2040:, by which two species competing for the same ecological niche cannot coexist with one another for a long period of time without either differentiating or one dominating over the other in the long run. In addition to the different skull morphology, the two taxa may have also been able to coexist due to the sheer size of the Pebas Mega-Wetlands they inhabited, as this ecosystem stretched over most of northern South America during the Middle Miocene. This reason may also prevent the two taxa from being in direct competition over nesting grounds and basking spaces.
578:
2371:
2168:
143:
121:
2100:
511:
2087:, some of the largest freshwater turtles alive, which are known to fight for dominance in overlapping territories. This hypothesis is supported by the presence of a deep, elongated scar along the left horn of CIAAP-2002-01, which could have been left by the horn of a rival male that engaged it in combat. The authors further suggest that in
634:. However, these estimates do not compensate for the large embayment present at the front of the shell. A more precise body mass estimate might be achieved by calculating the average between the results of weight estimates based on midline length and parasagittal length. Applying this method yields a weight of 1,145 kg for the largest
553:
can reach a straight midline length of greater than 2 meters with a low-arched profile. The nodular contours on the surface are irregular and the frontal margin of the shell is characterized by a deep notch flanked by large horns in male specimens. These horns are deeply grooved, suggesting that they
541:
bear a deep concavity at their center. In this view the premaxillae form most of the anteromedial edge of the skull, meeting each other towards the middle of the skull and narrowing just before the deep concavity. In front view, the premaxillae form the bottom margin of the bony nostrils, tapering as
2120:
shallow anal notch. The inner nuchal notch, anterior expansion of the peripherals 1 and 2, irregular nodular contours, inner contact between the 7th and 8th costals and the relative size of the plastral lobes and their arrangement (except for the pectorals) remains relatively consistent with size.
2119:
had been described. The discovery of a specimen with a carapace length smaller than 1 meter gives an insight into the changes the animal undergoes while reaching maturity. In addition to its small size, the animal is identified as a juvenile to young adult based on the absence of large horns and
330:, about 13 to 5 million years ago. Male specimens are known to have possessed bony horns growing from the front edges of the shell and the discovery of the fossil of a young adult shows that the carapace of these turtles flattens with age. A fossil skull described in 2021 indicates that
2233:, into the Late Miocene before eventually dying out during the Early Pliocene like much of the large crocodilian fauna of the Miocene wetlands. Besides the aforementioned reptiles the waterways of Late Miocene South America were also inhabited by fish, including
2091:
the males may have been the larger sex, similar to the condition seen in the modern podocnemids. However other traditionally sexually dimorphic traits of the turtle shell, such as a deeper anal notch or a xiphiplastral concavity, have not yet been observed in
2183:. The Wetlands provided favorable conditions to the native reptilian fauna, with several lineages of crocodilians reaching enormous sizes during the Mid to Late Miocene and also diversifying in ecology. Some of the enormous crocodilians that coexisted with
2127:
is its height. With age the shell of the turtle grows significantly flatter, while the nuchal region develops a pronounced upturn of its anterior margin and peripheral 1, creating a wide and deep anteromedial embayment of the carapace. The 2nd and 3rd
2154:
change of the vertebral scutes means that phylogenetic coding using the width of the vertebral scutes in relation to the pleural scutes should be treated with care due to the variable nature of these features as shown by
693:
established previously by Joyce and colleagues (2021), 268 characters across 104 species of turtles. The analysis was run once with all taxa and once with a focus on
Podocnemidoidae, removing all other taxa safe for
2209:
large body-size, putting pressure on the animal through predation. Bite marks have been found on
Colombian and Venezuelan specimens and an isolated tooth was found attached to the ventral surface of CIAAP-2002-01.
448:
and formally described by Cadena and colleagues in 2021, including the first definitive skull remains as well as the first remains of a juvenile or early adult specimen (carapace length under 1 meter). The
2032:
proposed to have been deploying a vacuum feeding strategy combined with a strong bite supported by tooth-like structures of the maxilla, while a more durophagous-omnivorous diet has been suggested for
641:
The evolution of such an enormous size may have been multi-facetted and caused by a combination of factors including pressure from predators, habitat size and favorable climatic conditions, although
346:
630:
was estimated based on the straight carapace length, with calculations indicating a weight of 871 kg for CIAAP-2002-01 and 744 kg for MCZ(P)-4376, the former largest known specimen of
2055:
in maintaining its large body size. Furthermore, Cadena and colleagues also highlight the role of turtles as seed-dispersers in modern-day
Amazonia, consuming fruit of palms for example (
2698:
Hoorn, C.; Wesselingh, F. P.; ter Steege, H.; Bermudez, M. A.; Mora, A.; Sevink, J.; Sanmartin, I.; Sanchez-Meseguer, A.; Anderson, C. L.; Figueiredo, J. P.; Jaramillo, C. (2010-11-11).
2083:
trait exclusively found in males, suggesting them to have been used similar to the horns and antlers found in artiodactyls. Among extant turtles similar behavior can be found in
600:
is the largest known species of freshwater turtle currently known to science, with several specimens reaching a carapace length exceeding 2 meters. The largest specimen of
2179:
was part of an interconnected series of lakes, rivers, swamps and marshes that drained into the
Caribbean known as the Pebas Mega-Wetlands, which included the Colombian
2514:
Cadena, E.-A.; Scheyer, T.M.; Carrillo-Briceño, J.D.; Sánchez, R.; Aguilera-Socorro, O.A.; Vanegas, A.; Pardo, M.; Hansen, D.M.; Sánchez-Villagra, M.R. (12 Feb 2020).
2047:
may have been very diverse and broad, possibly including molluscs and other hard shelled prey as well as vertebrate prey as suggested by Meylan and colleagues for
525:
is roughly triangular in top view and the edges of the jaws converge at the front of the snout in a straight edge. The skull is dorsally extremely inflated by the
2771:
2075:
specimens indicates that they were not used as a defense mechanism. However, their forward-facing position on the carapace may indicate that they were used in
417:, which have been ongoing since 1994. The material includes a relatively complete carapace that set a new maximum size for the genus and was designated as the
1065:
480:
is a combination of "stupendous", meaning extremely impressive, and the Latin word "emys" for freshwater turtle. The species name meanwhile honors the
799:
2051:. At its size it would have been easily capable of consuming various fish, snakes and small crocodilians. A broad dietary width would have helped
2881:
2649:
Ferreira, G. S.; Nascimento, E. R.; Cadena, E. A.; Cozzuol, M. A.; Farina, B. M.; Pacheco, M. L. A. F.; Rizzutto, M. A.; Langer, M. C. (2024).
775:
5137:
3854:
3765:
1774:
726:, now recognized as a distinct taxon, nested at the base of Erymnochelyinae. Similar results were later recovered in the 2024 description of
708:
499:
3773:
3527:
1757:
1432:
1351:
847:
751:
5177:
2651:"The latest freshwater giants: a new Peltocephalus (Pleurodira: Podocnemididae) turtle from the Late Pleistocene of the Brazilian Amazon"
1456:
1384:
1279:
1255:
2020:
Following the 2021 research of Cadena and colleagues, the Pebas Mega-Wetlands housed at least two species of giant side-necked turtles:
5172:
1334:
1791:
5162:
2868:
2150:. The 5th vertebral scute meanwhile belongs the longest and widest of the series in adults while keeping its trapezoid shape. This
2205:, some species of which reaching lengths of over 10 meters. Some of these crocodilians may have played a role in the evolution of
2028:. Despite their similar size (both sporting a carapace length greater than 2 meters), they vary greatly in skull morphology, with
413:. In February 2020, Cadena and colleagues published a paper describing material discovered during the routine excavations in the
5242:
2059:), seasonally sometimes in great quantities, even if they are not typically part of their standard diet. With its wide gape,
5107:
Turtles of the World 2017 Update: Annotated
Checklist and Atlas of Taxonomy, Synonymy, Distribution, and Conservation Status
2947:
5147:
5105:
2775:
2037:
5222:
4980:
4063:
2392:
2279:
609:
5237:
5167:
1303:
374:
533:. Following the prefrontals and orbits the skull slopes down drastically before ascending again through the
5217:
4957:
562:
is notably thickened and upturned. The surface is smooth to striated or lightly pitted. The margins of the
5232:
5116:
3639:
3023:
2356:
690:
681:. In their 2021 publication, Cadena and colleagues again attempted to determine the relationship between
481:
142:
5227:
5212:
5202:
3372:
1565:
718:
5207:
2516:"The anatomy, paleobiology, and evolutionary relationships of the largest extinct side-necked turtle"
2076:
1048:
1017:
646:
3005:
5197:
5157:
5152:
4018:
2940:
696:
2461:
sp. nov. (Pleurodira, Podocnemididae) from the
Miocene-Pliocene of the Solimões Formation, Brazil"
645:
temporal range indicates that it managed to survive through times of global cooling following the
5192:
5142:
4874:
2266:
2456:
2328:
would have inhabited a floodplain or lacustrine environment with savannahs and gallery forests.
