Meiolaniidae - Knowledge

1387:. 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. 3131:, 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 2880:. 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 6371: 2991: 3000: 2899:. 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 6364: 2943:. 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. 1253: 1139: 1093: 945: 137: 3112: 2908: 1453: 3172:

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

1062: 1026: 1873: 6410: 1464: 3143:, 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 1198: 899: 560: 3095:

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

3070:, 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. 1226: 1171: 123: 704: 1491:. 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 781:. 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 4458: 2964:. 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 2345:

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

793:, 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 567:

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

2459:. 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. 4430: 3176:, 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. 2903:

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.

2972:, 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. 1607:

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).

620:, 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 1444:. 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. 1900:

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

2968:. 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 3932: 2372:

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.

1079:, 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. 3207:

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.

3653: 1328:(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 3602: 542:. 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 4215:

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.

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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)".

4678:"Comparative neuroanatomy of extinct horned turtles (Meiolaniidae) and extant terrestrial turtles (Testudinidae), with comments on the palaeobiological implications of selected endocranial features" 3014:

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

3168:, 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. 3049:

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

532:, 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 4717:

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".

753:, 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 488:. 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 1888:

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.

727:. 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 1427:

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

4385: 3445:"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)" 436:, which contains more primitive turtles species lacking the distinctive morphology of meiolaniids, known from the Early Cretaceous-Paleocene of South America and Australia. 3160:

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

3271: 4307:; Tennyson, Alan J. D.; Hand, Suzanne J.; Godthelp, Henk; Scofield, R. Paul (2011). "Terrestrial Turtle Fossils from New Zealand Refloat Moa's Ark". 6490: 1411:

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).

1640:, the fragmentary nature of the former makes it somewhat of a wildcard in phylogenetic analysis. It has been recovered as either nesting alongside 6592: 4799: 1522:

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

6370: 4431:"The enigmatic palaeoecology and palaeobiogeography of the giant, horned, fossil turtles of Australasia: a review and reanalysis of the data" 3706: 3321: 3022:

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

3157: 2352:), placing it in Cryptodira, the group that includes most living turtles and tortoises. While Boulenger agreed with the identification of 3806:"The Cervical and Caudal Vertebrae of the Cryptodiran Turtle, Melolania platyceps, from the Pleistocene of Lord Howe Island, Australia" 3136:

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

6363: 4751: 4559: 4580:"A fossil egg clutch from the stem turtle Meiolania platyceps : implications for the evolution of turtle reproductive biology" 1632:. The absence of an accessory grinding surface in the jaws also identifies it as a sister taxon to the other meiolaniids. Although 4344: 3892:

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

3775:"The cranial morphology of the extinct horned turtle, Meiolania platyceps, from the Pleistocene of Lord Howe Island, Australia" 3598: 3192: 3031: 1664:

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

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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

1317: 3389:(1859). "Description of Some Remains of a Gigantic Land-Lizard (Megalania Prisca, Owen) from Australia". 757:, the researcher concluded that Roth's turtle represented the same species, but placed both in the genus 6529: 4396: 2230: 2072: 1321: 1309: 652: 629: 584:(great roamer) to the chimeric material from the mainland, he subsequently named the Lord Howe material 5137: 1472:

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

1325: 1296:(New South Wales) seem to belong to an intermediate taxon, combining the flattened horns of taxa like 1289: 777:

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

6477: 6011: 5927: 4579: 4178: 4047: 3981: 3933:"Warkalania, a New Meiolaniid Turtle from the Tertiary Riversleigh Deposits of Queensland, Australia" 3358: 1383:

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.

3079: 2705: 1705: 741: 154: 6267: 6025: 5969: 5948: 5885: 5829: 5690: 5683: 5514: 5414: 5123: 5073: 4363: 4103:"The Wyandotte Local Fauna: A New, Dated, Pleistocene Vertebrate Fauna from Northern Queensland" 2907: 2895:, several competing ideas exist in regards to their further dispersal across the islands of the 1585:

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

1395:

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

6534: 6468: 6308: 6039: 5711: 5697: 5648: 5634: 5194: 3187:

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

2645: 2420: 1452: 1399:, while the flattened B horns extend to the sides and back. Little is known about the horns of 6516: 5736: 4747: 4599: 4555: 4450: 4377: 4367: 4083: 4065: 4009: 3928: 3867: 3702: 3317: 3096: 3087: 2915: 2892: 2374: 2254: 2035: 1917:

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

972: 964:

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

593: 6521: 6508: 5745: 5641: 5613: 5578: 5530: 4626: 3654:"On some extinct reptiles from Patagonia, of the genera Miolania, Dinilysia, and Genyodectes" 1577:, have been estimated to have reached a carapace length of 0.7 m (2 ft 4 in). 635:

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

6053: 6018: 5997: 5281: 4722: 4697: 4689: 4627:"The Galapagos tortoises in their relation to the whaling industry: a study of old logbooks" 4591: 4528: 4520: 4485: 4442: 4359: 4316: 4304: 4273: 4265: 4225: 4186: 4073: 4055: 3999: 3989: 3963: 3903: 3859: 3813: 3786: 3694: 3668: 3619: 3575: 3534: 3526: 3466: 3456: 3398: 3366: 3294: 3196: 3067: 3063: 3043: 2858: 2369: 1356: 1067: 732: 529: 493: 417: 300: 5606: 1624:

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

580:, believing it to be a small relative of the mainland specimen. As Owen had given the name 6213: 5477: 5465: 5318: 4973: 4125: 4102: 1543: 1403:

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

789:

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

746: 712: 395: 4182: 4051: 3985: 3362: 6388: 6122: 6101: 5976: 5836: 5571: 5444: 5174: 5146: 5113: 4078: 4035: 4004: 3967: 3672: 3184: 3124: 2961: 2874: 2866: 2854: 2519: 2361: 1872: 1746: 1335:

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

968:

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

693: 433: 282: 240: 6551: 6149: 6136: 5862: 5507: 5493: 5457: 5399: 5369: 5201: 5103: 5093: 5083: 4956: 4934: 4908: 4865: 4808: 4677: 4237: 3514: 3188: 2948: 2896: 2862: 2621: 2452: 2432: 2402: 2387: 2379: 2365: 1340: 1031: 750: 697: 516: 421: 376: 214: 80: 39: 4611: 4497: 4474:"Reassessment of the phylogenetic interrelationships of basal turtles (Testudinata)" 4328: 4287: 4198: 3852:"The postcranial morphology of Meiolania platyceps and a review of the Meiolaniidae" 3623: 3548: 3115:

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

1431:

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

6403: 6398: 6287: 6253: 6237: 6194: 5941: 5892: 5752: 5564: 5334: 5235: 5053: 5043: 5033: 4994: 4983: 3851: 3805: 3386: 3340: 3173: 2383: 1880:, are among the most important factors in determining their internal relationships. 1804: 1763: 1722: 1599: 1463: 1336: 882: 832: 559: 553: 539: 447: 318: 266: 253: 4726: 4595: 4229: 3774: 4489: 4269: 4190: 3156:

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

3030:

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

6462: 6301: 6278: 6173: 6129: 6108: 5857: 5676: 5662: 5655: 5620: 5599: 5378: 5348: 5016: 5004: 4693: 4676:

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

3698: 3515:"A review of the fossil record of Gondwanan turtles of the clade Meiolaniformes" 3344: 2969: 2936: 2456: 1629: 1621: 1508: 1504: 1396: 1371: 1360: 1348: 1166:

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

1108:, the Wyandotte species remains unnamed and is at times tentatively assigned to 489: 477: 425: 413: 388: 387:, they are not closely related to any living species of turtle, and lie outside 380: 227: 55: 3034:, who discovered the aforementioned leg bones, considered the possibility that 6393: 6201: 6187: 6180: 6166: 5990: 5962: 5955: 5871: 5787: 5592: 5585: 5486: 5341: 5263: 5187: 5160: 4881: 4838: 4524: 3907: 3298: 2753: 2533: 2498: 2483: 2426: 2342: 2338: 2206: 1987: 1787: 1559: 1236: 918:, its exact phylogenetic position is unclear due to the fragmentary material. 807: 648: 485: 466: 384: 365: 350: 100: 65: 6453: 4603: 4454: 4381: 4069: 3871: 3603:"Notes on the extinct Chelonian Meiolania, with a record of a new occurrence" 6315: 6159: 6094: 5983: 5934: 5522: 5422: 5392: 5242: 3994: 3165: 3091: 2965: 2940: 2924: 2853:

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

1573:

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

1500: 1355:(2011) reported on what may be the remains of a meiolaniid from the Miocene 1258: 1225: 1203: 1170: 1144: 1098: 950: 927: 904: 841: 736: 724: 625: 588:(small roamer). This has however led to some confusion, as the etymology of 549: 544: 472: 455: 405: 400: 372: 342: 171: 105: 49: 4446: 4087: 4013: 3580: 3402: 3371: 3314:

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

3038:

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

1581:

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

814:

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

3082:, feeding on a variety of low-growing plants and plant material including 1041:, Gaffney argues that the material is insufficient to distinguish it from 122: 17: 6447: 6115: 5669: 5548: 5500: 5430: 5256: 5249: 4832: 4650:"The ecology of Meiolania platyceps, a Pleistocene turtle from Australia" 4160:"Re-Description and Evolutionary Remarks on the Patagonian Horned Turtle 3940:

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

3530: 3161: 3120: 2877: 2729: 2357: 2348: 2182: 2158: 1963: 1939: 1531: 1181: 1122: 819: 656: 534: 481: 460: 441: 429: 334: 326: 191: 95: 90: 75: 70: 60: 43: 4740:

