586:. According to Sterli, this competition may explain the whereabouts or rather the absence of Ameghino's fossil, which had seemingly disappeared from research history following the initial short description. No subsequent authors figured, described or even compared the Ameghino skull to the Roth skull. Juliana Sterli offers two possible explanations for this. It is possible that Ameghino's skull was simply lost, however it is likewise a possibility that the Ameghino and Roth skulls are the same specimen and that Ameghino misattributed the discovery to his brother. This would explain the improbability of two well preserved skulls being discovered in such quick succession by different teams and also explain why no researcher ever figured the Ameghino skull or compared it to the Roth material. Subsequently, Sterli refers to the
1705:
4103:
1713:
and frill and sideways stabbing and the stresses such actions put on the skeleton. Generally, impacts with the snout, forehead and struggling using the notch between the frill and horns cause the least amount of stress, as opposed to caudal torsion of the frill and sideways stabs. Given the large amounts of stress caused by the latter two actions, such behavior was generally deemed unlikely. This means that although the frill would be large enough to protect the neck, it's unlikely to have been used that way given the stresses of caudal torsion that would affect the bone in such a scenario.
4096:
843:
maximum thickness of 1.5 cm (0.59 in). Of the ten recovered shell remains, only four have their position on the shell known. The largest fragment was likely located at the back edge of the carapace, correlating with the 8th costal plate (which cover the shell between its spine and edge) and the 10th & 11th peripheral plates (the bony plates forming the edge of the bony shell). In this region the shell appears to possess backwards directed spikes. Rather than holding its shape towards the back of the shell, the carapace of
379:
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122:
388:
4142:
1832:
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1728:. The team further notes that the trabecular bone is more prominent in the horns and frills, yet these elements are positioned in a way that makes them unsuited for headbutting, with the frill in particular being angled so that would make direct contact with the frill of a rival impossible. Additional points against headbutting can be found in the absence of
1593:, which at one point would have been found in both southern South America and Antarctica. Given the fact that South Tasmanian Sea formed between the Cretaceous and late Eocene, meiolaniids must have arrived in Australia by this time, as is confirmed by the fossil record. Antarctica and South America would remain connected until the opening of the
764:
if this actually represents how it appeared in life or if this was simply the result of distortion after the animals death. The E scales are small elements restricted to the side of the skull, just before the tympanic cavity. They are convex and form small, backward directed bosses. The H scales seen in derived meiolaniids appear to be missing in
1745:. Nevertheless, this does not rule out other aggressive behavior that was speculated upon in prior studies, with Degrange and colleagues specifically highlighting the animal's ability to bite. Though a directly defensive function of the frill is also deemed unlikely, it may have still served as a visual deterent when dealing with predators.
1651:
produce chemical secretions (musk glands, cloacal secretions and mental glands). While the presence of these glands cannot be observed in meiolaniid fossils, they may have been tied to the keen sense of smell inferred for them. Furthermore, chemical cues are associated with combat and aggression between modern turtles, especially during
365:. While Ameghino's name was retained for the animal, later publications were entirely based on Roth's material, as Ameghino neither diagnosed nor figured his fossil. The fact that Ameghino's material was never recovered after his initial communication has led some researchers to believe that the two specimens are one and the same.
1736:
and the fact that meiolaniids as a whole have highly ankylosed skulls, meaning the shock of an impact could not be absorbed by the sutures between bones. The team does however note that it is theoretically possible that such a function may have been instead filled by the scutes that covered the skull
909:
in the same position relative to other meiolaniid turtles, which is as the basalmost member and sister to all
Australasian taxa. This placement is consistent with both its appearance in time and its place of origins, as this genus is older than all named species from Australia and the western pacific
763:
however, this scale is among the largest, appearing as a rectangular element that takes up the entire skullroof between the D scales and thus preventing them from touching each other. It further stands out due to being concave, rather than bulging out like the D and G scales. However it is not clear
570:
remained in storage and publications released during this time were primarily based on the work of
Woodward, rather than first hand observation of the skull. The convoluted history and poor description of the early meiolaniid discoveries of South America eventually led to a full redescription of the
486:
Around the same time Moreno authored a short communication describing the material Roth reportedly uncovered on his second expedition. Although various postcranial remains were also described, the skull (specimen MLP 26–40) was the only element that was figured. Two years later Arthur Smith
Woodward
1712:
The potential use of the horns and frill has been a frequently considered but rarely studied matter, with the primary study on the matter being published in 2024. The publication analyzed a variety of hypothetical combat behaviors performed with the head, including headbutting, locking of the horns
1642:
or even the production of sound. An enhanced sense of smell is also supported by the size of the nasal cavity (cavum nasi proprium). It is even greater than in extant terrestrial turtles, which in turn have a greater nasal cavity than aquatic species. Although the possibility that the dorsal region
1650:
Leaving aside the potential for additional uses, the enhanced sense of smell inferred by the large nasal cavity could have served several functions from foraging to intraspecific communication. Particular focus is placed on the later, with modern turtles displaying a wide variety of glands used to
741:
was covered in a multitude of horn or boss-like scales readily identifiable in the fossil material. These scales were described with varying nomenclature by different authors, with
Woodward assigning them the numerals I - VII, while Simpson assigned letters to the individual scales. From Simpson's
759:. A striking feature that sets this turtle apart from its relatives is the size of the X scale. The X scale is a singular scale that is situated along the midline of the skull atop the skullroof. Typically, this is a rather small scale surrounded on all sides by the much larger D and G scales. In
746:
appear in pairs, the exception being scale A, X, Y and Z. Scale A in particular is part of the series of scales that form large horns and bosses along the back of the skull in all meiolaniids. While the region furthest back on the skull is covered by the A scale, which forms an "occipital crest".
665:, meaning the majority of sutures between the individual skull bones are fused and thus largely obscured. Only a few sutures are visible on the surface of the skull, primarily along the midline, however in ventral view (from the bottom) the connections between the bones are more clearly visible.
1740:
The team, led by
Federico J. Degrange, concludes that though the animal's sense of smell was likely significant during courtship, it is more likely that the horns and frill served as a display structure rather than being used in intraspecific combat. Such a display ritual may have incorporated
880:
was covered in a series of protective bony rings that overlapped one another. These rings were roughly hexagonal in shape, with two pairs of spiky protrusions emerging. The top pair was more robust and directed upwards, while the smaller, lower pair was directed upwards and sideways. Unlike in
842:
had a domed shell like modern tortoises, a hypothesis in part based on the angle between the dorsal process and the acromion of the shoulder blade, which resembles modern testudinids. The shell appears to be thickest towards the borders and thinnest towards the midline of the shell, reaching a
1638:. In these lizards the elongated vestibulum helps the animal to keep its nose free of sand, which may also occur during digging. However, the authors do not settle on a specific hypothesis and instead offer a variety of alternative ideas. These include an enhanced sense of smell,
755:, are thought to have been even larger in life. The C scales sit before the B scales and are the smallest of the three elements, as is typical for turtles of this family. They extend outward and down, covering part of the skullroof and the side of the skull while overhanging the
930:, also from the Neogene of Argentina, did not change this placement much. However, this is in part due to the fragmentary nature of the later, as its position within the family is uncertain, being placed either alongside the derived Australasian forms or in a basal
1678:
80-95° and in meiolaniids up to 115°. In terrestrial species, this wide angle between the semicircular canals servers to improve the stability of the head while walking. Given the similarities to modern turtles however, it is assumed that they were sensitive to
1785:. Following the synonymty, this interpretation is either wrong or the scales would combine into a single element in adults, as exemplified by the neotype. The X scale, which is the scale placed atop the midline of the skull, extends between the G scales in
1630:. Among the findings of the paper was that meiolaniids had an elongated vestibulum nasi. Although such a feature is associated with aquatic turtles today, correlating with a snorkel-like nose, it can also be interpreted as an adaptation for life in
2398:
776:
that covers the cutting surface turtles have in place of teeth. While the lower half of the snout tip is made up by the I scales, the top is covered by the unpaired scales Y and Z, extending from above the eyes to the elongated nasal bones.
1724:, headbutting is still considered an unlikely behavior. Part of the reasoning behind this is that in pachycephalosaurids the shock of impacts is in part observed by their enlarged dome-like head structures, which as such are not present in
1780:
neotype skull. Accordingly, the shape of the horns is also significantly downplayed compared to those of adult individuals. The large scales at the back of the skull for instance, Scales A, were initially interpreted as being separate in
1655:, which may entail various shell-based maneuvers. This is congruent with prior studies suggesting a combat function for the highly armored bodies of meiolaniid turtles, which feature differently shaped osteoderms and armored tails.
938:. The two phylogenetic trees below, after Gaffney, Archer & White and Sterli, de la Fuenta & Krause respectively, show the internal relationship of meiolaniids following the description of the two most recently named genera.
