Meiolaniidae - Knowledge (XXG)

1376:. In meiolaniids, the individual plates that form the skull are highly ankylosed, meaning they are fused with each other to a degree that typically makes it impossible to determine where one element ends and the other begins. Despite the absence of such sutures however, researchers can readily distinguish the different genera and species through the presence of marks left by the overlying scale areas, with are either present through faint grooves or raised ridges. These scale areas, at times also simply referred to as scales or scutes, are largely homologous with one another and can easily be compared. To simplify diagnosis and create a consistent naming scheme, these scale areas are labeled with capital letters, a system already used in a similar form during early research and later refined by Gaffney. Some consistent features of these scales include the presence of paired G and D scales covering the roof of the skull, a singular X scale sitting at the center of these scales which varies in size between basal and derived genera and unpaired Y and Z scales that sit between the eyes and over the nose. 3120:, which in turn suggests that they were not very vocal animals. This matches well with the enhanced sense of smell, which may have been used as a crucial part in communication. Modern turtles possess a variety of different scent glands, including musk glands, cloacal secretions and mental glands. While no glands are known from meiolaniids directly, the strong sense of smell could be an indicator that they frequently used chemical signals. One situation in which this may have come into play would be during courtship. Chemical signals can induce aggressive and combat behavior in modern tortoises, which may respond with a variety of shell-based maneuvers like pushing, ramming, knocking and in the case of meiolaniids the use of their the clubbed tail. The involvement of the horns and frills has at times also been considered for such a function, but a 2024 study on 2869:. It is thought that meiolaniids evolved from meiolaniiforms in the approximate region of where the continents South America, Antarctica and Australia connected prior to the separation of these landmasses in the Late Eocene. This would account for the immense distance that today separates the areas where these turtles have been found. The fossil record of that time period is however scant and little is known about the early history of meiolaniids. It is therefore not certain whether they originated in South America and dispersed towards Australia, dispersed from Australia into South America or even originated in Antarctica and spread from there. The best fossils derive from the middle Eocene of Argentina, where 6361: 2980: 2989: 2888:. Four primary hypothesis have been suggested for this. Some researchers, in particular those in support of an aquatic lifestyle, have proposed that meiolaniids actively crossed oceans to arrive on distant islands, either by swimming, wading or floating. However, modern research generally agrees that meiolaniids were terrestrial animals and the work by Brown and Moll specifically discusses the many aspects of meiolaniid anatomy that would be a detriment to such dispersal. According to their research, the comparably shallow shell of 6354: 2932:. Dispersal similar to that of modern giant tortoises has also been suggested by Sterli, who maintains that the overall similarities to modern tortoises may be enough to enable them to drift along with ocean currents. In addition to citing many of the same reasons that render active swimming unlikely (the insufficient buoyancy of the shell and heavy build), Brown and Moll argue that adults would struggle with finding rafts large enough, while juveniles would be easy prey to any marine or island predators. 1242: 1128: 1082: 934: 126: 3101: 2897: 1442: 3161:

drop in temperatures, allowing meiolaniids to survive past the Oligocene and into the Pleistocene to Holocene. Although several species of meiolaniids were present on Australia during the Pleistocene, it is not known what led to their extinction. The disappearance of the island populations meanwhile on the other hand has been discussed more commonly in publications. One hypothesis suggests that many meiolaniids fell victim to rising sea levels following the

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smell. While other possible uses of an enlarged nasal cavity are also considered, including sound production and thermoregulation, the benefits to the sense of smell is considered to be the most likely cause. Compared to this, smell plays a very minor rolle in the lives of aquatic turtles, which subsequently have a much smaller nasal cavity. The vestibulum of the nose is elongated and although this is associated with trunk-like structures in modern

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anchor-like clubbed tail have all been cited as being detrimental to an animal living in the water, as they would be a hinderance when the animal were to try and swim between islands or try to reach its head above water. Brown and Moll further criticize the methodology and sample size of Lichtig and Lucas specifically, pointing out that their publication worked with a single juvenile specimen, which was a composite and thus didn't reflect actual

1051: 1015: 1862: 6400: 1453: 3132:, which indicate that they made their nests in high moisture environments to prevent water evaporation from drying out the clutch. Suitable environments would include beaches, which is where the nests have been found on Lord Howe Island. Individual eggs were roughly spherical and measured 53.9 mm (2.12 in) across, making them comparable in size to those of modern giant tortoises. A single clutch of 1187: 888: 549: 3084:

impossible that meiolaniids could have browsed on occasion, even if it wasn't their preferred way of feeding. The mild climate of some of its habitats, such as Lord Howe Island, could suggest that they were periodically moving throughout the year in accordance with seasonally available food sources, possibly utilizing the enhanced sense of smell suggested by their large nasal cavities. Finally,

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see part of the chain submerged by water on one end and new land emerge on the other. Through this, a terrestrial animal may move over time from island to island, with its final distribution being much further off shore than where it started. Additionally, this could explain the precense of a relatively ancient lineage on what is a comparably young landmass. This has been suggested for

3059:, it could also be an adaptation towards keeping sand out of the nose as seen in modern lizards. This would be especially useful for meiolaniids living in arid regions or entering sandy areas such as beaches. Finally, the angle at which the inner ear is directed matches more closely with that of terrestrial tortoises, which are adapted to stabilizing the head while walking. 1215: 1160: 112: 693: 1480:. The individual rings appear to correlate directly with the vertebrae, meaning that each vertebra is surrounded by a singular ring that articulates with those before and after it. The sides of the ring form bony spikes, one smaller pair that faces towards the side and one larger pair that juts out more dorsally. Some forms, namely 770:. While these early publications largely treated Ameghino's and Roth's turtles as separate specimens, the former never provided a detailed diagnosis, description or even figure of his material. At the same time however, Ameghino claimed knowledge over where Roth's material originated. Recent research conducted on the history of 4447: 2953:. Sterli however argues that this model is limited in its ability to explain distribution, as many of the island chains meiolaniid remains were found on run parallel to mainland Australia, rather than moving away from it. Another hypothesis ties the distribution of island meiolaniids to the continent 2334:

or an entirely separate, independent group. Many of the problems responsible for this varying placement can be found in the incompleteness of meiolaniid remains and their highly derived nature. After meiolaniids were recognized as turtles, Huxley suggested they were related to modern snapping turtles

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Aside from the large horns present on the skulls, meiolaniids are also characterized by their heavily armored tails. It is believed that the entirety of the tail in meiolaniids was covered in bony rings flanked by at least two pairs of spikes. Such bone rings are known from even the most basal genus,

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As meiolaniid fossils are often found in the form of broken horn cores and tail rings, much of the collected material is only present in the form of fragmentary remains too scrappy to be named or even assigned to any existing species. Due to this, much of meiolaniid diversity is only known to science

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is known primarily from limb elements that show clear signs of butchering and burning. However, the absence of skulls, horns or tail rings has led some researchers to question if this turtle was actually a meiolaniid or some other, unrelated type of turtle. Sterli further points at several anatomical

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The wide and oftentimes isolated nature of meiolaniid distribution means that their extinction was not a singular event but rather the combination of several factors that gradually caused their disappearance from different landmasses. Meiolaniids probably disappeared from South America at some point

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butchering turtles on the island of Vanuatu. However, there are multiple logistic problems that decrease the probability of this having happened. Adults with their great size and clubbed tails may have been difficult and even dangerous to transport, while juveniles would take a substantial amount of

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The most prominent scale areas are those designated A to C in order from the back-most area to the front-most pair. These scale areas, commonly referred to as horns or horn cores due to their size and shape, are very pronounced and highly distinct in the individual genera and even species. Generally

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suggests that at least in said genus, the horns likely served more of a display function. This is due to the fact that certain behaviors, like sideways stabbing using the horns, causes a great deal of stress on the bone, while headbutting is discounted due to the lack of direct adaptations for such

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where available. Part of the reason for this is the limited range of motion provided by their neck and the heavy skulls, which are not suited for an animal that would have to consistently keep its head raised to feed. Instead, the neck was much more built for side to side movement. Still, it is not

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s skeleton, such as the domed shell, robust forelimbs and anatomy of the shoulder girdle, all compare favorable to terrestrial tortoises rather than aquatic terrapins or turtles. Features such as the osteoderm-covered limbs, limited range of motion of the neck, large and heavy skulls and the almost

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of Lord Howe Island. However, the name is retained regardless due to its importance for communication, making it easier to clarify which island's turtles are referred to. Furthermore, Gaffney concurs that it may have been a "biological species", meaning it could have been genetically distinct given

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The final hypothesis, and the one favored by Brown and Moll, proposes that meiolaniids primarily arrived on distant islands through travel over land. Among these, one possible explanation can be found in the "escalator hopscotch" model. According to this, an island chain may undergo a process that

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Fossil discoveries made since them have drastically changed this however. Several genera of Mesozoic turtles have been found to share similarities to meiolaniids, giving crucial insight into the potential origin of the group. The first instance of this was recognized as early 1987, when Ckhikvadzé

782:, providing the most extensive look at this taxon to date. This detailed look at the type species ran in tandem with several studies examining meiolaniid fossils from other localities. 1992 saw the description of three new meiolaniid taxa in the span of a single year, consisting of the new species 556:

By 1884 better recorded fossil discoveries had been made on Lord Howe Island, with multiple shipments being sent to Owen in London. Again, the material had been correctly identified as having belonged to turtles by local collectors and researchers, but was then misattributed to lizards by Owen. It

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Australasian meiolaniids meanwhile would fare better, in part due to the continent they inhabited not being as stationary. While South America generally remained in the same place, Australia would continuously drift northward, entering higher latitudes and subsequently compensating for the global

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Amphichelydia. During the 1970s Amphichelydia fell out of use, with groups previously included in it being split among pleurodires and cryptodires. Gaffney at the time argued that meiolaniids were not just cryptodires, but eucryptodires, placing them as a sister group to today's snapping turtles,

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specimens also highlights how variable these turtles can be, as some individuals show clearly defined B horns while others have them no larger than the C horns. The reason for this is currently unknown, but sexual dimorphism is considered to be unlikely given how these horn morphs are distributed

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No new species were named between 1938 and the 1990s. Instead, the vast quantity of fossil material collected on Lord Howe Island led to a series of major publications penned by Eugene S. Gaffney, now renowned for his work on this group. Split across three papers published in 1983, 1985 and 1996,

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is another hypothesis that has been suggested and would propose that meiolaniids dispersed when animals stuck on natural rafts were washed to distant islands. Multiple reports exist of giant tortoises coming ashore far away from their place of origin after severe storms, with one particular case

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and the Wyandotte species. Both were estimated to have reached a similar weight and the latter was estimated to have reached a carapace length of up to 2 m (6 ft 7 in). Notably, these length estimates are restricted to the bony shell and do not factor in the combined length of the

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has a long and complex history strongly tied to the rivalry between the researchers that named and described it. The subsequent confusion extends to the type locality, for which contradictory information exists, however, recent research suggests that the fossil likely originated in the Sarmiento

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An additional point in favor of terrestrial life was recovered when the nasal cavities and inner ears of several meiolaniids were analysed. The study found that meiolaniids had enlarged nasal cavities, larger than even those of modern tortoises, which could be indicative of an enhanced sense of

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Meiolaniids were large animals, with the bigger species reaching total lengths of perhaps up to 2–3 m (6.6–9.8 ft). Meiolaniid remains can easily be identified by their skulls, which are covered in distinctive scale patterns and formed elaborate head crests and horns that vary greatly

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Bauer and Vindum (1990) on the other hand suggest that rather than spreading naturally, the last meiolaniids were helped in their dispersal by humans bringing them along as a food source. Historically, tortoises have been used as living provisions by seafarers and evidence exists of the native

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such as pigs and rats that were a bigger threat to the species than humans, as they would raid nests and prey on juvenile turtles. The fossil record suggests that turtles disappeared from the island only 300 years after humans arrived. However, it is unclear how much this actually impacts the

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The research history of meiolaniids is long and at times complicated, with especially the early years suffering from poor records, incorrect identifications and loss of information. Some of the earliest supposed discoveries made by western scientists are said to date to the middle of the 19th

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were synonymized with one another. What followed was a long, uninterrupted period of fossil collection on Lord Howe Island, providing a massive quantity of fossil material. Although excavations were productive, this time period was relatively uneventful in regards to taxonomy, with the only

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speaking, the A horn is a singular element located at the back of the skull that ranges from forming a large, frill-like structure to an almost vestigial shelf. The B scales are paired and appear more horn-like in their morphology, while the C horns are typically reduced and knob-like.