5187:
4846:
3646:
3604:
3244:
2912:
2798:
2213:
As the Pebas System began to disappear with the onset of the transcontinental Amazon
Drainage,
2886:
529:
that make up a large area of the front region of the skull, forming a vertical wall above the
3590:
3092:
2907:
2899:
2836:
2700:"Amazonia Through Time: Andean Uplift, Climate Change, Landscape Evolution, and Biodiversity"
2180:
728:
450:
2222:
396:
5182:
4832:
3498:
3122:
2855:
2711:
2603:
2527:
381:
and various other bones. The fossils were unearthed by a paleontological excavation of the
8:
4817:
3176:
2933:
2376:
1139:
577:
563:
2715:
2675:
2650:
2607:
2531:
537:. The orbits are relatively small and oriented to the sides. When viewed from below the
4224:
4189:
4056:
3795:
2753:
2626:
2591:
2550:
2515:
1122:
1098:
653:, with several other related forms being known to have possessed gigantic proportions.
382:
262:
137:
2590:
Cadena, E.A.; Link, A.; Cooke, S.B.; Stroik, L.K.; Vanegas, A.F.; Tallman, M. (2021).
4767:
3750:
3702:
3693:
3576:
3153:
2894:
2757:
2745:
2737:
2680:
2631:
2555:
2297:
2218:
2080:
414:
366:
2592:"New insights on the anatomy and ontogeny of the largest extinct freshwater turtles"
4559:
3947:
3922:
3883:
3831:
3562:
3107:
2727:
2719:
2670:
2662:
2621:
2611:
2545:
2535:
2475:
2343:
993:
350:
2616:
2132:
grow narrower as the animal matures from juvenile to adult, similar to the extant
677:. However, this position was influenced by the inclusion of material belonging to
5007:
4655:
4384:
4352:
3989:
3898:
3674:
3260:
2239:
712:. The single most parsimonious tree resulting from the second analysis recovered
674:
526:
4902:
4892:
4789:
4639:
4479:
4181:
4126:
3868:
3803:
3343:
3334:
3303:
3211:
3100:
3014:
2422:
2244:
2167:
605:
445:
307:
214:
377:(P)-4376. This specimen preserves much of the carapace alongside a fragmented
5131:
5082:
4965:
4865:
4854:
4825:
4739:
4623:
4607:
4567:
4501:
4411:
4376:
4320:
4304:
4288:
4262:
4250:
4234:
4212:
4204:
4011:
3722:
3555:
3416:
3363:
3311:
2821:
2741:
2699:
2338:
2320:
2312:
2146:
902:
662:
534:
425:. Venezuela also yielded fossils of a lower jaw, which has been used to lump
319:
71:
2723:
649:. Lastly, the giant size could have a phylogenetic link and be ancestral to
5017:
4942:
4918:
4435:
4312:
4296:
4272:
4173:
4118:
4042:
3958:
3932:
3846:
3757:
3736:
3660:
3625:
3435:
3401:
3268:
3184:
3130:
2749:
2684:
2666:
2635:
2559:
2540:
2140:
2129:
2036:. This difference in diet and feeding strategy would be in accordance with
1716:
969:
567:
559:
378:
2480:
2115:
Prior to the 2021 study of Cadena and colleagues, only adult specimens of
492:
constitutes a feminine generic name. The name was eventually corrected to
120:
4950:
4796:
4781:
4703:
4695:
4687:
4663:
4543:
4527:
4511:
4427:
4419:
4368:
4344:
4336:
4328:
4196:
4148:
4071:
3996:
3861:
3817:
3743:
3729:
3681:
3538:
3456:
3394:
3387:
3380:
3295:
3237:
3229:
3146:
3138:
3085:
3075:
2847:
2830:
2307:
2253:
2201:
2195:
2189:
2084:
1740:
1637:
46:
2732:
5074:
4987:
4973:
4934:
4711:
4631:
4599:
4583:
4551:
4489:
4462:
4443:
4360:
4165:
4085:
4078:
4026:
4003:
3913:
3780:
3512:
3319:
3252:
3169:
3161:
3064:
3055:
2873:
2284:
2226:
2134:
2099:
1661:
1613:
1408:
878:
686:
613:
538:
453:
also yielded the remains of an adult female as well as more fossils of
400:
300:
204:
91:
56:
570:
of the carapace are relatively thin. In overall shape the carapace of
5066:
4926:
4910:
4839:
4803:
4753:
4746:
4671:
4647:
4591:
4535:
4242:
4034:
3982:
3940:
3653:
3611:
3519:
3505:
3491:
3484:
3477:
3470:
3442:
3287:
3115:
2302:
2056:
1833:
1685:
1224:
823:
409:
386:
154:
96:
40:
34:
2792:
2513:
510:
310:. It is the largest freshwater turtle known to have existed, with a
4881:
4810:
4730:
4719:
4615:
4280:
4157:
4049:
3891:
3876:
3838:
3787:
3713:
3618:
3597:
3583:
3569:
3425:
3352:
3221:
3200:
3192:
2974:
2815:
2349:
2292:
2258:
2248:
2151:
2079:. Cadena and colleagues hypothesize that the horns may have been a
919:
702:
618:
546:
442:
422:
418:
354:
327:
311:
174:
86:
81:
66:
61:
51:
30:
473:
was contested and the former was re-established as a valid genus.
5025:
4774:
4679:
4575:
4519:
4099:
4092:
3974:
3824:
3810:
3667:
3632:
3547:
3449:
2989:
2980:
2697:
2274:
2234:
555:
362:
323:
293:
184:
101:
76:
2860:
395:
was described by
Bocquentin and Melo based on material from the
4760:
4403:
2995:
2968:
2956:
2648:
2316:
2230:
404:
315:
303:
194:
164:
3967:
3034:
2925:
2288:
586:
530:
369:. Wood also described several other specimens he referred to
358:
296:
2063:
would qualify as a megafaunal frugivore and seed disperser.
558:
sheath. In addition to these horns, the front margin of the
3463:
345:
was first named in 1976 by Roger C. Wood based on specimen
623:
the largest known turtle, (carapace length 2.20 meters).
461:
known in association with a carapace and new fossils of
2123:
One of the most significant changes of the carapace of
2589:
604:
is CIAAP-2002-01, an almost complete carapace with a
421:, meaning the specimen is of the opposite sex of the
2366:
608:
length of 2.86 meters. This exceeds the size of the
500:
International Code of
Zoological Nomenclature (ICZN)
488:, as coined by Wood, is grammatically incorrect, as
271:
247:
2772:"Giant fossil matamata turtles (matamatas part V)"
2175:During the Middle Miocene, the area inhabited by
433:in the 2020 study. The authors likewise consider
5129:
566:peripheral bones are moderately scalloped. The
441:. However, more fossils were discovered in the
2454:
665:by Wood, later studies consistently recovered
5104:Phylogenetic arrangement of turtles based on
3342:
2941:
716:as an early branching member of a clade with
4147:
3074:
3013:
2948:
2934:
2217:persisted in the wetlands of the northern
669:as a podocnemidid turtle instead. In 2020
119:
2731:
2674:
2625:
2615:
2549:
2539:
2479:
2324:. At least within the Solimões Formation
647:middle Miocene climatic transition (MMCT)
2450:
2448:
2446:
2444:
2166:
2098:
576:
509:
2455:Bocquentin, Jean; Melo, Janira (2006).
2311:. Further aquatic vertebrates included
2199:and large-bodied gharials of the genus
5130:
2353:, a giant sea turtle of the Cretaceous
3022:
2929:
2797:
2796:
2585:
2583:
2581:
2579:
2577:
2575:
2573:
2571:
2569:
2509:
2507:
2505:
2503:
2501:
2499:
2497:
2495:
2493:
2491:
2441:
2416:
2414:
2412:
2410:
337:
5138:Prehistoric turtles of South America
2420:
2066:
661:Although initially believed to be a
2468:Revista Brasileira de Paleontologia
2162:
2103:Hypothetical ontogenetic series of
673:was recovered as a basal member of
457:. With definitive skull remains of
389:in 1972. In 2006 a second species,
13:
5178:Pliocene reptiles of South America
2642:
2566:
2488:
2407:
322:, in rocks dating from the Middle
14:
5254:
5173:Miocene reptiles of South America
2385:
2369:
141:
44:
5163:Miocene genus first appearances
5050:
4470:
2295:found in the same deposits are
2015:
2010:
498:in 2021 in accordance with the
2955:
2764:
2691:
2280:Paleohoplias assisbrasiliensis
2187:included the enormous caimain
505:
1:
5243:Fossil taxa described in 1976
2617:10.1016/j.heliyon.2021.e08591
2427:, the world's largest turtle"
2362:
2071:The absence of horns on most
1304:Erymnochelys madagascariensis
2397:Wood 1976 (sideneck turtle)"
2171:Middle Miocene South America
656:
318:have been found in northern
7:
5117:List of Testudines families
2357:Largest prehistoric animals
2331:
2110:
574:is longer than it is wide.