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

1415:

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

6495: 6087: 6032: 6004: 5355: 4847: 4702: 4533: 3539: 3471: 2928: 2873:, the majority of their known diversity could be found on the southern 1648:

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

1293: 999: 823: 774: 720: 703: 597: 409: 201: 110: 85: 4320: 4278: 4060: 3461: 3444: 3410: 2455:

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

1224: 1169: 971: 528:

century, with writings suggesting that various locals and visitors of

6331: 5537: 4856: 4826: 3863: 3817: 3691:

Origin, Evolution and Biogeographic History of South American Turtles

837: 504:

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

452: 391: 368: 181: 6424: 3790: 1672:

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

3689:

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

773:

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

6080: 3191:. However, there are issues with this idea. While widespread, the 3110: 3083: 2906: 2891:

While the early distribution of the family is easily explained by

1871: 1384: 1370: 702: 558: 552:, Owen came to believe that the bones represented a type of giant 4126:"A meiolaniid turtle from the Pleistocene of Northern Queensland" 3127:. The inner ear suggests that meiolaniids were more sensitive to 1332:, which is clearly differentiated through the horn core anatomy. 1292:

of Queensland. Remains found in Early Miocene Canadian Lead near

6482: 4777: 3164:

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

2857:, a group of turtles that likely evolved during or prior to the 1503:

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

1344: 1217:, another Argentinian meiolaniid, was found to be a juvenile of 827: 745:, a large meiolaniid turtle he claimed was found by his brother 432:

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

6428: 4819: 4781: 3250:(Woodward), a Horned Turtle from the Pleistocene of Queensland" 1487:, and surrounds the entire circumference of the tail in it and 3200:

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

1598:

Phylogenetic analysis consistently recovers Meiolaniidae as a

2960:, as Lord Howe Island is a volcanic island situated atop the 1308:. Other continental remains were found in the Late Oligocene 739:. Ameghino published a short communication in which he names 439:

Meiolaniidae includes a total of five different genera, with

1679:, the phylogenetic tree matches with prior work by Gaffney. 1558:

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

749:. Simultaneously, Woodward received material from collector 480:. The group is believed to have evolved on the continent of 1884:

Creating a phylogenetic tree for the individual species of

1440:

across specimen. The most reduced horns can be observed in

1005:

Known from archaeological sites on Vanuatu and Fiji, ?

616:

published a paper correcting Owen and naming the material

596:

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

4654:

New Mexico Museum of Natural History and Science Bulletin

3568:

Philosophical Transactions of the Royal Society of London

3391:

Philosophical Transactions of the Royal Society of London

1660:

is the second basalmost genus. It is clearly united with

3564:"On Parts of the Skeleton of Meiolania platyceps (Owen)" 538:

was discovered in Queensland and sent to paleontologist

492:

based on the discovery of turtle remains as part of the

4473: 3179:

Evidence for hunting may be found in the case of ?

910:

An early, but relatively poorly understood meiolaniid,

3966:; Hawkins, S.; Bedford, S.; Spriggs, M. (2010-08-16). 130:

Restoration of the head of various meiolaniid species

4578:

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

3443:

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

2424:

and meiolaniids together, as did subsequent authors.

1562:

that covered the limbs have been recovered from both

568:

was based on this material that Owen named the genus

484:

prior to its split into South America, Australia and

398:. They are best known from the last surviving genus, 6050: 5854: 5733: 5441: 5366: 5315: 5278: 5171: 5134: 5119: 5109: 5099: 5089: 5079: 5069: 5059: 5049: 5039: 5029: 5000: 4990: 4979: 4969: 4551:

The Rise of Reptiles. 320 Million Years of Evolution

4218:

Alcheringa: An Australasian Journal of Palaeontology

3895:

Alcheringa: An Australasian Journal of Palaeontology

2406:

and meiolaniids in a single group. In 2000 Hirayama

1320:

of the Riversleigh (Queensland), the Middle Miocene

818:

representing an uncertain member of this group from

364:

is an extinct family of large, probably herbivorous

6437: 6330: 6277: 6211: 6147: 6049: 5853: 5732: 5547: 5476: 5440: 5365: 5314: 5277: 5225: 5170: 5133: 5014: 4954: 4907: 4880: 4036:"Ancient tortoise hunting in the southwest Pacific" 3746:"Crossochelys, Eocene horned turtle from Patagonia" 765:). Later finds in the area would produce the taxon 1542:, instead bearing more resemblance to that of the 671:. Woodward placed it in the same genus, naming it 3519:Bulletin of the Peabody Museum of Natural History 3693:. Springer Earth System Sciences. pp. 1–5. 1876:The scale areas of meiolaniids, as seen here in 4158:Sterli, Juliana; de la Fuente, Marcelo (2011). 3724:"New reptiles from the Eocene of South America" 3661:Proceedings of the Zoological Society of London 3350:Philosophical Transactions of the Royal Society 1271:is the oldest named meiolaniid from Australia. 1010:features that do not match meiolaniid anatomy. 592:was never specified in the actual publication. 4554:. Johns Hopkins University Press. p. 53. 3438: 3436: 3434: 3432: 3430: 3428: 3426: 3424: 3422: 3420: 2869:. Although some fossil evidence may suggest a 4793: 1419:, and the B horns face straight to the side. 735:, were competing in a fashion similar to the 8: 4766:: CS1 maint: multiple names: authors list ( 4401:: CS1 maint: multiple names: authors list ( 600:", a notion later contested in the works of 4364:10.3854/crm.5.000e.fossil.checklist.v1.2015 4164:Ameghino, 1899 (Testudinata, Meiolaniidae)" 6425: 6072: 5449: 5179: 5022: 4962: 4951: 4886: 4877: 4816: 4800: 4786: 4778: 4429:Lauren E. Brown, Don Moll (October 2019). 4299: 4297: 3058:proportions, much less those of an adult. 135: 121: 31: 4701: 4682:Zoological Journal of the Linnean Society 4532: 4277: 4077: 4059: 4003: 3993: 3931:; Archer, Michael; White, Arthur (1992). 3579: 3538: 3470: 3460: 3449:Zoological Journal of the Linnean Society 3370: 3316:. Bloomington: Indiana University Press. 2911:Range of meiolaniids in the South Pacific 1112:. It has some of the largest horns among 3923: 3921: 3919: 3917: 1534:is not quite as high as seen in today's 855: 4671: 4669: 4667: 4391:from the original on 21 September 2022. 4251: 4249: 4247: 3508: 3506: 3504: 3502: 3272:"Notes on the Extinct Reptilian Genera 3217: 2331: 1644:at the base of Meiolaniidae, alongside 394:, having diverged from them around the 4759: 4573: 4571: 4424: 4422: 4420: 4418: 4416: 4414: 4412: 4394: 4210: 4208: 4153: 4151: 4149: 4147: 4145: 4143: 4029: 4027: 4025: 4023: 3957: 3955: 3953: 3887: 3885: 3883: 3881: 3845: 3843: 3841: 3839: 3837: 3835: 3833: 3831: 3829: 3827: 3768: 3766: 3744:Simpson, G.G.; Williams, C.S. (1938). 3684: 3682: 3500: 3498: 3496: 3494: 3492: 3490: 3488: 3486: 3484: 3482: 3287:Annals and Magazine of Natural History 2141: 1922: 1620:is further supported by comparing the 1075:is the type and best known species of 675:in Owen's honour. Shortly afterwards, 3639:Imprenta la Libertad (Author Edition) 3593: 3591: 3335: 3333: 3235: 3233: 3231: 3229: 3227: 3225: 3223: 3221: 3078:Meiolaniids are thought to have been 2939:, at times emanciated and covered in 1316:(South Australia), the Early Miocene 7: 4124:Gaffney, E.S.; McNamara, G. (1990). 3139:Several egg clutches are known from 2368:. Boulenger would find support from 416:, and insular species that lived on 4648:Lichtig, A.J.; Lucas, S.G. (2018). 4548:Hans-Dieter Sues (August 6, 2019). 4478:Journal of Systematic Palaeontology 4258:Journal of Systematic Palaeontology 2745: 2721: 2697: 2637: 2613: 2606: 2556: 2549: 2525: 2476: 2469: 2462: 2246: 2222: 2198: 2174: 2150: 2143: 2051: 2027: 2003: 1979: 1955: 1931: 1924: 1779: 1738: 1697: 1689: 1682: 1407:, if with more rounded tips. While 1324:(South Australia) and the Pliocene 688:species named in this period being 500:between genera. While some such as 4719:Journal of Vertebrate Paleontology 4625:Townsend, Charles Haskins (1925). 4584:Journal of Vertebrate Paleontology 4461:from the original on 18 June 2022. 4171:Journal of Vertebrate Paleontology 3673:10.1111/j.1469-7998.1901.tb08537.x 1375:The horns of a juvenile and adult 811:, a new genus with reduced horns. 25: 2436:followed in 2007 and 2011, while 1037:A potentially dubious species of 6409: 6408: 6369: 6362: 4746:. Elsevier Science. p. 30. 4130:Memoirs of the Queensland Museum 4107:Memoirs of the Queensland Museum 3611:Records of the Australian Museum 3107:Social behavior and reproduction 3090:and perhaps the fallen fruit of 2998: 2989: 2410:expanded on this idea, grouping 1462: 1451: 1251: 1196: 1137: 1091: 1060: 1024: 992: 943: 897: 572:in 1886 to include two species, 158: 3624:10.3853/j.0067-1975.14.1925.844 3193:Pleistocene overkill hypothesis 2388:pig-nosed and softshell turtles 769:, now thought to be a juvenile 371:with heavily armored heads and 6593:Taxa named by Richard Lydekker 2865:stretching as far back as the 836:being a second genus from the 655:officially split the chimeric 1: 4727:10.1080/02724634.2024.2357315 4596:10.1080/02724634.2016.1223685 4352:Chelonian Research Monographs 4230:10.1080/03115518.2016.1224441 383:. Though once believed to be 4490:10.1080/14772019.2011.558928 4270:10.1080/14772019.2012.708674 4191:10.1080/02724634.2011.615685 2442:was described in 2012. Both 1919: 1656:. As far as stable taxa go, 1628:and compares favorably with 1152:Teenage Mutant Ninja Turtles 3699:10.1007/978-3-319-00518-8_1 3204:was actually a meiolaniid. 1304:with the recurved horns of 6614: 3973:Proc. Natl. Acad. Sci. USA 3016:Allan Riverstone McCulloch 2380:pond turtles and tortoises 1921: 956:Named for its short neck, 6383: 6359: 6075: 5452: 5182: 5025: 4965: 4950: 4931: 4889: 4876: 4815: 4743:Galapagos Giant Tortoises 4694:10.1093/zoolinnean/zlw024 4525:10.1163/18759866-07903002 4472:Anquetin, Jérémy (2012). 3908:10.1080/03115519208619035 3810:American Museum Novitates 3731:American Museum Novitates 3312:Molnar, Ralph E. (2004). 3299:10.1080/00222938809460686 3257:American Museum Novitates 2935:arriving on the coast of 2931:. Other examples include 2921:Galapagos giant tortoises 2871:cosmopolitan distribution 2767: 2750: 2743: 2726: 2719: 2702: 2695: 2659: 2642: 2635: 2618: 2611: 2604: 2578: 2561: 2554: 2547: 2530: 2523: 2496: 2481: 2474: 2467: 2268: 2251: 2244: 2227: 2220: 2203: 2196: 2179: 2172: 2155: 2148: 2069: 2056: 2049: 2032: 2025: 2008: 2001: 1984: 1977: 1960: 1953: 1936: 1929: 1818: 1801: 1784: 1777: 1760: 1743: 1736: 1719: 1702: 1695: 1687: 1616:. The basal condition of 1540:galapagos giant tortoises 970: 960:is the oldest species of 924: 797:from mainland Australia, 761:(likely a misspelling of 313: 308: 155:Scientific classification 153: 143: 134: 129: 120: 34: 27:Extinct family of turtles 4513:Contributions to Zoology 3270:Woodward, A. S. (1888). 2622:Patagoniaemys gasparinae 1805:Gaffneylania auricularis 1764:Gaffneylania auricularis 1723:Gaffneylania auricularis 889:Gaffneylania auricularis 548:and the foot bones of a 458:and the remaining taxa, 4101:McNamara, G.C. (1990). 