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show scale areas similar to those characteristic for meiolaniids and several other anatomical features have been observed uniting these Mesozoic turtles with meiolaniids. Sterli and de la Fuente conclude that the presence of well defined scale areas present on the skull may have been

2448:. Their analysis recovers meiolaniids as deeply nested in a group of primarily Gondwanan turtles they named Meiolaniformes, which contradictory to the previously held opinion indidcates that meiolaniids sit on a branch of turtles that lies outside of the Pleurodira Cryptodira clade. 4419: 3165:, which drastically cut down the available space on many islands. There are however issues to this hypothesis, as not all islands were equally affected by this change in sea levels. Human hunting is another suggestion made to explain the disappearance of the last meiolaniids. 2892:

would be less buoyant than those of modern giant tortoises, the tail would function as an anchor and the heavy head and restrictive range of motion would be an inconvenience when trying to raise the head to breathe. According to them, meiolaniids would likely drown in water.

2961:, which have been interpreted by Worthy and colleagues to represent a potential meiolaniid. Little is known about this form, but it is argued that the presence of a large terrestrial tortoise dispels the hypothesis that New Zealand was entirely flooded in this time period. 1596:

is consistently found to be the basalmost meiolaniid, sitting at the base of the tree as a sister to all Australasian forms. This matches its geographic range and age, which clearly separates it from younger meiolaniids. Some of the seemingly ancestral scale conditions of

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Rhodin, Anders G. J.; Thomson, Scott; Georgalis, Georgios L.; Karl, Hans Volker; Danilov, Igor G.; Takahashi, Akio; de la Fuente, Marcelo SaulIcon ; Bourque, Jason; Delfino, Massimo; Bour, Roger; Iverson, John B.; Shaffer, Bradley H.; van Dijk, Peter Paul (2015).

609:, to which Huxley further assigned the Queensland skull. Owen meanwhile, who had received more material from Australia, slightly amended his prior research. While now also recognizing some turtle affinities, Owen maintained that there was a connection to lizards, with 1433:. Although all three horn types are still present and distinct, they are much more reduced and form neither a large frill nor pronounced B horns, instead only appearing as a relatively subtle ridge extending from behind the eyes to the back of the head. 1889:

species, a trait that has been proven to be highly variable even within a single species. Doing so regardless would yield the following results, with the groupings being entirely based on the length to width ratio of the horns. In 2015, Sterli recovers

2957:. In this scenario, meiolaniids were possibly more widespread across this continent and were eventually restricted to isolated island ecosystems once the continent was submerged by the sea. This could find support in turtle remains discovered on 3921: 2361:

and meiolaniids were generally viewed as pleurodires for the following decades. Anderson and Simpson both suggested that meiolaniids were part of neither group, instead declaring them descendants of early turtles and placing them in the

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the large distance between the Walpole population and those of New Caledonia and Australia. This notion would later be echoed by Sterli, who reasoned that the two populations would have been unable to maintain gene flow between another.

1068:, known from several hundred individuals found during almost uninterrupted collection on Lord Howe Island. It is the only meiolaniid of which the entire skeleton is known and thus one of the main sources for information on this group. 3196:

The youngest confirmed meiolaniid remains were recovered from Pindai Caves and have been dated to 1720 ± 70 years BP (160–300 AD) via uncalibrated radiocarbon dating and 1820–1419 years BP (130–531 AD) through calibrated 14C dating.

3642: 1317:(Queensland). Some of these may have beend alongside named genera, indicating that two or more meiolaniids could be found in the same environment. The indeterminate Riversleigh meiolaniid for instance likely coexisted with 3591: 531:. Although the fossils was correctly identified by its collector, G. F. Bennett, Owen instead believed the skull to have belonged to a type of lizard. Combining the skull with the vertebrae of the giant monitor lizard 4204:

Poropat, Stephen F.; Kool, Lesley; Vickers-Rich, Patricia; Rich, Thomas H. (2017). "Oldest meiolaniid turtle remains from Australia: Evidence from the Eocene Kerosene Creek Member of the Rundle Formation, Queensland".

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in the form of various fossils designated Meiolaniidae indeterminate. However, even if fragmentary, this material nonetheless shows that members of this group were diverse and widespread throughout Cenozoic Australia.

3172:, which preserves signs of being butchered by human settlers on Vanuatu. The material of this turtle, consisting primarily of the meaty limbs, were discovered in the remains of a human settlement dating to 2,800 1591:

group with well resolved internal relationships. Among the most important features in this are the different scale areas, which provide the majority of characters used in phylogenetic analysis of this group.

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Owen's identification was soon criticized by other scientists in London, who agreed with the Australian researchers in that these remains were actually those of turtles, not lizards. Just one year after

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Sterli, Juliana; de la Fuente, Marcelo S. (2013). "New evidence from the Palaeocene of Patagonia (Argentina) on the evolution and palaeo-biogeography of Meiolaniformes (Testudinata, new taxon name)".

4667:"Comparative neuroanatomy of extinct horned turtles (Meiolaniidae) and extant terrestrial turtles (Testudinidae), with comments on the palaeobiological implications of selected endocranial features" 3003:

The lifestyle of meiolaniids has historically often been questioned. Even during the earliest discoveries on Lord Howe Island, the idea that they were marine animals was proposed by scientists like

3157:, meiolaniids disappeared entirely. Turtles would be absent from Patagonia for the next 15 million years until the late Oligocene to early Miocene, when testudinids began to settle the region. 3038:

Despite the reoccurring notions of semi-aquatic or even aquatic habits in meiolaniids, most historic and contemporary research favors an exclusively terrestrial lifestyle. Multiple elements of

521:, situated off the eastern coast of Australia, discovered the remains of large turtles. The first well supported finds came just prior to the 1880s, when a large skull of what is now known as 4706:

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".

742:, who had discovered a strikingly similar animal. Roth's find was first figured in a communication by Moreno and was later described in greater detail by Woodward. Having heard of Ameghino's 477:. For this reason it is speculated that meiolaniids were also present on the latter, although no fossils of them have yet been found there. Furthermore, meiolaniids may have been present on 1877:

is theoretically possible, however as discussed by Gaffney, the results of doing so are highly questionable. Only two species would be complete enough to provide valuable characters, as

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While their internal relationships are relatively well understood, their relation to other turtles has long remained elusive. Throughout their history, they've been variable considered

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While their lifestyle was long debated, current research indicates that they were terrestrial herbivores with a keen sense of smell that may have used their heavily armored bodies in

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and the wyandotte species is only represented through horn cores. This renders the morphology of the B horns the only way to possibly determine relationships within the established

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shows the least elaborate head gear among meiolaniids, with the horns and shields seen in other genera being reduced to a continuous shelf of horns that spans the back of the head.

716:. The precise history of these events is however poorly understood due to a large amount of conflicting information. At the time, two rivaling groups of paleontologists, one led by 1416:

this area is so reduced its even described as being vestigial. The B horns on the other hand are typically well developed and conical rather than flattened. Typically the horns of

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this club is larger, formed by four distinct elements. The spikes seen on the prior tail rings continue onto the tail club, where they typically decrease in size towards the end.

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Sterli, Juliana (2010). "Phylogenetic relationships among extinct and extant turtles: The position of Pleurodira and the effects of the fossils on rooting crown-group turtles".

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had cow-like, recurved horns. They also had long tails that were covered in spiked rings of bones that, at least in some genera, transitioned into a tail club towards the tip.

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as the most recent genus represents an extreme in regards to this gradual reduction of the A horn, with the structure only forming a small shelf at the back of the skull. In

4374: 3434:"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)" 425:, which contains more primitive turtles species lacking the distinctive morphology of meiolaniids, known from the Early Cretaceous-Paleocene of South America and Australia. 3149:

put pressure on the turtles native to Patagonia, which failed to cope with the changing conditions. This effect was not exclusive to meiolaniids and also wiped out the

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time until they were big enough for consumption. Furthermore, the slow growth cycle would render these turtles an overall unsustainable food source in the long run.

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possibly representing a relative to both reptile groups. For this new clade, Owen coined the name Ceratosauria, unaware the name was already occupied by a group of

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The oldest unnamed meiolaniid from Australia, known based on shell remains, osteoderms and a tail ring, dates to the Late Eocene and has been discovered in the

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was named in honor of Eugene S. Gaffney for his contributions to the understanding of this group. While geographically close and having similar B-horns to

3260: 4296:; Tennyson, Alan J. D.; Hand, Suzanne J.; Godthelp, Henk; Scofield, R. Paul (2011). "Terrestrial Turtle Fossils from New Zealand Refloat Moa's Ark". 6480: 1400:

lived long after these Eocene forms, its horn structure mirrors the South American genera and likely indicates that this is the basal condition. In

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Hawkins, Stuart; Worthy, Trevor H.; Bedford, Stuart; Spriggs, Matthew; Clark, Geoffrey; Irwin, Geoff; Best, Simon; Kirch, Patrick (December 2016).

1629:, the fragmentary nature of the former makes it somewhat of a wildcard in phylogenetic analysis. It has been recovered as either nesting alongside 6582: 4788: 1511:

Other parts of the skeleton are harder to compare due to the incomplete nature of most meiolaniids, with much of the information stemming from

6360: 4420:"The enigmatic palaeoecology and palaeobiogeography of the giant, horned, fossil turtles of Australasia: a review and reanalysis of the data" 3695: 3310: 3011:

was a marine turtle that died while coming ashore to lay eggs. While McCulloch's hypothesis was quickly dismissed following the discovery of

3146: 2341:), placing it in Cryptodira, the group that includes most living turtles and tortoises. While Boulenger agreed with the identification of 3795:"The Cervical and Caudal Vertebrae of the Cryptodiran Turtle, Melolania platyceps, from the Pleistocene of Lord Howe Island, Australia" 3125:

behavior and the fact that contact between frills in a head-on collision would be impossible given the animals likely range of motion.

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has been used to inferr different types of behavior for meiolaniids, especially in regards to intraspecific communication and possibly

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Parallel to the later stages of this initial burst of revisions, the remains of a third meiolaniid were discovered in 1898 across the

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as a turtle, he proposed it was a member of Pleurodira, the side-necked turtles, which today include Southern Hemisphere groups like

6353: 4740: 4548: 4569:"A fossil egg clutch from the stem turtle Meiolania platyceps : implications for the evolution of turtle reproductive biology" 1621:. The absence of an accessory grinding surface in the jaws also identifies it as a sister taxon to the other meiolaniids. Although 4333: 3881:

Megirian, D. (1992). "Meiolania brevicollissp. Nov. (Testudines: Meiolaniidae): A new horned turtle from the Australian Miocene".

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includes the enormous A scale area and the more laterally directed B horns, both traits shared with the basalmost Australian form

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is the basal-most meiolaniid of Australasia. This is evident through the anatomy of its horns, which bears closer resemblance to

3764:"The cranial morphology of the extinct horned turtle, Meiolania platyceps, from the Pleistocene of Lord Howe Island, Australia" 3587: 3181: 3020: 1653:

due to the second accessory ridge, the broad head and the partially separated internal nares. However, it is excluded from the

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into monitor lizard, marsupial and turtle remains, with the name being constrained to the lizard. While this marked the end of

1352:. However, as the remains do not represent the characteristic horns or tail rings, the affinities of this form may change. 2877:

have been found, with the discovery of an isolated tail ring confirming the group's presence in Eocene Australia as well.

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head, neck and long tail. This may indicate that meiolaniids could have reached lengths of up to 3 m (9.8 ft).

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is commonly criticized and a controversial idea among researchers. In the case of the Vanuatu turtles, it may have been

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Pictured below is the phylogenetic tree recovered in Sterli, de la Fuente and Krause in 2015. Other than the wildcard

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suggests that there never were two specimens, and that Ameghino simply missattributed the Roth skull to his brother.

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as a lizard, Woodward agreed with Owen in that the skull from the mainland clearly belonged to an animal related to

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One of the earliest interpretations of meiolaniids depicted them as large lizards similar to today's Thorny Devils.

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Ameghino, F. (1899). "Sinopsis geológico-paleontológica de la Argentina. Suplemento (adiciones y correcciones)".

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has suggested a herbivorous to omnivorous diet, which would match what has been inferred for other meiolaniids.

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the club appears to be made from two segments that are fused with each other and form a spiked sheath, while in

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and the overall morphology of the legs, which is robust with blunt toes, also supports terrestrial locomotion.

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Most researchers today agree that meiolaniids were terrestrial animals that fed on low growing vegetation.

