
4467:
4466:
4461:
4460:
4458:
4456:
4452:
4449:
4447:
4443:
4433:
4432:
4428:
4426:
4425:
4420:
4418:
4417:
4413:
4411:
4410:
4405:
4403:
4402:
4398:
4396:
4395:
4391:
4389:
4388:
4383:
4381:
4380:
4375:
4373:
4372:
4368:
4366:
4365:
4364:Malacochersus
4361:
4359:
4358:
4354:
4352:
4351:
4347:
4345:
4344:
4340:
4338:
4337:
4332:
4330:
4329:
4324:
4322:
4321:
4317:
4315:
4314:
4310:
4308:
4307:
4302:
4300:
4299:
4294:
4292:
4291:
4287:
4285:
4284:
4280:
4278:
4277:
4273:
4271:
4270:
4266:
4264:
4263:
4259:
4257:
4256:
4255:Aldabrachelys
4252:
4251:
4249:
4247:
4243:
4237:
4236:
4232:
4231:
4229:
4227:
4222:
4216:
4215:
4211:
4209:
4208:
4204:
4202:
4201:
4197:
4195:
4194:
4190:
4188:
4187:
4183:
4181:
4180:
4176:
4174:
4173:
4169:
4167:
4166:
4162:
4160:
4159:
4155:
4153:
4152:
4148:
4146:
4145:
4141:
4139:
4138:
4137:Leucocephalon
4134:
4132:
4131:
4127:
4125:
4124:
4120:
4118:
4117:
4113:
4111:
4110:
4106:
4104:
4103:
4099:
4097:
4096:
4092:
4090:
4089:
4088:Banhxeochelys
4084:
4082:
4081:
4077:
4076:
4074:
4072:
4068:
4062:
4061:
4056:
4054:
4053:
4049:
4047:
4046:
4042:
4040:
4039:
4035:
4033:
4032:
4028:
4026:
4025:
4021:
4019:
4018:
4014:
4012:
4011:
4007:
4005:
4004:
4000:
3998:
3997:
3993:
3991:
3990:
3986:
3984:
3983:
3979:
3977:
3976:
3972:
3970:
3969:
3964:
3963:
3961:
3959:
3955:
3952:
3950:
3949:Testudinoidea
3946:
3936:
3935:
3931:
3929:
3928:
3924:
3922:
3921:
3917:
3915:
3914:
3909:
3907:
3906:
3902:
3901:
3899:
3897:
3896:Kinosternidae
3893:
3887:
3886:
3882:
3881:
3879:
3877:
3873:
3870:
3868:
3864:
3854:
3853:
3848:
3846:
3845:
3844:Protosphargis
3840:
3838:
3837:
3832:
3830:
3829:
3824:
3822:
3821:
3816:
3815:
3813:
3809:
3803:
3802:
3797:
3795:
3794:
3789:
3787:
3786:
3781:
3779:
3778:
3774:
3772:
3771:
3766:
3764:
3763:
3758:
3756:
3755:
3750:
3749:
3747:
3745:
3741:
3735:
3734:
3729:
3727:
3726:
3721:
3719:
3718:
3713:
3711:
3710:
3706:
3704:
3703:
3698:
3696:
3695:
3690:
3688:
3687:
3683:
3681:
3680:
3675:
3673:
3672:
3667:
3665:
3664:
3659:
3657:
3656:
3652:
3650:
3649:
3644:
3642:
3641:
3637:
3635:
3634:
3629:
3627:
3626:
3622:
3620:
3619:
3614:
3613:
3611:
3609:
3605:
3602:
3600:(Sea turtles)
3598:
3594:
3591:
3589:
3585:
3569:
3568:
3562:
3559:
3557:
3553:
3550:
3548:
3544:
3541:
3539:
3535:
3529:
3525:
3523:
3519:
3516:
3514:
3510:
3508:
3504:
3502:
3498:
3497:
3494:
3490:
3483:
3478:
3476:
3471:
3469:
3464:
3463:
3460:
3448:
3447:Tortoiseshell
3445:
3443:
3440:
3438:
3435:
3433:
3430:
3428:
3427:Jackson ratio
3425:
3423:
3420:
3418:
3415:
3413:
3412:Chelonitoxism
3410:
3408:
3405:
3404:
3401:
3392:
3387:
3385:
3380:
3378:
3373:
3372:
3369:
3363:
3361:
3360:New Scientist
3357:
3354:
3351:
3350:
3346:
3338:
3334:
3328:
3325:
3320:
3313:
3306:
3303:
3291:
3284:
3277:
3274:
3270:
3266:
3262:
3256:
3253:
3248:
3244:
3237:
3234:
3229:
3225:
3221:
3217:
3213:
3209:
3202:
3199:
3194:
3190:
3186:
3184:Proganochelys
3183:
3176:
3173:
3168:
3164:
3160:
3156:
3152:
3148:
3144:
3140:
3136:
3132:
3125:
3118:
3116:
3112:
3107:
3103:
3099:
3095:
3091:
3087:
3083:
3079:
3075:
3071:
3067:
3063:
3056:
3049:
3046:
3041:
3037:
3033:
3029:
3025:
3021:
3017:
3013:
3009:
3005:
2998:
2991:
2988:
2983:
2979:
2975:
2971:
2967:
2963:
2959:
2955:
2951:
2947:
2940:
2937:
2932:
2928:
2924:
2920:
2916:
2912:
2905:
2902:
2897:
2893:
2889:
2885:
2880:
2875:
2871:
2867:
2860:
2853:
2850:
2845:
2841:
2837:
2831:
2827:
2820:
2817:
2812:
2808:
2804:
2800:
2796:
2792:
2788:
2784:
2780:
2776:
2772:
2768:
2761:
2758:
2753:
2749:
2745:
2741:
2737:
2733:
2729:
2725:
2721:
2717:
2710:
2707:
2702:
2696:
2688:
2684:
2680:
2678:9780253025074
2674:
2670:
2663:
2660:
2655:
2651:
2646:
2641:
2637:
2633:
2632:Palaeontology
2629:
2622:
2620:
2618:
2616:
2614:
2612:
2608:
2603:
2599:
2595:
2591:
2587:
2583:
2576:
2573:
2568:
2564:
2560:
2556:
2552:
2548:
2544:
2540:
2533:
2531:
2527:
2522:
2518:
2514:
2508:
2504:
2497:
2494:
2489:
2485:
2481:
2477:
2473:
2469:
2465:
2461:
2454:
2451:
2446:
2442:
2438:
2434:
2430:
2426:
2422:
2418:
2414:
2410:
2403:
2396:
2394:
2390:
2385:
2379:
2375:
2368:
2365:
2360:
2353:
2350:
2345:
2339:
2331:
2324:
2321:
2316:
2309:
2306:
2301:
2297:
2293:
2289:
2285:
2281:
2277:
2273:
2266:
2263:
2258:
2252:
2244:
2240:
2235:
2230:
2226:
2222:
2218:
2211:
2208:
2203:
2197:
2189:
2185:
2181:
2177:
2172:
2167:
2163:
2159:
2152:
2145:
2142:
2136:
2131:
2127:
2123:
2119:
2112:
2109:
2096:
2092:
2086:
2083:
2070:
2066:
2059:
2056:
2051:
2047:
2043:
2039:
2035:
2031:
2027:
2023:
2019:
2015:
2008:
2005:
2000:
1996:
1992:
1988:
1983:
1978:
1974:
1970:
1966:
1962:
1958:
1951:
1948:
1943:
1939:
1935:
1931:
1927:
1923:
1919:
1915:
1907:
1905:
1901:
1896:
1892:
1888:
1884:
1879:
1874:
1870:
1866:
1862:
1858:
1851:
1848:
1843:
1839:
1834:
1829:
1825:
1821:
1816:
1811:
1807:
1803:
1799:
1795:
1791:
1784:
1782:
1778:
1773:
1769:
1765:
1761:
1757:
1753:
1749:
1745:
1738:
1731:
1728:
1723:
1719:
1714:
1709:
1705:
1701:
1697:
1690:
1687:
1683:
1677:
1674:
1670:
1666:
1660:
1657:
1652:
1648:
1644:
1640:
1636:
1632:
1628:
1624:
1620:
1616:
1609:
1606:
1601:
1597:
1592:
1587:
1583:
1579:
1575:
1568:
1565:
1559:
1556:
1552:
1546:
1544:
1542:
1538:
1533:
1529:
1525:
1521:
1517:
1513:
1506:
1499:
1496:
1491:
1487:
1480:
1477:
1472:
1468:
1461:
1458:
1452:
1449:
1445:
1441:
1437:
1433:
1429:
1423:
1420:
1409:
1405:
1398:
1395:
1390:
1384:
1373:
1369:
1365:
1358:
1356:
1347:
1345:
1343:
1341:
1337:
1333:
1327:
1325:
1323:
1321:
1319:
1317:
1315:
1311:
1307:
1301:
1299:
1297:
1293:
1288:
1284:
1279:
1274:
1270:
1266:
1262:
1255:
1252:
1246:
1242:
1239:
1238:
1234:
1228:
1221:
1216:
1212:
1205:
1200:
1198:
1196:
1192:
1188:
1185:; inadequate
1184:
1180:
1176:
1168:
1166:
1164:
1160:
1156:
1152:
1147:
1142:
1137:
1135:
1133:
1129:
1128:
1127:Proganochelys
1122:
1119:
1115:
1114:
1108:
1106:
1105:
1100:
1095:
1093:
1092:
1087:
1079:
1077:
1075:
1071:
1067:
1063:
1058:
1056:
1052:
1048:
1044:
1043:
1038:
1030:
1024:
1017:
1015:
1013:
1012:
1007:
1006:
1001:
993:
988:
986:
984:
980:
976:
972:
967:
965:
961:
957:
953:
949:
945:
943:
938:
937:
931:
930:
920:
916:
909:
907:
899:
896:
893:
890:
889:
885:
882:
879:
876:
875:
871:
868:
865:
862:
861:
857:
854:
851:
848:
847:
843:
840:
837:
834:
833:
829:
826:
823:
820:
819:
815:
812:
809:
806:
805:
801:
798:
795:
792:
791:
787:
784:
781:
778:
777:
773:
770:
767:
764:
763:
759:
756:
753:
750:
749:
745:
742:
739:
736:
735:
731:
728:
725:
723:Zangerl 1969
722:
721:
715:
711:
708:
703:
699:
695:
691:
682:
675:
673:
671:
667:
663:
658:
656:
644:
641:
640:
636:
633:
632:
628:
625:
624:
620:
617:
616:
612:
609:
608:
604:
601:
600:
596:
593:
592:
589:
588:
587:
581:
576:
575:
568:
561:
552:
545:
531:
529:
525:
521:
517:
513:
508:
506:
501:
499:
495:
491:
487:
483:
478:
475:
471:
467:
463:
462:Odontochelys,
459:
455:
451:
446:
443:
437:
435:
431:
427:
423:
419:
415:
405:
404:
396:
388:
387:
379:
369:
368:
363:
362:
351:
343:
341:
339:
338:
333:
329:
324:
322:
318:
314:
310:
306:
302:
298:
294:
293:dermal plates
290:
286:
282:
278:
262:
261:
255:
248:
246:
242:
239:
234:
230:
228:
219:
218:
213:
212:
207:
206:
200:
196:
192:
190:
186:
182:
178:
174:
169:
164:
161:
157:
152:
149:
145:
141:
137:
134:
126:
121:
114:
112:
110:
109:tortoiseshell
106:
101:
97:
95:
90:
86:
82:
71:
67:
60:
51:
45:
32:
19:
5638:
5613:
5605:
5597:
5556:
5548:
5526:
5518:
5511:
5504:
5496:
5490:Erymnochelys
5488:
5481:
5475:Cerrejonemys
5473:
5465:
5457:
5451:Brontochelys
5449:
5441:
5433:
5412:
5405:
5385:
5377:
5370:
5365:Pseudemydura
5363:
5356:
5348:
5341:
5334:
5327:
5320:
5312:
5305:
5298:
5291:
5284:
5277:
5270:
5250:
5242:
5234:
5226:
5218:
5210:
5202:
5194:
5186:
5178:
5170:
5162:
5154:
5146:
5138:
5130:
5122:
5114:
5106:
5098:
5090:
5082:
5074:
5066:
5058:
5050:
5042:
5020:
4974:
4968:Prionochelys
4966:
4958:
4950:
4942:
4934:
4915:
4907:
4899:
4891:
4883:
4875:
4867:
4859:
4851:
4845:Cratochelone
4843:
4835:
4829:Bouliachelys
4827:
4819:
4811:
4805:Alienochelys
4803:
4781:
4773:
4765:
4743:
4735:
4727:
4720:
4712:
4706:Chelydropsis
4704:
4696:
4688:
4657:
4651:Basilochelys
4649:
4630:
4623:
4616:
4609:
4602:
4594:
4587:
4580:
4575:Khunnuchelys
4573:
4565:
4557:
4549:
4542:
4534:
4527:
4520:
4513:
4505:
4498:
4491:Trionychidae
4478:
4471:
4465:Allaeochelys
4463:
4429:
4422:
4416:Stigmochelys
4414:
4407:
4399:
4392:
4385:
4379:Megalochelys
4377:
4369:
4362:
4355:
4348:
4341:
4334:
4326:
4318:
4311:
4304:
4296:
4290:Cylindraspis
4288:
4281:
4274:
4269:Centrochelys
4267:
4260:
4253:
4246:Testudinidae
4235:Platysternon
4233:
4214:Vijayachelys
4212:
4205:
4198:
4193:Rhinoclemmys
4191:
4184:
4177:
4170:
4163:
4158:Melanochelys
4156:
4149:
4142:
4135:
4128:
4121:
4114:
4107:
4100:
4093:
4086:
4078:
4058:
4050:
4043:
4036:
4029:
4022:
4015:
4008:
4001:
3994:
3987:
3980:
3973:
3968:Acherontemys
3966:
3934:Sternotherus
3932:
3925:
3918:
3911:
3903:
3883:
3850:
3842:
3834:
3826:
3818:
3801:Psephophorus
3799:
3791:
3783:
3775:
3768:
3760:
3752:
3731:
3723:
3717:Pacifichelys
3715:
3707:
3700:
3692:
3686:Lepidochelys
3684:
3677:
3669:
3663:Gigantatypus
3661:
3655:Eretmochelys
3653:
3646:
3638:
3631:
3623:
3616:
3597:Chelonioidea
3565:
3517:
3441:
3359:
3336:
3327:
3318:
3305:
3293:. Retrieved
3289:
3276:
3255:
3246:
3242:
3236:
3211:
3207:
3201:
3185:
3181:
3175:
3134:
3130:
3065:
3061:
3048:
3007:
3003:
2990:
2949:
2945:
2939:
2914:
2910:
2904:
2869:
2865:
2852:
2825:
2819:
2770:
2766:
2760:
2719:
2715:
2709:
2668:
2662:
2635:
2631:
2585:
2581:
2575:
2542:
2538:
2502:
2496:
2466:(1): 31–80.
2463:
2459:
2453:
2412:
2408:
2373:
2367:
2358:
2352:
2329:
2323:
2314:
2308:
2275:
2271:
2265:
2251:cite journal
2224:
2220:
2210:
2196:cite journal
2161:
2157:
2144:
2125:
2121:
2111:
2099:. Retrieved
2094:
2085:
2073:. Retrieved
2069:the original
2058:
2017:
2013:
2007:
1964:
1960:
1950:
1917:
1913:
1878:10138/223805
1860:
1856:
1850:
1797:
1793:
1750:(2): 47–58.