482:National Geographic Society
278:Bocquentin & Melo, 2006
10:
5259:
2301:(a fossil relative of the
1566:Peltocephalus dumerilianus
903:Lapparentemys vilavilensis
800:Portezueloemys patagonicus
719:Peltocephalus dumerilianus
306:, belonging to the family
5148:Prehistoric turtle genera
5100:
5057:
5043:
5005:
4891:
4864:
4729:
4499:
4477:
4461:
4394:
4260:
4222:
4140:
4109:
3957:
3921:
3912:
3712:
3691:
3537:
3424:
3415:
3362:
3333:
3278:
3210:
3063:
3054:
3032:
3031:
3004:
2963:
2805:
1830:
1823:
1788:
1771:
1754:
1737:
1730:
1713:
1706:
1699:
1682:
1675:
1658:
1651:
1634:
1627:
1610:
1603:
1586:
1579:
1562:
1555:
1470:
1453:
1446:
1429:
1422:
1405:
1398:
1381:
1374:
1348:
1331:
1324:
1317:
1300:
1293:
1276:
1269:
1252:
1245:
1238:
1221:
1214:
1207:
1136:
1119:
1112:
1095:
1088:
1062:
1049:Podocnemis sextuberculata
1045:
1038:
1031:
1018:Podocnemis erythrocephala
1014:
1007:
990:
983:
966:
959:
952:
916:
899:
892:
875:
868:
861:
844:
837:
820:
813:
796:
789:
772:
765:
748:
741:
407:, also home to the giant
353:portion of a large sized
334:was a generalist feeder.
326:to the very start of the
268:
261:
244:
239:
138:Scientific classification
136:
127:
118:
23:
2425:Stupendemys geographicus
709:Platychelys oberndorferi
697:Proganochelys quenstedti
691:morphological characters
486:Stupendemys geographicus
314:over 2 meters long. Its
16:Extinct genus of turtles
2724:10.1126/science.1194585
2267:South American lungfish
2105:Stupendemys geographica
1614:Mogharemys blackenhorni
1590:Stupendemys geographica
970:Cerrejonemys wayuunaiki
776:Hamadachelys escuilliei
687:pan-pleurodiran turtles
592:
560:nuchal-peripheral bones
495:Stupendemys geographica
2667:10.1098/rsbl.2024.0010
2541:10.1126/sciadv.aay4593
2342:, a giant prehistoric
2172:
2107:
739:Podocnemidoidae
703:Notoemys laticentralis
589:
518:
437:to be synonymous with
5223:Honda Group, Colombia
2908:Paleobiology Database
2481:10.4072/rbp.2006.2.02
2181:La Victoria Formation
2170:
2102:
1717:Brontochelys gaffneyi
1457:Dacquemys paleomorpha
1433:Turkanemys pattersoni
950:Podocnemididae
879:Peiropemys mezzalirai
848:Cambaremys langertoni
729:Peltocephalus maturin
580:
513:
451:La Victoria Formation
357:with associated left
5238:Fossils of Venezuela
5168:Pliocene extinctions
2421:Wood, R. C. (1976).
2291:. Other turtles and
2271:Lepidosiren paradoxa
2263:Colossoma macropomum
2077:intraspecific combat
1741:Lemurchelys diasphax
1686:Bairdemys healeyorum
1385:Neochelys fajumensis
1225:Caninemys tridentata
484:. However, the name
5218:Fossils of Colombia
2716:2010Sci...330..927H
2608:2021Heliy...708591C
2532:2020SciA....6.4593C
2377:Paleontology portal
1792:Piramys auffenbergi
1775:Stereogenys cromeri
1758:Shweboemys pilgrimi
1662:Latentemys plowdeni
1638:Cordichelys antiqua
1409:Carbonemys cofrinii
1352:Papoulemys laurenti
1140:Podocnemis unifilis
1066:Podocnemis bassleri
5233:Pliocene Venezuela
4225:Nanhsiungchelyidae
4190:Macrocephalochelys
2459:Stupendemys souzai
2321:"Anhinga" fraileyi
2254:Acregoliath rancii
2223:Solimões Formation
2173:
2108:
2081:sexually dimorphic
1335:Kenyemys williamsi
1280:Neochelys franzeni
1256:Neochelys arenarum
1123:Podocnemis lewyana
1099:Podocnemis expansa
590:
554:were covered by a
519:
465:, the referral of
397:Solimões Formation
392:Stupendemys souzai
383:Harvard University
338:History and naming
5228:Miocene Venezuela
5213:Pliocene Colombia
5203:Fossils of Brazil
5125:
5124:
5096:
5095:
5092:
5091:
5039:
5038:
5035:
5034:
4457:
4456:
4453:
4452:
4136:
4135:
3908:
3907:
3411:
3410:
3329:
3328:
3050:
3049:
3046:
3045:
3042:
3041:
2923:
2922:
2895:Open Tree of Life
2799:Taxon identifiers
2710:(6006): 927–931.
2298:Chelus columbiana
2283:) and freshwater
2219:Urumaco Formation
2067:Sexual dimorphism
2007:
2006:
1998:
1997:
1989:
1988:
1980:
1979:
1971:
1970:
1962:
1961:
1953:
1952:
1944:
1943:
1935:
1934:
1926:
1925:
1917:
1916:
1908:
1907:
1899:
1898:
1890:
1889:
1881:
1880:
1872:
1871:
1863:
1862:
1854:
1853:
1845:
1844:
1812:
1811:
1803:
1802:
1544:
1543:
1535:
1534:
1526:
1525:
1517:
1516:
1508:
1507:
1499:
1498:
1490:
1489:
1481:
1480:
1363:
1362:
1196:
1195:
1187:
1186:
1178:
1177:
1169:
1168:
1160:
1159:
1151:
1150:
1077:
1076:
940:
939:
931:
930:
612:-specimen of the
415:Urumaco Formation
373:, which includes
367:cervical vertebra
285:
284:
279:
255:
235:
5250:
5208:Miocene Colombia
5055:
5054:
5048:
5047:
4560:Chupacabrachelys
4475:
4474:
4468:
4467:
4145:
4144:
3923:Carettochelyidae
3919:
3918:
3422:
3421:
3340:
3339:
3072:
3071:
3061:
3060:
3029:
3028:
3020:
3019:
3011:
3010:
2950:
2943:
2936:
2927:
2926:
2916:
2915:
2903:
2902:
2890:
2889:
2877:
2876:
2864:
2863:
2851:
2850:
2841:
2840:
2839:
2826:
2825:
2824:
2794:
2793:
2787:
2786:
2784:
2783:
2774:. Archived from
2768:
2762:
2761:
2735:
2695:
2689:
2688:
2678:
2646:
2640:
2639:
2629:
2619:
2587:
2564:
2563:
2553:
2543:
2520:Science Advances
2511:
2486:
2485:
2483:
2465:
2452:
2439:
2438:
2418:
2405:
2404:
2389:
2379:
2374:
2373:
2372:
2344:softshell turtle
2163:Paleoenvironment
2130:vertebral scutes
2085:snapping turtles
1826:
1825:
1733:
1732:
1709:
1708:
1702:
1701:
1678:
1677:
1654:
1653:
1630:
1629:
1606:
1605:
1582:
1581:
1558:
1557:
1449:
1448:
1425:
1424:
1401:
1400:
1377:
1376:
1327:
1326:
1320:
1319:
1296:
1295:
1272:
1271:
1248:
1247:
1241:
1240:
1217:
1216:
1210:
1209:
1115:
1114:
1091:
1090:
1041:
1040:
1034:
1033:
1010:
1009:
994:Podocnemis vogli
986:
985:
962:
961:
955:
954:
895:
894:
871:
870:
864:
863:
840:
839:
824:Bauremys elegans
816:
815:
792:
791:
768:
767:
752:Brasilemys josai
744:
743:
736:
735:
542:they move down.