847:
forms a ditch at its rear-most point, creating an embayment bordered by the final supracaudal scale. The recovered pattern of sutures and scale sulci is similar to that of modern tortoises, but could not be compared to the shell of
1962:
768:. The F scales cover the region above the eye sockets, and the K and J scales the side of the skull from its back up to the beak region. The beak itself is covered by scale I, which can be divided into scale I1 and scale I2 in
530:
in 1938, reasoning that
Ameghino's material was not described thoroughly enough to serve as the holotype. In the same paper the researchers also officially described additional meiolaniid remains from the Eocene parts of the
1695:
was a terrestrial animal like modern tortoises. This is the most commonly inferred lifestyle for meiolaniid turtles and generally favoured over the aquatic model, which is occasionally suggested but not widely accepted.
623:
specifying the Cañadón Blanco locality. However the mammal fauna of said locality is more in line with an Eocene to
Oligocene age, which matches Ameghino's claim that Roth's material stems from Middle Eocene strata near
507:, Ameghino seemingly concurred with Woodward, using the combination himself in later publications. However, Sterli raises the possibility that rather than accepting synonymity, Ameghino never meant to call his material
1683:
and not especially vocal animals, relying more heavily on smell. However as significant changes in the inner ear anatomy may often be subtle, a larger sample size would be needed to make more concrete observations for
606:
remains were found at and the general ambiguity around whether or not
Ameghino's and Roth's skulls are distinct has led to a lot of confusion in subsequent years. Ameghino named the Guaranitic Formation in the
471:, didn't diagnose the species or even figured the fossil material. The imprecise information was not an accident, but the result of the rivalry between Ameghino and Moreno. Their rivalry, similar to the
2423:
Poropat, S. F.; Kool, L.; Vickers-Rich, P.; Rich, T. H. (2016). "Oldest meiolaniid turtle remains from
Australia: evidence from the Eocene Kerosene Creek Member of the Rundle Formation, Queensland".
2355:"Comparative neuroanatomy of extinct horned turtles (Meiolaniidae) and extant terrestrial turtles (Testudinidae), with comments on the palaeobiological implications of selected endocranial features"
306:
was a robust and heavily armored terrestrial turtle with large, horn like scales covering its head and a tail encased by rings of bone. This heavily armored build may have served the animal during
487:
published a more detailed paper, in which he illustrated the material more extensively and assigned it to the same species mentioned previously by
Ameghino. However, rather than using the name
2490:
Degrange, F.J.; Nieto, M.N.; Sterli, J.; Vlachos, E. (2024). "Biomechanical skull performance in the meiolaniid Niolamia argentina (Testudinata: Meiolaniidae) using Finite Element Analysis".
1776:, makes it possible to identify several changes the animal underwent in the process of reaching adulthood. AMNH 3161 represents a very young individual, less than a fourth the size of the
742:
work, Gaffney refined and altered the nomenclature, which has since then become the standard nomenclature for the scale areas and horns of meiolaniid turtles. Most scales on the skull of
873:, however this estimate is not a certain one due to the fact that only fragmentary shell remains were known. Subsequently, future discoveries could confirm or contradict these results.
1589:
in their distribution, with Antarctica in particular serving as a connective bridge between the fauna of South America and Australasia. Meiolaniids derive from the primarily Gondwanan
834:
is known, researchers were nonetheless able to determine several aspects of its shape. Although the largest fossils of the shell are flattened, this is believed to have been caused by
636:, very much unlike the sediments described by Woodward and similar to what is found in the Eocene to Miocene Sarmiento Formation. This is supported by the discovery of additional
582:
In a later publication, Sterli would further comment on the complex history of this taxon and the impact of the intense rivalry between Ameghino and Moreno on the history of
2458:
Brown, L.E.; Moll, D. (2019). "The enigmatic palaeoecology and palaeobiogeography of the giant, horned, fossil turtles of Australasia: a review and reanalysis of the data".
2264:"A new turtle from the Palaeogene of Patagonia (Argentina) sheds new light on the diversity and evolution of the bizarre clade of horned turtles (Meiolaniidae, Testudinata)"
1597:
during the Eocene-Oligocene, by which point the South American meiolaniids had not only dispersed into Patagonia but already gone extinct. The cause of the extinction of
318:
likely had a great sense of smell but only low frequency hearing, indicating that these animals communicated more through chemical signals and smells than through sound.
448:. According to Woodward, Moreno tasked Roth with a follow-up expedition, which recovered additional remains in the form of a carapace, skull and associated mandible.
1903:
922:, typically recovered as the basalmost of the Australasian forms, as it shares multiple traits including the laterally directed horns and large A scale area with
3645:
464:
711:, which makes up approximately 60% of its volume. The vestibulum nasi is elongated, which today is only seen in turtles with snorkel like noses such as the
314:, though such encounters likely did not involve the horns and frill, which are thought to serve more of a display function. Scans of the skull reveal that
2869:
1666:
meanwhile is rather conservative and more in-line with what is seen in terrestrial turtles, which have a wide angle between the anterior and posterior
3743:
3659:
3955:
3022:
2947:
4235:
4026:
3977:
3631:
3498:
3450:
3359:
3117:
3036:
3029:
2940:
2885:
2795:
1222:
1205:
747:
This crest appears as a large, upward directed frill with a deep notch along its middle. The B scales, which cover the horn cores formed by the
3992:
3652:
3610:
3554:
3526:
3491:
3436:
3059:
661:
is a nearly completely preserved skull, which is the most diagnostic element in meiolaniid turtles. Like in its relatives, the skull is highly
4304:
3962:
3638:
3575:
3547:
3540:
3512:
3505:
3289:
3139:
809:. Overall, the anatomy of the shoulder blade has been taken to indicate a terrestrial lifestyle due to its similarity with modern tortoises.
1618:
s lifestyle. One exception to this is a 2017 study dealing with the neuroanatomy of meiolaniids, which described the endocranial anatomy of
3999:
3757:
3701:
3680:
3617:
3561:
3422:
3415:
3246:
3146:
2855:
2805:
612:
4040:
3771:
3443:
3429:
3380:
3366:
2926:
1262:
4359:
2552:
4309:
3477:
3373:
3345:
3310:
3262:
751:, appear as large triangular elements directed to the side of the animal. These horns, which are similar to those of the Australian
4222:
3750:
3729:
483:
in America, saw the two intentionally hide records from one another, eventually leading to the complete loss of some information.
3338:
2531:
1647:
dinosaurs, is not dismissed outright, it is considered relatively unlikely given the much more simple structure in meiolaniids.
4354:
644:) from the Sarmiento Formation. Subsequently, later research favors the hypothesis that these turtles lived during the Eocene.
2241:
353:
were seemingly first uncovered by Santiago Roth in 1889, with a well preserved skull found only shortly afterwards. The name
579:
individual, with the distinguishing features simply representing individual variation or traits that would change with age.
4102:
1704:
4294:
1557:
The presence of these two genera marks them as part of the Austral biogeogeographical kingdom, which includes southern
1601:
and other South American meiolaniids was likely the gradually cooling and drying climate of middle Eocene Patagonia.
2057:
4095:
2315:
2146:
2013:
1995:
Ameghino, F. (1899). "Sinopsis geológico-paleontológica de la Argentina. Suplemento (adiciones y correcciones)".
4349:
4344:
426:. Woodward writes that he was able to study the fossil itself in early 1899 after the material was sent from
4364:
4339:
859:
has been estimated based on comparison between shell fragments and the general proportions of the related
2524:
571:
neotype in 2011, authored by Juliana Sterli and Marcelo de la Fuente. Like Gaffney, they too argue that
467:. However, little information was given by Ameghino at the time, as the researcher didn't established a
1741:
differnet ritualized movements of the head based on the range of motion observed from the more recent
2399:"Warkalania, a New Meiolaniid Turtle from the Tertiary Riversleigh Deposits of Queensland, Australia"
423:
121:
1691:
Generally, both the enhanced sense of smell and the anatomy of the inner ear support the idea that
673:, lacking any divisions in its internal structure. This sets the genus apart from later taxa like
619:. Woodward wrote that Roth's specimen meanwhile came from Cretaceous aged red sandstone, with the
4329:
4299:
4289:
1820:", however this is likely simply the result of the sutures not yet being closed in the juvenile.