1747:
1743:
1730:
1703:
1699:
1689:
1676:
1668:
1659:
1618:
1614:
1608:
1581:
1577:
1567:
1558:
1550:
1515:
1511:
1498:
1489:
1485:
1479:
1470:
1466:
1460:
1451:
1443:
1439:
1435:
1422:
1411:. Retrieved
1407:
1397:
1383:cite journal
1372:the original
1367:
1363:
1354:
1331:
1305:
1268:
1264:
1254:
1172:
1148:
1146:
1125:
1123:
1111:
1109:
1102:
1096:
1089:
1083:
1074:Eunotosaurus
1073:
1062:Eunotosaurus
1061:
1059:
1055:Odontochelys
1054:
1050:
1047:Eunotosaurus
1046:
1042:Eunotosaurus
1040:
1034:
1009:
1003:
997:
968:
963:
959:
955:
951:
947:
940:
934:
927:
924:
905:
802:supracaudal
799:supracaudal
796:postcentral
712:
707:Thayer's law
701:
687:
670:Oracle bones
666:plastromancy
659:
652:
585:
515:
511:
509:
502:
479:
473:
470:E. africanus
469:
465:
461:
458:E. africanus
457:
453:
449:
447:
442:endochondral
438:
411:
401:
384:
365:
359:
335:
325:
304:
274:
258:
243:
235:
231:
223:
215:
209:
203:
193:
168:evolutionary
165:
153:
139:
130:
124:
102:
98:
81:turtle shell
80:
78:
5528:Stupendemys
5483:Cordichelys
5329:Mesoclemmys
5314:Hydromedusa
5236:Taphrosphys
5228:Sankuchemys
5220:Rhothonemys
5196:Polysternon
5076:Cearachelys
5060:Azabbaremys
5044:Araiochelys
4960:Ctenochelys
4952:Bashuchelys
4901:Rhinochelys
4877:Ocepechelon
4869:Notochelone
4861:Iserosaurus
4729:Macrochelys
4681:Chelydridae
4604:Palaeoamyda
4529:Cyclanorbis
4394:Psammobates
4387:Oligopherus
4350:Indotestudo
4298:Cymatholcus
4276:Chelonoidis
4262:Astrochelys
4071:Geoemydidae
3989:Deirochelys
3927:Staurotypus
3920:Kinosternon
3913:Hoplochelys
3828:Peritresius
3777:Dermochelys
3770:Cosmochelys
3762:Corsochelys
3679:Itilochelys
3671:Glarichelys
3618:Allopleuron
3608:Cheloniidae
3547:Superfamily
3422:Gular scute
2588:: 124–131.
2545:(1): 1–14.
2158:Development
2075:12 February
1967:(1): 5211.
1161:, and poor
1091:Pappochelys
1051:Pappochelys
1005:Bradysaurus
919:Development
816:intergular
813:intergular
807:intergular
740:precentral
668:. See also
597:= intergul
512:gular scute
477:specimens.
474:Pappochelys
466:Pappochelys
454:Pappochelys
426:osteoblasts
422:exoskeleton
227:Pelomedusid
148:Cryptodires
144:Pleurodires
94:dermal bone
70:cryptodires
66:Pleurodires
5664:Categories
5607:Caririemys
5520:Podocnemis
5506:Latentemys
5467:Carbonemys
5407:Pelomedusa
5244:Ummulisani
5164:Kurmademys
5132:Inaechelys
5116:Galianemys
5084:Chedighaii
5022:Araripemys
4995:Pleurodira
4976:Toxochelys
4893:Protostega
4698:Chelydrops
4618:Pelodiscus
4611:Pelochelys
4559:Gilmoremys
4536:Cycloderma
4446:Trionychia
4313:Geochelone
4306:Floridemys
4172:Notochelys
4060:Wilburemys
4031:Malaclemys
3885:Dermatemys
3852:Puppigerus
3785:Eosphargis
3702:Miocaretta
3694:Mexichelys
3588:Cryptodira
3528:Testudines
3489:Testudines
3437:Pyramiding
3269:072169327X
3249:: 273–277.
2844:1037017014
2521:1037017014
2101:5 November
1413:2021-12-21
1247:References
1195:vitamin D3
1169:Pyramiding
1132:ankylosaur
1000:osteoderms
981:including
872:abdominal
869:abdominal
866:abdominal
863:abdominal
757:vertebral
751:vertebral
726:Carr 1952
662:divination
594:intergular
496:, male or
434:periosteum
432:, and the
332:armadillos
309:soft shell
181:locomotion
133:keratinous
5647:See also
5599:Caribemys
5459:Caninemys
5443:Bauruemys
5372:Rheodytes
5336:Myuchelys
5279:Chelodina
5204:Puentemys
5180:Nigeremys
5124:Ilatardia
5068:Bothremys
4775:Basilemys
4589:Nilssonia
4567:Hutchemys
4515:Axestemys
4473:Anosteira
4328:Hadrianus
4186:Pangshura
4144:Malayemys
4109:Geoclemys
4052:Trachemys
4045:Terrapene
4038:Pseudemys
4024:Graptemys
4017:Glyptemys
4010:Emydoidea
4003:Actinemys
3975:Chrysemys
3820:Euclastes
3648:Eochelone
3499:Kingdom:
3228:133893011
3193:263164288
3090:0028-0836
2982:205243837
2795:0028-0836
2744:0028-0836
2695:cite book
2687:962141060
2654:0031-0239
2480:0077-8923
2429:1552-5007
2338:cite book
2128:: 23–25.