3995:10.1073/pnas.1005780107 3652:Woodward, A.S. (1901). 3240:Gaffney, E. S. (1992). 3042:lived much like modern 2933:Aldabra giant tortoises 2849:Evolution and dispersal 1892:is likely a synonym of 1636:likely lived alongside 1536:aldabra giant tortoises 1206:(Sarmiento Formation?) 641:George Albert Boulenger 4447:10.33256/hj29.4.252263 4435:Herpetological Journal 3850:Gaffney, E.S. (1996). 3804:Gaffney, E.S. (1985). 3773:Gaffney, E.S. (1983). 3722:Simpson, G.G. (1937). 3641:. La Plata, Argentina. 3581:10.1098/rstb.1888.0007 3403:10.1098/rstl.1859.0002 3372:10.1098/rstl.1886.0015 3116: 2923:swept out at sea by a 2912: 2754:Warkalania carinaminor 2663:Kallokibotion bajazidi 2646:Otwayemys cunicularius 2565:Mongolochelys efremovi 1881: 1788:Warkalania carinaminor 1380: 1351:. Furthermore, Worthy 1261:(Riversleigh Station) 1243:Warkalania carinaminor 1229: 1174: 976: 907:(Sarmiento Formation) 716: 618:Ceratochelys sthenurus 564: 6530:Paleobiology Database 3114: 3026:limbs, the idea that 2910: 2582:Peligrochelys walshae 2231:Meiolania brevicollis 2073:Meiolania brevicollis 1909:as sister taxa, with 1875: 1526:itself. The shell of 1374: 1280:Indeterminate remains 1228: 1215:Crossochelys corniger 1173: 975: 935:Meiolania brevicollis 795:Meiolania brevicollis 767:Crossochelys corniger 706: 653:Arthur Smith Woodward 630:Othniel Charles Marsh 562: 3856:Bulletin of the AMNH 3779:Bulletin of the AMNH 3753:Bulletin of the AMNH 3531:10.3374/014.056.0102 3018:, who believed that 1913:being the basalmost 1546:. Shell elements of 1318:Carl Creek Limestone 1247:Oligocene - Miocene 1104:The largest form of 707:A 1914 depiction of 523:History of discovery 513:intraspecific combat 6583:Pleistocene turtles 5802:Protoxinjiangchelys 4183:2011JVPal..31.1210S 4052:2016NatSR...638317H 3986:2010PNAS..10715512W 3980:(35): 15512–15516. 3513:Sterli, J. (2015). 3363:1886RSPT..177..471O 3343:(January 1, 1886). 3141:Meiolania platyceps 3129:low frequency sound 2958:Meiolania platyceps 2771:Meiolania platyceps 2534:Chubutemys copelloi 2272:Meiolania platyceps 2012:Meiolania platyceps 1822:Meiolania platyceps 1579:Meiolania platyceps 1524:Meiolania platyceps 1474:Meiolania platyceps 1437:Meiolania platyceps 1073:Meiolania platyceps 1052:Meiolania platyceps 857: 791:Meiolania platyceps 729:Florentino Ameghino 614:Thomas Henry Huxley 147:Meiolania platyceps 6563:Cretaceous turtles 4162:Niolamia argentina 4040:Scientific Reports 3929:Gaffney, Eugene S. 3246:, a new name for " 3181:Meiolania damelipi 3125:courtship behavior 3117: 3101:Meiolania damelipi 2913: 2706:Niolamia argentina 2058:Wyandotte species 1882: 1706:Niolamia argentina 1499:, also preserve a 1381: 1322:Wipajiri Formation 1310:Etadunna Formation 1230: 1188:Niolamia argentina 1175: 1101:(Wyandotte Creek) 1084:Wyandotte species 984:Meiolania damelipi 977: 856: 816:Meiolania damelipi 801:as a new name for 755:Niolamia argentina 742:Niolamia argentina 717: 565: 6578:Oligocene turtles 6545: 6544: 6517:Open Tree of Life 6431:Taxon identifiers 6422: 6421: 6379: 6378: 6357: 6356: 6353: 6352: 6326: 6325: 5737:Xinjiangchelyidae 5728: 5727: 5273: 5272: 5138:Australochelyidae 4946: 4945: 4942: 4941: 4927: 4926: 4321:10.1643/CH-10-113 4305:Worthy, Trevor H. 4061:10.1038/srep38317 3708:978-3-319-00517-1 3562:Owen, R. (1888). 3462:10.1111/zoj.12252 3323:978-0-253-34374-1 3097:isotopic analysis 3088:herbaceous plants 3068:softshell turtles 3064:pig-nosed turtles 2893:continental drift 2845: 2844: 2836: 2835: 2827: 2826: 2818: 2817: 2809: 2808: 2800: 2799: 2791: 2790: 2782: 2781: 2683: 2682: 2674: 2673: 2593: 2592: 2509: 2508: 2375:wastebasket taxon 2335: 2334: 2328: 2327: 2319: 2318: 2310: 2309: 2301: 2300: 2292: 2291: 2283: 2282: 2255:Meiolania mackayi 2138: 2137: 2129: 2128: 2120: 2119: 2111: 2110: 2102: 2101: 2093: 2092: 2084: 2083: 2036:Meiolania mackayi 1869: 1868: 1860: 1859: 1851: 1850: 1842: 1841: 1833: 1832: 1575:Meiolania mackayi 1277: 1276: 1133:Late Pleistocene 1034:(Walpole Island) 1016:Meiolania mackayi 731:and the other by 594:Eugene S. Gaffney 515:, perhaps during 476:being endemic to 428:and possibly the 404:, which lived in 359: 358: 304: 16:(Redirected from 6605: 6588:Pliocene turtles 6538: 6537: 6525: 6524: 6512: 6511: 6499: 6498: 6486: 6485: 6473: 6472: 6471: 6458: 6457: 6456: 6426: 6412: 6411: 6374: 6373: 6367: 6366: 6073: 6069: 6068: 6054:Thalassochelydia 6052: 5856: 5735: 5450: 5443: 5368: 5317: 5282:Sichuanchelyidae 5280: 5180: 5173: 5136: 5121: 5111: 5101: 5091: 5081: 5071: 5061: 5051: 5041: 5031: 5023: 5002: 4992: 4981: 4971: 4963: 4952: 4887: 4878: 4871: 4870: 4817: 4802: 4795: 4788: 4779: 4772: 4771: 4765: 4757: 4737: 4731: 4730: 4714: 4708: 4707: 4705: 4673: 4662: 4661: 4645: 4639: 4638: 4622: 4616: 4615: 4575: 4566: 4565: 4545: 4539: 4538: 4536: 4508: 4502: 4501: 4469: 4463: 4462: 4426: 4407: 4406: 4400: 4392: 4390: 4349: 4339: 4333: 4332: 4301: 4292: 4291: 4281: 4253: 4242: 4241: 4212: 4203: 4202: 4177:(6): 1210–1229. 4168: 4155: 4138: 4137: 4121: 4115: 4114: 4098: 4092: 4091: 4081: 4063: 4031: 4018: 4017: 4007: 3997: 3959: 3948: 3947: 3937: 3925: 3912: 3911: 3889: 3876: 3875: 3847: 3822: 3821: 3801: 3795: 3794: 3770: 3761: 3760: 3750: 3741: 3735: 3734: 3728: 3719: 3713: 3712: 3686: 3677: 3676: 3658: 3649: 3643: 3642: 3634: 3628: 3627: 3607: 3595: 3586: 3585: 3583: 3559: 3553: 3552: 3542: 3510: 3477: 3476: 3474: 3464: 3440: 3415: 3414: 3383: 3377: 3376: 3374: 3337: 3328: 3327: 3309: 3303: 3302: 3284: 3267: 3261: 3260: 3254: 3248:Meiolania" oweni 3237: 3197:invasive species 3044:snapping turtles 3032:Charles Anderson 3002: 2993: 2859:Early Cretaceous 2746: 2722: 2698: 2638: 2614: 2607: 2557: 2550: 2526: 2477: 2470: 2463: 2370:Richard Lydekker 2247: 2223: 2199: 2175: 2151: 2144: 2052: 2028: 2004: 1980: 1956: 1932: 1925: 1920: 1780: 1739: 1698: 1690: 1683: 1466: 1455: 1357:St Bathans Fauna 1326:Chinchilla Sands 1314:Namba Formations 1290:Rundle Formation 1257: 1255: 1254: 1202: 1200: 1199: 1147:(Darling Downs) 1143: 1141: 1140: 1097: 1095: 1094: 1068:Lord Howe Island 1066: 1064: 1063: 1030: 1028: 1027: 998: 996: 995: 953:(Camfield Beds) 949: 947: 946: 903: 901: 900: 858: 733:Francisco Moreno 530:Lord Howe Island 494:St Bathans Fauna 418:Lord Howe Island 299: 294: 281: 163: 162: 139: 125: 115: 52: 38:Temporal range: 32: 21: 6613: 6612: 6608: 6607: 6606: 6604: 6603: 6602: 6598:Extinct turtles 6573:Miocene turtles 6548: 6547: 6546: 6541: 6533: 6528: 6520: 6515: 6507: 6502: 6494: 6489: 6481: 6476: 6467: 6466: 6461: 6452: 6451: 6446: 6433: 6423: 6418: 6375: 6368: 6361: 6349: 6322: 6273: 6214:Plesiochelyidae 6207: 6143: 6058: 6057: 6056: 6045: 6012:Tienfucheloides 5928:Hongkongochelys 5861: 5849: 5724: 5543: 5478:Pleurosternidae 5472: 5466:Helochelydridae 5436: 5361: 5319:Helochelydridae 5310: 5269: 5221: 5166: 5129: 5019: 5010: 4974:Eorhynchochelys 4959: 4938: 4923: 4903: 4872: 4822: 4821: 4811: 4806: 4776: 4775: 4758: 4754: 4739: 4738: 4734: 4716: 4715: 4711: 4675: 4674: 4665: 4647: 4646: 4642: 4624: 4623: 4619: 4590:(6): e1223685. 