1643:
of the nasal cavity could have aided in vocalisation or thermoregulation, based on comparison to
1534:, most other members of the family were endemic to Neogene to Quaternary Australasia, including
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4266:
4162:
4145:
2517:
1279:
519:
4240:
4261:
4253:
1963:"On some extinct reptiles from Patagonia, of the genera Miolania, Dinilysia, and Genyodectes"
480:
415:
4324:
4314:
4209:
1922:
1721:
307:
632:. Analysis of the sediments present in the neotype's nasal cavity revealed it to be white
629:
8:
4074:
3533:
2628:
1851:
1667:
1434:
532:
476:
452:
436:
401:
358:
222:
1926:
869:. The result yielded an estimated shell length of 1.2 m (3 ft 11 in) for
4319:
4200:
2440:
2120:
1981:
1938:
793:
are described, as both the left and right preserve the elongated dorsal process of the
253:
116:
566:
While meiolaniid research received a considerable boost under Gaffney, the fossils of
4248:
3468:
2444:
2394:
2354:
2237:
2199:
2090:
1817:
1670:. In modern tortoises this angle is approximately 100°, in more aquatic turtles like
1652:
1527:
720:
544:
311:
2184:"The postcranial morphology of Meiolania platyceps and a review of the Meiolaniidae"
2124:
2034:
2017:
1942:
611:
as the place of origin of his fossils, claiming the material was collected near the
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3013:
2495:
2467:
2432:
2374:
2366:
2285:
2275:
2229:
2191:
2183:
2110:
2102:
2029:
1977:
1930:
1837:
1639:
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of meiolaniid turtles due to its presence in Neogene South America. In contrast to
716:
405:
2499:
2436:
1585:. Animals of this biogeographical kingdom were heavily affected by the breakup of
3945:
3209:
3197:
3050:
2705:
2353:
Paulina-Carabajal, A.; Sterli, J.; Georgi, J.; Poropat, S.F.; Kear, B.P. (2017).
1934:
1675:
820:
790:
756:
620:
608:
460:
362:
346:
an important piece in the evolutionary history and origin of this turtle family.
2370:
2233:
2091:"A review of the fossil record of Gondwanan turtles of the clade Meiolaniformes"
324:
is one of only two named meiolaniid turtles from South America, the other being
4120:
3854:
3833:
3708:
3568:
3303:
3176:
2906:
2878:
2845:
1590:
1357:
1160:
748:
504:
378:
4283:
4185:
3881:
3868:
3594:
3239:
3225:
3189:
3131:
3101:
2933:
2835:
2825:
2815:
2688:
2666:
2640:
2597:
2540:
2203:
2018:"Notes on the extinct Chelonian Meiolania, with a record of a new occurrence"
1644:
1594:
1558:
865:
419:
296:
176:
61:
801:
at an obtuse angle (greater than 90°), which resembles what is also seen in
361:
who briefly wrote about what was said to be another skull discovered by his
4135:
4130:
4019:
3985:
3969:
3926:
3673:
3624:
3484:
3296:
3066:
2967:
2958:
2785:
2775:
2765:
2726:
2715:
2471:
1416:
1374:
1332:
1301:
893:
were fully formed and lacked the opened underside seen in the more derived
827:
806:
773:
708:
625:
441:
326:
299:
202:
4194:
4033:
4010:
3905:
3861:
3840:
3589:
3408:
3394:
3387:
3352:
3331:
3110:
3080:
2748:
2736:
2406:
The Beagle, Records of the Northern Territory Museum of Arts and Sciences
2106:
1904:"Re-Description and Evolutionary Remarks on the Patagonian Horned Turtle
1806:
1729:
1671:
1578:
700:
692:
339:
189:
36:
4227:
4125:
3933:
3919:
3912:
3898:
3722:
3694:
3687:
3603:
3519:
3324:
3317:
3218:
3073:
2995:
2919:
2892:
2613:
2570:
2379:
2290:
2115:
1680:
1582:
1562:
1399:
1174:
1008:
838:, crushing the bone during preservation. Instead, it is suggested that
688:
683:, in which the internal nares are partially or fully divided by a bony
616:
547:
reasoned for the contrary in his 1996 review of meiolaniids, declaring
335:
81:
46:
2280:
2263:
789:
is also known from a variety of postcranial remains. Elements of both
4047:
3891:
3826:
3715:
3666:
3254:
3154:
3124:
2974:
2226:
Origin, Evolution and Biogeographic History of South American Turtles
2195:
1659:
1570:
1083:
1066:
1049:
1032:
835:
726:
712:
662:
472:
445:
427:
133:
86:
4156:
2352:
1753:
418:
was sent a photograph of a meiolaniid tail ring by fossil collector
4179:
3847:
3401:
3280:
3232:
3162:
2981:
2564:
2224:
de la Fuente, M. S.; Sterli, J.; Maniel, I (2014). "Introduction".
1586:
1574:
1550:
was the only form found outside this region until the discovery of
984:
931:
798:
675:
591:
468:
153:
76:
71:
56:
51:
41:
602:
The purposeful lack of information on the locality that the first
387:
3819:
3764:
3736:
3087:
2579:
1635:
797:, which extends upwards in turtles. This process connects to the
794:
587:
523:
331:
330:. Given that this family is primarily distributed throughout the
289:
163:
103:
91:
66:
4214:
2509:
4063:
3269:
2588:
2558:
2324:(Woodward), a Horned Turtle from the Pleistocene of Queensland"
1631:
1566:
684:
431:
143:
357:
was however not coined by Roth or any of his contacts, but by
3812:
1793:. The fact that the X scales in derived meiolaniids, such as
812:
670:
292:
691:
forms a small projection that extends beyond the end of the
302:
turtle with a long and complex history. Like its relatives,
2422:
633:
558:
522:
and Coleman S. Williams established Roth's material as the
2489:
1716:
Though impacts using the forehead are less stressfull and
2261:
2223:
1812:
Among the features once considered to be diagnostic for
772:. This region was likely covered by an extension of the
695:, however this is nowhere near as well developed as in
535:(then named the Casamayor Formation), which they named
2262:
Sterli, J.; de la Fuente, M.S.; Krause, J.M. (2015).
3782:
3586:
3465:
3173:
3098:
3047:
3010:
2903:
2866:
2851:
2841:
2831:
2821:
2811:
2801:
2791:
2781:
2771:
2761:
2732:
2722:
2711:
2701:
2425:
Alcheringa: An Australasian Journal of Palaeontology
1827:
1720:
does share the presence of trabecular bone with the
2058:"Crossochelys, Eocene horned turtle from Patagonia"
1546:and a variety of unnamed forms. For over 100 years
562:
The Neotype skull as illustrated in Woodward (1901)
2393:
1901:
2095:Bulletin of the Peabody Museum of Natural History
4281:
2228:. Springer Earth System Sciences. pp. 1–5.
1614:Few studies directly deal with the specifics of
2257:
2255:
2253:
2147:"New reptiles from the Eocene of South America"
2055:
1970:Proceedings of the Zoological Society of London
1902:Sterli, Juliana; de la Fuente, Marcelo (2011).
539:. Although Simpson argued that the material of
434:, allowing him to compare it to the remains of
400:was heavily influenced by the rivalry between
2525:
2686:
2250:
2012:
852:, in which the number of scales is unknown.
518:The naming issue was somewhat resolved when
1908:Ameghino, 1899 (Testudinata, Meiolaniidae)"
905:Phylogenetic analysis consistently recover
2532:
2518:
1956:
1954:
1952:
451:That same year Argentinian paleontologist
102:
2612:
2457:
2397:; Archer, Michael; White, Arthur (1992).
2378:
2359:Zoological Journal of the Linnean Society
2289:
2279:
2268:Zoological Journal of the Linnean Society
2114:
2033:
2348:
2346:
2344:
2342:
2340:
1994:
1960:
1768:and AMNH 3161, formerly the holotype of
1752:
1708:Male gopher tortoises engaging in combat
1703:
1699:
811:
725:
557:
2313:
2181:
2144:
1949:
1609:
1512:
876:Like in other meiolaniids, the tail of
511:in the first place and also misspelled
4282:
2485:
2483:
2481:
2309:
2307:
2305:
2303:
2301:
2088:
2056:Simpson, G.G.; Williams, C.S. (1938).
1897:
1895:
1893:
1891:
1889:
1887:
1885:
1883:
1134:
943:
495:based on an accidental misspelling of
4161:
4160:
2551:
2513:
2387:
2337:
2219:
2217:
2215:
2213:
2177:
2175:
2173:
2171:
2169:
2167:
2165:
2163:
2140:
2138:
2136:
2134:
2084:
2082:
2080:
2078:
2051:
2049:
2047:
2045:
2008:
2006:
1997:Imprenta la Libertad (Author Edition)
1881:
1879:
1877:
1875:
1873:
1871:
1869:
1867:
1865:
1863:
737:Like other meiolaniids, the skull of
368:
4305:Prehistoric turtles of South America
1772:and now considered to be a juvenile
1526:provides important insight into the
1518:
863:as well as those of the stem-turtle
2478:
2451:
2416:
2298:
805:and modern tortoises of the family
669:possesses a single opening for the
597:
491:, Woodward created the combination
459:for a skull he claimed his brother
13:
2492:Journal of Vertebrate Paleontology
2210:
2160:
2131:
2075:
2042:
2003:
1988:
1982:10.1111/j.1469-7998.1901.tb08537.x
1915:Journal of Vertebrate Paleontology
1860:
1801:, forms an extension like that of
1764:Comparison between the neotype of
914:and the physically much different
652:
14:
4376:
4360:Taxa named by Florentino Ameghino
2539:
4310:Eocene reptiles of South America
4141:
4140:
4101:
4094:
2022:Records of the Australian Museum
1844:
1830:
1809:in later members of the family.