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is domed rather than flattened, one of several traits indicative of a terrestrial lifestyle. However, the

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and the Wyandotte species, which have the proportionally largest horns. However, the large sample size of

1306: 3378:(1859). "Description of Some Remains of a Gigantic Land-Lizard (Megalania Prisca, Owen) from Australia". 746:, the researcher concluded that Roth's turtle represented the same species, but placed both in the genus 6519: 4385: 2219: 2061: 1310: 1298: 641: 618: 573:(great roamer) to the chimeric material from the mainland, he subsequently named the Lord Howe material 5127: 1461:

Multiple elements of the caudal armor are known, including an anterior ring (top) and club (bottom) of

1314: 1285:(New South Wales) seem to belong to an intermediate taxon, combining the flattened horns of taxa like 1278: 766:

as Ameghino's skull could not be found. This highlights one of the key sources of confusion regarding

6467: 6001: 5917: 4568: 4167: 4036: 3970: 3922:"Warkalania, a New Meiolaniid Turtle from the Tertiary Riversleigh Deposits of Queensland, Australia" 3347: 1372:

The most defining feature of meiolaniids is the presence of clearly defined scale areas covering the

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eggs seems to have consisted of ten eggs, which were organized into two layers of a single nest.

3068: 2694: 1694: 730: 143: 6257: 6015: 5959: 5938: 5875: 5819: 5680: 5673: 5504: 5404: 5150: 5113: 5063: 4352: 4092:"The Wyandotte Local Fauna: A New, Dated, Pleistocene Vertebrate Fauna from Northern Queensland" 2896: 2884:, several competing ideas exist in regards to their further dispersal across the islands of the 1574:

was estimated at 1.2 m (3 ft 11 in). The largest sizes were seemingly reached by

1384:

possesses the most elaborate A horn, which forms a structure somewhat resembling the frill of a

6524: 6458: 6298: 6029: 5701: 5687: 5638: 5624: 5184: 3176:

and shows clear cut marks, fractures and even burns all indicative of human consumption by the

2634: 2409: 1441: 1388:, while the flattened B horns extend to the sides and back. Little is known about the horns of 6506: 5726: 4736: 4588: 4544: 4439: 4366: 4356: 4072: 4054: 3998: 3917: 3856: 3691: 3306: 3085: 3076: 2904: 2881: 2363: 2243: 2024: 1906:

species. The Wyandotte species was not used in this analysis due to it being too fragmentary.

961: 953:

and among the most complete. It shows more slender and more strongly curved horns compared to

582: 6511: 6498: 5735: 5631: 5603: 5568: 5520: 4615: 3643:"On some extinct reptiles from Patagonia, of the genera Miolania, Dinilysia, and Genyodectes" 1566:, have been estimated to have reached a carapace length of 0.7 m (2 ft 4 in). 624:

In spite of Owen's conviction, more and more researchers published on the turtle identity of

6043: 6008: 5987: 5271: 4711: 4686: 4678: 4616:"The Galapagos tortoises in their relation to the whaling industry: a study of old logbooks" 4580: 4517: 4509: 4474: 4431: 4348: 4305: 4293: 4262: 4254: 4214: 4175: 4062: 4044: 3988: 3978: 3952: 3892: 3848: 3802: 3775: 3683: 3657: 3608: 3564: 3523: 3515: 3455: 3445: 3387: 3355: 3283: 3185: 3056: 3052: 3032: 2847: 2358: 1345: 1056: 721: 518: 482: 406: 289: 5596: 1613:

to other turtle groups, as the intrapterygoid appears more "primitive" compared to that of

569:, believing it to be a small relative of the mainland specimen. As Owen had given the name 6203: 5467: 5455: 5308: 4962: 4114: 4091: 1532: 1392:

meanwhile, however the singular known B horn indicates that it may have looked similar to

778:

Gaffney described in great detail the skull, vertebrae and finally the shell and limbs of

735: 701: 384: 4171: 4040: 3974: 3351: 6378: 6112: 6091: 5966: 5826: 5561: 5434: 5164: 5136: 5103: 4067: 4024: 3993: 3956: 3661: 3173: 3113: 2950: 2863: 2855: 2843: 2508: 2350: 1861: 1735: 1324:

Indeterminate remains from islands have been discovered in the Pleistocene to Holocene

957:

and further is clearly geographically separate, as it was found on mainland Australia.

682: 422: 271: 229: 6541: 6139: 6126: 5852: 5497: 5483: 5447: 5389: 5359: 5191: 5093: 5083: 5073: 4945: 4923: 4897: 4854: 4797: 4666: 4226: 3503: 3177: 2937: 2885: 2851: 2610: 2441: 2421: 2391: 2376: 2368: 2354: 1329: 1020: 739: 686: 505: 410: 365: 203: 69: 28: 4600: 4486: 4463:"Reassessment of the phylogenetic interrelationships of basal turtles (Testudinata)" 4317: 4276: 4187: 3841:"The postcranial morphology of Meiolania platyceps and a review of the Meiolaniidae" 3612: 3537: 3104:

Meiolaniids may have used their tails in combat with other members of their species.

1420:

are recurved, resembling the horns of bovines like cows. This is most pronounced in

6393: 6388: 6277: 6243: 6227: 6184: 5931: 5882: 5742: 5554: 5324: 5225: 5043: 5033: 5023: 4983: 4972: 3840: 3794: 3375: 3329: 3162: 2372: 1869:, are among the most important factors in determining their internal relationships. 1793: 1752: 1711: 1588: 1452: 1325: 871: 821: 548: 542: 528: 436: 307: 255: 242: 4715: 4584: 4218: 3763: 4478: 4258: 4179: 3145:

during the middle Eocene. The gradually cooling of Earth's climate following the

3019:

was connected to water would still appear periodically in the following century.

6452: 6291: 6268: 6163: 6119: 6098: 5847: 5666: 5652: 5645: 5610: 5589: 5368: 5338: 5006: 4993: 4682: 4665:

Paulina-Carabajal, A.; Sterli, J.; Georgi, J.; Poropat, S.F.; Kear, B.P (2017).

3687: 3504:"A review of the fossil record of Gondwanan turtles of the clade Meiolaniformes" 3333: 2958: 2925: 2445: 1618: 1610: 1497: 1493: 1385: 1360: 1349: 1337: 1155:

was among the largest meiolaniids, rivaling the contemporary Wyandotte species.

1097:, the Wyandotte species remains unnamed and is at times tentatively assigned to 478: 466: 414: 402: 377: 376:, they are not closely related to any living species of turtle, and lie outside 369: 216: 44: 3023:, who discovered the aforementioned leg bones, considered the possibility that 6383: 6191: 6177: 6170: 6156: 5980: 5952: 5945: 5861: 5777: 5582: 5575: 5476: 5331: 5253: 5177: 4870: 4827: 4513: 3896: 3287: 2742: 2522: 2487: 2472: 2415: 2331: 2327: 2195: 1976: 1776: 1548: 1225: 907:, its exact phylogenetic position is unclear due to the fragmentary material. 796: 637: 474: 455: 373: 354: 339: 89: 54: 6443: 4592: 4443: 4370: 4058: 3860: 3592:"Notes on the extinct Chelonian Meiolania, with a record of a new occurrence" 6305: 6149: 6084: 5973: 5924: 5512: 5412: 5382: 5232: 3983: 3154: 3080: 2954: 2929: 2913: 2842:

According to research by Sterli and colleagues, meiolaniids derive from the

1562:

Meiolaniids were large and robust animals. Even the smaller species, namely

1489: 1344:(2011) reported on what may be the remains of a meiolaniid from the Miocene 1247: 1214: 1192: 1159: 1133: 1087: 939: 916: 893: 830: 725: 713: 614: 577:(small roamer). This has however led to some confusion, as the etymology of 538: 533: 461: 444: 394: 389: 361: 331: 160: 94: 38: 4435: 4076: 4002: 3569: 3391: 3360: 3303:

Dragons in the dust: the paleobiology of the giant monitor lizard Megalania

3027:

was semi-aquatic and more recently, Lichtig and Lucas (2018) proposed that

1570:

could have reached a carapace length of 1 m (3 ft 3 in) and

803:

Only two new taxa have been named since this boom in the 1990s, with ?

3071:, feeding on a variety of low-growing plants and plant material including 1030:, Gaffney argues that the material is insufficient to distinguish it from 111: 6437: 6105: 5659: 5538: 5490: 5420: 5246: 5239: 4821: 4639:"The ecology of Meiolania platyceps, a Pleistocene turtle from Australia" 4149:"Re-Description and Evolutionary Remarks on the Patagonian Horned Turtle 3929:

The Beagle, Records of the Northern Territory Museum of Arts and Sciences

3519: 3150: 3109: 2866: 2718: 2346: 2337: 2171: 2147: 1952: 1928: 1520: 1170: 1111: 808: 645: 523: 470: 449: 430: 418: 323: 315: 180: 84: 79: 64: 59: 49: 32: 4729:

James Gibbs, Linda Cayot, Washington Tapia Aguilera (November 7, 2020).

1404:

too the A horn forms an enlarged frill, even if less pronounced than in

6485: 6077: 6022: 5994: 5345: 4836: 4691: 4522: 3528: 3460: 2917: 2862:, the majority of their known diversity could be found on the southern 1637:

at the base of the Australasian clade and as a derived genus alongside

1282: 988: 812: 763: 709: 692: 586: 398: 190: 99: 74: 4309: 4267: 4049: 3450: 3433: 3399: 2444:

for all turtles, and was simply lost and re-evolved repeatedly in the

1213: 1158: 960: 517:

century, with writings suggesting that various locals and visitors of

6321: 5527: 4845: 4815: 3852: 3806: 3680:

Origin, Evolution and Biogeographic History of South American Turtles

826: 493:

had massive frills and sideways facing, flattened horns, others like

441: 380: 357: 170: 6414: 3779: 1661:

clade due to the size of the A horns and the shape of the D scales.

3678:

de la Fuente, M. S.; Sterli, J.; Maniel, I (2014). "Introduction".

762:

and around the same time the Roth skull was elevated to the genus'

6070: 3180:. However, there are issues with this idea. While widespread, the 3099: 3072: 2895: 2880:

While the early distribution of the family is easily explained by

1860: 1373: 1359: 691: 547: 541:, Owen came to believe that the bones represented a type of giant 4115:"A meiolaniid turtle from the Pleistocene of Northern Queensland" 3116:. The inner ear suggests that meiolaniids were more sensitive to 1321:, which is clearly differentiated through the horn core anatomy. 1281:

of Queensland. Remains found in Early Miocene Canadian Lead near

6472: 4766: 3153:

found in the region. While chelids managed to survive at higher

2846:, a group of turtles that likely evolved during or prior to the 1492:

that tips the end of the tail and has been compared to those of

1333: 1206:, another Argentinian meiolaniid, was found to be a juvenile of 816: 734:, a large meiolaniid turtle he claimed was found by his brother 421:

for the latter. Meiolaniids are part of the broader grouping of

6418: 4808: 4770: 3239:(Woodward), a Horned Turtle from the Pleistocene of Queensland" 1476:, and surrounds the entire circumference of the tail in it and 3189:

extinction date of meiolaniids, as it is not certain if ?

1587:

Phylogenetic analysis consistently recovers Meiolaniidae as a

2949:, as Lord Howe Island is a volcanic island situated atop the 1297:. Other continental remains were found in the Late Oligocene 728:. Ameghino published a short communication in which he names 428:

Meiolaniidae includes a total of five different genera, with

1668:, the phylogenetic tree matches with prior work by Gaffney. 1547:

all show that the back of the carapace had a serrated edge.

738:. Simultaneously, Woodward received material from collector 469:. The group is believed to have evolved on the continent of 1873:

Creating a phylogenetic tree for the individual species of

1429:

across specimen. The most reduced horns can be observed in

994:

Known from archaeological sites on Vanuatu and Fiji, ?

605:

published a paper correcting Owen and naming the material

585:

would later suggest that "-lania" actually translated to "

4643:

New Mexico Museum of Natural History and Science Bulletin

3557:

Philosophical Transactions of the Royal Society of London

3380:

Philosophical Transactions of the Royal Society of London

1649:

is the second basalmost genus. It is clearly united with

3553:"On Parts of the Skeleton of Meiolania platyceps (Owen)" 527:

was discovered in Queensland and sent to paleontologist

481:

based on the discovery of turtle remains as part of the

4462: 3168:

Evidence for hunting may be found in the case of ?

899:

An early, but relatively poorly understood meiolaniid,

3955:; Hawkins, S.; Bedford, S.; Spriggs, M. (2010-08-16). 119:

Restoration of the head of various meiolaniid species

4567:

Lawver, Daniel R.; Jackson, Frankie D. (2016-11-01).

3432:

Sterli, J.; de la Fuente, M.S.; Krause, J.M. (2015).