4577: 4576: 4569: 4562: 4547: 4546: 4542: 4510: 4509: 4505: 4471: 4470: 4466: 4428: 4427: 4410: 4393: 4388: 4374: 4347: 4341: 4340: 4336: 4303: 4302: 4295: 4255: 4254: 4245: 4214: 4213: 4206: 4166: 4157: 4156: 4141: 4123: 4122: 4118: 4100: 4099: 4095: 4033: 4032: 4021: 3961: 3960: 3951: 3935: 3927: 3926: 3915: 3891: 3890: 3879: 3849: 3848: 3825: 3803: 3802: 3798: 3772: 3771: 3764: 3748: 3743: 3742: 3738: 3726: 3721: 3720: 3716: 3709: 3688: 3687: 3680: 3656: 3651: 3650: 3646: 3636: 3635: 3631: 3605: 3597: 3596: 3589: 3561: 3560: 3556: 3512: 3511: 3480: 3442: 3441: 3418: 3385: 3384: 3380: 3339: 3338: 3331: 3324: 3311: 3310: 3306: 3282: 3269: 3268: 3264: 3252: 3239: 3238: 3219: 3214: 3153: 3109: 3076: 3012: 3011: 3010: 3009: 3005: 3004: 3003: 2995: 2994: 2983: 2978: 2919:telling of two 2851: 2846: 2837: 2828: 2819: 2810: 2801: 2792: 2783: 2684: 2675: 2594: 2510: 2329: 2320: 2311: 2302: 2293: 2284: 2139: 2130: 2121: 2112: 2103: 2094: 2085: 1870: 1861: 1852: 1843: 1834: 1596: 1544:gopher tortoise 1480: 1479: 1478: 1477: 1469: 1468: 1467: 1458: 1457: 1456: 1369: 1282: 1252: 1250: 1197: 1195: 1138: 1136: 1092: 1090: 1061: 1059: 1025: 1023: 1002:(Efate Island) 993: 991: 944: 942: 898: 896: 854: 844:in addition to 803:Meiolania oweni 713:Heinrich Harder 673:Meiolania oweni 525: 396:Middle Jurassic 298: 292: 279: 157: 116: 114: 113: 108: 103: 98: 93: 88: 83: 78: 73: 68: 63: 58: 47: 46: 36: 28: 23: 22: 15: 12: 11: 5: 6611: 6609: 6601: 6600: 6595: 6590: 6585: 6580: 6575: 6570: 6568:Eocene turtles 6565: 6560: 6558:Meiolaniformes 6550: 6549: 6543: 6542: 6540: 6539: 6526: 6513: 6500: 6487: 6474: 6459: 6443: 6441: 6435: 6434: 6429: 6420: 6419: 6417: 6416: 6406: 6401: 6396: 6391: 6389:Angolachelonia 6384: 6381: 6380: 6377: 6376: 6360: 6358: 6355: 6354: 6351: 6350: 6348: 6347: 6346: 6345: 6336: 6334: 6328: 6327: 6324: 6323: 6321: 6320: 6312: 6305: 6298: 6291: 6283: 6281: 6275: 6274: 6272: 6271: 6264: 6257: 6250: 6242: 6234: 6227: 6224:Craspedochelys 6219: 6217: 6209: 6208: 6206: 6205: 6198: 6191: 6184: 6177: 6170: 6163: 6155: 6153: 6145: 6144: 6142: 6141: 6133: 6126: 6123:Pelobatochelys 6119: 6112: 6105: 6102:Hispaniachelys 6098: 6091: 6084: 6076: 6070: 6047: 6046: 6044: 6043: 6036: 6029: 6022: 6015: 6008: 6001: 5994: 5987: 5980: 5977:Manchurochelys 5973: 5966: 5959: 5952: 5945: 5938: 5931: 5924: 5917: 5910: 5903: 5896: 5889: 5882: 5875: 5867: 5865: 5851: 5850: 5848: 5847: 5840: 5837:Xinjiangchelys 5833: 5826: 5819: 5812: 5805: 5798: 5791: 5784: 5777: 5770: 5763: 5756: 5749: 5741: 5739: 5730: 5729: 5726: 5725: 5723: 5722: 5715: 5708: 5701: 5694: 5687: 5680: 5673: 5666: 5659: 5652: 5645: 5638: 5631: 5624: 5617: 5610: 5603: 5596: 5589: 5582: 5575: 5568: 5561: 5553: 5551: 5545: 5544: 5542: 5541: 5534: 5527: 5519: 5511: 5504: 5497: 5490: 5482: 5480: 5474: 5473: 5471: 5470: 5462: 5453: 5447: 5445:Paracryptodira 5438: 5437: 5435: 5434: 5427: 5419: 5411: 5404: 5396: 5389: 5382: 5374: 5372: 5363: 5362: 5360: 5359: 5352: 5345: 5338: 5331: 5323: 5321: 5312: 5311: 5309: 5308: 5301: 5294: 5291:Laurasichersis 5286: 5284: 5275: 5274: 5271: 5270: 5268: 5267: 5260: 5253: 5246: 5239: 5231: 5229: 5223: 5222: 5220: 5219: 5216:Trapalcochelys 5212: 5205: 5198: 5191: 5183: 5177: 5175:Meiolaniformes 5168: 5167: 5165: 5164: 5157: 5150: 5147:Australochelys 5142: 5140: 5131: 5130: 5128: 5127: 5117: 5114:Proterochersis 5107: 5097: 5087: 5077: 5067: 5057: 5047: 5037: 5026: 5020: 5015: 5012: 5011: 5009: 5008: 4998: 4988: 4977: 4966: 4960: 4955: 4948: 4947: 4944: 4943: 4940: 4939: 4932: 4929: 4928: 4925: 4924: 4922: 4921: 4920: 4919: 4913: 4911: 4905: 4904: 4902: 4901: 4900: 4899: 4890: 4884: 4874: 4873: 4869: 4868: 4859: 4850: 4841: 4835: 4829: 4820: 4813: 4812: 4807: 4805: 4804: 4797: 4790: 4782: 4774: 4773: 4752: 4732: 4709: 4688:(4): 930–950. 4663: 4640: 4617: 4567: 4560: 4540: 4503: 4464: 4441:(4): 252–263. 4408: 4373:978-0965354097 4372: 4334: 4293: 4264:(7): 835–852. 4243: 4224:(2): 231–239. 4204: 4139: 4116: 4093: 4019: 3962:White, A. W.; 3949: 3913: 3877: 3823: 3796: 3762: 3736: 3714: 3707: 3678: 3667:(2): 169–184. 3644: 3629: 3618:(4): 223–242. 3587: 3554: 3478: 3455:(3): 519–548. 3416: 3378: 3329: 3322: 3304: 3262: 3216: 3215: 3213: 3210: 3158:Eocene Optimum 3152: 3149: 3108: 3105: 3075: 3072: 3007: 3006: 2997: 2996: 2988: 2987: 2986: 2985: 2984: 2982: 2979: 2977: 2974: 2962:Lord Howe Rise 2875:supercontinent 2867:Early Jurassic 2855:Meiolaniformes 2850: 2847: 2843: 2842: 2839: 2838: 2834: 2833: 2830: 2829: 2825: 2824: 2821: 2820: 2816: 2815: 2812: 2811: 2807: 2806: 2803: 2802: 2798: 2797: 2794: 2793: 2789: 2788: 2785: 2784: 2780: 2779: 2776: 2775: 2766: 2763: 2762: 2759: 2758: 2749: 2744: 2742: 2739: 2738: 2735: 2734: 2730:Ninjemys oweni 2725: 2720: 2718: 2715: 2714: 2711: 2710: 2701: 2696: 2694: 2690: 2689: 2686: 2685: 2681: 2680: 2677: 2676: 2672: 2671: 2668: 2667: 2658: 2655: 2654: 2651: 2650: 2641: 2636: 2634: 2631: 2630: 2627: 2626: 2617: 2612: 2610: 2605: 2603: 2600: 2599: 2596: 2595: 2591: 2590: 2587: 2586: 2577: 2574: 2573: 2570: 2569: 2560: 2555: 2553: 2548: 2546: 2543: 2542: 2539: 2538: 2529: 2524: 2522: 2520:Meiolaniformes 2516: 2515: 2512: 2511: 2507: 2506: 2503: 2502: 2495: 2492: 2491: 2488: 2487: 2480: 2475: 2473: 2468: 2466: 2461: 2333: 2332: 2330: 2326: 2325: 2322: 2321: 2317: 2316: 2313: 2312: 2308: 2307: 2304: 2303: 2299: 2298: 2295: 2294: 2290: 2289: 2286: 2285: 2281: 2280: 2277: 2276: 2267: 2264: 2263: 2260: 2259: 2250: 2245: 2243: 2240: 2239: 2236: 2235: 2226: 2221: 2219: 2216: 2215: 2212: 2211: 2202: 2197: 2195: 2192: 2191: 2188: 2187: 2178: 2173: 2171: 2168: 2167: 2164: 2163: 2154: 2149: 2147: 2142: 2140: 2136: 2135: 2132: 2131: 2127: 2126: 2123: 2122: 2118: 2117: 2114: 2113: 2109: 2108: 2105: 2104: 2100: 2099: 2096: 2095: 2091: 2090: 2087: 2086: 2082: 2081: 2078: 2077: 2068: 2065: 2064: 2061: 2060: 2055: 2050: 2048: 2045: 2044: 2041: 2040: 2031: 2026: 2024: 2021: 2020: 2017: 2016: 2007: 2002: 2000: 1997: 1996: 1993: 1992: 1983: 1978: 1976: 1973: 1972: 1969: 1968: 1959: 1954: 1952: 1949: 1948: 1945: 1944: 1935: 1930: 1928: 1923: 1911:M. brevicollis 1867: 1866: 1863: 1862: 1858: 1857: 1854: 1853: 1849: 1848: 1845: 1844: 1840: 1839: 1836: 1835: 1831: 1830: 1827: 1826: 1817: 1814: 1813: 1810: 1809: 1800: 1797: 1796: 1793: 1792: 1783: 1778: 1776: 1773: 1772: 1769: 1768: 1759: 1756: 1755: 1752: 1751: 1747:Ninjemys oweni 1742: 1737: 1735: 1732: 1731: 1728: 1727: 1718: 1715: 1714: 1711: 1710: 1701: 1696: 1694: 1688: 1686: 1681: 1595: 1592: 1471: 1470: 1461: 1460: 1459: 1450: 1449: 1448: 1447: 1446: 1433:M. brevicollis 1425:M. brevicollis 1368: 1365: 1281: 1278: 1275: 1274: 1272: 1262: 1248: 1245: 1240: 1232: 1231: 1222: 1207: 1193: 1190: 1185: 1177: 1176: 1167: 1150:Named for the 1148: 1134: 1131: 1129:Ninjemys oweni 1126: 1118: 1117: 1102: 1088: 1085: 1081: 1080: 1070: 1057: 1054: 1048: 1047: 1035: 1021: 1018: 1012: 1011: 1003: 989: 986: 979: 978: 969: 958:M. brevicollis 954: 940: 937: 932: 922: 921: 919: 908: 894: 891: 886: 878: 877: 874: 871: 868: 865: 862: 853: 850: 694:Walpole Island 628:as defined by 602:Juliana Sterli 524: 