1789:, while no such split occurs in
640:fossils (initially described as
414:In 1898, British paleontologist
386:
377:
120:
34:
2035:10.3853/j.0067-1975.14.1925.844
1604:
1805:suggests that this could be a
647:
1:
4355:Fossil taxa described in 1899
2500:10.1080/02724634.2024.2357315
2437:10.1080/03115518.2016.1224441
1823:
780:
657:The best preserved fossil of
1935:10.1080/02724634.2011.615685
1816:was the presence of a "true
940:
910:islands. The bridge between
900:
543:was not that of a juvenile,
274:Simpson & Williams, 1938
7:
2234:10.1007/978-3-319-00518-8_1
1748:
926:. The later description of
10:
4381:
942:
785:In addition to the skull,
4295:Prehistoric turtle genera
4169:
4115:
4091:
4062:
4009:
3943:
3879:
3807:
3781:
3585:
3464:
3279:
3208:
3184:
3172:
3097:
3046:
3009:
2957:
2914:
2902:
2865:
2757:
2746:
2697:
2682:
2663:
2639:
2621:
2608:
2547:
2371:10.1093/zoolinnean/zlw024
2331:American Museum Novitates
2154:American Museum Novitates
1431:
1413:
1396:
1389:
1371:
1354:
1347:
1329:
1312:
1305:
1276:
1259:
1252:
1245:
1219:
1202:
1195:
1188:
1171:
1164:
1148:
1141:
1080:
1063:
1046:
1029:
1022:
1005:
998:
981:
974:
957:
950:
424:Francisco Pascasio Moreno
259:
252:
235:
230:
117:Scientific classification
115:
110:
101:
23:
1417:Gaffneylania auricularis
1375:Gaffneylania auricularis
1333:Gaffneylania auricularis
396:The research history of
16:Extinct genus of turtles
2314:Gaffney, E. S. (1992).
1961:Woodward, A.S. (1901).
1688:and other meiolaniids.
918:is somewhat bridged by
855:The carapace length of
687:. Above the nares, the
2472:10.33256/hj29.4.252263
2460:Herpetological Journal
2182:Gaffney, E.S. (1996).
2145:Simpson, G.G. (1937).
1999:. La Plata, Argentina.
1761:
1709:
1400:Warkalania carinaminor
1280:Kallokibotion bajazidi
1263:Otwayemys cunicularius
1206:Mongolochelys efremovi
1009:Warkalania carinaminor
823:
734:
563:
520:George Gaylord Simpson
349:Remains attributed to
4350:Golfo San Jorge Basin
4262:Paleobiology Database
1756:
1707:
1700:Function of the horns
1223:Peligrochelys walshae
1033:Meiolania brevicollis
815:
729:
594:in this publication.
561:
537:Crossochelys corniger
481:Othniel Charles Marsh
416:Arthur Smith Woodward
271:Crossochelys corniger
4345:Fossils of Argentina
2188:Bulletin of the AMNH
2107:10.3374/014.056.0102
2065:Bulletin of the AMNH
1681:low frequency sounds
1610:Lifestyle and senses
889:, the tail rings of
707:is dominated by the
575:is simply a younger
551:a junior synonym of
499:. Despite declaring
465:Guaranitic Formation
440:, a turtle named by
308:intraspecific combat
4365:Sarmiento Formation
4340:Paleogene Argentina
3534:Protoxinjiangchelys
2089:Sterli, J. (2015).
1927:2011JVPal..31.1210S
1852:Paleontology portal
1732:on the holotype of
1722:pachycephalosaurids
1668:semicircular canals
1435:Meiolania platyceps
1175:Chubutemys copelloi
1050:Meiolania platyceps
885:, and more like in
803:Meiolania platyceps
533:Sarmiento Formation
515:in his prior work.
477:Edward Drinker Cope
453:Florentino Ameghino
437:Meiolania platyceps
422:, who worked under
402:Florentino Ameghino
359:Florentino Ameghino
111:Skull and mandible
2395:Gaffney, Eugene S.
2320:, a new name for "
1906:Niolamia argentina
1807:paedomorphic trait
1762:
1710:
1316:Niolamia argentina
961:Niolamia argentina
826:While no complete
824:
735:
732:Niolamia argentina
564:
493:Miolania argentina
463:discovered in the
457:Niolamia argentina
369:History and naming
263:Miolania argentina
4277:
4276:
4249:Open Tree of Life
4163:Taxon identifiers
4154:
4153:
4111:
4110:
4089:
4088:
4085:
4084:
4058:
4057:
3469:Xinjiangchelyidae
3460:
3459:
3005:
3004:
2870:Australochelyidae
2678:
2677:
2674:
2673:
2659:
2658:
2281:10.1111/zoj.12252
2243:978-3-319-00517-1
1818:temporal fenestra
1528:paleobiogeography
1519:Paleobiogeography
1516:
1515:
1509:
1508:
1500:
1499:
1491:
1490:
1482:
1481:
1473:
1472:
1464:
1463:
1455:
1454:
1446:
1445:
1291:
1290:
1234:
1233:
1150:Crown Testudines
1131:
1130:
1122:
1121:
1113:
1112:
1104:
1103:
1095:
1094:
1084:Meiolania mackayi
721:softshell turtles
630:Lake Colhué Huapí
545:Eugene S. Gaffney
281:
280:
275:
267:
246:
226:
4372:
4270:
4269:
4257:
4256:
4244:
4243:
4231:
4230:
4218:
4217:
4205:
4204:
4203:
4190:
4189:
4188:
4158:
4157:
4144:
4143:
4106:
4105:
4099:
4098:
3805:
3804:
3801:
3800:
3786:Thalassochelydia
3784:
3588:
3467:
3182:
3181:
3175:
3100:
3049:
3014:Sichuanchelyidae
3012:
2912:
2911:
2905:
2868:
2853:
2843:
2833:
2823:
2813:
2803:
2793:
2783:
2773:
2763:
2755:
2754:
2734:
2724:
2713:
2703:
2695:
2694:
2684:
2683:
2619:
2618:
2610:
2609:
2603:
2602:
2549:
2548:
2534:
2527:
2520:
2511:
2510:
2504:
2503:
2487:
2476:
2475:
2455:
2449:
2448:
2420:
2414:
2413:
2403:
2391:
2385:
2384:
2382:
2350:
2335:
2334:
2328:
2322:Meiolania" oweni
2311:
2296:
2295:
2293:
2283:
2259:
2248:
2247:
2221:
2208:
2207:
2179:
2158:
2157:
2151:
2142:
2129:
2128:
2118:
2086:
2073:
2072:
2062:
2053:
2040:
2039:
2037:
2010:
2001:
2000:
1992:
1986:
1985:
1967:
1958:
1947:
1946:
1921:(6): 1210–1229.