2413:

and meiolaniids together, as did subsequent authors.

1551:

that covered the limbs have been recovered from both

557:

was based on this material that Owen named the genus

473:

prior to its split into South America, Australia and

387:. They are best known from the last surviving genus, 6040: 5844: 5723: 5431: 5356: 5305: 5268: 5161: 5124: 5109: 5099: 5089: 5079: 5069: 5059: 5049: 5039: 5029: 5019: 4989: 4979: 4968: 4958: 4540:

The Rise of Reptiles. 320 Million Years of Evolution

4207:

Alcheringa: An Australasian Journal of Palaeontology

3884:

Alcheringa: An Australasian Journal of Palaeontology

2395:

and meiolaniids in a single group. In 2000 Hirayama

1309:

of the Riversleigh (Queensland), the Middle Miocene

807:

representing an uncertain member of this group from

353:

is an extinct family of large, probably herbivorous

6427: 6320: 6267: 6201: 6137: 6039: 5843: 5722: 5537: 5466: 5430: 5355: 5304: 5267: 5215: 5160: 5123: 5004: 4943: 4896: 4869: 4025:"Ancient tortoise hunting in the southwest Pacific" 3735:"Crossochelys, Eocene horned turtle from Patagonia" 754:). Later finds in the area would produce the taxon 1531:, instead bearing more resemblance to that of the 660:. Woodward placed it in the same genus, naming it 3508:Bulletin of the Peabody Museum of Natural History 3682:. Springer Earth System Sciences. pp. 1–5. 1865:The scale areas of meiolaniids, as seen here in 4147:Sterli, Juliana; de la Fuente, Marcelo (2011). 3713:"New reptiles from the Eocene of South America" 3650:Proceedings of the Zoological Society of London 3339:Philosophical Transactions of the Royal Society 1260:is the oldest named meiolaniid from Australia. 999:features that do not match meiolaniid anatomy. 581:was never specified in the actual publication. 4543:. Johns Hopkins University Press. p. 53. 3427: 3425: 3423: 3421: 3419: 3417: 3415: 3413: 3411: 3409: 2858:. Although some fossil evidence may suggest a 4782: 1408:, and the B horns face straight to the side. 724:, were competing in a fashion similar to the 8: 4755:: CS1 maint: multiple names: authors list ( 4390:: CS1 maint: multiple names: authors list ( 589:", a notion later contested in the works of 4353:10.3854/crm.5.000e.fossil.checklist.v1.2015 4153:Ameghino, 1899 (Testudinata, Meiolaniidae)" 6415: 6062: 5439: 5169: 5012: 4951: 4940: 4875: 4866: 4805: 4789: 4775: 4767: 4418:Lauren E. Brown, Don Moll (October 2019). 4288: 4286: 3047:proportions, much less those of an adult. 124: 110: 20: 4690: 4671:Zoological Journal of the Linnean Society 4521: 4266: 4066: 4048: 3992: 3982: 3920:; Archer, Michael; White, Arthur (1992). 3568: 3527: 3459: 3449: 3438:Zoological Journal of the Linnean Society 3359: 3305:. Bloomington: Indiana University Press. 2900:Range of meiolaniids in the South Pacific 1101:. It has some of the largest horns among 3912: 3910: 3908: 3906: 1523:is not quite as high as seen in today's 844: 4660: 4658: 4656: 4380:from the original on 21 September 2022. 4240: 4238: 4236: 3497: 3495: 3493: 3491: 3261:"Notes on the Extinct Reptilian Genera 3206: 2320: 1633:at the base of Meiolaniidae, alongside 383:, having diverged from them around the 4748: 4562: 4560: 4413: 4411: 4409: 4407: 4405: 4403: 4401: 4383: 4199: 4197: 4142: 4140: 4138: 4136: 4134: 4132: 4018: 4016: 4014: 4012: 3946: 3944: 3942: 3876: 3874: 3872: 3870: 3834: 3832: 3830: 3828: 3826: 3824: 3822: 3820: 3818: 3816: 3757: 3755: 3733:Simpson, G.G.; Williams, C.S. (1938). 3673: 3671: 3489: 3487: 3485: 3483: 3481: 3479: 3477: 3475: 3473: 3471: 3276:Annals and Magazine of Natural History 2130: 1911: 1609:is further supported by comparing the 1064:is the type and best known species of 664:in Owen's honour. Shortly afterwards, 3628:Imprenta la Libertad (Author Edition) 3582: 3580: 3324: 3322: 3224: 3222: 3220: 3218: 3216: 3214: 3212: 3210: 3067:Meiolaniids are thought to have been 2928:, at times emanciated and covered in 1305:(South Australia), the Early Miocene 7: 4113:Gaffney, E.S.; McNamara, G. (1990). 3128:Several egg clutches are known from 2357:. Boulenger would find support from 405:, and insular species that lived on 4637:Lichtig, A.J.; Lucas, S.G. (2018). 4537:Hans-Dieter Sues (August 6, 2019). 4467:Journal of Systematic Palaeontology 4247:Journal of Systematic Palaeontology 2734: 2710: 2686: 2626: 2602: 2595: 2545: 2538: 2514: 2465: 2458: 2451: 2235: 2211: 2187: 2163: 2139: 2132: 2040: 2016: 1992: 1968: 1944: 1920: 1913: 1768: 1727: 1686: 1678: 1671: 1396:, if with more rounded tips. While 1313:(South Australia) and the Pliocene 677:species named in this period being 489:between genera. While some such as 4708:Journal of Vertebrate Paleontology 4614:Townsend, Charles Haskins (1925). 4573:Journal of Vertebrate Paleontology 4450:from the original on 18 June 2022. 4160:Journal of Vertebrate Paleontology 3662:10.1111/j.1469-7998.1901.tb08537.x 1364:The horns of a juvenile and adult 800:, a new genus with reduced horns. 14: 2425:followed in 2007 and 2011, while 1026:A potentially dubious species of 6399: 6398: 6359: 6352: 4735:. Elsevier Science. p. 30. 4119:Memoirs of the Queensland Museum 4096:Memoirs of the Queensland Museum 3600:Records of the Australian Museum 3096:Social behavior and reproduction 3079:and perhaps the fallen fruit of 2987: 2978: 2399:expanded on this idea, grouping 1451: 1440: 1240: 1185: 1126: 1080: 1049: 1013: 981: 932: 886: 561:in 1886 to include two species, 147: 3613:10.3853/j.0067-1975.14.1925.844 3182:Pleistocene overkill hypothesis 2377:pig-nosed and softshell turtles 758:, now thought to be a juvenile 360:with heavily armored heads and 6583:Taxa named by Richard Lydekker 2854:stretching as far back as the 825:being a second genus from the 644:officially split the chimeric 1: 4716:10.1080/02724634.2024.2357315 4585:10.1080/02724634.2016.1223685 4341:Chelonian Research Monographs 4219:10.1080/03115518.2016.1224441 372:. Though once believed to be 4479:10.1080/14772019.2011.558928 4259:10.1080/14772019.2012.708674 4180:10.1080/02724634.2011.615685 2431:was described in 2012. Both 1908: 1645:. As far as stable taxa go, 1617:and compares favorably with 1141:Teenage Mutant Ninja Turtles 3688:10.1007/978-3-319-00518-8_1 3193:was actually a meiolaniid. 1293:with the recurved horns of 6604: 3962:Proc. Natl. Acad. Sci. USA 3005:Allan Riverstone McCulloch 2369:pond turtles and tortoises 1910: 945:Named for its short neck, 6373: 6349: 6065: 5442: 5172: 5015: 4954: 4939: 4920: 4878: 4865: 4804: 4732:Galapagos Giant Tortoises 4683:10.1093/zoolinnean/zlw024 4514:10.1163/18759866-07903002 4461:Anquetin, Jérémy (2012). 3897:10.1080/03115519208619035 3799:American Museum Novitates 3720:American Museum Novitates 3301:Molnar, Ralph E. (2004). 3288:10.1080/00222938809460686 3246:American Museum Novitates 2924:arriving on the coast of 2920:. Other examples include 2910:Galapagos giant tortoises 2860:cosmopolitan distribution 2756: 2739: 2732: 2715: 2708: 2691: 2684: 2648: 2631: 2624: 2607: 2600: 2593: 2567: 2550: 2543: 2536: 2519: 2512: 2485: 2470: 2463: 2456: 2257: 2240: 2233: 2216: 2209: 2192: 2185: 2168: 2161: 2144: 2137: 2058: 2045: 2038: 2021: 2014: 1997: 1990: 1973: 1966: 1949: 1942: 1925: 1918: 1807: 1790: 1773: 1766: 1749: 1732: 1725: 1708: 1691: 1684: 1676: 1605:. The basal condition of 1529:galapagos giant tortoises 959: 949:is the oldest species of 913: 786:from mainland Australia, 750:(likely a misspelling of 302: 297: 144:Scientific classification 142: 132: 123: 118: 109: 23: 16:Extinct family of turtles 4502:Contributions to Zoology 3259:Woodward, A. S. (1888). 2611:Patagoniaemys gasparinae 1794:Gaffneylania auricularis 1753:Gaffneylania auricularis 1712:Gaffneylania auricularis 878:Gaffneylania auricularis 537:and the foot bones of a 447:and the remaining taxa, 4090:McNamara, G.C. (1990). 3984:10.1073/pnas.1005780107 3641:Woodward, A.S. (1901). 3229:Gaffney, E. S. (1992). 3031:lived much like modern 2922:Aldabra giant tortoises 2838:Evolution and dispersal 1881:is likely a synonym of 1625:likely lived alongside 1525:aldabra giant tortoises 1195:(Sarmiento Formation?) 630:George Albert Boulenger 4436:10.33256/hj29.4.252263 4424:Herpetological Journal 3839:Gaffney, E.S. (1996). 3793:Gaffney, E.S. (1985). 3762:Gaffney, E.S. (1983). 3711:Simpson, G.G. (1937). 3630:. La Plata, Argentina. 