Ankylosaurus - Knowledge

129: 1364: 1159: 6005: 2903: 2602: 1404:(armor plates on the neck), but these are known only from fragments, making their exact arrangement uncertain. Carpenter suggested that when seen from above, the plates would have been paired, creating an inverted V-shape across the neck, with the midline gap probably being filled with small ossicles (round bony scutes) to allow for movement. He believed the width of this armor belt was too wide to have fitted solely on the neck, and that it covered the base of the neck and continued onto the shoulder region. Arbour and the Canadian paleontologist 2457: 973: 756: 537: 1355: 854: 2649: 6000: 1469: 645: 2365: 1641: 1632: 154: 4927: 1663: 432:, with a broad, robust body. It had a wide, low skull, with two horns pointing backward from the back of the head, and two horns below these that pointed backward and down. Unlike other ankylosaurs, its nostrils faced sideways rather than towards the front. The front part of the jaws was covered in a beak, with rows of small, leaf-shaped teeth farther behind it. It was covered in armor plates, or osteoderms, with bony half-rings covering the neck, and had a large 6028: 2556: 1427:

body, with four or five transverse rows separated by creases in the skin. The osteoderms on the flanks would probably have had a more square outline than those on the back. There may have been four longitudinal rows of osteoderms on the flanks. Unlike some basal ankylosaurs and many nodosaurs, ankylosaurids do not appear to have had co-ossified pelvic shields above their hips. Some osteoderms without keels may have been placed above the hip region of

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weight. In the cervical half-rings, the underlying bone band developed outgrowths connecting it with the underlying osteoderms, which simultaneously fused to each other. On the skull, the middle bone plates first ossified at the snout and the rear rim, with ossification gradually extending towards the middle regions. On the rest of the body, ossification progressed from the neck backward in the direction of the tail.

4915: 2444:) had jaws with low mechanical advantage, whereas those of earlier relatives were high to moderate. These late ankylosaurids also had tooth occlusion and complex biphasal jaw mechanisms, features shared with some Late Cretaceous nodosaurids, but those instead have jaws with high mechanical advantage. This indicates that while the two groups converged in some features, the nodosaurs had higher relative 2484:, not the nostrils, and that reduction in weight was minimal, as the spaces only accounted for a small percent of the skull volume. He also considered a gland unlikely and noted that the sinuses may not have had any specific function. It has also been suggested that the respiratory passages were used to perform a mammal-like treatment of inhaled air, based on the presence and arrangement of 1707:. They were widespread and inhabited a broad range of environments. As more complete specimens and new genera have been discovered, theories about ankylosaurian interrelatedness have become more complex, and hypotheses have often changed between studies. In addition to Ankylosauridae, Ankylosauria has been divided into the families Nodosauridae, and sometimes 959:, its skull is still larger than those of any other ankylosaurins. A few other ankylosaurs reached about 6 m (20 ft) in length. Because the vertebrae of AMNH 5214 are not significantly larger than those of other ankylosaurines, Arbour and Mallon considered their upper range estimate of nearly 10 meters (33 ft) for large 1564:. The last seven tail vertebrae formed the "handle" of the tail club. These vertebrae were in contact, with no cartilage between them, and were sometimes co-ossified, which made them immobile. Ossified tendons attached to the vertebrae in front of the tail club, and these features together helped strengthen it. The interlocked 673:, which was unknown at the time. Brown also reconstructed the armor plates in parallel rows running down the back; this arrangement was purely hypothetical. Brown's reconstruction became highly influential, and restorations of the animal based on his diagram were published as late as the 1980s. In a 1908 review of Brown's 1089:(internal nostrils), and the air passage was looped. The maxillae expanded to the sides, giving the impression of a bulge, which may have been due to the sinuses inside. The maxillae had a ridge that may have been the attachment site for fleshy cheeks; the presence of cheeks in ornithischians is controversial, but some 924: in) wide, and Carpenter estimated that it measured about 5.4 m (17 ft 9 in) long and about 1.4 m (4 ft 7 in) tall at the hips. The English paleontologist Roger B. J. Benson and colleagues estimated the weight for AMNH 5214 at 4.78 metric tons (5.27 short tons) in 2014. 1116:

was low in proportion to its length, and, when seen from the side, the tooth row was almost straight instead of arched. The mandibles are completely preserved only in the smallest specimen (AMNH 5214) and are about 41 centimeters (16 in) long. The incomplete mandible of the largest specimen (CMN

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Reconstructions of ankylosaur forelimb musculature made by Coombs in 1978 suggest that the forelimbs bore the majority of the animal's weight, and were adapted for high force delivery on the front feet, possibly for food gathering. In addition, Coombs suggested that ankylosaurs may have been capable

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sp., is also found in the same formations, but according to Carpenter, the range of the two genera does not seem to have overlapped. Their remains have so far not been found in the same localities, and the nodosaur appears to have inhabited the lowlands. The narrower muzzle of the nodosaur suggests