1912:
1899:
1854:
1849:
1848:
1847:
1840:
1838:Argentina portal
1835:
1834:
1833:
1640:thermoregulation
1634:based on extant
1392:
1391:
1350:
1349:
1308:
1307:
1255:
1254:
1248:
1247:
1198:
1197:
1191:
1190:
1167:
1166:
1144:
1143:
1137:
1136:
1025:
1024:
1001:
1000:
977:
976:
953:
952:
946:
945:
941:
791:shoulder girdles
717:pig-nosed turtle
598:Age and locality
455:coined the name
406:Francisco Moreno
390:
381:
273:
265:
244:
221:
214:
201:
125:
124:
106:
96:
33:
29:Temporal range:
21:
20:
4380:
4379:
4375:
4374:
4373:
4371:
4370:
4369:
4280:
4279:
4278:
4273:
4265:
4260:
4252:
4247:
4239:
4234:
4226:
4221:
4213:
4208:
4199:
4198:
4193:
4184:
4183:
4178:
4165:
4155:
4150:
4107:
4100:
4093:
4081:
4054:
4005:
3946:Plesiochelyidae
3939:
3875:
3790:
3789:
3788:
3777:
3744:Tienfucheloides
3660:Hongkongochelys
3593:
3581:
3456:
3275:
3210:Pleurosternidae
3204:
3198:Helochelydridae
3168:
3093:
3051:Helochelydridae
3042:
3001:
2953:
2898:
2861:
2751:
2742:
2706:Eorhynchochelys
2691:
2670:
2655:
2635:
2604:
2554:
2553:
2543:
2538:
2508:
2507:
2488:
2479:
2456:
2452:
2421:
2417:
2401:
2392:
2388:
2351:
2338:
2326:
2312:
2299:
2260:
2251:
2244:
2222:
2211:
2180:
2161:
2149:
2143:
2132:
2087:
2076:
2060:
2054:
2043:
2011:
2004:
1993:
1989:
1965:
1959:
1950:
1910:
1900:
1861:
1850:
1845:
1843:
1836:
1831:
1829:
1826:
1751:
1702:
1676:plesiochelydids
1612:
1607:
1521:
1510:
1501:
1492:
1483:
1474:
1465:
1456:
1447:
1292:
1235:
1132:
1123:
1114:
1105:
1096:
903:
821:Heinrich Harder
783:
757:tympanic cavity
749:squamosal bones
655:
653:Skull and horns
650:
621:La Plata Museum
609:Chubut Province
600:
412:
411:
410:
409:
393:
392:
391:
383:
382:
371:
220:
212:
199:
119:
97:
95:
94:
89:
84:
79:
74:
69:
64:
59:
54:
49:
44:
39:
31:
30:
27:
17:
12:
11:
5:
4378:
4368:
4367:
4362:
4357:
4352:
4347:
4342:
4337:
4332:
4330:Bartonian life
4327:
4322:
4317:
4312:
4307:
4302:
4300:Eocene turtles
4297:
4292:
4290:Meiolaniformes
4275:
4274:
4272:
4271:
4258:
4245:
4232:
4219:
4206:
4191:
4175:
4173:
4167:
4166:
4152:
4151:
4149:
4148:
4138:
4133:
4128:
4123:
4121:Angolachelonia
4116:
4113:
4112:
4109:
4108:
4092:
4090:
4087:
4086:
4083:
4082:
4080:
4079:
4078:
4077:
4068:
4066:
4060:
4059:
4056:
4055:
4053:
4052:
4044:
4037:
4030:
4023:
4015:
4013:
4007:
4006:
4004:
4003:
3996:
3989:
3982:
3974:
3966:
3959:
3956:Craspedochelys
3951:
3949:
3941:
3940:
3938:
3937:
3930:
3923:
3916:
3909:
3902:
3895:
3887:
3885:
3877:
3876:
3874:
3873:
3865:
3858:
3855:Pelobatochelys
3851:
3844:
3837:
3834:Hispaniachelys
3830:
3823:
3816:
3808:
3802:
3779:
3778:
3776:
3775:
3768:
3761:
3754:
3747:
3740:
3733:
3726:
3719:
3712:
3709:Manchurochelys
3705:
3698:
3691:
3684:
3677:
3670:
3663:
3656:
3649:
3642:
3635:
3628:
3621:
3614:
3607:
3599:
3597:
3583:
3582:
3580:
3579:
3572:
3569:Xinjiangchelys
3565:
3558:
3551:
3544:
3537:
3530:
3523:
3516:
3509:
3502:
3495:
3488:
3481:
3473:
3471:
3462:
3461:
3458:
3457:
3455:
3454:
3447:
3440:
3433:
3426:
3419:
3412:
3405:
3398:
3391:
3384:
3377:
3370:
3363:
3356:
3349:
3342:
3335:
3328:
3321:
3314:
3307:
3300:
3293:
3285:
3283:
3277:
3276:
3274:
3273:
3266:
3259:
3251:
3243:
3236:
3229:
3222:
3214:
3212:
3206:
3205:
3203:
3202:
3194:
3185:
3179:
3177:Paracryptodira
3170:
3169:
3167:
3166:
3159:
3151:
3143:
3136:
3128:
3121:
3114:
3106:
3104:
3095:
3094:
3092:
3091:
3084:
3077:
3070:
3063:
3055:
3053:
3044:
3043:
3041:
3040:
3033:
3026:
3023:Laurasichersis
3018:
3016:
3007:
3006:
3003:
3002:
3000:
2999:
2992:
2985:
2978:
2971:
2963:
2961:
2955:
2954:
2952:
2951:
2948:Trapalcochelys
2944:
2937:
2930:
2923:
2915:
2909:
2907:Meiolaniformes
2900:
2899:
2897:
2896:
2889:
2882:
2879:Australochelys
2874:
2872:
2863:
2862:
2860:
2859:
2849:
2846:Proterochersis
2839:
2829:
2819:
2809:
2799:
2789:
2779:
2769:
2758:
2752:
2747:
2744:
2743:
2741:
2740:
2730:
2720:
2709:
2698:
2692:
2687:
2680:
2679:
2676:
2675:
2672:
2671:
2664:
2661:
2660:
2657:
2656:
2654:
2653:
2652:
2651:
2645:
2643:
2637:
2636:
2634:
2633:
2632:
2631:
2622:
2616:
2606:
2605:
2601:
2600:
2591:
2582:
2573:
2567:
2561:
2545:
2544:
2537:
2536:
2529:
2522:
2514:
2506:
2505:
2477:
2466:(4): 252–263.
2450:
2431:(2): 231–239.
2415:
2386:
2365:(4): 930–950.
2336:
2297:
2274:(3): 519–548.
2249:
2242:
2209:
2159:
2130:
2074:
2041:
2028:(4): 223–242.
2002:
1987:
1976:(2): 169–184.
1948:
1858:
1857:
1856:
1855:
1841:
1825:
1822:
1750:
1747:
1701:
1698:
1611:
1608:
1606:
1603:
1591:Meiolaniformes
1520:
1517:
1514:
1513:
1511:
1507:
1506:
1503:
1502:
1498:
1497:
1494:
1493:
1489:
1488:
1485:
1484:
1480:
1479:
1476:
1475:
1471:
1470:
1467:
1466:
1462:
1461:
1458:
1457:
1453:
1452:
1449:
1448:
1444:
1443:
1440:
1439:
1430:
1427:
1426:
1423:
1422:
1412:
1409:
1408:
1405:
1404:
1395:
1390:
1388:
1385:
1384:
1381:
1380:
1370:
1367:
1366:
1363:
1362:
1358:Ninjemys oweni
1353:
1348:
1346:
1343:
1342:
1339:
1338:
1328:
1325:
1324:
1321:
1320:
1311:
1306:
1304:
1298:
1297:
1294:
1293:
1289:
1288:
1285:
1284:
1275:
1272:
1271:
1268:
1267:
1258:
1253:
1251:
1246:
1244:
1241:
1240:
1237:
1236:
1232:
1231:
1228:
1227:
1218:
1215:
1214:
1211:
1210:
1201:
1196:
1194:
1189:
1187:
1184:
1183:
1180:
1179:
1170:
1165:
1163:
1161:Meiolaniformes
1157:
1156:
1153:
1152:
1147:
1142:
1140:
1135:
1133:
1129:
1128:
1125:
1124:
1120:
1119:
1116:
1115:
1111:
1110:
1107:
1106:
1102:
1101:
1098:
1097:
1093:
1092:
1089:
1088:
1079:
1076:
1075:
1072:
1071:
1062:
1059:
1058:
1055:
1054:
1045:
1042:
1041:
1038:
1037:
1028:
1023:
1021:
1018:
1017:
1014:
1013:
1004:
999:
997:
994:
993:
990:
989:
985:Ninjemys oweni
980:
975:
973:
970:
969:
966:
965:
956:
951:
949:
944:
902:
899:
782:
779:
654:
651:
649:
646:
599:
596:
505:junior synonym
395:
394:
385:
384:
376:
375:
374:
373:
372:
370:
367:
297:South American
279:
278:
277:
276:
268:
266:Woodward, 1901
257:
256:
250:
249:
248:
247:
245:Ameghino, 1899
233:
232:
228:
227:
210:
206:
205:
197:
193:
192:
187:
180:
179:
174:
167:
166:
161:
157:
156:
151:
147:
146:
141:
137:
136:
131:
127:
126:
113:
112:
108:
107:
99:
98:
90:
85:
80:
75:
70:
65:
60:
55:
50:
45:
40:
35:
28:
15:
9:
6:
4:
3:
2:
4377:
4366:
4363:
4361:
4358:
4356:
4353:
4351:
4348:
4346:
4343:
4341:
4338:
4336:
4335:Lutetian life
4333:
4331:
4328:
4326:
4323:
4321:
4318:
4316:
4313:
4311:
4308:
4306:
4303:
4301:
4298:
4296:
4293:
4291:
4288:
4287:
4285:
4268:
4263:
4259:
4255:
4250:
4246:
4242:
4237:
4233:
4229:
4224:
4220:
4216:
4211:
4207:
4202:
4196:
4192:
4187:
4181:
4177:
4176:
4174:
4172:
4168:
4164:
4159:
4147:
4139:
4137:
4134:
4132:
4129:
4127:
4124:
4122:
4118:
4117:
4114:
4104:
4097:
4076:
4072:
4071:
4070:
4069:
4067:
4065:
4061:
4050:
4049:
4045:
4043:
4042:
4038:
4036:
4035:
4031:
4029:
4028:
4027:Brachyopsemys
4024:
4022:
4021:
4017:
4016:
4014:
4012:
4008:
4002:
4001:
3997:
3995:
3994:
3990:
3988:
3987:
3983:
3980:
3979:
3978:Jurassichelon
3975:
3972:
3971:
3967:
3965:
3964:
3960:
3958:
3957:
3953:
3952:
3950:
3947:
3942:
3936:
3935:
3931:
3929:
3928:
3924:
3922:
3921:
3917:
3915:
3914:
3910:
3908:
3907:
3903:
3901:
3900:
3896:
3894:
3893:
3889:
3888:
3886:
3883:
3882:Eurysternidae
3878:
3871:
3870:
3869:Protostegidae
3866:
3864:
3863:
3859:
3857:
3856:
3852:
3850:
3849:
3845:
3843:
3842:
3838:
3836:
3835:
3831:
3829:
3828:
3824:
3822:
3821:
3817:
3815:
3814:
3810:
3809:
3806:
3803:
3798:
3794:
3787:
3780:
3774:
3773:
3769:
3767:
3766:
3762:
3760:
3759:
3755:
3753:
3752:
3748:
3746:
3745:
3741:
3739:
3738:
3734:
3732:
3731:
3727:
3725:
3724:
3720:
3718:
3717:
3713:
3711:
3710:
3706:
3704:
3703:
3699:
3697:
3696:
3692:
3690:
3689:
3685:
3683:
3682:
3678:
3676:
3675:
3671:
3669:
3668:
3664:
3662:
3661:
3657:
3655:
3654:
3650:
3648:
3647:
3643:
3641:
3640:
3636:
3634:
3633:
3632:Changmachelys
3629:
3627:
3626:
3622:
3620:
3619:
3615:
3613:
3612:
3608:
3606:
3605:
3601:
3600:
3598:
3596:
3595:Macrobaenidae
3591:
3584:
3578:
3577:
3573:
3571:
3570:
3566:
3564:
3563:
3559:
3557:
3556:
3552:
3550:
3549:
3545:
3543:
3542:
3538:
3536:
3535:
3531:
3529:
3528:
3524:
3522:
3521:
3517:
3515:
3514:
3510:
3508:
3507:
3503:
3501:
3500:
3499:Chengyuchelys
3496:
3494:
3493:
3489:
3487:
3486:
3482:
3480:
3479:
3475:
3474:
3472:
3470:
3463:
3453:
3452:
3451:Trinitichelys
3448:
3446:
3445:
3441:
3439:
3438:
3434:
3432:
3431:
3427:
3425:
3424:
3420:
3418:
3417:
3413:
3411:
3410:
3406:
3404:
3403:
3399:
3397:
3396:
3392:
3390:
3389:
3385:
3383:
3382:
3378:
3376:
3375:
3371:
3369:
3368:
3364:
3362:
3361:
3360:Gehennachelys
3357:
3355:
3354:
3350:
3348:
3347:
3343:
3341:
3340:
3336:
3334:
3333:
3329:
3327:
3326:
3322:
3320:
3319:
3315:
3313:
3312:
3308:
3306:
3305:
3301:
3299:
3298:
3294:
3292:
3291:
3287:
3286:
3284:
3282:
3278:
3272:
3271:
3267:
3265:
3264:
3260:
3257:
3256:
3252:
3249:
3248:
3244:
3242:
3241:
3240:Pleurosternon
3237:
3235:
3234:
3230:
3228:
3227:
3226:Dorsetochelys
3223:
3221:
3220:
3216:
3215:
3213:
3211:
3207:
3200:
3199:
3195:
3192:
3191:
3190:Compsemydidae
3187:
3186:
3183:
3180:
3178:
3171:
3165:
3164:
3160:
3157:
3156:
3152:
3149:
3148:
3144:
3142:
3141:
3137:
3134:
3133:
3132:Kallokibotion
3129:
3127:
3126:
3122:
3120:
3119:
3118:Calissounemys
3115:
3113:
3112:
3108:
3107:
3105:
3103:
3102:Compsemydidae
3096:
3090:
3089:
3085:
3083:
3082:
3078:
3076:
3075:
3071:
3069:
3068:
3064:
3062:
3061:
3057:
3056:
3054:
3052:
3045:
3039:
3038:
3037:Sichuanchelys
3034:
3032:
3031:
3030:Mongolochelys
3027:
3025:
3024:
3020:
3019:
3017:
3015:
3008:
2998:
2997:
2993:
2991:
2990:
2986:
2984:
2983:
2979:
2977:
2976:
2972:
2970:
2969:
2965:
2964:
2962:
2960:
2956:
2950:
2949:
2945:
2943:
2942:
2941:Peligrochelys
2938:
2936:
2935:
2934:Patagoniaemys
2931:
2929:
2928:
2924:
2922:
2921:
2917:
2916:
2913:
2910:
2908:
2901:
2895:
2894:
2890:
2888:
2887:
2886:Palaeochersis
2883:
2881:
2880:
2876:
2875:
2873:
2871:
2864:
2858:
2857:
2850:
2848:
2847:
2840:
2838:
2837:
2836:Proganochelys
2830:
2828:
2827:
2826:Kayentachelys
2820:
2818:
2817:
2816:Kallokibotion
2810:
2808:
2807:
2800:
2798:
2797:
2796:Heckerochelys
2790:
2788:
2787:
2780:
2778:
2777:
2770:
2768:
2767:
2760:
2759:
2756:
2753:
2750:
2745:
2739:
2738:
2731:
2729:
2728:
2721:
2718:
2717:
2710:
2708:
2707:
2700:
2699:
2696:
2693:
2690:
2689:Pantestudines
2685:
2681:
2669:
2668:
2667:Pantestudines
2662:
2649:
2648:
2647:
2646:
2644:
2642:
2641:Pantestudines
2638:
2630:
2626:
2625:
2624:
2623:
2620:
2617:
2615:
2611:
2607:
2599:
2598:Archelosauria
2595:
2592:
2590:
2586:
2583:
2581:
2577:
2574:
2572:
2568:
2566:
2562:
2560:
2556:
2555:
2550:
2546:
2542:
2541:Pantestudines
2535:
2530:
2528:
2523:
2521:
2516:
2515:
2512:
2501:
2497:
2493:
2486:
2484:
2482:
2473:
2469:
2465:
2461:
2454:
2446:
2442:
2438:
2434:
2430:
2426:
2419:
2411:
2407:
2400:
2396:
2390:
2381:
2376:
2372:
2368:
2364:
2360:
2356:
2349:
2347:
2345:
2343:
2341:
2333:(3049): 1–10.