3570:10.1098/rstb.1888.0007 3392:10.1098/rstl.1859.0002 3361:10.1098/rstl.1886.0015 3105: 2912:swept out at sea by a 2901: 2743:Warkalania carinaminor 2652:Kallokibotion bajazidi 2635:Otwayemys cunicularius 2554:Mongolochelys efremovi 1870: 1777:Warkalania carinaminor 1369: 1340:. Furthermore, Worthy 1250:(Riversleigh Station) 1232:Warkalania carinaminor 1218: 1163: 965: 896:(Sarmiento Formation) 705: 607:Ceratochelys sthenurus 553: 6520:Paleobiology Database 3103: 3015:limbs, the idea that 2899: 2571:Peligrochelys walshae 2220:Meiolania brevicollis 2062:Meiolania brevicollis 1898:as sister taxa, with 1864: 1515:itself. The shell of 1363: 1269:Indeterminate remains 1217: 1204:Crossochelys corniger 1162: 964: 924:Meiolania brevicollis 784:Meiolania brevicollis 756:Crossochelys corniger 695: 642:Arthur Smith Woodward 619:Othniel Charles Marsh 551: 3845:Bulletin of the AMNH 3768:Bulletin of the AMNH 3742:Bulletin of the AMNH 3520:10.3374/014.056.0102 3007:, who believed that 1902:being the basalmost 1535:. Shell elements of 1307:Carl Creek Limestone 1236:Oligocene - Miocene 1093:The largest form of 696:A 1914 depiction of 512:History of discovery 502:intraspecific combat 6573:Pleistocene turtles 5792:Protoxinjiangchelys 4172:2011JVPal..31.1210S 4041:2016NatSR...638317H 3975:2010PNAS..10715512W 3969:(35): 15512–15516. 3502:Sterli, J. (2015). 3352:1886RSPT..177..471O 3332:(January 1, 1886). 3130:Meiolania platyceps 3118:low frequency sound 2947:Meiolania platyceps 2760:Meiolania platyceps 2523:Chubutemys copelloi 2261:Meiolania platyceps 2001:Meiolania platyceps 1811:Meiolania platyceps 1568:Meiolania platyceps 1513:Meiolania platyceps 1463:Meiolania platyceps 1426:Meiolania platyceps 1062:Meiolania platyceps 1041:Meiolania platyceps 846: 780:Meiolania platyceps 718:Florentino Ameghino 603:Thomas Henry Huxley 136:Meiolania platyceps 6553:Cretaceous turtles 4151:Niolamia argentina 4029:Scientific Reports 3918:Gaffney, Eugene S. 3235:, a new name for " 3170:Meiolania damelipi 3114:courtship behavior 3106: 3090:Meiolania damelipi 2902: 2695:Niolamia argentina 2047:Wyandotte species 1871: 1695:Niolamia argentina 1488:, also preserve a 1370: 1311:Wipajiri Formation 1299:Etadunna Formation 1219: 1177:Niolamia argentina 1164: 1090:(Wyandotte Creek) 1073:Wyandotte species 973:Meiolania damelipi 966: 845: 805:Meiolania damelipi 790:as a new name for 744:Niolamia argentina 731:Niolamia argentina 706: 554: 6568:Oligocene turtles 6535: 6534: 6507:Open Tree of Life 6421:Taxon identifiers 6412: 6411: 6369: 6368: 6347: 6346: 6343: 6342: 6316: 6315: 5727:Xinjiangchelyidae 5718: 5717: 5263: 5262: 5128:Australochelyidae 4935: 4934: 4931: 4930: 4916: 4915: 4310:10.1643/CH-10-113 4294:Worthy, Trevor H. 4050:10.1038/srep38317 3697:978-3-319-00517-1 3551:Owen, R. (1888). 3451:10.1111/zoj.12252 3312:978-0-253-34374-1 3086:isotopic analysis 3077:herbaceous plants 3057:softshell turtles 3053:pig-nosed turtles 2882:continental drift 2834: 2833: 2825: 2824: 2816: 2815: 2807: 2806: 2798: 2797: 2789: 2788: 2780: 2779: 2771: 2770: 2672: 2671: 2663: 2662: 2582: 2581: 2498: 2497: 2364:wastebasket taxon 2324: 2323: 2317: 2316: 2308: 2307: 2299: 2298: 2290: 2289: 2281: 2280: 2272: 2271: 2244:Meiolania mackayi 2127: 2126: 2118: 2117: 2109: 2108: 2100: 2099: 2091: 2090: 2082: 2081: 2073: 2072: 2025:Meiolania mackayi 1858: 1857: 1849: 1848: 1840: 1839: 1831: 1830: 1822: 1821: 1564:Meiolania mackayi 1266: 1265: 1122:Late Pleistocene 1023:(Walpole Island) 1005:Meiolania mackayi 720:and the other by 583:Eugene S. Gaffney 504:, perhaps during 465:being endemic to 417:and possibly the 393:, which lived in 348: 347: 293: 6595: 6578:Pliocene turtles 6528: 6527: 6515: 6514: 6502: 6501: 6489: 6488: 6476: 6475: 6463: 6462: 6461: 6448: 6447: 6446: 6416: 6402: 6401: 6364: 6363: 6357: 6356: 6063: 6059: 6058: 6044:Thalassochelydia 6042: 5846: 5725: 5440: 5433: 5358: 5307: 5272:Sichuanchelyidae 5270: 5170: 5163: 5126: 5111: 5101: 5091: 5081: 5071: 5061: 5051: 5041: 5031: 5021: 5013: 4991: 4981: 4970: 4960: 4952: 4941: 4876: 4867: 4860: 4859: 4806: 4791: 4784: 4777: 4768: 4761: 4760: 4754: 4746: 4726: 4720: 4719: 4703: 4697: 4696: 4694: 4662: 4651: 4650: 4634: 4628: 4627: 4611: 4605: 4604: 4564: 4555: 4554: 4534: 4528: 4527: 4525: 4497: 4491: 4490: 4458: 4452: 4451: 4415: 4396: 4395: 4389: 4381: 4379: 4338: 4328: 4322: 4321: 4290: 4281: 4280: 4270: 4242: 4231: 4230: 4201: 4192: 4191: 4166:(6): 1210–1229. 4157: 4144: 4127: 4126: 4110: 4104: 4103: 4087: 4081: 4080: 4070: 4052: 4020: 4007: 4006: 3996: 3986: 3948: 3937: 3936: 3926: 3914: 3901: 3900: 3878: 3865: 3864: 3836: 3811: 3810: 3790: 3784: 3783: 3759: 3750: 3749: 3739: 3730: 3724: 3723: 3717: 3708: 3702: 3701: 3675: 3666: 3665: 3647: 3638: 3632: 3631: 3623: 3617: 3616: 3596: 3584: 3575: 3574: 3572: 3548: 3542: 3541: 3531: 3499: 3466: 3465: 3463: 3453: 3429: 3404: 3403: 3372: 3366: 3365: 3363: 3326: 3317: 3316: 3298: 3292: 3291: 3273: 3256: 3250: 3249: 3243: 3237:Meiolania" oweni 3226: 3186:invasive species 3033:snapping turtles 3021:Charles Anderson 2991: 2982: 2848:Early Cretaceous 2735: 2711: 2687: 2627: 2603: 2596: 2546: 2539: 2515: 2466: 2459: 2452: 2359:Richard Lydekker 2236: 2212: 2188: 2164: 2140: 2133: 2041: 2017: 1993: 1969: 1945: 1921: 1914: 1909: 1769: 1728: 1687: 1679: 1672: 1455: 1444: 1346:St Bathans Fauna 1315:Chinchilla Sands 1303:Namba Formations 1279:Rundle Formation 1246: 1244: 1243: 1191: 1189: 1188: 1136:(Darling Downs) 1132: 1130: 1129: 1086: 1084: 1083: 1057:Lord Howe Island 1055: 1053: 1052: 1019: 1017: 1016: 987: 985: 984: 942:(Camfield Beds) 938: 936: 935: 892: 890: 889: 847: 722:Francisco Moreno 519:Lord Howe Island 483:St Bathans Fauna 407:Lord Howe Island 288: 283: 270: 152: 151: 128: 114: 104: 41: 27:Temporal range: 21: 6603: 6602: 6598: 6597: 6596: 6594: 6593: 6592: 6588:Extinct turtles 6563:Miocene turtles 6538: 6537: 6536: 6531: 6523: 6518: 6510: 6505: 6497: 6492: 6484: 6479: 6471: 6466: 6457: 6456: 6451: 6442: 6441: 6436: 6423: 6413: 6408: 6365: 6358: 6351: 6339: 6312: 6263: 6204:Plesiochelyidae 6197: 6133: 6048: 6047: 6046: 6035: 6002:Tienfucheloides 5918:Hongkongochelys 5851: 5839: 5714: 5533: 5468:Pleurosternidae 5462: 5456:Helochelydridae 5426: 5351: 5309:Helochelydridae 5300: 5259: 5211: 5156: 5119: 5009: 5000: 4963:Eorhynchochelys 4948: 4927: 4912: 4892: 4861: 4811: 4810: 4800: 4795: 4765: 4764: 4747: 4743: 4728: 4727: 4723: 4705: 4704: 4700: 4664: 4663: 4654: 4636: 4635: 4631: 4613: 4612: 4608: 4579:(6): e1223685. 4566: 4565: 4558: 4551: 4536: 4535: 4531: 4499: 4498: 4494: 4460: 4459: 4455: 4417: 4416: 4399: 4382: 4377: 4363: 4336: 4330: 4329: 4325: 4292: 4291: 4284: 4244: 4243: 4234: 4203: 4202: 4195: 4155: 4146: 4145: 4130: 4112: 4111: 4107: 4089: 4088: 4084: 4022: 4021: 4010: 3950: 3949: 3940: 3924: 3916: 3915: 3904: 3880: 3879: 3868: 3838: 3837: 3814: 3792: 3791: 3787: 3761: 3760: 3753: 3737: 3732: 3731: 3727: 3715: 3710: 3709: 3705: 3698: 3677: 3676: 3669: 3645: 3640: 3639: 3635: 3625: 3624: 3620: 3594: 3586: 3585: 3578: 3550: 3549: 3545: 3501: 3500: 3469: 3431: 3430: 3407: 3374: 3373: 3369: 3328: 3327: 3320: 3313: 3300: 3299: 3295: 3271: 3258: 3257: 3253: 3241: 3228: 3227: 3208: 3203: 3142: 3098: 3065: 3001: 3000: 2999: 2998: 2994: 2993: 2992: 2984: 2983: 2972: 2967: 2908:telling of two 2840: 2835: 2826: 2817: 2808: 2799: 2790: 2781: 2772: 2673: 2664: 2583: 2499: 2318: 2309: 2300: 2291: 2282: 2273: 2128: 2119: 2110: 2101: 2092: 2083: 2074: 1859: 1850: 1841: 1832: 1823: 1585: 1533:gopher tortoise 1469: 1468: 1467: 1466: 1458: 1457: 1456: 1447: 1446: 1445: 1358: 1271: 1241: 1239: 1186: 1184: 1127: 1125: 1081: 1079: 1050: 1048: 1014: 1012: 991:(Efate Island) 982: 980: 933: 931: 887: 885: 843: 833:in addition to 792:Meiolania oweni 702:Heinrich Harder 662:Meiolania oweni 514: 385:Middle Jurassic 287: 281: 268: 146: 105: 103: 102: 97: 92: 87: 82: 77: 72: 67: 62: 57: 52: 47: 36: 35: 25: 17: 12: 11: 5: 6601: 6599: 6591: 6590: 6585: 6580: 6575: 6570: 6565: 6560: 6558:Eocene turtles 6555: 6550: 6548:Meiolaniformes 6540: 6539: 6533: 6532: 6530: 6529: 6516: 6503: 6490: 6477: 6464: 6449: 6433: 6431: 6425: 6424: 6419: 6410: 6409: 6407: 6406: 6396: 6391: 6386: 6381: 6379:Angolachelonia 6374: 6371: 6370: 6367: 6366: 6350: 6348: 6345: 6344: 6341: 6340: 6338: 6337: 6336: 6335: 6326: 6324: 6318: 6317: 6314: 6313: 6311: 6310: 6302: 6295: 6288: 6281: 6273: 6271: 6265: 6264: 6262: 6261: 6254: 6247: 6240: 6232: 6224: 6217: 6214:Craspedochelys 6209: 6207: 6199: 6198: 6196: 6195: 6188: 6181: 6174: 6167: 6160: 6153: 6145: 6143: 6135: 6134: 6132: 6131: 6123: 6116: 6113:Pelobatochelys 6109: 6102: 6095: 6092:Hispaniachelys 6088: 6081: 6074: 6066: 6060: 6037: 6036: 6034: 6033: 6026: 6019: 6012: 6005: 5998: 5991: 5984: 5977: 5970: 5967:Manchurochelys 5963: 5956: 5949: 5942: 5935: 5928: 5921: 5914: 5907: 5900: 5893: 5886: 5879: 5872: 5865: 5857: 5855: 