277:
273:
253:
249:
233:
226:
146:
145:
123:
113:
43:
29:Temporal range:
21:
20:
5258:
5257:
5253:
5252:
5251:
5249:
5248:
5247:
5198:Pliocene Brazil
5158:Miocene turtles
5153:Extinct turtles
5128:
5127:
5126:
5121:
5088:
5031:
5008:Sahonachelyidae
5001:
4887:
4860:
4725:
4656:Phosphatochelys
4495:
4449:
4390:
4385:Terlinguachelys
4353:Pneumatoarthrus
4256:
4218:
4132:
4105:
3953:
3904:
3708:
3687:
3675:Siebenrockiella
3533:
3407:
3358:
3325:
3274:
3261:Mesodermochelys
3206:
3067:
3038:
3000:
2959:
2954:
2924:
2919:
2911:
2906:
2898:
2893:
2885:
2880:
2872:
2867:
2859:
2854:
2846:
2844:
2835:
2834:
2829:
2820:
2819:
2814:
2801:
2791:
2790:
2781:
2779:
2770:
2769:
2765:
2696:
2692:
2655:Biology Letters
2647:
2643:
2588:
2567:
2526:(7): eaay4593.
2512:
2489:
2463:
2453:
2442:
2419:
2408:
2391:
2390:
2386:
2375:
2370:
2368:
2365:
2334:
2240:Phractocephalus
2165:
2113:
2069:
2018:
2013:
2008:
1999:
1990:
1981:
1972:
1963:
1954:
1945:
1936:
1927:
1918:
1909:
1900:
1891:
1882:
1873:
1864:
1855:
1846:
1813:
1804:
1545:
1536:
1527:
1518:
1509:
1500:
1491:
1482:
1364:
1197:
1188:
1179:
1170:
1161:
1152:
1078:
941:
932:
920:Pricemys caiera
675:Erymnochelyinae
659:
595:
508:
363:scapulacoracoid
340:
232:
224:
140:
114:
112:
111:
110:
109:
104:
99:
94:
89:
84:
79:
74:
69:
64:
59:
54:
49:
39:9.0–7.246
38:
37:
27:
17:
12:
11:
5:
5256:
5246:
5245:
5240:
5235:
5230:
5225:
5220:
5215:
5210:
5205:
5200:
5195:
5193:Miocene Brazil
5190:
5185:
5180:
5175:
5170:
5165:
5160:
5155:
5150:
5145:
5143:Podocnemididae
5140:
5123:
5122:
5120:
5119:
5112:
5111:
5110:. † = extinct.
5101:
5098:
5097:
5094:
5093:
5090:
5089:
5087:
5086:
5078:
5070:
5061:
5059:
5052:
5045:
5041:
5040:
5037:
5036:
5033:
5032:
5030:
5029:
5021:
5012:
5010:
5003:
5002:
5000:
4999:
4991:
4984:
4977:
4969:
4961:
4954:
4946:
4938:
4930:
4922:
4914:
4906:
4903:Albertwoodemys
4897:
4895:
4893:Podocnemididae
4889:
4888:
4886:
4885:
4878:
4870:
4868:
4862:
4861:
4859:
4858:
4850:
4843:
4836:
4829:
4821:
4814:
4807:
4800:
4793:
4790:Lomalatachelys
4785:
4778:
4771:
4764:
4757:
4750:
4743:
4735:
4733:
4727:
4726:
4724:
4723:
4715:
4707:
4699:
4691:
4683:
4675:
4667:
4659:
4651:
4643:
4640:Labrostochelys
4635:
4627:
4619:
4611:
4603:
4595:
4587:
4579:
4571:
4563:
4555:
4547:
4539:
4531:
4523:
4515:
4506:
4504:
4497:
4496:
4494:
4493:
4484:
4482:
4480:Araripemydidae
4472:
4465:
4459:
4458:
4455:
4454:
4451:
4450:
4448:
4447:
4439:
4431:
4423:
4415:
4407:
4398:
4396:
4392:
4391:
4389:
4388:
4380:
4372:
4364:
4356:
4348:
4340:
4332:
4324:
4316:
4308:
4300:
4292:
4284:
4276:
4267:
4265:
4258:
4257:
4255:
4254:
4246:
4238:
4229:
4227:
4220:
4219:
4217:
4216:
4208:
4200:
4193:
4185:
4182:Emarginachelys
4177:
4169:
4161:
4153:
4151:
4142:
4138:
4137:
4134:
4133:
4131:
4130:
4127:Sinaspideretes
4122:
4113:
4111:
4107:
4106:
4104:
4103:
4096:
4089:
4082:
4075:
4067:
4060:
4053:
4046:
4038:
4030:
4022:
4015:
4007:
4000:
3993:
3986:
3978:
3971:
3963:
3961:
3955:
3954:
3952:
3951:
3944:
3936:
3927:
3925:
3916:
3910:
3909:
3906:
3905:
3903:
3902:
3895:
3887:
3880:
3872:
3865:
3858:
3850:
3842:
3835:
3828:
3821:
3814:
3807:
3804:Hesperotestudo
3799:
3791:
3784:
3777:
3769:
3761:
3754:
3747:
3740:
3733:
3726:
3718:
3716:
3710:
3709:
3707:
3706:
3698:
3696:
3694:Platysternidae
3689:
3688:
3686:
3685:
3678:
3671:
3664:
3657:
3650:
3643:
3636:
3629:
3622:
3615:
3608:
3601:
3594:
3587:
3580:
3573:
3566:
3559:
3551:
3543:
3541:
3535:
3534:
3532:
3531:
3523:
3516:
3509:
3502:
3495:
3488:
3481:
3474:
3467:
3460:
3453:
3446:
3439:
3430:
3428:
3419:
3413:
3412:
3409:
3408:
3406:
3405:
3398:
3391:
3384:
3376:
3368:
3366:
3360:
3359:
3357:
3356:
3348:
3346:
3344:Dermatemydidae
3337:
3335:Kinosternoidea
3331:
3330:
3327:
3326:
3324:
3323:
3315:
3307:
3304:Procolpochelys
3299:
3291:
3282:
3280:
3276:
3275:
3273:
3272:
3264:
3256:
3248:
3241:
3233:
3225:
3216:
3214:
3212:Dermochelyidae
3208:
3207:
3205:
3204:
3196:
3188:
3180:
3173:
3165:
3157:
3150:
3142:
3134:
3126:
3119:
3111:
3104:
3101:Carolinochelys
3096:
3089:
3080:
3078:
3069:
3058:
3052:
3051:
3048:
3047:
3044:
3043:
3040:
3039:
3026:
3017:
3008:
3002:
3001:
2999:
2998:
2992:
2983:
2977:
2971:
2964:
2961:
2960:
2953:
2952:
2945:
2938:
2930:
2921:
2920:
2918:
2917:
2904:
2891:
2878:
2865:
2852:
2842:
2827:
2811:
2809:
2803:
2802:
2789:
2788:
2763:
2690:
2641:
2602:(12): e08591.
2565:
2487:
2474:(2): 187–192.
2440:
2406:
2383:
2382:
2381:
2380:
2364:
2361:
2360:
2359:
2354:
2346:
2333:
2330:
2315:and the large
2313:river dolphins
2245:Callichthyidae
2193:, the bizarre
2164:
2161:
2112:
2109:
2068:
2065:
2017:
2014:
2012:
2009:
2005:
2004:
2001:
2000:
1996:
1995:
1992:
1991:
1987:
1986:
1983:
1982:
1978:
1977:
1974:
1973:
1969:
1968:
1965:
1964:
1960:
1959:
1956:
1955:
1951:
1950:
1947:
1946:
1942:
1941:
1938:
1937:
1933:
1932:
1929:
1928:
1924:
1923:
1920:
1919:
1915:
1914:
1911:
1910:
1906:
1905:
1902:
1901:
1897:
1896:
1893:
1892:
1888:
1887:
1884:
1883:
1879:
1878:
1875:
1874:
1870:
1869:
1866:
1865:
1861:
1860:
1857:
1856:
1852:
1851:
1848:
1847:
1843:
1842:
1839:
1838:
1834:Bairdemys spp.