2332:
2325:
2323:
2319:
2310:
2308:
2306:
2304:
2302:
2292:
2287:
2282:
2277:
2273:
2269:
2265:
2258:
2256:
2254:
2245:
2239:
2235:
2231:
2227:
2220:
2218:
2216:
2214:
2205:
2201:
2197:
2193:
2189:
2185:
2178:
2176:
2174:
2172:
2170:
2168:
2166:
2164:
2155:
2148:
2141:
2139:
2137:
2135:
2126:
2122:
2117:
2112:
2108:
2104:
2100:
2096:
2092:
2085:
2083:
2081:
2079:
2070:
2066:
2059:
2052:
2050:
2048:
2046:
2036:
2031:
2027:
2023:
2019:
2015:
2009:
2007:
1998:
1991:
1983:
1979:
1975:
1971:
1964:
1957:
1955:
1953:
1944:
1940:
1936:
1932:
1928:
1924:
1920:
1916:
1909:
1907:
1898:
1896:
1894:
1892:
1890:
1888:
1886:
1884:
1882:
1880:
1878:
1876:
1874:
1872:
1870:
1868:
1866:
1864:
1859:
1853:
1842:
1839:
1828:
1821:
1819:
1815:
1810:
1808:
1804:
1800:
1796:
1792:
1788:
1784:
1779:
1775:
1771:
1767:
1760:
1755:
1746:
1744:
1738:
1735:
1731:
1727:
1723:
1719:
1714:
1706:
1697:
1694:
1689:
1687:
1682:
1677:
1673:
1669:
1665:
1661:
1656:
1654:
1648:
1646:
1645:ornithischian
1641:
1637:
1633:
1629:
1625:
1621:
1617:
1602:
1600:
1596:
1595:Drake Passage
1592:
1588:
1584:
1580:
1576:
1572:
1568:
1564:
1560:
1559:South America
1555:
1553:
1549:
1545:
1541:
1537:
1533:
1529:
1525:
1505:
1504:
1496:
1495:
1487:
1486:
1478:
1477:
1469:
1468:
1460:
1459:
1451:
1450:
1442:
1441:
1438:
1437:
1436:
1429:
1428:
1425:
1424:
1421:
1419:
1418:
1411:
1410:
1407:
1406:
1403:
1402:
1401:
1394:
1393:
1387:
1386:
1383:
1382:
1379:
1377:
1376:
1369:
1368:
1365:
1364:
1361:
1360:
1359:
1352:
1351:
1345:
1344:
1341:
1340:
1337:
1335:
1334:
1327:
1326:
1323:
1322:
1319:
1318:
1317:
1310:
1309:
1303:
1300:
1299:
1296:
1295:
1287:
1286:
1283:
1282:
1281:
1274:
1273:
1270:
1269:
1266:
1265:
1264:
1257:
1256:
1250:
1249:
1243:
1242:
1239:
1238:
1230:
1229:
1226:
1225:
1224:
1217:
1216:
1213:
1212:
1209:
1208:
1207:
1200:
1199:
1193:
1192:
1186:
1185:
1182:
1181:
1178:
1177:
1176:
1169:
1168:
1162:
1159:
1158:
1155:
1154:
1151:
1146:
1145:
1139:
1138:
1127:
1126:
1118:
1117:
1109:
1108:
1100:
1099:
1091:
1090:
1087:
1086:
1085:
1078:
1077:
1074:
1073:
1070:
1069:
1068:
1067:Meiolania sp.
1061:
1060:
1057:
1056:
1053:
1052:
1051:
1044:
1043:
1040:
1039:
1036:
1035:
1034:
1027:
1026:
1020:
1019:
1016:
1015:
1012:
1011:
1010:
1003:
1002:
996:
995:
992:
991:
988:
987:
986:
979:
978:
972:
971:
968:
967:
964:
963:
962:
955:
954:
948:
947:
939:
937:
933:
929:
925:
921:
917:
913:
908:
898:
896:
892:
888:
884:
879:
874:
872:
868:
867:
866:Proganochelys
862:
858:
853:
851:
846:
841:
837:
833:
829:
822:
818:
814:
810:
808:
804:
800:
796:
792:
788:
778:
775:
771:
767:
762:
758:
754:
750:
745:
740:
733:
730:Tail ring of
728:
724:
722:
718:
714:
710:
706:
702:
698:
694:
690:
686:
682:
678:
677:
672:
668:
664:
660:
645:
643:
639:
635:
631:
627:
622:
618:
614:
610:
605:
595:
593:
589:
585:
580:
578:
574:
569:
560:
556:
554:
550:
546:
542:
538:
534:
529:
525:
521:
516:
514:
510:
506:
502:
498:
494:
490:
484:
482:
478:
474:
470:
466:
462:
458:
454:
449:
447:
443:
439:
438:
433:
429:
425:
421:
420:Santiago Roth
417:
407:
403:
399:
389:
380:
366:
364:
360:
356:
352:
347:
345:
342:, this makes
341:
337:
333:
329:
328:
323:
319:
317:
313:
309:
305:
301:
298:
294:
291:
287:
286:
272:
269:
264:
261:
260:
258:
255:
251:
243:
242:
237:
236:
234:
229:
224:
219:
218:
211:
208:
207:
204:
198:
195:
194:
191:
188:
185:
182:
181:
178:
177:Pantestudines
175:
172:
169:
168:
165:
162:
159:
158:
155:
152:
149:
148:
145:
142:
139:
138:
135:
132:
129:
128:
123:
118:
114:
109:
105:
100:
93:
88:
83:
78:
73:
68:
63:
58:
53:
48:
43:
38:
26:
22:
19:
4170:
4136:Perichelydia
4131:Mesochelydia
4046:
4039:
4032:
4025:
4020:Angolachelys
4018:
3998:
3993:Portlandemys
3991:
3986:Plesiochelys
3984:
3976:
3970:Hylaeochelys
3968:
3961:
3954:
3932:
3927:Palaeomedusa
3925:
3918:
3911:
3904:
3897:
3890:
3867:
3860:
3853:
3846:
3839:
3832:
3825:
3818:
3811:
3796:
3792:
3770:
3763:
3756:
3749:
3742:
3735:
3728:
3721:
3714:
3707:
3700:
3693:
3686:
3679:
3674:Jeholochelys
3672:
3665:
3658:
3653:Galvechelone
3651:
3644:
3637:
3630:
3625:Aurorachelys
3623:
3616:
3611:Appalachemys
3609:
3602:
3574:
3567:
3560:
3555:Tienfuchelys
3553:
3546:
3539:
3532:
3527:Phunoichelys
3525:
3518:
3511:
3504:
3497:
3492:Camerochelys
3490:
3485:Brodiechelys
3483:
3476:
3449:
3442:
3437:Stygiochelys
3435:
3428:
3421:
3414:
3407:
3400:
3393:
3386:
3379:
3372:
3365:
3358:
3351:
3344:
3337:
3330:
3323:
3316:
3309:
3302:
3297:Arvinachelys
3295:
3288:
3268:
3261:
3253:
3245:
3238:
3231:
3224:
3217:
3196:
3188:
3161:
3153:
3145:
3138:
3130:
3123:
3116:
3109:
3086:
3079:
3072:
3067:Helochelydra
3065:
3060:Aragochersis
3058:
3035:
3028:
3021:
2994:
2988:
2987:
2980:
2973:
2968:Gaffneylania
2966:
2959:Meiolaniidae
2946:
2939:
2932:
2925:
2918:
2891:
2884:
2877:
2854:
2844:
2834:
2824:
2814:
2804:
2794:
2786:Eileanchelys
2784:
2776:Condorchelys
2774:
2766:Chinlechelys
2764:
2735:
2727:Odontochelys
2725:
2716:Eunotosaurus
2714:
2704:
2665:
2593:
2584:
2575:
2491:
2463:
2459:
2453:
2428:
2424:
2418:
2409:
2405:
2389:
2362:
2358:
2330:
2321:
2317:
2271:
2267:
2225:
2187:
2153:
2101:(1): 21–45.
2098:
2094:
2068:
2064:
2025:
2021:
2014:Anderson, C.
1996:
1990:
1973:
1969:
1918:
1914:
1905:
1814:Crossochelys
1813:
1811:
1803:Crossochelys
1802:
1798:
1794:
1790:
1787:Crossochelys
1786:
1783:Crossochelys
1782:
1777:
1773:
1770:Crossochelys
1769:
1765:
1763:
1758:
1757:Ontogeny of
1742:
1739:
1733:
1725:
1717:
1715:
1711:
1692:
1690:
1685:
1663:
1657:
1649:
1627:
1624:Gaffneylania
1623:
1619:
1615:
1613:
1605:Paleobiology
1598:
1556:
1552:Gaffneylania
1551:
1547:
1543:
1539:
1535:
1531:
1523:
1522:
1433:
1432:
1415:
1414:
1398:
1397:
1373:
1372:
1356:
1355:
1331:
1330:
1315:
1314:
1313:
1302:Meiolaniidae
1278:
1277:
1261:
1260:
1221:
1220:
1204:
1203:
1173:
1172:
1149:
1082:
1081:
1065:
1064:
1048:
1047:
1031:
1030:
1007:
1006:
983:
982:
960:
959:
958:
935:
928:Gaffneylania
927:
923:
919:
915:
911:
906:
904:
894:
890:
886:
882:
877:
875:
870:
864:
860:
856:
854:
849:
844:
839:
831:
825:
816:
807:Testudinidae
802:
786:
784:
774:rhamphotheca
769:
765:
760:
752:
743:
738:
736:
731:
709:nasal cavity
704:
696:
680:
674:
666:
658:
656:
642:Crossochelys
641:
637:
626:Lake Musters
603:
601:
583:
581:
576:
573:Crossochelys
572:
567:
565:
552:
549:Crossochelys
548:
541:Crossochelys
540:
536:
527:
517:
512:
508:
500:
496:
492:
488:
485:
456:
450:
442:Richard Owen
435:
413:
397:
354:
350:
348:
343:
327:Gaffneylania
325:
321:
320:
315:
303:
284:
283:
282:
270:
262:
241:N. argentina
240:
239:
216:
215:
203:Meiolaniidae
183:
170:
24:
18:
4325:Casamayoran
4315:Divisaderan
4195:Wikispecies
4034:Leyvachelys
4011:Sandownidae
3963:Globochelus
3906:Eurysternum
3862:Thalassemys
3841:Neusticemys
3639:Dracochelys
3590:Sinemydidae
3576:Yanduchelys
3548:Siamochelys
3541:Shartegemys
3513:Kalasinemys
3506:Jastmelchyi
3409:Plesiobaena
3395:Palatobaena
3388:Neurankylus
3353:Gamerabaena
3332:Denazinemys
3290:Arundelemys
3140:Peltochelys
3111:Berruchelus
3081:Naomichelys
2749:Testudinata
2737:Pappochelys
2412:(1): 35–48.