5841: 5840: 5838: 5837: 5830: 5827:Xinjiangchelys 5823: 5816: 5809: 5802: 5795: 5788: 5781: 5774: 5767: 5760: 5753: 5746: 5739: 5731: 5729: 5720: 5719: 5716: 5715: 5713: 5712: 5705: 5698: 5691: 5684: 5677: 5670: 5663: 5656: 5649: 5642: 5635: 5628: 5621: 5614: 5607: 5600: 5593: 5586: 5579: 5572: 5565: 5558: 5551: 5543: 5541: 5535: 5534: 5532: 5531: 5524: 5517: 5509: 5501: 5494: 5487: 5480: 5472: 5470: 5464: 5463: 5461: 5460: 5452: 5443: 5437: 5435:Paracryptodira 5428: 5427: 5425: 5424: 5417: 5409: 5401: 5394: 5386: 5379: 5372: 5364: 5362: 5353: 5352: 5350: 5349: 5342: 5335: 5328: 5321: 5313: 5311: 5302: 5301: 5299: 5298: 5291: 5284: 5281:Laurasichersis 5276: 5274: 5265: 5264: 5261: 5260: 5258: 5257: 5250: 5243: 5236: 5229: 5221: 5219: 5213: 5212: 5210: 5209: 5206:Trapalcochelys 5202: 5195: 5188: 5181: 5173: 5167: 5165:Meiolaniformes 5158: 5157: 5155: 5154: 5147: 5140: 5137:Australochelys 5132: 5130: 5121: 5120: 5118: 5117: 5107: 5104:Proterochersis 5097: 5087: 5077: 5067: 5057: 5047: 5037: 5027: 5016: 5010: 5005: 5002: 5001: 4999: 4998: 4987: 4977: 4966: 4955: 4949: 4944: 4937: 4936: 4933: 4932: 4929: 4928: 4921: 4918: 4917: 4914: 4913: 4911: 4910: 4909: 4908: 4902: 4900: 4894: 4893: 4891: 4890: 4889: 4888: 4879: 4873: 4863: 4862: 4858: 4857: 4848: 4839: 4830: 4824: 4818: 4809: 4802: 4801: 4796: 4794: 4793: 4786: 4779: 4771: 4763: 4762: 4741: 4721: 4698: 4677:(4): 930–950. 4652: 4629: 4606: 4556: 4549: 4529: 4492: 4453: 4430:(4): 252–263. 4397: 4362:978-0965354097 4361: 4323: 4282: 4253:(7): 835–852. 4232: 4213:(2): 231–239. 4193: 4128: 4105: 4082: 4008: 3951:White, A. W.; 3938: 3902: 3866: 3812: 3785: 3751: 3725: 3703: 3696: 3667: 3656:(2): 169–184. 3633: 3618: 3607:(4): 223–242. 3576: 3543: 3467: 3444:(3): 519–548. 3405: 3367: 3318: 3311: 3293: 3251: 3205: 3204: 3202: 3199: 3147:Eocene Optimum 3141: 3138: 3097: 3094: 3064: 3061: 2996: 2995: 2986: 2985: 2977: 2976: 2975: 2974: 2973: 2971: 2968: 2966: 2963: 2951:Lord Howe Rise 2864:supercontinent 2856:Early Jurassic 2844:Meiolaniformes 2839: 2836: 2832: 2831: 2828: 2827: 2823: 2822: 2819: 2818: 2814: 2813: 2810: 2809: 2805: 2804: 2801: 2800: 2796: 2795: 2792: 2791: 2787: 2786: 2783: 2782: 2778: 2777: 2774: 2773: 2769: 2768: 2765: 2764: 2755: 2752: 2751: 2748: 2747: 2738: 2733: 2731: 2728: 2727: 2724: 2723: 2719:Ninjemys oweni 2714: 2709: 2707: 2704: 2703: 2700: 2699: 2690: 2685: 2683: 2679: 2678: 2675: 2674: 2670: 2669: 2666: 2665: 2661: 2660: 2657: 2656: 2647: 2644: 2643: 2640: 2639: 2630: 2625: 2623: 2620: 2619: 2616: 2615: 2606: 2601: 2599: 2594: 2592: 2589: 2588: 2585: 2584: 2580: 2579: 2576: 2575: 2566: 2563: 2562: 2559: 2558: 2549: 2544: 2542: 2537: 2535: 2532: 2531: 2528: 2527: 2518: 2513: 2511: 2509:Meiolaniformes 2505: 2504: 2501: 2500: 2496: 2495: 2492: 2491: 2484: 2481: 2480: 2477: 2476: 2469: 2464: 2462: 2457: 2455: 2450: 2322: 2321: 2319: 2315: 2314: 2311: 2310: 2306: 2305: 2302: 2301: 2297: 2296: 2293: 2292: 2288: 2287: 2284: 2283: 2279: 2278: 2275: 2274: 2270: 2269: 2266: 2265: 2256: 2253: 2252: 2249: 2248: 2239: 2234: 2232: 2229: 2228: 2225: 2224: 2215: 2210: 2208: 2205: 2204: 2201: 2200: 2191: 2186: 2184: 2181: 2180: 2177: 2176: 2167: 2162: 2160: 2157: 2156: 2153: 2152: 2143: 2138: 2136: 2131: 2129: 2125: 2124: 2121: 2120: 2116: 2115: 2112: 2111: 2107: 2106: 2103: 2102: 2098: 2097: 2094: 2093: 2089: 2088: 2085: 2084: 2080: 2079: 2076: 2075: 2071: 2070: 2067: 2066: 2057: 2054: 2053: 2050: 2049: 2044: 2039: 2037: 2034: 2033: 2030: 2029: 2020: 2015: 2013: 2010: 2009: 2006: 2005: 1996: 1991: 1989: 1986: 1985: 1982: 1981: 1972: 1967: 1965: 1962: 1961: 1958: 1957: 1948: 1943: 1941: 1938: 1937: 1934: 1933: 1924: 1919: 1917: 1912: 1900:M. brevicollis 1856: 1855: 1852: 1851: 1847: 1846: 1843: 1842: 1838: 1837: 1834: 1833: 1829: 1828: 1825: 1824: 1820: 1819: 1816: 1815: 1806: 1803: 1802: 1799: 1798: 1789: 1786: 1785: 1782: 1781: 1772: 1767: 1765: 1762: 1761: 1758: 1757: 1748: 1745: 1744: 1741: 1740: 1736:Ninjemys oweni 1731: 1726: 1724: 1721: 1720: 1717: 1716: 1707: 1704: 1703: 1700: 1699: 1690: 1685: 1683: 1677: 1675: 1670: 1584: 1581: 1460: 1459: 1450: 1449: 1448: 1439: 1438: 1437: 1436: 1435: 1422:M. brevicollis 1414:M. brevicollis 1357: 1354: 1270: 1267: 1264: 1263: 1261: 1251: 1237: 1234: 1229: 1221: 1220: 1211: 1196: 1182: 1179: 1174: 1166: 1165: 1156: 1139:Named for the 1137: 1123: 1120: 1118:Ninjemys oweni 1115: 1107: 1106: 1091: 1077: 1074: 1070: 1069: 1059: 1046: 1043: 1037: 1036: 1024: 1010: 1007: 1001: 1000: 992: 978: 975: 968: 967: 958: 947:M. brevicollis 943: 929: 926: 921: 911: 910: 908: 897: 883: 880: 875: 867: 866: 863: 860: 857: 854: 851: 842: 839: 683:Walpole Island 617:as defined by 591:Juliana Sterli 513: 510: 423:Meiolaniformes 346: 345: 344: 343: 335: 327: 319: 311: 300: 299: 295: 294: 279: 275: 274: 272:Meiolaniformes 266: 259: 258: 253: 246: 245: 240: 233: 232: 230:Rhaptochelydia 227: 220: 219: 214: 207: 206: 201: 194: 193: 188: 184: 183: 178: 174: 173: 168: 164: 163: 158: 154: 153: 140: 139: 130: 129: 121: 120: 116: 115: 107: 106: 98: 93: 88: 83: 78: 73: 68: 63: 58: 53: 48: 43: 42: 37:48–0.003 26: 15: 13: 10: 9: 6: 4: 3: 2: 6600: 6589: 6586: 6584: 6581: 6579: 6576: 6574: 6571: 6569: 6566: 6564: 6561: 6559: 6556: 6554: 6551: 6549: 6546: 6545: 6543: 6526: 6521: 6517: 6513: 6508: 6504: 6500: 6495: 6491: 6487: 6482: 6478: 6474: 6469: 6465: 6460: 6454: 6450: 6445: 6439: 6435: 6434: 6432: 6430: 6426: 6422: 6417: 6405: 6397: 6395: 6392: 6390: 6387: 6385: 6382: 6380: 6376: 6375: 6372: 6362: 6355: 6334: 6330: 6329: 6328: 6327: 6325: 6323: 6319: 6308: 6307: 6303: 6301: 6300: 6296: 6294: 6293: 6289: 6287: 6286: 6285:Brachyopsemys 6282: 6280: 6279: 6275: 6274: 6272: 6270: 6266: 6260: 6259: 6255: 6253: 6252: 6248: 6246: 6245: 6241: 6238: 6237: 6236:Jurassichelon 6233: 6230: 6229: 6225: 6223: 6222: 6218: 6216: 6215: 6211: 6210: 6208: 6205: 6200: 6194: 6193: 6189: 6187: 6186: 6182: 6180: 6179: 6175: 6173: 6172: 6168: 6166: 6165: 6161: 6159: 6158: 6154: 6152: 6151: 6147: 6146: 6144: 6141: 6140:Eurysternidae 6136: 6129: 6128: 6127:Protostegidae 6124: 6122: 6121: 6117: 6115: 6114: 6110: 6108: 6107: 6103: 6101: 6100: 6096: 6094: 6093: 6089: 6087: 6086: 6082: 6080: 6079: 6075: 6073: 6072: 6068: 6067: 6064: 6061: 6056: 6052: 6045: 6038: 6032: 6031: 6027: 6025: 6024: 6020: 6018: 6017: 6013: 6011: 6010: 6006: 6004: 6003: 5999: 5997: 5996: 5992: 5990: 5989: 5985: 5983: 5982: 5978: 5976: 5975: 5971: 5969: 5968: 5964: 5962: 5961: 5957: 5955: 5954: 5950: 5948: 5947: 5943: 5941: 5940: 5936: 5934: 5933: 5929: 5927: 5926: 5922: 5920: 5919: 5915: 5913: 5912: 5908: 5906: 5905: 5901: 5899: 5898: 5894: 5892: 5891: 5890:Changmachelys 5887: 5885: 5884: 5880: 5878: 5877: 5873: 5871: 5870: 5866: 5864: 5863: 5859: 5858: 5856: 5854: 5853:Macrobaenidae 5849: 5842: 5836: 5835: 5831: 5829: 5828: 5824: 5822: 5821: 5817: 5815: 5814: 5810: 5808: 5807: 5803: 5801: 5800: 5796: 5794: 5793: 5789: 5787: 5786: 5782: 5780: 5779: 5775: 5773: 5772: 5768: 5766: 5765: 5761: 5759: 5758: 5757:Chengyuchelys 5754: 5752: 5751: 5747: 5745: 5744: 5740: 5738: 5737: 5733: 5732: 5730: 5728: 5721: 5711: 5710: 5709:Trinitichelys 5706: 5704: 5703: 5699: 5697: 5696: 5692: 5690: 5689: 5685: 5683: 5682: 5678: 5676: 5675: 5671: 5669: 5668: 5664: 5662: 5661: 5657: 5655: 5654: 5650: 5648: 5647: 5643: 5641: 5640: 5636: 5634: 5633: 5629: 5627: 5626: 5622: 5620: 5619: 5618:Gehennachelys 5615: 5613: 5612: 5608: 5606: 5605: 5601: 5599: 5598: 5594: 5592: 5591: 5587: 5585: 5584: 5580: 5578: 5577: 5573: 5571: 5570: 5566: 5564: 5563: 5559: 5557: 5556: 5552: 5550: 5549: 5545: 5544: 5542: 5540: 5536: 5530: 5529: 5525: 5523: 5522: 5518: 5515: 5514: 5510: 5507: 5506: 5502: 5500: 5499: 5498:Pleurosternon 5495: 5493: 5492: 5488: 5486: 5485: 5484:Dorsetochelys 5481: 5479: 5478: 5474: 5473: 5471: 5469: 5465: 5458: 5457: 5453: 5450: 5449: 5448:Compsemydidae 5445: 5444: 5441: 5438: 5436: 5429: 5423: 5422: 5418: 5415: 5414: 5410: 5407: 5406: 5402: 5400: 5399: 5395: 5392: 5391: 5390:Kallokibotion 5387: 5385: 5384: 5380: 5378: 5377: 5376:Calissounemys 5373: 5371: 5370: 5366: 5365: 5363: 5361: 5360:Compsemydidae 5354: 5348: 5347: 5343: 5341: 5340: 5336: 5334: 5333: 5329: 5327: 5326: 5322: 5320: 5319: 5315: 5314: 5312: 5310: 5303: 5297: 5296: 5295:Sichuanchelys 5292: 5290: 5289: 5288:Mongolochelys 5285: 5283: 5282: 5278: 5277: 5275: 5273: 5266: 5256: 5255: 5251: 5249: 5248: 5244: 5242: 5241: 5237: 5235: 5234: 5230: 5228: 5227: 5223: 5222: 5220: 5218: 5214: 5208: 5207: 5203: 5201: 5200: 5199:Peligrochelys 5196: 5194: 5193: 5192:Patagoniaemys 5189: 5187: 5186: 5182: 5180: 5179: 5175: 5174: 5171: 5168: 5166: 5159: 5153: 5152: 5148: 5146: 5145: 5144:Palaeochersis 5141: 5139: 5138: 5134: 5133: 5131: 5129: 5122: 5116: 5115: 5108: 5106: 5105: 5098: 5096: 5095: 5094:Proganochelys 5088: 5086: 5085: 5084:Kayentachelys 5078: 5076: 5075: 5074:Kallokibotion 5068: 5066: 5065: 5058: 5056: 5055: 5054:Heckerochelys 5048: 5046: 5045: 5038: 5036: 5035: 5028: 5026: 5025: 5018: 5017: 5014: 5011: 5008: 5003: 4996: 4995: 4988: 4986: 4985: 4978: 4975: 4974: 4967: 4965: 4964: 4957: 4956: 4953: 4950: 4947: 4946:Pantestudines 4942: 4938: 4926: 4925: 4924:Pantestudines 4919: 4906: 4905: 4904: 4903: 4901: 4899: 4898:Pantestudines 4895: 4887: 4883: 4882: 4881: 4880: 4877: 4874: 4872: 4868: 4864: 4856: 4855:Archelosauria 4852: 4849: 4847: 4843: 4840: 4838: 4834: 4831: 4829: 4825: 4823: 4819: 4817: 4813: 4812: 4807: 4803: 4799: 4798:Pantestudines 4792: 4787: 4785: 4780: 4778: 4773: 4772: 4769: 4758: 4752: 4744: 4742:9780128175552 4738: 4734: 4733: 4725: 4722: 4717: 4713: 4709: 4702: 4699: 4693: 4688: 4684: 4680: 4676: 4672: 4668: 4661: 4659: 4657: 4653: 4648: 4644: 4640: 4633: 4630: 4625: 4621: 4617: 4610: 4607: 4602: 4598: 4594: 4590: 4586: 4582: 4578: 4574: 4570: 4563: 4561: 4557: 4552: 4550:9781421428680 4546: 4542: 4541: 4533: 4530: 4524: 4519: 4515: 4511: 4508:(3): 93–106. 