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seems to have been an active defensive weapon, capable of producing enough of an impact to break the bones of an assailant. The tendons of the tail were partially ossified and were not very elastic, allowing great force to be transmitted to the club when it was used as a weapon. Coombs suggested in

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was unique among ankylosaurs, and had undergone an "extreme" transformation compared to its relatives. The snout was arched and truncated at the front, and the nostrils were elliptical and were directed downward and outward, unlike in all other known ankylosaurids where they faced obliquely forward

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and been effective weapons. The study also found that while adult ankylosaurid tail clubs were capable of breaking bones, those of juveniles were not. Despite the feasibility of tail-swinging, the researchers could not determine whether ankylosaurids used their clubs for defense against potential

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was composed of two large osteoderms, with a row of small osteoderms at the midline, and two small osteoderms at the tip; these osteoderms obscured the last tail vertebra. As only the tail club of specimen AMNH 5214 is known, the range of variation between individuals is unknown. The tail club of

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The first osteoderms behind the second cervical half-ring would have been similar in shape to those in the first half-ring, and the osteoderms on the back probably decreased in diameter hindwards. The largest osteoderms were probably arranged in transverse and longitudinal rows across most of the

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include a relatively large, hexagonal (or diamond-shaped) nasal caputegulum at the front of the snout between the nostrils, which had a loreal caputegulum on each side, an anterior and posterior supraorbital caputegulum above each orbit, and a ridge of nuchal caputegulae at the back of the skull.

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in a squatting posture and with a huge tail club being dragged over the ground. Modern reconstructions show the animal with a more upright limb posture and with the tail held off the ground. Likewise, large spines projecting sideways from the body (similar to those of nodosaurid ankylosaurs) are

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In 1993, Tony Thulborn proposed that the tail club of ankylosaurids primarily acted as a decoy for the head, as he thought the tail too short and inflexible to have an effective reach; the "dummy head" would lure a predator close to the tail, where it could be struck. Carpenter has rejected this

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of different ages found that during ontogenetic development, the ribs of juvenile ankylosaurs fused with their vertebrae. The forelimbs strongly increased in robustness while the hindlimbs did not become larger relative to the rest of the skeleton, further evidence that the arms bore most of the

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disagreed with Carpenter's interpretation in 2015 and pointed out that the cervical half-ring fragments of the holotype specimen did not fit together in the way proposed by Carpenter (though this could be due to breakage). They instead suggested that the fragments represented the remains of two

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concluded that despite the large size of ankylosaur skulls, the associated musculature was relatively weak. He also thought jaw movement was limited to up and down movements. Extrapolating from this, Haas suggested that ankylosaurs ate relatively soft non-abrasive vegetation. Later research on

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was its armor, consisting of knobs and plates of bone known as osteoderms, or scutes, embedded in the skin. These have not been found in articulation, so their exact placement on the body is unknown, though inferences can be made based on related animals, and various configurations have been

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were expanded to the sides of the premaxilla bones, to a larger extent than seen in other ankylosaurs. Large loreal caputegulae—strap-like, side osteoderms of the snout—completely roofed the enlarged opening of the nostrils, giving a bulbous appearance. The nostrils also had an intranarial

2718:. The type specimen is from the Hell Creek Formation of Montana, while other specimens have been found in the Lance and Ferris Formations in Wyoming, the Scollard Formation in Alberta, and the Frenchman Formation in Saskatchewan, all of which date to the end of the Cretaceous. 2617:, they were embedded directly into the bone tissue, a feature unique to ankylosaurids. This would have provided the ankylosaurids with an armor covering that was both lightweight and highly durable, being resistant to breakage and penetration by the teeth of predators. The 2729:

it is known from, and the distribution of its remains suggests that it was ecologically rare, or restricted to the uplands of the formations, where it would have been less likely to fossilize, rather than the coastal lowlands. Another ankylosaur, a nodosaur referred to as

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and Jordan Mallon redescribed the genus in light of newer ankylosaur discoveries, including elements of the holotype that had not been previously mentioned in the literature (such as parts of the skull and the cervical half-rings). They concluded that though

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for these, he refrained from doing so, as the range of variation in the species was not completely documented. He also raised the possibility that the two teeth associated with the holotype specimen perhaps did not belong to it, as they were found in

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were smoother. Many smaller osteoderms and ossicles probably occupied the space between the larger ones, as in other ankylosaurids. The osteoderms covering the body were very flat, though with a low keel at one margin. In contrast, the nodosaurid

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between skull elements, which is common for adult ankylosaurs. The caputegulum pattern of the skull was variable between specimens, though some details are shared. The caputegulae are named according to their position on the skull, and those of

2535:, as the nostrils tend to be reduced or the premaxilla extended. In addition, though the widely separated nostrils may have allowed for stereo-olfaction (where each nostril senses smells from different directions), as has been proposed for the 1004:(eye sockets) were almost round to slightly oval and did not face directly sideways because the skull tapered towards the front. The braincase was short and robust, as in other ankylosaurines. Crests above the orbits merged into the upper 963:

too long, and suggested a length of 8 m (26 ft) instead. Arbour and Mallon estimated a weight of 4.78 t (5.27 short tons) for AMNH 5214, and tentatively estimated the weight of CMN 8880 at 7.95 t (8.76 short tons).