2050:205243837
1991:2041-1723
1934:1552-5015
1887:1552-5015
1824:0027-8424
1671:45:701-+.
1651:206519888
1175:tortoises
1143:Shell rot
1118:Guangling
1101:, China,
1066:fossorial
858:pectoral
855:pectoral
852:pectoral
849:pectoral
788:marginal
785:marginal
782:marginal
779:marginal
737:cervical
626:abdominal
486:abdominal
418:gastralia
311:turtles,
229:turtles.
160:epidermis
5414:Pelusios
5379:Rhinemys
5350:Platemys
5343:Phrynops
5263:Chelidae
5252:Zolhafah
5148:Jainemys
4813:Archelon
4690:Chelydra
4582:Lissemys
4424:Stylemys
4409:Solitudo
4371:Manouria
4320:Gopherus
4283:Chersina
4151:Mauremys
4130:Heosemys
4123:Hardella
4116:Geoemyda
4102:Cyclemys
3958:Emydidae
3905:Claudius
3754:Arabemys
3733:Tasbacka
3725:Syllomus
3640:Chelonia
3538:Suborder
3522:Diapsida
3513:Reptilia
3507:Chordata
3505:Phylum:
3501:Animalia
3432:Plastron
3407:Carapace
3319:Cabi.Org
3159:19037315
3098:19037315
3032:19037315
2974:26106865
2888:27426515
2803:26331544
2752:29609847
2602:28570929
2567:10939665
2559:21210938
2488:84532271
2437:15968684
2300:88095099
2243:28992039
2180:25053434
2095:SeaWorld
2042:26106865
1999:25376734
1942:24898540
1895:25074288
1842:27114549
1772:25901314
1764:11341674
1722:23787042
1643:19590000
1600:26391496
1492:: 82–86.
1432:plastron
1428:carapace
1287:28267966
1235:See also
1159:abrasion
1151:bacteria
1138:Diseases
1086:Triassic
1011:Anthodon
910:Ontogeny
886:femoral
883:femoral
880:femoral
877:femoral
844:humeral
841:humeral
838:humeral
835:humeral
768:lateral
765:pleural
760:central
754:central
698:chelonii
618:pectoral
482:pectoral
344:Plastron
313:pig-nose
277:carapace
249:Carapace
185:friction
140:plastron
5558:Sokatra
5307:Emydura
5212:Rosasia
5108:Foxemys
5052:Arenila
4632:Trionyx
4625:Rafetus
4551:Dogania
4507:Apalone
4431:Testudo
4357:Kinixys
4343:Homopus
4200:Sacalia
4179:Orlitia
4165:Morenia
4080:Batagur
3982:Clemmys
3709:Natator
3625:Caretta
3526:Order:
3511:Class:
3295:17 July
3167:4405644
3139:Bibcode
3106:4405644
3070:Bibcode
3040:4405644
3012:Bibcode
2954:Bibcode
2919:Bibcode
2896:3935231
2811:4401555
2775:Bibcode
2724:Bibcode
2445:2484583
2280:Bibcode
2188:7737357
2022:Bibcode
1969:Bibcode
1833:4868452
1802:Bibcode
1623:Bibcode
1615:Science
1532:1443917
1193:and/or
1187:calcium
1183:protein
1179:pyramid
1099:Guizhou
1037:Permian
774:costal
771:costal
746:nuchal
743:nuchal
694:keratin
664:called
634:femoral
621:= pect
610:humeral
494:concave
490:species
430:osteoid
301:keratin
279:is the
238:keratin
179:during
85:turtles
5590:
5583:
5576:
5300:Elusor
5293:Elseya
5286:Chelus
5003:
4936:Adocus
4927:
4673:
4642:
4522:Chitra
4224:
3811:
3556:Family
3267:
3226:
3191:
3165:
3157:
3131:Nature
3104:
3096:
3088:
3062:Nature