2380:11336/63435
2291:11336/41594
2116:11336/21194
1730:pathologies
1579:New Zealand
701:endocranium
693:premaxillae
648:Description
404:(left) and
340:Australasia
190:Testudinata
4284:Categories
4126:Ankylopoda
4119:See also:
4075:Testudines
4064:Testudines
4000:Tropidemys
3934:Parachelys
3920:Idiochelys
3913:Hydropelta
3899:Chelonides
3758:Xiaochelys
3723:Osteopygis
3702:Macrobaena
3695:Liaochelys
3688:Kirgizemys
3681:Judithemys
3618:Asiachelys
3604:Anatolemys
3562:Undjulemys
3520:Larachelus
3423:Saxochelys
3416:Protobaena
3325:Chisternon
3318:Cedrobaena
3247:Riodevemys
3219:Dinochelys
3147:Riodevemys
3074:Helochelys
2996:Warkalania
2920:Chubutemys
2893:Waluchelys
2856:Spoochelys
2806:Indochelys
2650:see below↓
2629:Sauropsida
2614:Sauropsida
2571:Sauropsida
1824:References
1799:Warkalania
1672:geoemydids
1583:New Guinea
1563:Antarctica
1536:Warkalania
781:Postcrania
689:nasal bone
617:Rio Chubut
613:Rio Sehuen
336:Quaternary
300:meiolaniid
4320:Mustersan
4048:Solnhofia
4041:Sandownia
3892:Achelonia
3827:Euryaspis
3772:Yumenemys
3716:Ordosemys
3667:Hoyasemys
3444:Thescelus
3430:Scabremys
3381:Lakotemys
3367:Goleremys
3255:Selenemys
3155:Selenemys
3125:Compsemys
2975:Meiolania
2927:Otwayemys
2557:Kingdom:
2445:131795055
2204:0003-0090
2196:2246/1670
1743:Meiolania
1737:in life.
1660:inner ear
1653:courtship
1628:Meiolania
1616:Niolamia'
1571:Australia
1544:Meiolania
916:Meiolania
901:Phylogeny
895:Meiolania
883:Meiolania
861:Meiolania
850:Meiolania
836:taphonomy
713:mata mata
681:Meiolania
663:ankylosed
513:Meiolania
497:Meiolania
473:Bone Wars
446:Australia
428:Argentina
312:courtship
140:Kingdom:
134:Eukaryota
4201:Niolamia
4180:Wikidata
4171:Niolamia
4146:Category
3848:Owadowia
3402:Peckemys
3281:Baenidae
3233:Glyptops
3163:Tongemys
2989:Niolamia
2982:Ninjemys
2580:Diapsida
2565:Chordata
2563:Phylum:
2559:Animalia
2318:Ninjemys
2125:83799914
2016:(1925).
1943:83503956
1795:Ninjemys
1791:Niolamia
1778:Niolamia
1774:Niolamia
1766:Niolamia
1759:Niolamia
1749:Ontogeny
1734:Niolamia
1726:Niolamia
1718:Niolamia
1693:Niolamia
1686:Niolamia
1664:Niolamia
1620:Niolamia
1599:Niolamia
1587:Gondwana
1575:Tasmania
1565:, south
1548:Niolamia
1540:Ninjemys
1532:Niolamia
1524:Niolamia
936:Niolamia
932:polytomy
924:Niolamia
920:Ninjemys
912:Niolamia
907:Niolamia
891:Niolamia
887:Ninjemys
878:Niolamia
871:Niolamia
857:Niolamia
845:Niolamia
840:Niolamia
832:Niolamia
817:Niolamia
799:acromion
787:Niolamia
770:Niolamia
766:Niolamia
761:Niolamia
753:Ninjemys
744:Niolamia
739:Niolamia
705:Niolamia
697:Ninjemys
676:Ninjemys
667:Niolamia
659:Niolamia
638:Niolamia
604:Niolamia
592:holotype
584:Niolamia
577:Niolamia
568:Niolamia
553:Niolamia
528:Niolamia
509:Niolamia
501:Niolamia
489:Niolamia
475:between
469:holotype
408:(right).
398:Niolamia
355:Niolamia
351:Niolamia
344:Niolamia
322:Niolamia
316:Niolamia
304:Niolamia
285:Niolamia
254:Synonyms
231:Species
223:Ameghino
217:Niolamia
196:Family:
164:Reptilia
154:Chordata
150:Phylum:
144:Animalia
130:Domain:
25:Niolamia
4254:4132521
4241:1193798
4228:4818917
4215:4522670
4186:Q148454
3820:Cyrtura
3799:, 2021)
3765:Yakemys
3737:Sinemys
3646:Gallica
3478:Annemys
3374:Hayemys
3346:Eubaena
3311:Boremys
3263:Toremys
3088:Solemys
2569:Class:
2190:(229).
1923:Bibcode
1636:lizards
1632:deserts
795:scapula
590:as the
588:neotype
524:neotype
363:brother
332:Neogene
310:during
290:extinct
209:Genus:
160:Class:
3797:et al.
3795:Joyce
3751:Wuguia
3730:Oxemys
3270:Uluops
2589:Sauria
2443:
2240:
2202:
2156:(927).
2123:
1941:
1567:Africa
699:. The
685:septum
461:Carlos
432:London
288:is an
225:, 1899
32:Eocene
4267:37645
4236:IRMNG
3813:Aplax
3793:sensu
3339:Edowa
3304:Baena
2594:Clade
2585:Clade
2576:Clade
2441:S2CID
2402:(PDF)
2327:(PDF)
2150:(PDF)
2121:S2CID
2061:(PDF)
1966:(PDF)
1939:S2CID
1911:(PDF)
934:with
828:shell
671:nares
444:from
293:genus
184:Clade
171:Clade
4223:GBIF
4073:see
2627:see
2238:ISBN
2200:ISSN
2071:(5).
1797:and
1674:and
1658:The
1626:and
1581:and
719:and
679:and
634:tuff
628:and
615:and
479:and
334:and
37:PreꞒ
4210:EoL
2496:doi
2468:doi
2433:doi
2375:hdl
2367:doi
2363:180
2286:hdl
2276:doi
2272:174
2230:doi
2192:hdl
2111:hdl
2103:doi
2030:doi
1978:doi
1931:doi
1662:of
830:of
819:by
703:of
526:of
430:to
338:of
295:of
4286::
4264::
4251::
4238::
4225::
4212::
4197::
4182::
2596::
2587::
2578::
2494:.
2480:^
2464:29
2462:.
2439:.
2429:41
2427:.
2408:.
2404:.
2373:.
2361:.
2357:.
2339:^
2329:.
2300:^
2284:.
2270:.
2266:.
2252:^
2236:.
2212:^
2198:.
2186:.
2162:^
2152:.
2133:^
2119:.
2109:.
2099:56
2097:.
2093:.
2077:^
2069:74
2067:.
2063:.
2044:^
2026:14
2024:.
2020:.
2005:^
1974:70
1972:.
1968:.
1951:^
1937:.
1929:.
1919:31
1917:.
1913:.
1862:^
1622:,
1577:,
1573:,
1569:,
1561:,
1554:.
1542:,
1538:,
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897:.
723:.
715:,
555:.
503:a
186::
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87:Pg
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3880:"
3872:?
3791:(
3783:†
3592:/
3587:†
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3258:?
3250:?
3201:?
3193:?
3174:†
3158:?
3150:?
3135:?
3099:†
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3011:†
2904:†
2867:†
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2842:†
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2802:†
2792:†
2782:†
2772:†
2762:†
2733:†
2723:†
2719:?
2712:†
2702:†
2533:e
2526:t
2519:v
2502:.
2498::
2474:.
2470::
2447:.
2435::
2410:9
2383:.
2377::
2369::
2316:"
2294:.
2288::
2278::
2246:.
2232::
2206:.
2194::
2127:.
2113::
2105::
2038:.
2032::
1984:.
1980::
1945:.
1933::
1925::
238:†
213:†
200:†
92:N
82:K
77:J
72:T
67:P
62:C
57:D
52:S
47:O
42:Ꞓ
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