1829:
1824:
1822:
1819:
1818:
1815:
1814:
1810:
1809:
1806:
1805:
1801:
1800:
1797:
1796:
1787:
1784:
1783:
1780:
1779:
1770:
1767:
1766:
1763:
1762:
1753:
1750:
1749:
1746:
1745:
1736:
1731:
1729:
1726:
1725:
1722:
1721:
1712:
1707:
1705:
1700:
1698:
1695:
1694:
1691:
1690:
1681:
1676:
1674:
1671:
1670:
1667:
1666:
1657:
1652:
1650:
1647:
1646:
1643:
1642:
1633:
1628:
1626:
1623:
1622:
1619:
1618:
1609:
1604:
1602:
1599:
1598:
1595:
1594:
1585:
1580:
1578:
1575:
1574:
1571:
1570:
1561:
1556:
1554:
1551:
1550:
1547:
1546:
1542:
1541:
1538:
1537:
1533:
1532:
1529:
1528:
1524:
1523:
1520:
1519:
1515:
1514:
1511:
1510:
1506:
1505:
1502:
1501:
1497:
1496:
1493:
1492:
1488:
1487:
1484:
1483:
1479:
1478:
1475:
1474:
1469:
1466:
1465:
1462:
1461:
1452:
1447:
1445:
1442:
1441:
1438:
1437:
1428:
1423:
1421:
1418:
1417:
1414:
1413:
1404:
1399:
1397:
1394:
1393:
1390:
1389:
1380:
1375:
1373:
1370:
1369:
1366:
1365:
1361:
1360:
1357:
1356:
1347:
1344:
1343:
1340:
1339:
1330:
1325:
1323:
1318:
1316:
1313:
1312:
1309:
1308:
1299:
1294:
1292:
1289:
1288:
1285:
1284:
1275:
1270:
1268:
1265:
1264:
1261:
1260:
1251:
1246:
1244:
1239:
1237:
1234:
1233:
1230:
1229:
1220:
1215:
1213:
1208:
1206:
1203:
1202:
1199:
1198:
1194:
1193:
1190:
1189:
1185:
1184:
1181:
1180:
1176:
1175:
1172:
1171:
1167:
1166:
1163:
1162:
1158:
1157:
1154:
1153:
1149:
1148:
1145:
1144:
1135:
1132:
1131:
1128:
1127:
1118:
1113:
1111:
1108:
1107:
1104:
1103:
1094:
1089:
1087:
1084:
1083:
1080:
1079:
1075:
1074:
1071:
1070:
1061:
1058:
1057:
1054:
1053:
1044:
1039:
1037:
1032:
1030:
1027:
1026:
1023:
1022:
1013:
1008:
1006:
1003:
1002:
999:
998:
989:
984:
982:
979:
978:
975:
974:
965:
960:
958:
953:
951:
947:
946:
943:
942:
938:
937:
934:
933:
929:
928:
925:
924:
915:
912:
911:
908:
907:
898:
893:
891:
888:
887:
884:
883:
874:
869:
867:
862:
860:
857:
856:
853:
852:
843:
838:
836:
833:
832:
829:
828:
819:
814:
812:
809:
808:
805:
804:
795:
790:
788:
785:
784:
781:
780:
771:
766:
764:
761:
760:
757:
756:
747:
742:
740:
734:
658:
655:
626:The weight of
594:
591:
507:
504:
446:Tatacoa Desert
439:S. geographica
339:
336:
308:Podocnemididae
299:of freshwater
283:
282:
281:
280:
266:
265:
259:
258:
257:
256:
251:S. geographica
242:
241:
237:
236:
222:
218:
217:
215:Podocnemididae
212:
208:
207:
202:
198:
197:
192:
188:
187:
182:
178:
177:
172:
168:
167:
162:
158:
157:
152:
148:
147:
134:
133:
128:A young adult
125:
124:
116:
115:
107:
106:
105:
100:
95:
90:
85:
80:
75:
70:
65:
60:
55:
50:
45:
28:
15:
9:
6:
4:
3:
2:
5255:
5244:
5241:
5239:
5236:
5234:
5231:
5229:
5226:
5224:
5221:
5219:
5216:
5214:
5211:
5209:
5206:
5204:
5201:
5199:
5196:
5194:
5191:
5189:
5188:Montehermosan
5186:
5184:
5181:
5179:
5176:
5174:
5171:
5169:
5166:
5164:
5161:
5159:
5156:
5154:
5151:
5149:
5146:
5144:
5141:
5139:
5136:
5135:
5133:
5118:
5114:
5113:
5109:
5108:
5103:
5102:
5099:
5085:
5084:
5083:Tacuarembemys
5079:
5077:
5076:
5071:
5069:
5068:
5063:
5062:
5060:
5056:
5053:
5049:
5046:
5042:
5028:
5027:
5022:
5020:
5019:
5018:Sahonachelys
5014:
5013:
5011:
5009:
5004:
4998:
4997:
4992:
4990:
4989:
4985:
4983:
4982:
4981:Peltocephalus
4978:
4976:
4975:
4970:
4968:
4967:
4966:Lapparentemys
4962:
4960:
4959:
4955:
4953:
4952:
4947:
4945:
4944:
4939:
4937:
4936:
4931:
4929:
4928:
4923:
4921:
4920:
4915:
4913:
4912:
4907:
4905:
4904:
4899:
4898:
4896:
4894:
4890:
4884:
4883:
4879:
4877:
4876:
4872:
4871:
4869:
4867:
4866:Pelomedusidae
4863:
4857:
4856:
4855:Yaminuechelys
4851:
4849:
4848:
4844:
4842:
4841:
4837:
4835:
4834:
4830:
4828:
4827:
4826:Prochelidella
4822:
4820:
4819:
4815:
4813:
4812:
4808:
4806:
4805:
4801:
4799:
4798:
4794:
4792:
4791:
4786:
4784:
4783:
4779:
4777:
4776:
4772:
4770:
4769:
4765:
4763:
4762:
4758:
4756:
4755:
4751:
4749:
4748:
4744:
4742:
4741:
4740:Acanthochelys
4737:
4736:
4734:
4732:
4728:
4722:
4721:
4716:
4714:
4713:
4708:
4706:
4705:
4700:
4698:
4697:
4692:
4690:
4689:
4684:
4682:
4681:
4676:
4674:
4673:
4668:
4666:
4665:
4660:
4658:
4657:
4652:
4650:
4649:
4644:
4642:
4641:
4636:
4634:
4633:
4628:
4626:
4625:
4624:Kinkonychelys
4620:
4618:
4617:
4612:
4610:
4609:
4608:Itapecuruemys
4604:
4602:
4601:
4596:
4594:
4593:
4588:
4586:
4585:
4580:
4578:
4577:
4572:
4570:
4569:
4568:Eotaphrosphys
4564:
4562:
4561:
4556:
4554:
4553:
4548:
4546:
4545:
4540:
4538:
4537:
4532:
4530:
4529:
4524:
4522:
4521:
4516:
4514:
4513:
4508:
4507:
4505:
4503:
4502:Bothremydidae
4498:
4492:
4491:
4486:
4485:
4483:
4481:
4476:
4473:
4469:
4466:
4464:
4460:
4446:
4445:
4440:
4438:
4437:
4432:
4430:
4429:
4424:
4422:
4421:
4416:
4414:
4413:
4412:Argillochelys
4408:
4406:
4405:
4400:
4399:
4397:
4393:
4387:
4386:
4381:
4379:
4378:
4377:Santanachelys
4373:
4371:
4370:
4365:
4363:
4362:
4357:
4355:
4354:
4349:
4347:
4346:
4341:
4339:
4338:
4333:
4331:
4330:
4325:
4323:
4322:
4321:Desmatochelys
4317:
4315:
4314:
4309:
4307:
4306:
4305:Calcarichelys
4301:
4299:
4298:
4293:
4291:
4290:
4289:Atlantochelys
4285:
4283:
4282:
4277:
4275:
4274:
4269:
4268:
4266:
4264:
4263:Protostegidae
4259:
4253:
4252:
4251:Jiangxichelys
4247:
4245:
4244:
4239:
4237:
4236:
4235:Anomalochelys
4231:
4230:
4228:
4226:
4221:
4215:
4214:
4213:Protochelydra
4209:
4207:
4206:
4205:Planiplastron
4201:
4199:
4198:
4194:
4192:
4191:
4186:
4184:
4183:
4178:
4176:
4175:
4170:
4168:
4167:
4162:
4160:
4159:
4155:
4154:
4152:
4150:
4146:
4143:
4139:
4129:
4128:
4123:
4121:
4120:
4115:
4114:
4112:
4108:
4102:
4101:
4097:
4095:
4094:
4090:
4088:
4087:
4083:
4081:
4080:
4076:
4074:
4073:
4068:
4066:
4065:
4061:
4059:
4058:
4054:
4052:
4051:
4047:
4045:
4044:
4039:
4037:
4036:
4031:
4029:
4028:
4023:
4021:
4020:
4016:
4014:
4013:
4012:Drazinderetes
4008:
4006:
4005:
4001:
3999:
3998:
3994:
3992:
3991:
3987:
3985:
3984:
3979:
3977:
3976:
3972:
3970:
3969:
3965:
3964:
3962:
3960:
3956:
3950:
3949:
3948:Carettochelys
3945:
3943:
3942:
3937:
3935:
3934:
3929:
3928:
3926:
3924:
3920:
3917:
3915:
3911:
3901:
3900:
3896:
3894:
3893:
3888:
3886:
3885:
3881:
3879:
3878:
3873:
3871:
3870:
3866:
3864:
3863:
3859:
3857:
3856:
3851:
3849:
3848:
3843:
3841:
3840:
3836:
3834:
3833:
3832:Malacochersus
3829:
3827:
3826:
3822:
3820:
3819:
3815:
3813:
3812:
3808:
3806:
3805:
3800:
3798:
3797:
3792:
3790:
3789:
3785:
3783:
3782:
3778:
3776:
3775:
3770:
3768:
3767:
3762:
3760:
3759:
3755:
3753:
3752:
3748:
3746:
3745:
3741:
3739:
3738:
3734:
3732:
3731:
3727:
3725:
3724:
3723:Aldabrachelys
3720:
3719:
3717:
3715:
3711:
3705:
3704:
3700:
3699:
3697:
3695:
3690:
3684:
3683:
3679:
3677:
3676:
3672:
3670:
3669:
3665:
3663:
3662:
3658:
3656:
3655:
3651:
3649:
3648:
3644:
3642:
3641:
3637:
3635:
3634:
3630:
3628:
3627:
3623:
3621:
3620:
3616:
3614:
3613:
3609:
3607:
3606:
3605:Leucocephalon
3602:
3600:
3599:
3595:
3593:
3592:
3588:
3586:
3585:
3581:
3579:
3578:
3574:
3572:
3571:
3567:
3565:
3564:
3560:
3558:
3557:
3556:Banhxeochelys
3552:
3550:
3549:
3545:
3544:
3542:
3540:
3536:
3530:
3529:
3524:
3522:
3521:
3517:
3515:
3514:
3510:
3508:
3507:
3503:
3501:
3500:
3496:
3494:
3493:
3489:
3487:
3486:
3482:
3480:
3479:
3475:
3473:
3472:
3468:
3466:
3465:
3461:
3459:
3458:
3454:
3452:
3451:
3447:
3445:
3444:
3440:
3438:
3437:
3432:
3431:
3429:
3427:
3423:
3420:
3418:
3417:Testudinoidea
3414:
3404:
3403:
3399:
3397:
3396:
3392:
3390:
3389:
3385:
3383:
3382:
3377:
3375:
3374:
3370:
3369:
3367:
3365:
3364:Kinosternidae
3361:
3355:
3354:
3350:
3349:
3347:
3345:
3341:
3338:
3336:
3332:
3322:
3321:
3316:
3314:
3313:
3312:Protosphargis
3308:
3306:
3305:
3300:
3298:
3297:
3292:
3290:
3289:
3284:
3283:
3281:
3277:
3271:
3270:
3265:
3263:
3262:
3257:
3255:
3254:
3249:
3247:
3246:
3242:
3240:
3239:
3234:
3232:
3231:
3226:
3224:
3223:
3218:
3217:
3215:
3213:
3209:
3203:
3202:
3197:
3195:
3194:
3189:
3187:
3186:
3181:
3179:
3178:
3174:
3172:
3171:
3166:
3164:
3163:
3158:
3156:
3155:
3151:
3149:
3148:
3143:
3141:
3140:
3135:
3133:
3132:
3127:
3125:
3124:
3120:
3118:
3117:
3112:
3110:
3109:
3105:
3103:
3102:
3097:
3095:
3094:
3090:
3088:
3087:
3082:
3081:
3079:
3077:
3073:
3070:
3068:(Sea turtles)
3066:
3062:
3059:
3057:
3053:
3037:
3036:
3030:
3027:
3025:
3021:
3018:
3016:
3012:
3009:
3007:
3003:
2997:
2993:
2991:
2987:
2984:
2982:
2978:
2976:
2972:
2970:
2966:
2965:
2962:
2958:
2951:
2946:
2944:
2939:
2937:
2932:
2931:
2928:
2914:
2909:
2905:
2901:
2896:
2892:
2888:
2883:
2879:
2875:
2870:
2866:
2862:
2857:
2853:
2849:
2843:
2838:
2832:
2828:
2823:
2817:
2813:
2812:
2810:
2808:
2804:
2800:
2795:
2778:on 2011-10-11
2777:
2773:
2767:
2759:
2755:
2751:
2747:
2743:
2739:
2734:
2729:
2725:
2721:
2717:
2713:
2709:
2705:
2701:
2694:
2686:
2682:
2677:
2672:
2668:
2664:
2660:
2656:
2652:
2645:
2637:
2633:
2628:
2623:
2618:
2613:
2609:
2605:
2601:
2597:
2593:
2586:
2584:
2582:
2580:
2578:
2576:
2574:
2572:
2570:
2561:
2557:
2552:
2547:
2542:
2537:
2533:
2529:
2525:
2521:
2517:
2510:
2508:
2506:
2504:
2502:
2500:
2498:
2496:
2494:
2492:
2482:
2477:
2473:
2469:
2462:
2460:
2451:
2449:
2447:
2445:
2436:
2432:
2428:
2426:
2417:
2415:
2413:
2411:
2402:
2398:
2396:
2388:
2384:
2378:
2367:
2358:
2355:
2352:
2351:
2347:
2345:
2341:
2340:
2339:Drazinderetes
2336:
2335:
2329:
2327:
2323:
2322:
2318:
2314:
2310:
2309:
2304:
2300:
2299:
2294:
2290:
2286:
2282:
2281:
2276:
2272:
2268:
2264:
2260:
2256:
2255:
2250:
2246:
2242:
2241:
2236:
2232:
2228:
2224:
2220:
2216:
2211:
2208:
2204:
2203:
2198:
2197:
2192:
2191:
2186:
2182:
2178:
2169:
2160:
2158:
2153:
2149:
2148:
2147:Peltocephalus
2143:
2142:
2137:
2136:
2131:
2126:
2121:
2118:
2106:
2101:
2097:
2095:
2090:
2086:
2082:
2078:
2074:
2064:
2062:
2058:
2054:
2050:
2046:
2041:
2039:
2035:
2031:
2027:
2023:
2003:
2002:
1994:
1993:
1985:
1984:
1976:
1975:
1967:
1966:
1958:
1957:
1949:
1948:
1940:
1939:
1931:
1930:
1922:
1921:
1913:
1912:
1904:
1903:
1895:
1894:
1886:
1885:
1877:
1876:
1868:
1867:
1859:
1858:
1850:
1849:
1841:
1840:
1837:
1836:
1835:
1828:
1827:
1821:
1820:
1817:
1816:
1808:
1807:
1799:
1798:
1795:
1794:
1793:
1786:
1785:
1782:
1781:
1778:
1777:
1776:
1769:
1768:
1765:
1764:
1761:
1760:
1759:
1752:
1751:
1748:
1747:
1744:
1743:
1742:
1735:
1734:
1728:
1727:
1724:
1723:
1720:
1719:
1718:
1711:
1710:
1704:
1703:
1697:
1696:
1693:
1692:
1689:
1688:
1687:
1680:
1679:
1673:
1672:
1669:
1668:
1665:
1664:
1663:
1656:
1655:
1649:
1648:
1645:
1644:
1641:
1640:
1639:
1632:
1631:
1625:
1624:
1621:
1620:
1617:
1616:
1615:
1608:
1607:
1601:
1600:
1597:
1596:
1593:
1592:
1591:
1584:
1583:
1577:
1576:
1573:
1572:
1569:
1568:
1567:
1560:
1559:
1553:
1552:
1549:
1548:
1540:
1539:
1531:
1530:
1522:
1521:
1513:
1512:
1504:
1503:
1495:
1494:
1486:
1485:
1477:
1476:
1473:
1468:
1467:
1464:
1463:
1460:
1459:
1458:
1451:
1450:
1444:
1443:
1440:
1439:
1436:
1435:
1434:
1427:
1426:
1420:
1419:
1416:
1415:
1412:
1411:
1410:
1403:
1402:
1396:
1395:
1392:
1391:
1388:
1387:
1386:
1379:
1378:
1372:
1371:
1368:
1367:
1359:
1358:
1355:
1354:
1353:
1346:
1345:
1342:
1341:
1338:
1337:
1336:
1329:
1328:
1322:
1321:
1315:
1314:
1311:
1310:
1307:
1306:
1305:
1298:
1297:
1291:
1290:
1287:
1286:
1283:
1282:
1281:
1274:
1273:
1267:
1266:
1263:
1262:
1259:
1258:
1257:
1250:
1249:
1243:
1242:
1236:
1235:
1232:
1231:
1228:
1227:
1226:
1219:
1218:
1212:
1211:
1205:
1204:
1201:
1200:
1192:
1191:
1183:
1182:
1174:
1173:
1165:
1164:
1156:
1155:
1147:
1146:
1143:
1142:
1141:
1134:
1133:
1130:
1129:
1126:
1125:
1124:
1117:
1116:
1110:
1109:
1106:
1105:
1102:
1101:
1100:
1093:
1092:
1086:
1085:
1082:
1081:
1073:
1072:
1069:
1068:
1067:
1060:
1059:
1056:
1055:
1052:
1051:
1050:
1043:
1042:
1036:
1035:
1029:
1028:
1025:
1024:
1021:
1020:
1019:
1012:
1011:
1005:
1004:
1001:
1000:
997:
996:
995:
988:
987:
981:
980:
977:
976:
973:
972:
971:
964:
963:
957:
956:
949:
948:
945:
944:
936:
935:
927:
926:
923:
922:
921:
914:
913:
910:
909:
906:
905:
904:
897:
896:
890:
889:
886:
885:
882:
881:
880:
873:
872:
866:
865:
859:
858:
855:
854:
851:
850:
849:
842:
841:
835:
834:
831:
830:
827:
826:
825:
818:
817:
811:
810:
807:
806:
803:
802:
801:
794:
793:
787:
786:
783:
782:
779:
778:
777:
770:
769:
763:
762:
759:
758:
755:
754:
753:
746:
745:
738:
737:
733:
731:
730:
725:
722:at its base.