4507: 4503: 4496: 4493: 4488: 4484: 4480: 4476: 4472: 4468: 4464: 4457: 4454: 4449: 4445: 4441: 4437: 4433: 4429: 4425: 4421: 4414: 4412: 4410: 4408: 4406: 4404: 4402: 4398: 4393: 4387: 4376: 4372: 4368: 4364: 4358: 4354: 4350: 4346: 4342: 4335: 4327: 4324: 4319: 4315: 4311: 4307: 4303: 4299: 4295: 4289: 4287: 4283: 4278: 4274: 4269: 4264: 4260: 4256: 4252: 4248: 4241: 4239: 4237: 4233: 4228: 4224: 4220: 4216: 4212: 4208: 4200: 4198: 4194: 4189: 4185: 4181: 4177: 4173: 4169: 4165: 4161: 4154: 4152: 4143: 4141: 4139: 4137: 4135: 4133: 4129: 4124: 4120: 4116: 4109: 4106: 4101: 4097: 4093: 4086: 4083: 4078: 4074: 4069: 4064: 4060: 4056: 4051: 4046: 4042: 4038: 4034: 4030: 4026: 4019: 4017: 4015: 4013: 4009: 4004: 4000: 3995: 3990: 3985: 3980: 3976: 3972: 3968: 3964: 3963: 3958: 3954: 3953:Worthy, T. H. 3947: 3945: 3943: 3939: 3934: 3930: 3923: 3919: 3913: 3911: 3909: 3907: 3903: 3898: 3894: 3891:(2): 93–106. 3890: 3886: 3885: 3877: 3875: 3873: 3871: 3867: 3862: 3858: 3854: 3850: 3846: 3842: 3835: 3833: 3831: 3829: 3827: 3825: 3823: 3821: 3819: 3817: 3813: 3808: 3804: 3800: 3796: 3789: 3786: 3781: 3777: 3773: 3769: 3765: 3758: 3756: 3752: 3747: 3743: 3736: 3729: 3726: 3721: 3714: 3707: 3704: 3699: 3693: 3689: 3685: 3681: 3674: 3672: 3668: 3663: 3659: 3655: 3651: 3644: 3637: 3634: 3629: 3622: 3619: 3614: 3610: 3606: 3602: 3601: 3593: 3589: 3583: 3581: 3577: 3571: 3566: 3562: 3558: 3554: 3547: 3544: 3539: 3535: 3530: 3525: 3521: 3517: 3513: 3509: 3505: 3498: 3496: 3494: 3492: 3490: 3488: 3486: 3484: 3482: 3480: 3478: 3476: 3474: 3472: 3468: 3462: 3457: 3452: 3447: 3443: 3439: 3435: 3428: 3426: 3424: 3422: 3420: 3418: 3416: 3414: 3412: 3410: 3406: 3401: 3397: 3393: 3389: 3385: 3381: 3377: 3371: 3368: 3362: 3357: 3353: 3349: 3345: 3341: 3340: 3335: 3331: 3330:Owen, Richard 3325: 3323: 3319: 3314: 3308: 3304: 3297: 3294: 3289: 3285: 3281: 3277: 3270: 3268: 3264: 3255: 3252: 3248:(3049): 1–10. 3247: 3240: 3238: 3234: 3225: 3223: 3221: 3219: 3217: 3215: 3213: 3211: 3207: 3200: 3198: 3194: 3192: 3187: 3183: 3179: 3178:Lapita people 3175: 3171: 3166: 3164: 3158: 3156: 3152: 3148: 3139: 3137: 3135: 3131: 3126: 3123: 3119: 3115: 3111: 3108:Study on the 3102: 3095: 3093: 3091: 3087: 3082: 3078: 3074: 3070: 3062: 3060: 3058: 3054: 3048: 3046: 3041: 3036: 3034: 3030: 3026: 3022: 3018: 3014: 3010: 3006: 2990: 2981: 2969: 2964: 2962: 2960: 2956: 2952: 2948: 2942: 2939: 2938:Lapita people 2933: 2931: 2927: 2923: 2919: 2915: 2911: 2906: 2898: 2894: 2891: 2887: 2886:South Pacific 2883: 2878: 2876: 2872: 2868: 2865: 2861: 2857: 2853: 2852:ghost lineage 2849: 2845: 2837: 2830: 2829: 2821: 2820: 2812: 2811: 2803: 2802: 2794: 2793: 2785: 2784: 2776: 2775: 2767: 2766: 2763: 2762: 2761: 2754: 2753: 2750: 2749: 2746: 2745: 2744: 2737: 2736: 2730: 2729: 2726: 2725: 2722: 2721: 2720: 2713: 2712: 2706: 2705: 2702: 2701: 2698: 2697: 2696: 2689: 2688: 2682:Meiolaniidae 2681: 2680: 2677: 2676: 2668: 2667: 2659: 2658: 2655: 2654: 2653: 2646: 2645: 2642: 2641: 2638: 2637: 2636: 2629: 2628: 2622: 2621: 2618: 2617: 2614: 2613: 2612: 2605: 2604: 2598: 2597: 2591: 2590: 2587: 2586: 2578: 2577: 2574: 2573: 2572: 2565: 2564: 2561: 2560: 2557: 2556: 2555: 2548: 2547: 2541: 2540: 2534: 2533: 2530: 2529: 2526: 2525: 2524: 2517: 2516: 2510: 2507: 2506: 2503: 2502: 2494: 2493: 2490: 2489: 2483: 2482: 2479: 2478: 2475: 2474: 2468: 2467: 2461: 2460: 2454: 2453: 2449: 2447: 2443: 2442:plesiomorphic 2438: 2437:Peligrochelys 2434: 2433:Mongolochelys 2430: 2429: 2428:Peligrochelys 2424: 2423: 2422:Patagoniaemys 2418: 2417: 2412: 2411: 2406: 2402: 2401:Mongolochelys 2398: 2394: 2393: 2392:Kallokibotion 2388: 2387: 2386:Mongolochelys 2380: 2378: 2374: 2370: 2365: 2360: 2356: 2352: 2351:podocnemidids 2348: 2344: 2340: 2339: 2333: 2329: 2313: 2312: 2304: 2303: 2295: 2294: 2286: 2285: 2277: 2276: 2268: 2267: 2264: 2263: 2262: 2255: 2254: 2251: 2250: 2247: 2246: 2245: 2238: 2237: 2231: 2230: 2227: 2226: 2223: 2222: 2221: 2214: 2213: 2207: 2206: 2203: 2202: 2199: 2198: 2197: 2190: 2189: 2183: 2182: 2179: 2178: 2175: 2174: 2173: 2166: 2165: 2159: 2158: 2155: 2154: 2151: 2150: 2149: 2142: 2141: 2135: 2134: 2123: 2122: 2114: 2113: 2105: 2104: 2096: 2095: 2087: 2086: 2078: 2077: 2069: 2068: 2065: 2064: 2063: 2056: 2055: 2052: 2051: 2048: 2043: 2042: 2036: 2035: 2032: 2031: 2028: 2027: 2026: 2019: 2018: 2012: 2011: 2008: 2007: 2004: 2003: 2002: 1995: 1994: 1988: 1987: 1984: 1983: 1980: 1979: 1978: 1971: 1970: 1964: 1963: 1960: 1959: 1956: 1955: 1954: 1947: 1946: 1940: 1939: 1936: 1935: 1932: 1931: 1930: 1923: 1922: 1916: 1915: 1907: 1905: 1901: 1897: 1893: 1888: 1884: 1880: 1876: 1868: 1863: 1854: 1853: 1845: 1844: 1836: 1835: 1827: 1826: 1818: 1817: 1814: 1813: 1812: 1805: 1804: 1801: 1800: 1797: 1796: 1795: 1788: 1787: 1784: 1783: 1780: 1779: 1778: 1771: 1770: 1764: 1763: 1760: 1759: 1756: 1755: 1754: 1747: 1746: 1743: 1742: 1739: 1738: 1737: 1730: 1729: 1723: 1722: 1719: 1718: 1715: 1714: 1713: 1706: 1705: 1702: 1701: 1698: 1697: 1696: 1689: 1688: 1682:Meiolaniidae 1681: 1680: 1674: 1673: 1669: 1667: 1662: 1660: 1656: 1652: 1648: 1644: 1640: 1636: 1632: 1628: 1624: 1620: 1616: 1612: 1608: 1604: 1600: 1595: 1590: 1582: 1580: 1577: 1573: 1569: 1565: 1560: 1558: 1554: 1550: 1546: 1542: 1538: 1534: 1530: 1526: 1522: 1518: 1514: 1509: 1507: 1503: 1499: 1495: 1491: 1487: 1483: 1479: 1475: 1464: 1454: 1443: 1434: 1432: 1427: 1423: 1419: 1415: 1411: 1407: 1403: 1399: 1395: 1391: 1387: 1383: 1377: 1375: 1367: 1362: 1355: 1353: 1351: 1347: 1343: 1339: 1335: 1331: 1330:New Caledonia 1327: 1322: 1320: 1316: 1312: 1308: 1304: 1300: 1296: 1292: 1288: 1284: 1280: 1275: 1268: 1262: 1259: 1255: 1252: 1249: 1238: 1235: 1233: 1230: 1228: 1227: 1223: 1222: 1216: 1212: 1209: 1205: 1200: 1197: 1194: 1183: 1180: 1178: 1175: 1173: 1172: 1168: 1167: 1161: 1157: 1154: 1150: 1146: 1142: 1138: 1135: 1124: 1121: 1119: 1116: 1114: 1113: 1109: 1108: 1104: 1100: 1096: 1092: 1089: 1078: 1075: 1072: 1071: 1067: 1063: 1060: 1058: 1047: 1044: 1042: 1039: 1038: 1033: 1029: 1025: 1022: 1021:New Caledonia 1011: 1008: 1006: 1003: 1002: 997: 993: 990: 979: 976: 974: 970: 969: 963: 956: 952: 948: 944: 941: 930: 927: 925: 922: 920: 919: 918: 912: 909: 906: 902: 898: 895: 884: 881: 879: 876: 874: 873: 869: 868: 864: 861: 858: 855: 852: 849: 848: 840: 838: 836: 832: 828: 824: 823: 818: 814: 810: 806: 801: 799: 798: 793: 789: 785: 781: 775: 773: 769: 765: 761: 757: 753: 749: 745: 741: 740:Santiago Roth 737: 733: 732: 727: 723: 719: 715: 711: 703: 699: 694: 690: 688: 687:New Caledonia 684: 680: 676: 673:Australasian 671: 667: 663: 659: 655: 651: 647: 643: 639: 635: 631: 627: 622: 620: 616: 612: 608: 604: 600: 594: 592: 588: 584: 580: 576: 572: 568: 564: 560: 550: 546: 544: 540: 536: 535: 530: 526: 525: 520: 511: 509: 507: 506:mating season 503: 498: 496: 492: 486: 484: 480: 476: 472: 468: 464: 463: 458: 457: 452: 451: 446: 443: 439: 438: 433: 432: 426: 424: 420: 416: 412: 411:New Caledonia 408: 404: 400: 396: 392: 391: 386: 382: 379: 375: 371: 367: 366:South America 363: 362:clubbed tails 359: 356: 352: 342: 341: 336: 334: 333: 328: 326: 325: 320: 318: 317: 312: 310: 309: 304: 303: 301: 296: 291: 286: 280: 277: 276: 273: 267: 264: 261: 260: 257: 254: 251: 248: 247: 244: 241: 238: 235: 234: 231: 228: 225: 222: 221: 218: 215: 212: 209: 208: 205: 204:Pantestudines 202: 199: 196: 