3038:
3030:
3004:Nature
2980:
2972:
2946:Nature
2894:
2886:
2842:
2832:
2809:
2801:
2793:
2767:Nature
2750:
2742:
2716:Nature
2685:
2675:
2652:
2600:
2565:
2557:
2519:
2509:
2486:
2478:
2443:
2435:
2427:
2380:
2298:
2241:
2186:
2178:
2048:
2040:
2014:Nature
1997:
1989:
1940:
1932:
1893:
1885:
1840:
1830:
1822:
1770:
1762:
1720:
1649:
1641:
1598:
1530:
1512:Copeia
1285:
975:dermis
971:embryo
830:gular
824:gular
821:gular
690:scutes
676:Scutes
637:= fem
629:= abd
613:= hum
605:= gul
528:trowel
524:scutes
520:turtle
505:turtle
498:convex
428:, the
414:turtle
297:scutes
285:turtle
281:dorsal
266:Legend
177:turtle
173:turtle
156:turtle
136:scutes
50:Scutes
4596:Palea
4500:Amyda
4401:Pyxis
4095:Cuora
3567:Genus
3518:Clade
3442:Shell
3315:(PDF)
3286:(PDF)
3224:S2CID
3163:S2CID
3127:(PDF)
3102:S2CID
3058:(PDF)
3036:S2CID
3000:(PDF)
2978:S2CID
2892:S2CID
2862:(PDF)
2807:S2CID
2748:S2CID
2563:S2CID
2484:S2CID
2441:S2CID
2405:(PDF)
2296:S2CID
2184:S2CID
2154:(PDF)
2046:S2CID
1768:S2CID
1740:(PDF)
1647:S2CID
1528:JSTOR
1508:(PDF)
1375:(PDF)
1360:(PDF)
1155:fungi
1053:and
983:FGF10
900:anal
897:anal
894:anal
891:anal
645:= an
602:gular
518:on a
289:spine
89:order
87:(the
3996:Emys
3297:2019
3265:ISBN
3189:OCLC
3155:PMID
3094:PMID
3086:ISSN
3028:PMID
2970:PMID
2884:PMID
2840:OCLC
2830:ISBN
2799:PMID
2791:ISSN
2740:ISSN
2701:link
2683:OCLC
2673:ISBN
2650:ISSN
2598:PMID
2555:PMID
2517:OCLC
2507:ISBN
2476:ISSN
2433:PMID
2425:ISSN
2413:304B
2378:ISBN
2344:link
2257:link
2239:PMID
2202:link
2176:PMID
2103:2023
2077:2011
2038:PMID
1995:PMID
1987:ISSN
1938:PMID
1930:ISSN
1891:PMID
1883:ISSN
1838:PMID
1820:ISSN
1760:PMID
1718:PMID
1639:PMID
1596:PMID
1516:1977
1430:and
1389:link
1283:PMID
962:and
960:Pax1
956:Pax1
944:gene
939:and
936:PAX1
702:skin
642:anal
510:The
484:and
460:and
321:skin
305:keel
275:The
189:drag
187:and
79:The
3216:doi
3147:doi
3135:456
3078:doi
3066:456
3020:doi
3008:456
2962:doi
2950:523
2927:doi
2874:doi
2783:doi
2771:525
2732:doi
2720:379
2640:doi
2590:doi
2547:doi
2468:doi
2417:doi
2288:doi
2229:doi
2166:doi
2162:141
2130:doi
2126:150
2030:doi
2018:523
1977:doi
1922:doi
1918:324
1873:hdl
1865:doi
1861:324
1828:PMC
1810:doi
1798:113
1752:doi
1708:doi
1631:doi
1619:325
1586:doi
1520:doi
1273:doi
1191:UVB
1153:or
1116:of
1008:to
964:Shh
952:Shh
948:Shh
514:or
263:.
158:'s
5666::
3520::
3335:.
3317:.
3288:.
3245:.
3222:.
3212:30
3210:.
3187:.
3161:.
3153:.
3145:.
3133:.
3129:.
3114:^
3100:.
3092:.
3084:.
3076:.
3064:.
3060:.
3034:.
3026:.
3018:.
3006:.
3002:.
2976:.
2968:.
2960:.
2948:.
2925:.
2913:.
2890:.
2882:.
2870:26
2868:.
2864:.
2838:.
2805:.
2797:.
2789:.
2781:.
2769:.
2746:.
2738:.
2730:.
2718:.
2697:}}
2693:{{
2681:.