669:. In contrast to modern depictions, Brown's stegosaur-like reconstruction showed robust forelimbs, a strongly arched back, a pelvis with prongs projecting forwards from the ilium and pubis, as well as a short, drooping tail without a 2349: in) in length, respectively. The smaller tooth is heavily worn, leading Carpenter to suggest that ankylosaurids in general or at least the young did not swallow their food whole but employed some sort of chewing. Since adult 844:

is the best-known member of its group, it was bizarre in comparison to related ankylosaurs, and therefore not representative of the group. In spite of its familiarity, it is known from far fewer remains than its closest relatives.

2511:. Reconstructions of the inner ear suggest adaptation to hearing at low frequencies, such as the low-toned resonant sounds possibly produced by the nasal passages. They disputed the possibility that the looping is related to 1261:

had centra (or bodies) that were short relative to their width, and their neural spines were short and narrow. The dorsal vertebrae were tightly spaced, which limited the downwards movement of the back. The neural spines had

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Weishampel, D. B.; Barrett, P. M.; Coria, R. A.; Le Loeuff, J.; Xu X.; Zhao X.; Sahni, A.; Gomani, E. M. P.; Noto, C. R. (2004). "Dinosaur Distribution". In Weishampel, D. B.; Dodson, P.; Osmolska, H.. (eds.).

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had broad neural spines that increased in height towards the body. The front part of the neural spines had well-developed entheses, which was common among adult dinosaurs, and indicates the presence of large

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indicate that airflow was unidirectional (looping through the lungs during inhalation and exhalation), although it may also have been bidirectional in the posterior nasal chamber, with air directed past the

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had small, phylliform (leaf-shaped) teeth, which were compressed sideways. The teeth were mostly taller than they were wide, and were very small; their size in proportion to the skull meant that the jaws of

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means "fused" or "bent lizard", and the specific name means "great belly". A handful of specimens have been excavated to date, but a complete skeleton has not been discovered. Though other members of

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did little chewing of its food, it would have spent less time in the day foraging than an elephant. Based on the broadness of the ribcage, the digestion of unchewed food may have been facilitated by

575:. The specimen (found by collector Peter Kaisen) consisted of the upper part of a skull, two teeth, part of the shoulder girdle, cervical, dorsal, and caudal vertebrae, ribs, and more than thirty 775:

in Alberta, Canada. This specimen included a complete skull, mandibles, the first and only tail club known of this genus, as well as ribs, vertebrae, limb bones, and armor. In 1947 the American

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and nasal chambers in the snout may have been for heat and water balance or may have played a role in vocalization. The tail club is thought to have been used in defense against predators or in

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by paleontologist Tetsuto Miyashita and colleagues supported their function as a heat and water balancing system, noting the extensive blood vessel system and an increased surface area for the

1568:(articular processes) and neural spines of the handle vertebrae were U-shaped when seen from above, whereas those of most other ankylosaurids are V-shaped, which may be due to the handle of 835:

specimens were not scientifically described at length, though several paleontologists planned to do so until Carpenter redescribed the genus in 2004. In 2017 the Canadian paleontologists

2499:(used for heat and water exchange in modern animals). The researchers also supported the idea of the loops acting as a resonance chamber, comparable to the elongated nasal passages of 868:

dinosaur and possibly the largest ankylosaurid. In 2004 Carpenter estimated that the individual with the largest-known skull (specimen CMN 8880), which is 64.5 centimeters (2 ft

2527:. The enlarged olfactory region of ankylosaurids indicates a well-developed sense of smell. Though hindwards retraction of the nostrils is seen in aquatic animals and animals with a 4172: 2568:

would have limited fossorial activity. Ankylosaurs were likely to have been slow-moving and sluggish animals, though they may have been capable of quick movements when necessary.

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was a better example of how ankylosaurs would have appeared in life. The claim of synonymy was not accepted by other researchers, and the two genera are now considered distinct.

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would have been unable to knock down trees like modern elephants do. It was also incapable of chewing bark and thus unlikely to have practiced bark stripping. As an adult,

1604:(then only known from a partial skull and osteoderms) as part of the family. Due to the fragmentary condition of the remains, Brown was unable to fully distinguish between 1085:

were elongated and separated by a septum at the midline, which divided the inside of the snout into two mirrored halves. The nasal chambers had two openings, including the

1419:. Arbour and Mallon elaborated on this idea, describing the shape of these half-rings as "continuous U-shaped yokes" over the upper part of the neck, and suggested that 3271: 2609:

The osteoderms of ankylosaurids were thin in comparison to those of other ankylosaurs, and appear to have been strengthened by randomly distributed cushions of

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on the end of its tail. Bones in the skull and other parts of the body were fused, increasing their strength, and this feature is the source of the genus name.