521: 434:Meiolaniformes 357: 356: 355: 354: 346: 338: 330: 322: 311: 310: 306: 305: 290: 286: 285: 283:Meiolaniformes 277: 270: 269: 264: 257: 256: 251: 244: 243: 241:Rhaptochelydia 238: 231: 230: 225: 218: 217: 212: 205: 204: 199: 195: 194: 189: 185: 184: 179: 175: 174: 169: 165: 164: 151: 150: 141: 140: 132: 131: 127: 126: 118: 117: 109: 104: 99: 94: 89: 84: 79: 74: 69: 64: 59: 54: 53: 48:48–0.003 37: 26: 24: 14: 13: 10: 9: 6: 4: 3: 2: 6610: 6599: 6596: 6594: 6591: 6589: 6586: 6584: 6581: 6579: 6576: 6574: 6571: 6569: 6566: 6564: 6561: 6559: 6556: 6555: 6553: 6536: 6531: 6527: 6523: 6518: 6514: 6510: 6505: 6501: 6497: 6492: 6488: 6484: 6479: 6475: 6470: 6464: 6460: 6455: 6449: 6445: 6444: 6442: 6440: 6436: 6432: 6427: 6415: 6407: 6405: 6402: 6400: 6397: 6395: 6392: 6390: 6386: 6385: 6382: 6372: 6365: 6344: 6340: 6339: 6338: 6337: 6335: 6333: 6329: 6318: 6317: 6313: 6311: 6310: 6306: 6304: 6303: 6299: 6297: 6296: 6295:Brachyopsemys 6292: 6290: 6289: 6285: 6284: 6282: 6280: 6276: 6270: 6269: 6265: 6263: 6262: 6258: 6256: 6255: 6251: 6248: 6247: 6246:Jurassichelon 6243: 6240: 6239: 6235: 6233: 6232: 6228: 6226: 6225: 6221: 6220: 6218: 6215: 6210: 6204: 6203: 6199: 6197: 6196: 6192: 6190: 6189: 6185: 6183: 6182: 6178: 6176: 6175: 6171: 6169: 6168: 6164: 6162: 6161: 6157: 6156: 6154: 6151: 6150:Eurysternidae 6146: 6139: 6138: 6137:Protostegidae 6134: 6132: 6131: 6127: 6125: 6124: 6120: 6118: 6117: 6113: 6111: 6110: 6106: 6104: 6103: 6099: 6097: 6096: 6092: 6090: 6089: 6085: 6083: 6082: 6078: 6077: 6074: 6071: 6066: 6062: 6055: 6048: 6042: 6041: 6037: 6035: 6034: 6030: 6028: 6027: 6023: 6021: 6020: 6016: 6014: 6013: 6009: 6007: 6006: 6002: 6000: 5999: 5995: 5993: 5992: 5988: 5986: 5985: 5981: 5979: 5978: 5974: 5972: 5971: 5967: 5965: 5964: 5960: 5958: 5957: 5953: 5951: 5950: 5946: 5944: 5943: 5939: 5937: 5936: 5932: 5930: 5929: 5925: 5923: 5922: 5918: 5916: 5915: 5911: 5909: 5908: 5904: 5902: 5901: 5900:Changmachelys 5897: 5895: 5894: 5890: 5888: 5887: 5883: 5881: 5880: 5876: 5874: 5873: 5869: 5868: 5866: 5864: 5863:Macrobaenidae 5859: 5852: 5846: 5845: 5841: 5839: 5838: 5834: 5832: 5831: 5827: 5825: 5824: 5820: 5818: 5817: 5813: 5811: 5810: 5806: 5804: 5803: 5799: 5797: 5796: 5792: 5790: 5789: 5785: 5783: 5782: 5778: 5776: 5775: 5771: 5769: 5768: 5767:Chengyuchelys 5764: 5762: 5761: 5757: 5755: 5754: 5750: 5748: 5747: 5743: 5742: 5740: 5738: 5731: 5721: 5720: 5719:Trinitichelys 5716: 5714: 5713: 5709: 5707: 5706: 5702: 5700: 5699: 5695: 5693: 5692: 5688: 5686: 5685: 5681: 5679: 5678: 5674: 5672: 5671: 5667: 5665: 5664: 5660: 5658: 5657: 5653: 5651: 5650: 5646: 5644: 5643: 5639: 5637: 5636: 5632: 5630: 5629: 5628:Gehennachelys 5625: 5623: 5622: 5618: 5616: 5615: 5611: 5609: 5608: 5604: 5602: 5601: 5597: 5595: 5594: 5590: 5588: 5587: 5583: 5581: 5580: 5576: 5574: 5573: 5569: 5567: 5566: 5562: 5560: 5559: 5555: 5554: 5552: 5550: 5546: 5540: 5539: 5535: 5533: 5532: 5528: 5525: 5524: 5520: 5517: 5516: 5512: 5510: 5509: 5508:Pleurosternon 5505: 5503: 5502: 5498: 5496: 5495: 5494:Dorsetochelys 5491: 5489: 5488: 5484: 5483: 5481: 5479: 5475: 5468: 5467: 5463: 5460: 5459: 5458:Compsemydidae 5455: 5454: 5451: 5448: 5446: 5439: 5433: 5432: 5428: 5425: 5424: 5420: 5417: 5416: 5412: 5410: 5409: 5405: 5402: 5401: 5400:Kallokibotion 5397: 5395: 5394: 5390: 5388: 5387: 5386:Calissounemys 5383: 5381: 5380: 5376: 5375: 5373: 5371: 5370:Compsemydidae 5364: 5358: 5357: 5353: 5351: 5350: 5346: 5344: 5343: 5339: 5337: 5336: 5332: 5330: 5329: 5325: 5324: 5322: 5320: 5313: 5307: 5306: 5305:Sichuanchelys 5302: 5300: 5299: 5298:Mongolochelys 5295: 5293: 5292: 5288: 5287: 5285: 5283: 5276: 5266: 5265: 5261: 5259: 5258: 5254: 5252: 5251: 5247: 5245: 5244: 5240: 5238: 5237: 5233: 5232: 5230: 5228: 5224: 5218: 5217: 5213: 5211: 5210: 5209:Peligrochelys 5206: 5204: 5203: 5202:Patagoniaemys 5199: 5197: 5196: 5192: 5190: 5189: 5185: 5184: 5181: 5178: 5176: 5169: 5163: 5162: 5158: 5156: 5155: 5154:Palaeochersis 5151: 5149: 5148: 5144: 5143: 5141: 5139: 5132: 5126: 5125: 5118: 5116: 5115: 5108: 5106: 5105: 5104:Proganochelys 5098: 5096: 5095: 5094:Kayentachelys 5088: 5086: 5085: 5084:Kallokibotion 5078: 5076: 5075: 5068: 5066: 5065: 5064:Heckerochelys 5058: 5056: 5055: 5048: 5046: 5045: 5038: 5036: 5035: 5028: 5027: 5024: 5021: 5018: 5013: 5007: 5006: 4999: 4997: 4996: 4989: 4986: 4985: 4978: 4976: 4975: 4968: 4967: 4964: 4961: 4958: 4957:Pantestudines 4953: 4949: 4937: 4936: 4935:Pantestudines 4930: 4917: 4916: 4915: 4914: 4912: 4910: 4909:Pantestudines 4906: 4898: 4894: 4893: 4892: 4891: 4888: 4885: 4883: 4879: 4875: 4867: 4866:Archelosauria 4863: 4860: 4858: 4854: 4851: 4849: 4845: 4842: 4840: 4836: 4834: 4830: 4828: 4824: 4823: 4818: 4814: 4810: 4809:Pantestudines 4803: 4798: 4796: 4791: 4789: 4784: 4783: 4780: 4769: 4763: 4755: 4753:9780128175552 4749: 4745: 4744: 4736: 4733: 4728: 4724: 4720: 4713: 4710: 4704: 4699: 4695: 4691: 4687: 4683: 4679: 4672: 4670: 4668: 4664: 4659: 4655: 4651: 4644: 4641: 4636: 4632: 4628: 4621: 4618: 4613: 4609: 4605: 4601: 4597: 4593: 4589: 4585: 4581: 4574: 4572: 4568: 4563: 4561:9781421428680 4557: 4553: 4552: 4544: 4541: 4535: 4530: 4526: 4522: 4519:(3): 93–106. 4518: 4514: 4507: 4504: 4499: 4495: 4491: 4487: 4483: 4479: 4475: 4468: 4465: 4460: 4456: 4452: 4448: 4444: 4440: 4436: 4432: 4425: 4423: 4421: 4419: 4417: 4415: 4413: 4409: 4404: 4398: 4387: 4383: 4379: 4375: 4369: 4365: 4361: 4357: 4353: 4346: 4338: 4335: 4330: 4326: 4322: 4318: 4314: 4310: 4306: 4300: 4298: 4294: 4289: 4285: 4280: 4275: 4271: 4267: 4263: 4259: 4252: 4250: 4248: 4244: 4239: 4235: 4231: 4227: 4223: 4219: 4211: 4209: 4205: 4200: 4196: 4192: 4188: 4184: 4180: 4176: 4172: 4165: 4163: 4154: 4152: 4150: 4148: 4146: 4144: 4140: 4135: 4131: 4127: 4120: 4117: 4112: 4108: 4104: 4097: 4094: 4089: 4085: 4080: 4075: 4071: 4067: 4062: 4057: 4053: 4049: 4045: 4041: 4037: 4030: 4028: 4026: 4024: 4020: 4015: 4011: 4006: 4001: 3996: 3991: 3987: 3983: 3979: 3975: 3974: 3969: 3965: 3964:Worthy, T. H. 3958: 3956: 3954: 3950: 3945: 3941: 3934: 3930: 3924: 3922: 3920: 3918: 3914: 3909: 3905: 3902:(2): 93–106. 3901: 3897: 3896: 3888: 3886: 3884: 3882: 3878: 3873: 3869: 3865: 3861: 3857: 3853: 3846: 3844: 3842: 3840: 3838: 3836: 3834: 3832: 3830: 3828: 3824: 3819: 3815: 3811: 3807: 3800: 3797: 3792: 3788: 3784: 3780: 3776: 3769: 3767: 3763: 3758: 3754: 3747: 3740: 3737: 3732: 3725: 3718: 3715: 3710: 3704: 3700: 3696: 3692: 3685: 3683: 3679: 3674: 3670: 3666: 3662: 3655: 3648: 3645: 3640: 3633: 3630: 3625: 3621: 3617: 3613: 3612: 3604: 3600: 3594: 3592: 3588: 3582: 3577: 3573: 3569: 3565: 3558: 3555: 3550: 3546: 3541: 3536: 3532: 3528: 3524: 3520: 3516: 3509: 3507: 3505: 3503: 3501: 3499: 3497: 3495: 3493: 3491: 3489: 3487: 3485: 3483: 3479: 3473: 3468: 3463: 3458: 3454: 3450: 3446: 3439: 3437: 3435: 3433: 3431: 3429: 3427: 3425: 3423: 3421: 3417: 3412: 3408: 3404: 3400: 3396: 3392: 3388: 3382: 3379: 3373: 3368: 3364: 3360: 3356: 3352: 3351: 3346: 3342: 3341:Owen, Richard 3336: 3334: 3330: 3325: 3319: 3315: 3308: 3305: 3300: 3296: 3292: 3288: 3281: 3279: 3275: 3266: 3263: 3259:(3049): 1–10. 