721:
720:
715:
711:
710:
705:
704:
699:
698:
692:
688:
684:
680:
676:
672:
668:
664:
654:
652:
648:
644:
639:
637:
633:
629:
624:
622:
620:
615:
611:
607:
603:
599:
588:
584:
579:
575:
573:
569:
568:costal scutes
565:
561:
557:
552:
548:
543:
540:
536:
532:
528:
524:
521:The skull of
516:
512:
503:
501:
497:
496:
491:
487:
483:
479:
474:
472:
468:
464:
460:
456:
452:
447:
444:
440:
436:
432:
428:
424:
420:
416:
412:
411:
406:
402:
398:
394:
393:
388:
384:
380:
376:
372:
368:
364:
360:
356:
352:
348:
344:
335:
333:
329:
325:
321:
320:South America
317:
313:
309:
305:
302:
298:
295:
291:
290:
276:
270:
269:
267:
264:
260:
252:
246:
245:
243:
238:
231:
230:
223:
220:
219:
216:
213:
210:
209:
206:
203:
200:
199:
196:
193:
190:
189:
186:
183:
180:
179:
176:
173:
170:
169:
166:
163:
160:
159:
156:
153:
150:
149:
144:
139:
135:
131:
126:
122:
117:
103:
98:
93:
88:
83:
78:
73:
68:
63:
58:
53:
48:
42:
36:
32:
26:
22:
19:
5106:
5081:
5073:
5065:
5024:
5016:
4995:
4994:
4986:
4979:
4972:
4964:
4958:Erymnochelys
4956:
4949:
4943:Cerrejonemys
4941:
4933:
4925:
4919:Brontochelys
4917:
4909:
4901:
4880:
4873:
4853:
4845:
4838:
4833:Pseudemydura
4831:
4824:
4816:
4809:
4802:
4795:
4788:
4780:
4773:
4766:
4759:
4752:
4745:
4738:
4718:
4710:
4702:
4694:
4686:
4678:
4670:
4662:
4654:
4646:
4638:
4630:
4622:
4614:
4606:
4598:
4590:
4582:
4574:
4566:
4558:
4550:
4542:
4534:
4526:
4518:
4510:
4488:
4442:
4436:Prionochelys
4434:
4426:
4418:
4410:
4402:
4383:
4375:
4367:
4359:
4351:
4343:
4335:
4327:
4319:
4313:Cratochelone
4311:
4303:
4297:Bouliachelys
4295:
4287:
4279:
4273:Alienochelys
4271:
4249:
4241:
4233:
4211:
4203:
4195:
4188:
4180:
4174:Chelydropsis
4172:
4164:
4156:
4125:
4119:Basilochelys
4117:
4098:
4091:
4084:
4077:
4070:
4062:
4055:
4048:
4043:Khunnuchelys
4041:
4033:
4025:
4017:
4010:
4002:
3995:
3988:
3981:
3973:
3966:
3959:Trionychidae
3946:
3939:
3933:Allaeochelys
3931:
3897:
3890:
3884:Stigmochelys
3882:
3875:
3867:
3860:
3853:
3847:Megalochelys
3845:
3837:
3830:
3823:
3816:
3809:
3802:
3794:
3786:
3779:
3772:
3764:
3758:Cylindraspis
3756:
3749:
3742:
3737:Centrochelys
3735:
3728:
3721:
3714:Testudinidae
3703:Platysternon
3701:
3682:Vijayachelys
3680:
3673:
3666:
3661:Rhinoclemmys
3659:
3652:
3645:
3638:
3631:
3626:Melanochelys
3624:
3617:
3610:
3603:
3596:
3589:
3582:
3575:
3568:
3561:
3554:
3546:
3526:
3518:
3511:
3504:
3497:
3490:
3483:
3476:
3469:
3462:
3455:
3448:
3441:
3436:Acherontemys
3434:
3402:Sternotherus
3400:
3393:
3386:
3379:
3371:
3351:
3318:
3310:
3302:
3294:
3286:
3269:Psephophorus
3267:
3259:
3251:
3243:
3236:
3228:
3220:
3199:
3191:
3185:Pacifichelys
3183:
3175:
3168:
3160:
3154:Lepidochelys
3152:
3145:
3137:
3131:Gigantatypus
3129:
3123:Eretmochelys
3121:
3114:
3106:
3099:
3091:
3084:
3065:Chelonioidea
3033:
2985:
2806:
2780:. Retrieved
2776:the original
2766:
2733:10261/167010
2707:
2703:
2693:
2658:
2654:
2644:
2599:
2595:
2523:
2519:
2471:
2467:
2458:
2434:
2430:
2424:
2400:
2394:
2387:
2348:
2337:
2325:
2319:
2306:
2296:
2278:
2270:
2262:
2252:
2238:
2214:
2212:
2207:Stupendemys'
2206:
2200:
2194:
2188:
2184:
2176:
2174:
2156:
2145:
2141:Erymnochelys
2139:
2133:
2124:
2122:
2116:
2114:
2104:
2093:
2088:
2072:
2070:
2060:
2052:
2048:
2044:
2043:The diet of
2042:
2033:
2029:
2025:
2021:
2019:
2016:Paleoecology
2011:Paleobiology
1832:
1831:
1790:
1789:
1773:
1772:
1756:
1755:
1739:
1738:
1715:
1714:
1684:
1683:
1660:
1659:
1636:
1635:
1612:
1611:
1589:
1588:
1587:
1564:
1563:
1471:
1455:
1454:
1431:
1430:
1407:
1406:
1383:
1382:
1350:
1349:
1333:
1332:
1302:
1301:
1278:
1277:
1254:
1253:
1223:
1222:
1138:
1137:
1121:
1120:
1097:
1096:
1064:
1063:
1047:
1046:
1016:
1015:
992:
991:
968:
967:
918:
917:
901:
900:
877:
876:
846:
845:
822:
821:
798:
797:
774:
773:
750:
749:
727:
723:
717:
713:
707:
701:
695:
682:
678:
670:
666:
660:
650:
642:
640:
635:
631:
627:
625:
617:
606:parasagittal
601:
597:
596:
582:
571:
550:
544:
531:bony nostril
522:
520:
514:
494:
493:
489:
485:
477:
475:
470:
466:
462:
458:
454:
438:
434:
430:
426:
408:
391:
390:
370:
342:
341:
331:
288:
287:
286:
274:
250:
228:
227:
129:
24:
18:
5183:Huayquerian
4996:Stupendemys
4951:Cordichelys
4797:Mesoclemmys
4782:Hydromedusa
4704:Taphrosphys
4696:Sankuchemys
4688:Rhothonemys
4664:Polysternon
4544:Cearachelys
4528:Azabbaremys
4512:Araiochelys
4428:Ctenochelys
4420:Bashuchelys
4369:Rhinochelys
4345:Ocepechelon
4337:Notochelone
4329:Iserosaurus
4197:Macrochelys
4149:Chelydridae
4072:Palaeoamyda
3997:Cyclanorbis
3862:Psammobates
3855:Oligopherus
3818:Indotestudo
3766:Cymatholcus
3744:Chelonoidis
3730:Astrochelys
3539:Geoemydidae
3457:Deirochelys
3395:Staurotypus
3388:Kinosternon
3381:Hoplochelys
3296:Peritresius
3245:Dermochelys
3238:Cosmochelys
3230:Corsochelys
3147:Itilochelys
3139:Glarichelys
3086:Allopleuron
3076:Cheloniidae
3015:Superfamily
2837:Stupendemys
2831:Wikispecies
2807:Stupendemys
2401:Fossilworks
2395:Stupendemys
2326:Stupendemys
2308:Chelonoidis
2215:Stupendemys
2202:Gryposuchus
2196:Mourasuchus
2190:Purussaurus
2185:Stupendemys
2177:Stupendemys
2157:Stupendemys
2152:ontogenetic
2125:Stupendemys
2117:Stupendemys
2094:Stupendemys
2089:Stupendemys
2073:Stupendemys
2061:Stupendemys
2053:Stupendemys
2045:Stupendemys
2038:Gause's Law
2034:Stupendemys
2022:Stupendemys
1472:UCMP-42008
714:Stupendemys
683:Stupendemys
671:Stupendemys
667:Stupendemys
663:pelomedusid
651:Stupendemys
643:Stupendemys
636:Stupendemys
632:Stupendemys
628:Stupendemys
616:sea turtle
602:Stupendemys
598:Stupendemys
583:MCZ(P)-4376
581:Replica of
572:Stupendemys
551:Stupendemys
539:premaxillae
527:prefrontals
523:Stupendemys
515:Stupendemys
506:Description
490:Stupendemys
478:Stupendemys
471:Stupendemys
459:Stupendemys
431:Stupendemys
371:Stupendemys
343:Stupendemys
332:Stupendemys
301:side-necked
289:Stupendemys
229:Stupendemys
130:Stupendemys
25:Stupendemys
5132:Categories
5075:Caririemys
4988:Podocnemis
4974:Latentemys
4935:Carbonemys
4875:Pelomedusa
4712:Ummulisani
4632:Kurmademys
4600:Inaechelys
4584:Galianemys
4552:Chedighaii
4490:Araripemys
4463:Pleurodira
4444:Toxochelys
4361:Protostega
4166:Chelydrops
4086:Pelodiscus
4079:Pelochelys
4027:Gilmoremys
4004:Cycloderma
3914:Trionychia
3781:Geochelone
3774:Floridemys
3640:Notochelys
3528:Wilburemys
3499:Malaclemys
3353:Dermatemys
3320:Puppigerus
3253:Eosphargis
3170:Miocaretta
3162:Mexichelys
3056:Cryptodira
2996:Testudines
2957:Testudines
2782:2011-10-11
2363:References
2227:Acre State
2135:Podocnemis
689:using the
685:and other
638:specimen.