2648:.
2636:63
2634:.
2630:.
2610:^
2596:.
2586:45
2584:.
2561:.
2553:.
2543:13
2541:.
2529:^
2515:.
2482:.
2474:.
2464:31
2462:.
2439:.
2431:.
2423:.
2411:.
2407:.
2392:^
2340:}}
2336:{{
2294:.
2286:.
2276:49
2274:.
2253:}}
2249:{{
2237:.
2225:57
2223:.
2219:.
2198:}}
2194:{{
2182:.
2174:.
2160:.
2156:.
2124:.
2120:.
2093:.
2044:.
2036:.
2028:.
2016:.
1993:.
1985:.
1975:.
1963:.
1959:.
1936:.
1928:.
1916:.
1903:^
1889:.
1881:.
1871:.
1859:.
1836:.
1826:.
1818:.
1808:.
1796:.
1792:.
1780:^
1766:.
1758:.
1746:.
1742:.
1716:.
1704:23
1702:.
1698:.
1667:"
1645:.
1637:.
1629:.
1617:.
1594:.
1582:28
1580:.
1576:.
1553::.
1540:^
1526:.
1514:.
1510:.
1488:.
1471:31
1469:.
1406:.
1385:}}
1381:{{
1368:42
1366:.
1362:.
1339:^
1313:^
1295:^
1281:.
1269:27
1267:.
1263:.
1225:A
1189:,
1134:.
985:.
810:-
672:.
530:.
472:.
436:.
191:.
111:.
5612:†
5604:†
5596:†
5555:†
5547:†
5538:†
5525:†
5503:†
5495:†
5480:†
5472:†
5464:†
5456:†
5448:†
5440:†
5432:†
5384:†
5355:†
5319:†
5249:†
5241:†
5233:†
5225:†
5217:†
5209:†
5201:†
5193:†
5185:†
5177:†
5169:†
5161:†
5153:†
5145:†
5137:†
5129:†
5121:†
5113:†
5105:†
5097:†
5089:†
5081:†
5073:†
5065:†
5057:†
5049:†
5041:†
5032:†
5019:†
5010:†
4973:†
4965:†
4957:†
4949:†
4941:†
4933:†
4914:†
4906:†
4898:†
4890:†
4882:†
4874:†
4866:†
4858:†
4850:†
4842:†
4834:†
4826:†
4818:†
4810:†
4802:†
4793:†
4780:†
4772:†
4764:†
4755:†
4742:†
4734:†
4719:†
4711:†
4703:†
4695:†
4656:†
4648:†
4601:†
4572:†
4564:†
4556:†
4541:†
4512:†
4470:†
4462:†
4421:†
4406:†
4384:†
4376:†
4333:†
4325:†
4303:†
4295:†
4085:†
4057:†
3965:†
3910:†
3849:†
3841:†
3833:†
3825:†
3817:†
3798:†
3790:†
3782:†
3767:†
3759:†
3751:†
3730:†
3722:†
3714:†
3699:†
3691:†
3676:†
3668:†
3660:†
3645:†
3630:†
3615:†
3481:e
3474:t
3467:v
3390:e
3383:t
3376:v
3321:.
3299:.
3271:.
3247:7
3230:.
3218::
3195:.
3169:.
3149::
3141::
3108:.
3080::
3072::
3042:.
3022::
3014::
2984:.
2964::
2956::
2933:.
2929::
2921::
2915:5
2898:.
2876::
2846:.
2813:.
2785::
2777::
2754:.
2734::
2726::
2703:)
2689:.
2656:.
2642::
2604:.
2592::
2569:.
2549::
2523:.
2490:.
2470::
2447:.
2419::
2386:.
2346:)
2302:.
2290::
2282::
2259:)
2245:.
2231::
2204:)
2190:.
2168::
2138:.
2132::
2105:.
2079:.
2052:.
2032::
2024::
2001:.
1979::
1971::
1965:5
1944:.
1924::
1897:.
1875::
1867::
1844:.
1812::
1804::
1774:.
1754::
1748:3
1724:.
1710::
1653:.
1633::
1625::
1602:.
1588::
1534:.
1522::
1490:2
1416:.
1391:)
1308:.
1289:.
1275::
946:(
553:.
370:)
352:.
72:.
20:)
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