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drainage, Montana, part of the Hell Creek Formation. In addition to these five incomplete specimens, many other isolated osteoderms and teeth have been found.

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may indicate earth-moving behavior. These factors, as well as the low rate of tooth formation in ankylosaurs compared to other ornithischians, indicate that

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over the eyes may have provided additional protection for them. Carpenter suggested in 1982 that the heavily vascularized armor may also have had a role in

955: in) for CMN 8880, and 6.02 to 7.95 m (19 ft 9 in to 26 ft 1 in) for AMNH 5214. Though the latter is the smallest specimen of 2385:

indicates that forward and sideways jaw movement was possible in these animals, the skull being able to withstand considerable forces. A 2016 study of the

1012:"), which pointed backwards to the sides from the back of the skull. The crest and horn were probably separate elements originally, as seen in the related 2637:

1979 that several hindlimb muscles would have controlled the swinging of the tail, and that violent thrusts of the club would have been able to break the

6180: 1453:. Osteoderms with oval keels could have been placed on the upper side of the tail or the side of the limbs. Compressed, triangular osteoderms found with 4295:(Ornithischia: Ankylosauria) from the Upper Cretaceous Alagteeg Formation, Alag Teeg, Mongolia: implications for ontogenetic allometry in ankylosaurs". 1445:. Smaller, triangular osteoderms may have been present on the sides of the pelvis. Flattened, pointed plates resemble those on the sides of the tail of 428:

is estimated to have been between 6 and 8 m (20 and 26 ft) long and to have weighed between 4.8 and 8 t (5.3 and 8.8 short tons). It was

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Arbour, V.M.; Currie, P.J.; Badamgarav, D. (2014). "The ankylosaurid dinosaurs of the Upper Cretaceous Baruungoyot and Nemegt formations of Mongolia".

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may have had three or four transverse rows of circular osteoderms over the pelvic region, which were smaller than those on the rest of the body, as in

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Thompson, R. S.; Parish, J. C.; Maidment, S. C. R.; Barrett, P. M. (2012). "Phylogeny of the ankylosaurian dinosaurs (Ornithischia: Thyreophora)".

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was very broad in this part of the body. The caudal vertebrae had centra that were slightly amphicoelous, meaning they were concave on both sides.

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franchise, where they are depicted as attacking with their tails and running, abilities that have been criticized as unlikely by paleontologists.

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teeth exhibit wear on the face of the crown rather than on the tip of the crown, as in nodosaurid ankylosaurs. In 1982 Carpenter ascribed to baby

6121: 1100:(tooth sockets) in the maxilla. The tooth rows in the maxillae of this specimen are about 20 centimeters (7.9 in) long. Each alveolus had a 2777:

dividing western and eastern North America during the Cretaceous, a broad coastal plain extending westward from the seaway to the newly formed

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does not appear to have congregated in groups (though some ankylosaurs appear to have congregated when young). It is therefore improbable that

2440:. A 2023 study by paleontologist Antonio Ballell and colleagues found that North American ankylosaurids from the latest Cretaceous (including 2679:

showed signs of injury on both the pelvic and tail area and the club was found to be asymmetrical, a sign of being worn down by the strikes.

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is thought to have been a slow-moving animal, able to make quick movements when necessary. Its broad muzzle indicates it was a non-selective

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was shorter in relation to its body length than those of other ankylosaurids, or that it had the same proportions but with a smaller club.

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differed from those of the holotype specimen in some details, and though he expressed a "considerate temptation" to name a new species of

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In 2017, based on comparisons with more complete ankylosaurines, Arbour and Mallon estimated a length of 7.56 to 9.99 m (24 ft

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skull are smaller than those of the smallest skull in the absolute sense. Some teeth from behind in the tooth row curved backwards, and

992:. The skull was low and triangular in shape, and wider than it was long; the back of the skull was broad and low. The skull had a broad 4874: 128: 2715: 1596:, Ankylosauridae, typified by massive, triangular skulls, short necks, stiff backs, broad bodies, and osteoderms. He also classified 1386:(K–L). Right: elements of the holotype that were identified in 2017, including parts of the skull (A–D) and cervical half rings (E–H) 2318:. Certain invertebrates, which the small teeth may have been adapted for handling, could also have provided supplemental nutrition. 1457:

specimens may have been placed on the sides of the pelvis or the tail. Ovoid, keeled, and teardrop-shaped osteoderms are known from

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Vickaryous, M. K.; Maryanska, T.; Weishampel, D. B. (2004). "Ankylosauria". In Weishampel, D. B.; Dodson, P.; Osmólska, H. (eds.).

4561:"A new ankylosaurid from the Upper Cretaceous Nemegt Formation of Mongolia and implications for paleoecology of armoured dinosaurs" 1117:

8880) is the same length. AMNH 5214 has 35 dental alveoli in the left dentary bone () and 36 in the right, for a total of 71. The

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in 2004, and that all the specimens belonged to the same species, noting that the teeth of other ankylosaurs are highly variable.