3258: 3251: 3249: 3245: 3236: 3234: 3232: 3230: 3228: 3226: 3224: 3222: 3218: 3211: 3209: 3205: 3203: 3198: 3194: 3190: 3189:Lapita people 3186: 3182: 3177: 3175: 3169: 3167: 3163: 3159: 3150: 3148: 3146: 3142: 3137: 3134: 3130: 3126: 3122: 3119:Study on the 3113: 3106: 3104: 3102: 3098: 3093: 3089: 3085: 3081: 3073: 3071: 3069: 3065: 3059: 3057: 3052: 3047: 3045: 3041: 3037: 3033: 3029: 3025: 3021: 3017: 3001: 2992: 2980: 2975: 2973: 2971: 2967: 2963: 2959: 2953: 2950: 2949:Lapita people 2944: 2942: 2938: 2934: 2930: 2926: 2922: 2917: 2909: 2905: 2902: 2898: 2897:South Pacific 2894: 2889: 2887: 2883: 2879: 2876: 2872: 2868: 2864: 2863:ghost lineage 2860: 2856: 2848: 2841: 2840: 2832: 2831: 2823: 2822: 2814: 2813: 2805: 2804: 2796: 2795: 2787: 2786: 2778: 2777: 2774: 2773: 2772: 2765: 2764: 2761: 2760: 2757: 2756: 2755: 2748: 2747: 2741: 2740: 2737: 2736: 2733: 2732: 2731: 2724: 2723: 2717: 2716: 2713: 2712: 2709: 2708: 2707: 2700: 2699: 2693:Meiolaniidae 2692: 2691: 2688: 2687: 2679: 2678: 2670: 2669: 2666: 2665: 2664: 2657: 2656: 2653: 2652: 2649: 2648: 2647: 2640: 2639: 2633: 2632: 2629: 2628: 2625: 2624: 2623: 2616: 2615: 2609: 2608: 2602: 2601: 2598: 2597: 2589: 2588: 2585: 2584: 2583: 2576: 2575: 2572: 2571: 2568: 2567: 2566: 2559: 2558: 2552: 2551: 2545: 2544: 2541: 2540: 2537: 2536: 2535: 2528: 2527: 2521: 2518: 2517: 2514: 2513: 2505: 2504: 2501: 2500: 2494: 2493: 2490: 2489: 2486: 2485: 2479: 2478: 2472: 2471: 2465: 2464: 2460: 2458: 2454: 2453:plesiomorphic 2449: 2448:Peligrochelys 2445: 2444:Mongolochelys 2441: 2440: 2439:Peligrochelys 2435: 2434: 2433:Patagoniaemys 2429: 2428: 2423: 2422: 2417: 2413: 2412:Mongolochelys 2409: 2405: 2404: 2403:Kallokibotion 2399: 2398: 2397:Mongolochelys 2391: 2389: 2385: 2381: 2376: 2371: 2367: 2363: 2362:podocnemidids 2359: 2355: 2351: 2350: 2344: 2340: 2324: 2323: 2315: 2314: 2306: 2305: 2297: 2296: 2288: 2287: 2279: 2278: 2275: 2274: 2273: 2266: 2265: 2262: 2261: 2258: 2257: 2256: 2249: 2248: 2242: 2241: 2238: 2237: 2234: 2233: 2232: 2225: 2224: 2218: 2217: 2214: 2213: 2210: 2209: 2208: 2201: 2200: 2194: 2193: 2190: 2189: 2186: 2185: 2184: 2177: 2176: 2170: 2169: 2166: 2165: 2162: 2161: 2160: 2153: 2152: 2146: 2145: 2134: 2133: 2125: 2124: 2116: 2115: 2107: 2106: 2098: 2097: 2089: 2088: 2080: 2079: 2076: 2075: 2074: 2067: 2066: 2063: 2062: 2059: 2054: 2053: 2047: 2046: 2043: 2042: 2039: 2038: 2037: 2030: 2029: 2023: 2022: 2019: 2018: 2015: 2014: 2013: 2006: 2005: 1999: 1998: 1995: 1994: 1991: 1990: 1989: 1982: 1981: 1975: 1974: 1971: 1970: 1967: 1966: 1965: 1958: 1957: 1951: 1950: 1947: 1946: 1943: 1942: 1941: 1934: 1933: 1927: 1926: 1918: 1916: 1912: 1908: 1904: 1899: 1895: 1891: 1887: 1879: 1874: 1865: 1864: 1856: 1855: 1847: 1846: 1838: 1837: 1829: 1828: 1825: 1824: 1823: 1816: 1815: 1812: 1811: 1808: 1807: 1806: 1799: 1798: 1795: 1794: 1791: 1790: 1789: 1782: 1781: 1775: 1774: 1771: 1770: 1767: 1766: 1765: 1758: 1757: 1754: 1753: 1750: 1749: 1748: 1741: 1740: 1734: 1733: 1730: 1729: 1726: 1725: 1724: 1717: 1716: 1713: 1712: 1709: 1708: 1707: 1700: 1699: 1693:Meiolaniidae 1692: 1691: 1685: 1684: 1680: 1678: 1673: 1671: 1667: 1663: 1659: 1655: 1651: 1647: 1643: 1639: 1635: 1631: 1627: 1623: 1619: 1615: 1611: 1606: 1601: 1593: 1591: 1588: 1584: 1580: 1576: 1571: 1569: 1565: 1561: 1557: 1553: 1549: 1545: 1541: 1537: 1533: 1529: 1525: 1520: 1518: 1514: 1510: 1506: 1502: 1498: 1494: 1490: 1486: 1475: 1465: 1454: 1445: 1443: 1438: 1434: 1430: 1426: 1422: 1418: 1414: 1410: 1406: 1402: 1398: 1394: 1388: 1386: 1378: 1373: 1366: 1364: 1362: 1358: 1354: 1350: 1346: 1342: 1341:New Caledonia 1338: 1333: 1331: 1327: 1323: 1319: 1315: 1311: 1307: 1303: 1299: 1295: 1291: 1286: 1279: 1273: 1270: 1266: 1263: 1260: 1249: 1246: 1244: 1241: 1239: 1238: 1234: 1233: 1227: 1223: 1220: 1216: 1211: 1208: 1205: 1194: 1191: 1189: 1186: 1184: 1183: 1179: 1178: 1172: 1168: 1165: 1161: 1157: 1153: 1149: 1146: 1135: 1132: 1130: 1127: 1125: 1124: 1120: 1119: 1115: 1111: 1107: 1103: 1100: 1089: 1086: 1083: 1082: 1078: 1074: 1071: 1069: 1058: 1055: 1053: 1050: 1049: 1044: 1040: 1036: 1033: 1032:New Caledonia 1022: 1019: 1017: 1014: 1013: 1008: 1004: 1001: 990: 987: 985: 981: 980: 974: 967: 963: 959: 955: 952: 941: 938: 936: 933: 931: 930: 929: 923: 920: 917: 913: 909: 906: 895: 892: 890: 887: 885: 884: 880: 879: 875: 872: 869: 866: 863: 860: 859: 851: 849: 847: 843: 839: 835: 834: 829: 825: 821: 817: 812: 810: 809: 804: 800: 796: 792: 786: 784: 780: 776: 772: 768: 764: 760: 756: 752: 751:Santiago Roth 748: 744: 743: 738: 734: 730: 726: 722: 714: 710: 705: 701: 699: 698:New Caledonia 695: 691: 687: 684:Australasian 682: 678: 674: 670: 666: 662: 658: 654: 650: 646: 642: 638: 633: 631: 627: 623: 619: 615: 611: 605: 603: 599: 595: 591: 587: 583: 579: 575: 571: 561: 557: 555: 551: 547: 546: 541: 537: 536: 531: 522: 520: 518: 517:mating season 514: 509: 507: 503: 497: 495: 491: 487: 483: 479: 475: 474: 469: 468: 463: 462: 457: 454: 450: 449: 444: 443: 437: 435: 431: 427: 423: 422:New Caledonia 419: 415: 411: 407: 403: 402: 397: 393: 390: 386: 382: 378: 377:South America 374: 373:clubbed tails 370: 367: 363: 353: 352: 347: 345: 344: 339: 337: 336: 331: 329: 328: 323: 321: 320: 315: 314: 312: 307: 302: 297: 291: 288: 287: 284: 278: 275: 272: 271: 268: 265: 262: 259: 258: 255: 252: 249: 246: 245: 242: 239: 236: 233: 232: 229: 226: 223: 220: 219: 216: 215:Pantestudines 213: 210: 207: 206: 203: 200: 197: 196: 193: 190: 187: 186: 183: 180: 177: 176: 173: 170: 167: 166: 161: 156: 152: 149: 148: 142: 138: 133: 128: 124: 119: 112: 107: 102: 97: 92: 87: 82: 77: 72: 67: 62: 57: 51: 45: 41: 40:Middle Eocene 33: 30: 19: 6469:Meiolaniidae 6439:Meiolaniidae 6438: 6404:Perichelydia 6399:Mesochelydia 6314: 6307: 6300: 6293: 6288:Angolachelys 6286: 6266: 6261:Portlandemys 6259: 6254:Plesiochelys 6252: 6244: 6238:Hylaeochelys 6236: 6229: 6222: 6200: 6195:Palaeomedusa 6193: 6186: 6179: 6172: 6165: 6158: 6135: 6128: 6121: 6114: 6107: 6100: 6093: 6086: 6079: 6064: 6060: 6038: 6031: 6024: 6017: 6010: 6003: 5996: 5989: 5982: 5975: 5968: 5961: 5954: 5947: 5942:Jeholochelys 5940: 5933: 5926: 5921:Galvechelone 5919: 5912: 5905: 5898: 5893:Aurorachelys 5891: 5884: 5879:Appalachemys 5877: 5870: 5842: 5835: 5828: 5823:Tienfuchelys 5821: 5814: 5807: 5800: 5795:Phunoichelys 5793: 5786: 5779: 5772: 5765: 5760:Camerochelys 5758: 5753:Brodiechelys 5751: 5744: 5717: 5710: 5705:Stygiochelys 5703: 5696: 5689: 5682: 5675: 5668: 5661: 5654: 5647: 5640: 5633: 5626: 5619: 5612: 5605: 5598: 5591: 5584: 5577: 5570: 5565:Arvinachelys 5563: 5556: 5536: 5529: 5521: 5513: 5506: 5499: 5492: 5485: 5464: 5456: 5429: 5421: 5413: 5406: 5398: 5391: 5384: 5377: 5354: 5347: 5340: 5335:Helochelydra 5333: 5328:Aragochersis 5326: 5303: 5296: 5289: 5262: 5255: 5248: 5241: 5236:Gaffneylania 5234: 5227:Meiolaniidae 5226: 5214: 5207: 5200: 5193: 5186: 5159: 5152: 5145: 5122: 5112: 5102: 5092: 5082: 5072: 5062: 5054:Eileanchelys 5052: 5044:Condorchelys 5042: 5034:Chinlechelys 5032: 5003: 4995:Odontochelys 4993: 4984:Eunotosaurus 4982: 4972: 4933: 4861: 4852: 4843: 4742: 4735: 4718: 4712: 4685: 4681: 4657: 4653: 4643: 4634: 4630: 4620: 4587: 4583: 4550: 4543: 4516: 4512: 4506: 4481: 4477: 4467: 4438: 4434: 4397:cite journal 4355: 4351: 4337: 4312: 4308: 4261: 4257: 4221: 4217: 4174: 4170: 4161: 4133: 4129: 4119: 4110: 4106: 4096: 4046:(1): 38317. 