195: 192: 189: 186: 185: 182: 179: 176: 175: 172: 169: 166: 165: 162: 159: 156: 155: 150: 145: 141: 138: 137: 131: 127: 122: 117: 113: 108: 101: 96: 91: 86: 81: 76: 71: 66: 61: 56: 51: 46: 40: 34: 30: 29:Middle Eocene 22: 19: 6459:Meiolaniidae 6429:Meiolaniidae 6428: 6394:Perichelydia 6389:Mesochelydia 6304: 6297: 6290: 6283: 6278:Angolachelys 6276: 6256: 6251:Portlandemys 6249: 6244:Plesiochelys 6242: 6234: 6228:Hylaeochelys 6226: 6219: 6212: 6190: 6185:Palaeomedusa 6183: 6176: 6169: 6162: 6155: 6148: 6125: 6118: 6111: 6104: 6097: 6090: 6083: 6076: 6069: 6054: 6050: 6028: 6021: 6014: 6007: 6000: 5993: 5986: 5979: 5972: 5965: 5958: 5951: 5944: 5937: 5932:Jeholochelys 5930: 5923: 5916: 5911:Galvechelone 5909: 5902: 5895: 5888: 5883:Aurorachelys 5881: 5874: 5869:Appalachemys 5867: 5860: 5832: 5825: 5818: 5813:Tienfuchelys 5811: 5804: 5797: 5790: 5785:Phunoichelys 5783: 5776: 5769: 5762: 5755: 5750:Camerochelys 5748: 5743:Brodiechelys 5741: 5734: 5707: 5700: 5695:Stygiochelys 5693: 5686: 5679: 5672: 5665: 5658: 5651: 5644: 5637: 5630: 5623: 5616: 5609: 5602: 5595: 5588: 5581: 5574: 5567: 5560: 5555:Arvinachelys 5553: 5546: 5526: 5519: 5511: 5503: 5496: 5489: 5482: 5475: 5454: 5446: 5419: 5411: 5403: 5396: 5388: 5381: 5374: 5367: 5344: 5337: 5330: 5325:Helochelydra 5323: 5318:Aragochersis 5316: 5293: 5286: 5279: 5252: 5245: 5238: 5231: 5226:Gaffneylania 5224: 5217:Meiolaniidae 5216: 5204: 5197: 5190: 5183: 5176: 5149: 5142: 5135: 5112: 5102: 5092: 5082: 5072: 5062: 5052: 5044:Eileanchelys 5042: 5034:Condorchelys 5032: 5024:Chinlechelys 5022: 4992: 4984:Odontochelys 4982: 4973:Eunotosaurus 4971: 4961: 4922: 4850: 4841: 4832: 4731: 4724: 4707: 4701: 4674: 4670: 4646: 4642: 4632: 4623: 4619: 4609: 4576: 4572: 4539: 4532: 4505: 4501: 4495: 4470: 4466: 4456: 4427: 4423: 4386:cite journal 4344: 4340: 4326: 4301: 4297: 4250: 4246: 4210: 4206: 4163: 4159: 4150: 4122: 4118: 4108: 4099: 4095: 4085: 4035:(1): 38317. 4032: 4028: 3966: 3960: 3932: 3928: 3888: 3882: 3844: 3798: 3788: 3771: 3767: 3745: 3741: 3728: 3719: 3706: 3679: 3653: 3649: 3636: 3627: 3621: 3604: 3598: 3588:Anderson, C. 3560: 3556: 3546: 3514:(1): 21–45. 3511: 3507: 3441: 3437: 3383: 3379: 3370: 3343: 3337: 3302: 3296: 3282:(2): 85–89. 3279: 3275: 3266: 3265:, Owen, and 3262: 3254: 3245: 3236: 3232: 3195: 3190: 3169: 3167: 3163:last ice age 3159: 3143: 3133: 3129: 3127: 3121: 3107: 3089: 3066: 3049: 3044: 3039: 3037: 3028: 3024: 3016: 3012: 3008: 3002: 2965:Paleobiology 2946: 2943: 2934: 2903: 2889: 2879: 2875:Gaffneylania 2874: 2870: 2841: 2758: 2757: 2741: 2740: 2717: 2716: 2693: 2692: 2650: 2649: 2633: 2632: 2609: 2608: 2569: 2568: 2552: 2551: 2521: 2520: 2486: 2471: 2436: 2432: 2426: 2420: 2414: 2408: 2400: 2396: 2390: 2384: 2381: 2355:pelomedusids 2342: 2336: 2325: 2259: 2258: 2242: 2241: 2218: 2217: 2194: 2193: 2170: 2169: 2146: 2145: 2060: 2059: 2046: 2023: 2022: 1999: 1998: 1975: 1974: 1951: 1950: 1927: 1926: 1903: 1899: 1895: 1892:M. platyceps 1891: 1886: 1883:M. platyceps 1882: 1878: 1874: 1872: 1866: 1809: 1808: 1792: 1791: 1775: 1774: 1751: 1750: 1734: 1733: 1710: 1709: 1693: 1692: 1666:Gaffneylania 1665: 1663: 1658: 1654: 1650: 1646: 1642: 1638: 1634: 1630: 1626: 1623:Gaffneylania 1622: 1614: 1606: 1602: 1598: 1593: 1589:monophyletic 1586: 1575: 1571: 1567: 1563: 1561: 1557:Gaffneylania 1556: 1552: 1544: 1541:Gaffneylania 1540: 1536: 1516: 1512: 1510: 1505: 1501: 1485: 1481: 1477: 1473: 1470: 1462: 1430: 1425: 1421: 1417: 1413: 1409: 1405: 1401: 1397: 1393: 1390:Gaffneylania 1389: 1381: 1378: 1371: 1365: 1341: 1326:Pindai Caves 1323: 1318: 1294: 1290: 1286: 1276: 1272: 1257: 1253: 1231: 1224: 1207: 1203: 1198: 1176: 1169: 1152: 1148: 1144: 1117: 1110: 1102: 1099:M. platyceps 1098: 1094: 1076:Pleistocene 1065: 1061: 1045:Pleistocene 1040: 1032:M. platyceps 1031: 1027: 1009:Pleistocene 1004: 995: 972: 955:M. platyceps 954: 950: 946: 923: 915: 914: 904: 901:Gaffneylania 900: 877: 872:Gaffneylania 870: 834: 822:Gaffneylania 820: 804: 802: 795: 791: 787: 783: 779: 776: 771: 767: 759: 755: 751: 747: 743: 729: 707: 697: 678: 674: 669: 666:M. platyceps 665: 661: 657: 653: 649: 633: 625: 623: 610: 606: 598: 595: 578: 574: 570: 566: 563:M. platyceps 562: 558: 555: 543:thorny devil 532: 529:Richard Owen 522: 515: 499: 494: 490: 487: 460: 454: 448: 437:Gaffneylania 435: 429: 427: 388: 351:Meiolaniidae 350: 349: 338: 330: 322: 314: 308:Gaffneylania 306: 285:Meiolaniidae 284: 262: 256:Perichelydia 249: 243:Mesochelydia 236: 223: 210: 197: 134: 133:Skeleton of 24:Meiolaniidae 18: 6453:Wikispecies 6292:Leyvachelys 6269:Sandownidae 6221:Globochelus 6164:Eurysternum 6120:Thalassemys 6099:Neusticemys 5897:Dracochelys 5848:Sinemydidae 5834:Yanduchelys 5806:Siamochelys 5799:Shartegemys 5771:Kalasinemys 5764:Jastmelchyi 5667:Plesiobaena 5653:Palatobaena 5646:Neurankylus 5611:Gamerabaena 5590:Denazinemys 5548:Arundelemys 5398:Peltochelys 5369:Berruchelus 5339:Naomichelys 5007:Testudinata 4994:Pappochelys 4692:11336/63435 4523:11336/84233 3935:(1): 35–48. 3563:: 181–191. 3529:11336/21194 3461:11336/41594 3346:: 471–480. 3191:M. damelipi 2959:New Zealand 2926:East Africa 2446:crown group 2405:sinochelyds 2375:as well as 2373:sea turtles 2332:cryptodires 2328:pleurodires 1611:basicranium 1498:ankylosaurs 1494:glyptodonts 1386:ceratopsian 1356:Description 1350:New Zealand 1338:Tiga Island 1202:Formation. 996:M. damelipi 601:was named, 479:New Zealand 467:Australasia 415:Pleistocene 413:during the 403:Pleistocene 378:crown group 374:cryptodires 370:Australasia 364:known from 217:Testudinata 6542:Categories 6384:Ankylopoda 6377:See also: 6333:Testudines 6322:Testudines 6258:Tropidemys 6192:Parachelys 6178:Idiochelys 6171:Hydropelta 6157:Chelonides 6016:Xiaochelys 5981:Osteopygis 5960:Macrobaena 5953:Liaochelys 5946:Kirgizemys 5939:Judithemys 5876:Asiachelys 5862:Anatolemys 5820:Undjulemys 5778:Larachelus 5681:Saxochelys 5674:Protobaena 5583:Chisternon 5576:Cedrobaena 5505:Riodevemys 5477:Dinochelys 5405:Riodevemys 5332:Helochelys 5254:Warkalania 5178:Chubutemys 5151:Waluchelys 5114:Spoochelys 5064:Indochelys 4907:see below↓ 4886:Sauropsida 4871:Sauropsida 4828:Sauropsida 4649:: 363–368. 4347:: 11, 23. 4268:11336/3254 4125:(107–113). 4102:: 285–297. 3201:References 3140:Extinction 3081:palm trees 3040:Meiolania' 2488:Cryptodira 2473:Pleurodira 2416:Chubutemys 2196:Warkalania 1977:Warkalania 1896:M. mackayi 1879:M. mackayi 1655:Warkalania 1639:Warkalania 1619:sinemydids 1549:Osteoderms 1431:Warkalania 1319:Warkalania 1291:Warkalania 1258:Warkalania 1254:Warkalania 1226:Warkalania 797:Warkalania 679:M. mackayi 638:Pleurodira 475:Antarctica 456:Warkalania 440:native to 401:until the 381:Testudines 355:stem-group 340:Warkalania 6306:Solnhofia 6299:Sandownia 6150:Achelonia 6085:Euryaspis 6030:Yumenemys 5974:Ordosemys 5925:Hoyasemys 5702:Thescelus 5688:Scabremys 5639:Lakotemys 5625:Goleremys 5513:Selenemys 5413:Selenemys 5383:Compsemys 5233:Meiolania 5185:Otwayemys 4814:Kingdom: 4751:cite book 4626:: 55–135. 4620:Zoologica 4593:0272-4634 4444:0268-0130 4371:1088-7105 4304:: 72–76. 4227:131795055 4059:2045-2322 3861:0003-0090 3853:2246/1670 3807:2246/5279 3386:: 43–48. 3267:Meiolania 3263:Megalania 3155:latitudes 3134:Meiolania 3088:of ? 3045:Meiolania 3029:Meiolania 3025:Meiolania 3017:Meiolania 3013:Meiolania 3009:Meiolania 2970:Lifestyle 2955:Zealandia 2930:barnacles 2916:that hit 2914:hurricane 2890:Meiolania 2410:Otwayemys 2343:Meiolania 1904:Meiolania 1887:Meiolania 1875:Meiolania 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4837:Diapsida 4822:Chordata 4820:Phylum: 4816:Animalia 4601:88998996 4487:85295987 4473:: 3–45. 4448:Archived 4375:Archived 4318:84241716 4277:83804365 4188:83503956 4077:27922064 4003:20713711 3801:(2805). 3780:2246/978 3590:(1925). 3538:83799914 3233:Ninjemys 3122:Niolamia 3110:endocast 2871:Niolamia 2867:Gondwana 2383:grouped 2338:Chelydra 2172:Ninjemys 2148:Niolamia 1953:Ninjemys 1929:Niolamia 1647:Ninjemys 1635:Ninjemys 1631:Niolamia 1627:Niolamia 1607:Niolamia 1603:Ninjemys 1599:Niolamia 1594:Niolamia 1576:Ninjemys 1572:Niolamia 1537:Niolamia 1521:carapace 1502:Ninjemys 1482:Ninjemys 1478:Ninjemys 1474:Niolamia 1406:Niolamia 1402:Ninjemys 1398:Ninjemys 1394:Niolamia 1382:Niolamia 1366:Niolamia 1287:Niolamia 1208:Niolamia 1199:Niolamia 1171:Niolamia 1153:Ninjemys 1149:Niolamia 1145:Ninjemys 1112:Ninjemys 928:Miocene 905:Niolamia 853:Species 835:Niolamia 809:Holocene 788:Ninjemys 772:Niolamia 768:Niolamia 760:Niolamia 748:Miolania 698:Niolamia 670:M. minor 646:hypodigm 567:M. minor 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