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cervical half-rings, which formed two semi-circular plates of armor around the upper part of the neck, as in the closely related

2829:. An abundance of fossil leaves found at dozens of different sites indicates that the area was largely forested by small trees. 1026:

horns were present, which pointed backward and down. The horns may have originally been osteoderms that fused to the skull. The

6245: 6185: 4959: 3639:"Rates of Dinosaur Body Mass Evolution Indicate 170 Million Years of Sustained Ecological Innovation on the Avian Stem Lineage" 2673:, or both. Other studies have found evidence of ankylosaurids using their tail clubs for intraspecific combat. One specimen of 2468:

proposed that the complex sinuses and nasal cavities of ankylosaurs may have lightened the weight of the skull, housed a nasal

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Rybczynski, N.; Vickaryous, M. K. (2001). "Chapter 14: Evidence of Complex Jaw Movement in the Late Cretaceous Ankylosaurid,

3621: 3388: 3251: 3213: 1077:, which separated the nasal passage from the sinus. Each side of the snout had five sinuses, four of which expanded into the 3286: 3168: 6190: 4980: 698: 3735:"Baby dinosaurs from the Late Cretaceous Lance and Hell Creek formations and a description of a new species of theropod" 1280: 796:

skull, but it is damaged in places. A section of caudal vertebrae (specimen CCM V03) was discovered in the 1960s in the

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Scheyer, T. M.; Sander, P. M. (2004). "Histology of ankylosaur osteoderms: implications for systematics and function".

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ate fruit, which its small, cusp-like teeth and the shape of its beak seem well adapted for, compared to for example

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Arbour, V. M.; Currie, P. J. (2015). "Systematics, phylogeny and palaeobiogeography of the ankylosaurid dinosaurs".

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The skeletal reconstruction accompanying the 1908 description restored the missing parts in a fashion similar to

4399: 2389:(contact between the teeth) of ankylosaur specimens found that the ability for backwards (palinal) jaw movement 3945:"Diversity and convergences in the evolution of feeding adaptations in ankylosaurs (Dinosauria: Ornithischia)" 1143: 6230: 6225: 2933: 2329:

two very small teeth that originate from the Lance and Hell Creek Formations and measure 3.2 to 3.3 mm (

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would have eaten 60 kilograms (130 pounds) of ferns per day, similar to the amount of dry vegetation a large

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were large, their number ranging from six to eight on the front part of the tooth, and five to seven behind.

1612:. Having for comparison only a few, incomplete members of the family, he believed the group was part of the 5999: 2660:

A 2009 study estimated that ankylosaurids could swing their tails at 100 degrees laterally, and the mainly

2436:(eating both plant and animal matter). It may also (or alternatively) have dug in the ground for roots and 1255: 4264:

Coombs, W. (1979). "Osteology and myology of the hindlimb in the Ankylosauria (Reptillia, Ornithischia)".

1270:, which also overlapped some of the vertebrae. The ribs of the last four back vertebrae were fused to the 782:

and T. Potter Chamney collected a skull and mandible (specimen CMN 8880, formerly NMC 8880), 1 kilometer (

2377: 1363: 691:, which Williston had named in 1905. Williston also stated that a skeletal reconstruction of the related 153: 2940:

an "iconic" dinosaur in 2017, and noted that the World's Fair sculpture, as well as the American artist

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are incompletely known, the hindfeet probably had three toes, as is the case in advanced ankylosaurids.

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Carpenter, K. (2001). "Chapter 21: Phylogenetic Analysis of the Ankylosauria". In Carpenter, K. (ed.).

2197:), Arbour and Currie suggested that earlier North American ankylosaurids had gone extinct by the late 6200: 3637:

Benson, R. B. J.; Campione, N. E.; Carrano, M. T.; Mannion, P. D.; Sullivan, C.; et al. (2014).

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Brown, B. (1908). "The Ankylosauridae, a new family of armored dinosaurs from the Upper Cretaceous".

2670: 1325: in) in diameter to 35.5 cm (14 in) in length, and varied in shape. The osteoderms of 1086: 779: 467: 383:, about 68–66 million years ago, in western North America, making it among the last of the non-avian 3984:"A complex hyobranchial apparatus in a Cretaceous dinosaur and the antiquity of avian paraglossalia" 5050: 1147: 678: 2994:-like tail club (a feature nodosaurids did not have), including in a mural by the American artist 617:('lizard'). The name can be translated as "fused lizard", "stiff lizard", or "curved lizard". The 6205: 4814: 4069:"Divergent strategies in cranial biomechanics and feeding ecology of the ankylosaurian dinosaurs" 3467:

Brown 1908 (Ankylosauridae) from the Upper Cretaceous of the Western Interior of North America".

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Coombs, W. (1990). "Teeth and taxonomy in ankylosaurs". In Carpenter, K.; Currie, P. J. (eds.).