4043: 4039: 3977: 3971: 3943: 3939: 3899: 3893: 3855: 3809: 3799: 3782: 3778: 3756: 3752: 3739: 3730: 3717: 3690: 3664: 3660: 3647: 3638: 3632: 3615: 3609: 3599:Anderson, C. 3571: 3567: 3557: 3525:(1): 21–45. 3522: 3518: 3452: 3448: 3394: 3390: 3381: 3354: 3348: 3313: 3307: 3293:(2): 85–89. 3290: 3286: 3277: 3276:, Owen, and 3273: 3265: 3256: 3247: 3243: 3206: 3201: 3180: 3178: 3174:last ice age 3170: 3154: 3144: 3140: 3138: 3132: 3118: 3100: 3077: 3060: 3055: 3050: 3048: 3039: 3035: 3027: 3023: 3019: 3013: 2976:Paleobiology 2957: 2954: 2945: 2914: 2900: 2890: 2886:Gaffneylania 2885: 2881: 2852: 2769: 2768: 2752: 2751: 2728: 2727: 2704: 2703: 2661: 2660: 2644: 2643: 2620: 2619: 2580: 2579: 2563: 2562: 2532: 2531: 2497: 2482: 2447: 2443: 2437: 2431: 2425: 2419: 2411: 2407: 2401: 2395: 2392: 2366:pelomedusids 2353: 2347: 2336: 2270: 2269: 2253: 2252: 2229: 2228: 2205: 2204: 2181: 2180: 2157: 2156: 2071: 2070: 2057: 2034: 2033: 2010: 2009: 1986: 1985: 1962: 1961: 1938: 1937: 1914: 1910: 1906: 1903:M. platyceps 1902: 1897: 1894:M. platyceps 1893: 1889: 1885: 1883: 1877: 1820: 1819: 1803: 1802: 1786: 1785: 1762: 1761: 1745: 1744: 1721: 1720: 1704: 1703: 1677:Gaffneylania 1676: 1674: 1669: 1665: 1661: 1657: 1653: 1649: 1645: 1641: 1637: 1634:Gaffneylania 1633: 1625: 1617: 1613: 1609: 1604: 1600:monophyletic 1597: 1586: 1582: 1578: 1574: 1572: 1568:Gaffneylania 1567: 1563: 1555: 1552:Gaffneylania 1551: 1547: 1527: 1523: 1521: 1516: 1512: 1496: 1492: 1488: 1484: 1481: 1473: 1441: 1436: 1432: 1428: 1424: 1420: 1416: 1412: 1408: 1404: 1401:Gaffneylania 1400: 1392: 1389: 1382: 1376: 1352: 1337:Pindai Caves 1334: 1329: 1305: 1301: 1297: 1287: 1283: 1268: 1264: 1242: 1235: 1218: 1214: 1209: 1187: 1180: 1163: 1159: 1155: 1128: 1121: 1113: 1110:M. platyceps 1109: 1105: 1087:Pleistocene 1076: 1072: 1056:Pleistocene 1051: 1043:M. platyceps 1042: 1038: 1020:Pleistocene 1015: 1006: 983: 966:M. platyceps 965: 961: 957: 934: 926: 925: 915: 912:Gaffneylania 911: 888: 883:Gaffneylania 881: 845: 833:Gaffneylania 831: 815: 813: 806: 802: 798: 794: 790: 787: 782: 778: 770: 766: 762: 758: 754: 740: 718: 708: 689: 685: 680: 677:M. platyceps 676: 672: 668: 664: 660: 644: 636: 634: 621: 617: 609: 606: 589: 585: 581: 577: 574:M. platyceps 573: 569: 566: 554:thorny devil 543: 540:Richard Owen 533: 526: 510: 505: 501: 498: 471: 465: 459: 448:Gaffneylania 446: 440: 438: 399: 362:Meiolaniidae 361: 360: 349: 341: 333: 325: 319:Gaffneylania 317: 296:Meiolaniidae 295: 273: 267:Perichelydia 260: 254:Mesochelydia 247: 234: 221: 208: 145: 144:Skeleton of 35:Meiolaniidae 29: 6463:Wikispecies 6302:Leyvachelys 6279:Sandownidae 6231:Globochelus 6174:Eurysternum 6130:Thalassemys 6109:Neusticemys 5907:Dracochelys 5858:Sinemydidae 5844:Yanduchelys 5816:Siamochelys 5809:Shartegemys 5781:Kalasinemys 5774:Jastmelchyi 5677:Plesiobaena 5663:Palatobaena 5656:Neurankylus 5621:Gamerabaena 5600:Denazinemys 5558:Arundelemys 5408:Peltochelys 5379:Berruchelus 5349:Naomichelys 5017:Testudinata 5005:Pappochelys 4703:11336/63435 4534:11336/84233 3946:(1): 35–48. 3574:: 181–191. 3540:11336/21194 3472:11336/41594 3357:: 471–480. 3202:M. damelipi 2970:New Zealand 2937:East Africa 2457:crown group 2416:sinochelyds 2386:as well as 2384:sea turtles 2343:cryptodires 2339:pleurodires 1622:basicranium 1509:ankylosaurs 1505:glyptodonts 1397:ceratopsian 1367:Description 1361:New Zealand 1349:Tiga Island 1213:Formation. 1007:M. damelipi 612:was named, 490:New Zealand 478:Australasia 426:Pleistocene 424:during the 414:Pleistocene 389:crown group 385:cryptodires 381:Australasia 375:known from 228:Testudinata 6552:Categories 6394:Ankylopoda 6387:See also: 6343:Testudines 6332:Testudines 6268:Tropidemys 6202:Parachelys 6188:Idiochelys 6181:Hydropelta 6167:Chelonides 6026:Xiaochelys 5991:Osteopygis 5970:Macrobaena 5963:Liaochelys 5956:Kirgizemys 5949:Judithemys 5886:Asiachelys 5872:Anatolemys 5830:Undjulemys 5788:Larachelus 5691:Saxochelys 5684:Protobaena 5593:Chisternon 5586:Cedrobaena 5515:Riodevemys 5487:Dinochelys 5415:Riodevemys 5342:Helochelys 5264:Warkalania 5188:Chubutemys 5161:Waluchelys 5124:Spoochelys 5074:Indochelys 4918:see below↓ 4897:Sauropsida 4882:Sauropsida 4839:Sauropsida 4660:: 363–368. 4358:: 11, 23. 4279:11336/3254 4136:(107–113). 4113:: 285–297. 3212:References 3151:Extinction 3092:palm trees 3051:Meiolania' 2499:Cryptodira 2484:Pleurodira 2427:Chubutemys 2207:Warkalania 1988:Warkalania 1907:M. mackayi 1890:M. mackayi 1666:Warkalania 1650:Warkalania 1630:sinemydids 1560:Osteoderms 1442:Warkalania 1330:Warkalania 1302:Warkalania 1269:Warkalania 1265:Warkalania 1237:Warkalania 808:Warkalania 690:M. mackayi 649:Pleurodira 486:Antarctica 467:Warkalania 451:native to 412:until the 392:Testudines 366:stem-group 351:Warkalania 18:Meiolaniid 6316:Solnhofia 6309:Sandownia 6160:Achelonia 6095:Euryaspis 6040:Yumenemys 5984:Ordosemys 5935:Hoyasemys 5712:Thescelus 5698:Scabremys 5649:Lakotemys 5635:Goleremys 5523:Selenemys 5423:Selenemys 5393:Compsemys 5243:Meiolania 5195:Otwayemys 4825:Kingdom: 4762:cite book 4637:: 55–135. 4631:Zoologica 4604:0272-4634 4455:0268-0130 4382:1088-7105 4315:: 72–76. 4238:131795055 4070:2045-2322 3872:0003-0090 3864:2246/1670 3818:2246/5279 3397:: 43–48. 3278:Meiolania 3274:Megalania 3166:latitudes 3145:Meiolania 3099:of ? 3056:Meiolania 3040:Meiolania 3036:Meiolania 3028:Meiolania 3024:Meiolania 3020:Meiolania 2981:Lifestyle 2966:Zealandia 2941:barnacles 2927:that hit 2925:hurricane 2901:Meiolania 2421:Otwayemys 2354:Meiolania 1915:Meiolania 1898:Meiolania 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5250:Ninjemys 4848:Diapsida 4833:Chordata 4831:Phylum: 4827:Animalia 4612:88998996 4498:85295987 4484:: 3–45. 4459:Archived 4386:Archived 4329:84241716 4288:83804365 4199:83503956 4088:27922064 4014:20713711 3812:(2805). 3791:2246/978 3601:(1925). 3549:83799914 3244:Ninjemys 3133:Niolamia 3121:endocast 2882:Niolamia 2878:Gondwana 2394:grouped 2349:Chelydra 2183:Ninjemys 2159:Niolamia 1964:Ninjemys 1940:Niolamia 1658:Ninjemys 1646:Ninjemys 1642:Niolamia 1638:Niolamia 1618:Niolamia 1614:Ninjemys 1610:Niolamia 1605:Niolamia 1587:Ninjemys 1583:Niolamia 1548:Niolamia 1532:carapace 1513:Ninjemys 1493:Ninjemys 1489:Ninjemys 1485:Niolamia 1417:Niolamia 1413:Ninjemys 1409:Ninjemys 1405:Niolamia 1393:Niolamia 1377:Niolamia 1298:Niolamia 1219:Niolamia 1210:Niolamia 1182:Niolamia 1164:Ninjemys 1160:Niolamia 1156:Ninjemys 1123:Ninjemys 939:Miocene 916:Niolamia 864:Species 846:Niolamia 820:Holocene 799:Ninjemys 783:Niolamia 779:Niolamia 771:Niolamia 759:Miolania 709:Niolamia 681:M. minor 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