614:Cretaceous
556:keratinous
467:Caninemys'
401:Acre State
349:-244, the
254:Wood, 1976
234:Wood, 1976
205:Pleurodira
201:Suborder:
195:Testudines
5115:See also
5067:Caribemys
4927:Caninemys
4911:Bauruemys
4840:Rheodytes
4804:Myuchelys
4747:Chelodina
4672:Puentemys
4648:Nigeremys
4592:Ilatardia
4536:Bothremys
4243:Basilemys
4057:Nilssonia
4035:Hutchemys
3983:Axestemys
3941:Anosteira
3796:Hadrianus
3654:Pangshura
3612:Malayemys
3577:Geoclemys
3520:Trachemys
3513:Terrapene
3506:Pseudemys
3492:Graptemys
3485:Glyptemys
3478:Emydoidea
3471:Actinemys
3443:Chrysemys
3288:Euclastes
3116:Eochelone
2967:Kingdom:
2758:206528591
2742:0036-8075
2303:mata mata
2293:tortoises
2249:characids
2096:fossils.
2057:Arecaceae
2049:Caninemys
2030:Caninemys
2026:Caninemys
724:Caninemys
679:Caninemys
657:Phylogeny
564:posterior
549:of adult
535:parietals
476:The name
469:skull to
463:Caninemys
455:Caninemys
443:Colombian
435:S. souzai
427:Caninemys
410:Caninemys
387:Venezuela
275:S. souzai
161:Kingdom:
155:Eukaryota
132:specimen
35:Messinian
4882:Pelusios
4847:Rhinemys
4818:Platemys
4811:Phrynops
4731:Chelidae
4720:Zolhafah
4616:Jainemys
4281:Archelon
4158:Chelydra
4050:Lissemys
3892:Stylemys
3877:Solitudo
3839:Manouria
3788:Gopherus
3751:Chersina
3619:Mauremys
3598:Heosemys
3591:Hardella
3584:Geoemyda
3570:Cyclemys
3426:Emydidae
3373:Claudius
3222:Arabemys
3201:Tasbacka
3193:Syllomus
3108:Chelonia
3006:Suborder
2990:Diapsida
2981:Reptilia
2975:Chordata
2973:Phylum:
2969:Animalia
2845:BioLib:
2816:Wikidata
2750:21071659
2685:38471564
2676:10932709
2636:35005268
2560:32095528
2431:Breviora
2350:Archelon
2332:See also
2275:trahiras
2259:tambaqui
2257:and the
2251:such as
2237:such as
2221:and the
2111:Ontogeny
619:Archelon
547:carapace
423:holotype
419:allotype
379:plastron
355:carapace
328:Pliocene
312:carapace
263:Synonyms
240:Species
211:Family:
185:Reptilia
175:Chordata
171:Phylum:
165:Animalia
151:Domain:
31:Langhian
5026:Sokatra
4775:Emydura
4680:Rosasia
4576:Foxemys
4520:Arenila
4100:Trionyx
4093:Rafetus
4019:Dogania
3975:Apalone
3899:Testudo
3825:Kinixys
3811:Homopus
3668:Sacalia
3647:Orlitia
3633:Morenia
3548:Batagur
3450:Clemmys
3177:Natator
3093:Caretta
2994:Order:
2979:Class:
2900:3606100
2887:1038762
2874:4819015
2861:4522034
2822:Q134896
2712:Bibcode
2704:Science
2627:8717240
2604:Bibcode
2596:Heliyon
2551:7015691
2528:Bibcode
2437:: 1–31.
2265:), the
2235:catfish
324:Miocene
316:fossils
294:extinct
221:Genus:
191:Order:
181:Class:
108:↓
5058:
5051:
5044:
4768:Elusor
4761:Elseya
4754:Chelus
4471:
4404:Adocus
4395:
4141:
4110:
3990:Chitra
3692:
3279:
3024:Family
2756:
2748:
2740:
2683:
2673:
2634:
2624:
2558:
2548:
2317:darter
2305:) and
2289:sharks
2277:(e.g.
2231:Brazil
706:, and
610:Vienna
405:Brazil
365:and a
351:medial
304:turtle
292:is an
4064:Palea
3968:Amyda
3869:Pyxis
3563:Cuora
3035:Genus
2986:Clade
2913:37607
2882:IRMNG
2848:60931
2754:S2CID
2661:(3).
2464:(PDF)
587:Osaka
517:skull
429:into
359:femur
297:genus
3464:Emys
2869:GBIF
2746:PMID
2738:ISSN
2681:PMID
2632:PMID
2556:PMID
2287:and
2285:rays
2243:and
2144:and
2024:and
593:Size
545:The
347:MCNC
47:PreꞒ
2856:EoL
2728:hdl
2720:doi
2708:330
2671:PMC
2663:doi
2622:PMC
2612:doi
2546:PMC
2536:doi
2476:doi
2435:436
2273:),
2225:in
585:in
403:in
399:in
385:in
375:MCZ
5134::
2988::
2910::
2897::
2884::
2871::
2858::
2833::
2818::
2752:.
2744:.
2736:.
2726:.
2718:.
2706:.
2702:.
2679:.
2669:.
2659:20
2657:.
2653:.
2630:.
2620:.
2610:.
2598:.
2594:.
2568:^
2554:.
2544:.
2534:.
2522:.
2518:.
2490:^
2470:.
2466:.
2443:^
2433:.
2429:.
2409:^
2399:.
2393:"†
2247:,
2229:,
2159:.
2138:,
732:.
700:,
502:.
361:,
97:Pg
41:Ma
33:-
5080:†
5072:†
5064:†
5023:†
5015:†
5006:†
4993:†
4971:†
4963:†
4948:†
4940:†
4932:†
4924:†
4916:†
4908:†
4900:†
4852:†
4823:†
4787:†
4717:†
4709:†
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4693:†
4685:†
4677:†
4669:†
4661:†
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4645:†
4637:†
4629:†
4621:†
4613:†
4605:†
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4581:†
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3553:†
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2949:e
2942:t
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2785:.
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2687:.
2665::
2638:.
2614::
2606::
2600:7
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2478::
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2457:"
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2269:(
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621:,
272:†
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102:N
92:K
87:J
82:T
77:P
72:C
67:D
62:S
57:O
52:Ꞓ
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