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and can be distinguished from the teeth of other ankylosaurids based on their smooth sides. The

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Park JY, Lee YN, Kobayashi Y, Jacobs LL, Barsbold R, Lee HJ, Kim N, Song KY, Polcyn MJ (2021).

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formations, but it appears to have been rare in its environment. Although it lived alongside a

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by Arbour and Mallon in 2017, and though it was probably not a burrowing animal, the snout of

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of Wyoming in 1900. He mentioned these osteoderms (specimen AMNH 5866) in his description of

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and other Late Cretaceous North American ankylosaurids were grouped with Asian genera (in a

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Size of the smallest (AMNH 5214) and largest (CMN 8880) known specimens, compared to a human

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had armor plates that covered the cheek region, which may have been embedded in the flesh.

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criticized the skeletal reconstruction as being based on too few remains, and claimed that

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existed between 68 and 66 million years ago, in the final, or Maastrichtian, stage of the

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idea, as tail club shape is highly variable among ankylosaurids, even in the same genus.

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had high keels stretching from one margin to the other on the midline of its osteoderms.

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Currie, P. J.; Badamgarav, D.; Koppelhus, E. B.; Sissons, R.; Vickaryous, M. K. (2011).

4308: 4101: 4084: 4068: 3881: 3734: 3480: 2465: 1399: 5173: 4595: 4560: 4536: 4501: 4442: 4411: 4403: 4368: 4320: 4273: 4243: 4204: 3964: 3893: 3822: 3798:"Ankylosaurid dinosaur tail clubs evolved through stepwise acquisition of key features" 3797: 3778: 3712: 3665: 3638: 3359: 2868: 2864: 2763: 1303: 1246: 836: 768: 483: 148: 4465:(Ornithischia: Ankylosauridae) from the Late Cretaceous Oldman Formation of Alberta". 4337: 2310:

would consume. The requirements for nutrition could have been more effectively met if

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Coombs, W. (1978). "Forelimb muscles of the Ankylosauria (Reptilia, Ornithischia)".

3968: 3897: 3782: 2902: 2580:

specimen are blunter than those of the smallest specimen, which is also the case in

2515:(sense of smell) as the olfactory region is pushed to the sides of the main airway. 1206:(upper arm bone) of AMNH 5214 was short, very broad and about 54 cm (1 ft 5395: 5342: 5180: 4919: 4687:

Lofgren, D. F. (1997). "Hell Creek Formation". In Currie, P.J.; Padian, K. (eds.).

4590: 4580: 4531: 4521: 4482: 4395: 4358: 4312: 4199: 4191: 4096: 4088: 4020: 3995: 3956: 3920: 3885: 3850: 3817: 3809: 3770: 3759:

Coombs, W. P. (1978). "Theoretical aspects of cursorial adaptations in dinosaurs".

3704: 3660: 3650: 3587: 3511: 3484: 3349: 3205: 3149: 3044: 2638: 2622: 2618: 2473: 2448:, which suggests diverging jaw mechanics and dietary partitioning between the two. 2255: 1719:

Ankylosaurinae (members of which are called ankylosaurines) within Ankylosauridae.

1593: 1405: 1354: 1180:, and its hind limbs were longer than its forelimbs. In the holotype specimen, the 1043: 817: 499: 491: 380: 4170: 3960: 3708: 3502:

Osborn, H. F. (1905). "Tyrannosaurus and other Cretaceous carnivorous dinosaurs".

2357:

like in modern herbivorous lizards, which have several chambers in their enlarged

1068:

or upward. Additionally, the nostrils were not visible from the front because the

5860: 5824: 5764: 5730: 5723: 5616: 5557: 5429: 5326: 5242: 5202: 5035: 4712:

Breithaupt, B. H. (1997). "Lance Formation". In Currie, P.J.; Padian, K. (eds.).

4526: 3889: 3655: 3124: 2932:, perhaps due to a life-sized reconstruction of the animal being featured at the 2929: 2810: 2778: 2773:

fossils have been found represent different sections of the western shore of the

2711: 2565: 2358: 2190: 1801: 1653: 1613: 1281: 1130:

could accommodate more teeth than other ankylosaurines. The teeth of the largest

1121:

bone of the tip of the mandibles has not yet been found. Like other ankylosaurs,

1097: 1039: 1027: 1001: 711: 479: 442:

is a member of the family Ankylosauridae, and its closest relatives appear to be

392: 37: 4762:

Johnson, K. R. (1997). "Hell Creek Flora". In Currie, P. J.; Padian, K. (eds.).

3172: 1686: 1273: 1033: 648:

1908 skeletal reconstruction of the holotype, with missing parts restored after

5923: 5909: 5895: 5803: 5776: 5672: 5653: 5568: 5514: 5379: 5349: 5319: 5282: 5258: 4632: 4585: 4092: 2853: 2849: 2524: 2500: 2066: 2040: 1746: 1589: 1411: 1374: 1005: 865: 772: 744: 731: 670: 634: 626: 450: 444: 433: 429: 421: 272: 259: 2817:. In the Hell Creek Formation, many types of plants were supported, primarily 2456: 2364: 6169: 5886: 5796: 5789: 5741: 5685: 5632: 5598: 5499: 5445: 5420: 5402: 5305: 5251: 5224: 5216: 4737:

Eberth, D. A. (1997). "Edmonton Group". In Currie, P. J.; Padian, K. (eds.).

3592: 3571: 3004: 2978: 2874: 2859: 2601: 2421: 2274:

cropping, although not to the extent seen in some related genera, especially

2254:

Back vertebra with fused ribs of the holotype. The broad body housed a large

2194: 1921: 1760: 1736: 1708: 1704: 1667: 1640: 1631: 1135: 1030:-like cranial ornamentation on the surfaces of ankylosaurs skulls is called " 706: 584: 524: 512: 285: 79: 41: 4788: 2286:, they may have had a varied diet, including tough leaves and pulpy fruits. 1198:(a rectangular bone connected to the lower end of the scapula). It also had 792:

mile) north of where the 1910 specimen was found. This is the largest-known

6071: 5916: 5831: 5715: 5639: 5584: 5535: 5507: 5492: 5479: 5452: 5371: 5356: 5312: 5292: 5159: 5141: 5119: 5089: 5062: 5010: 4968: 4604: 4545: 4213: 4110: 3831: 3674: 2880: 2834: 2814: 2758:

was able to modify the landscape of its ecosystem in the way elephants do;

2504: 2485: 2417: 2369: 1935: 1818: 1620: 1530: in) tall. The club of the largest specimen may have been 57 cm ( 1263: 1105: 1074: 1014: 618: 604:, the stiffness produced by the fusion of bones in the skull and body, and 556: 553: 495: 406: 388: 305: 246: 214: 20: 4363: 2209:. Ankylosaurids thereafter recolonized North America from Asia during the 972: 755: 738:

in 1905. More recent examination has shown them to be similar to those of

6080: 5951: 5868: 5817: 5757: 5708: 5550: 5528: 5460: 5409: 5209: 5187: 5019: 4502:"Estimating impact forces of tail club strikes by ankylosaurid dinosaurs" 2838: 2759: 2626: 2283: 2090: 1784: 1696: 1662: 1616: 1468: 1441: 693: 659: 650: 518: 365: 230: 54: 4316: 2954:

with a tail club, following the first discovery of the feature in 1910.

2648: 853: 536: 6113: 5944: 5937: 5875: 5750: 5701: 5693: 5646: 5591: 5577: 5543: 5521: 5387: 5231: 5166: 4446: 4407: 4277: 4247: 3363: 2972: 2886: 2844: 2818: 2798: 2790: 2714:

Period. It was among the last dinosaur genera that appeared before the

2661: 2555: 2544: 2445: 2206: 2202: 1890: 1700: 1695:

Ankylosauria and Stegosauria are now grouped together within the clade

1286:(the structures that articulated the ribs with the vertebrae), and the 1118: 1023: 997: 687: 576: 377: 99: 64: 4000: 3983: 3924: 3813: 3572:"Unusual cranial and postcranial anatomy in the archetypal ankylosaur 1329:

were generally thin walled and hollowed on the underside. Compared to

988:(changes happening during decay and fossilization of the remains) and 984:

skulls differ in various details; this is thought to be the result of

5958: 5838: 5625: 5605: 5467: 5364: 5266: 4024: 3854: 3515: 3153: 2925: 2806: 2802: 2782: 2726: 2642: 2528: 2512: 2413: 2299: 2267: 2210: 1966: 1849: 1732: 1716: 1473: 1383: 1298: 1177: 1090: 1055: 985: 665: 601: 414: 165: 104: 48: 6042: 4486: 3845:

Osborn, H. F. (1923). "Two Lower Cretaceous dinosaurs of Mongolia".

3488: 3337: 2539:, little is known about this feature. The position of the orbits of 2250: 6065: 5965: 5001: 4992: 4461:

Carpenter, K. (1982). "Skeletal and dermal armor reconstruction of

3774: 3354: 2786: 2691: 2610: 2588:

variation (related to growth development). Studies of specimens of

2585: 2477: 2433: 2307: 2214: 2107: 1461:, and may have been placed on the forelimbs, like those known from 1258: 1251: 1199: 1195: 1162: 1109: 541: 384: 198: 185: 94: 89: 74: 69: 59: 1703:

age, and survived for 135 million years until disappearing in the

5853: 5845: 4291:

Burns, M; Tumanova, T; Currie, P (2015). "Postcrania of juvenile

4171:

Miyashita, T.; Arbour V. M.; Witmer L. M.; Currie, P. J. (2011).

2822: 2797:

and other Late Cretaceous ankylosaurs have been found had a warm

2675: 1873: 1287: 1203: 1181: 1101: 1078: 1069: 1009: 748:

osteoderms, which had been erroneously cataloged as belonging to

572: 109: 84: 6100: 3982:

Hill, R. V.; D'Emic, M. D.; Bever, G. S.; Norell, M. A. (2015).

4986: 4937: 4037: 3272:"The families of the ornithischian dinosaur order Ankylosauria" 2962:

present in many traditional depictions, but are not known from

2481: 2234: 2198: 1267: 1173: 644: 369: 175: 2833:

shared its environment with other dinosaurs that included the

2397:, including Late Cretaceous North American ankylosaurids like 641:('belly'), referring to the great width of the animal's body. 5810: 3867: 2826: 2536: 2469: 2437: 2295: 2279: 1423:

had six keeled osteoderms with oval bases on each half-ring.

1346:

had some smaller osteoderms with a keel across the midline.

1224:(thigh bone), also from AMNH 5214, was 67 cm (2 ft 1221: 663:, and Brown likened the result to the extinct armored mammal 611: 605: 594: 588: 361: 3636: 3074: 2699:

fossils have been discovered; the holotype is shown in red (

1572:

being wider. The larger width may indicate that the tail of

1465:, but it is unknown whether the hindlimbs bore osteoderms. 763:

In 1910, another AMNH expedition led by Brown discovered an

5972: 4789:"Cretaceous 'Hell Creek Faunal Facies'; Late Maastrichtian" 4400:

10.1671/0272-4634(2004)024[0874:hoaoif]2.0.co;2

4385: 3690: 3688: 3686: 3684: 3077: 3065: 3059: 3053: 2605:

Possible back osteoderm (holotype), in outer and inner view

2518:

According to Carpenter, the shape of the nasal chambers of

2508: 2291: 1623:, thereby placing the ankylosaurids in their own suborder. 1548:

was semicircular when seen from above, similar to those of

993: 417:

member of its group, despite having some unusual features.

16:

Ankylosaurid dinosaur genus from the Late Cretaceous Period

4618:

Thulborn, T. (1993). "Mimicry in ankylosaurid dinosaurs".

4177:

corroborated by X-ray computed tomographic reconstruction"

4015:

Haas, G. (1969). "On the jaw musculature of ankylosaurs".

3694: 3315:. London: A & C Black Publishers LTD. pp. 58–59. 2460:

Diagram showing nasal chambers inside the snout (holotype)

4173:"The internal cranial morphology of an armoured dinosaur 4067:

Ballell, Antonio; Mai, Bohao; Benton, Michael J. (2023).

3050: 2237:

record of North American ankylosaurids between the ages.

583:

the animal in 1908; the generic name is derived from the

3943:Ősi, A.; Prondvai, E.; Mallon, J.; Bodor, E. R. (2016). 3681: 1202:(connective tissue) for various muscle attachments. The 3193: 820:

within the nasal chambers. The American paleontologist

4558: 3942: 3910: 2928:

member of its group, and the best-known ankylosaur in

1739:

of the Ankylosaurinae conducted by Arbour and Currie:

759:

Excavation of AMNH 5214 (center, above the pick), 1910

4903: 3242:. McFarland & Company, Inc. Publishers. pp. 3089: 3080: 3071: 3056: 3047: 807:

pointed out that the teeth of two skulls assigned to

704:

Brown had collected 77 osteoderms while excavating a

600:('bent' or 'crooked'), referring to the medical term 4666:. University of California Press. pp. 517–606. 4060: 3062: 2476:. Carpenter rejected these hypotheses, arguing that 2270:. Its wide muzzle was adapted for non-selective low- 4637:. University of California Press. pp. 363–92. 4290: 4042:(Dinosauria: Thyreophora)". In K. Carpenter (ed.). 3068: 2233:. The theory explains a 30-million-year gap in the 1315:

proposed. The osteoderms ranged from 1 centimeter (

409:are represented by more extensive fossil material, 4626: 3169:"Dinosauria Translation and Pronunciation Guide A" 3146:Bulletin of the American Museum of Natural History 1238: in) long and very robust. While the feet of 1184:(shoulder blade) measures 61.5 cm (2 ft 1108:could be seen. Compared to other ankylosaurs, the 742:; it seems that Brown had compared them with some 4711: 3728: 3726: 3613:Dinosaur systematics: Approaches and perspectives 3458: 3456: 3454: 3452: 3450: 3448: 3446: 3444: 3442: 3440: 3438: 3436: 3434: 3432: 3430: 3428: 3426: 3424: 3422: 3420: 1378:(A–B) compared with half-ring fragments (C–J) of 1138:were usually flatter on one side than the other. 6167: 4284: 4122: 4120: 4066: 3418: 3416: 3414: 3412: 3410: 3408: 3406: 3404: 3402: 3400: 3335: 3616:. Cambridge University Press. pp. 269–79. 3605: 3603: 2564:diggers, though the hoof-like structure of the 2531:, it is unlikely either possibility applies to 726:also expressed this view when he described the 5034: 4761: 4705: 4686: 4046:. Indiana University Press. pp. 299–317. 3723: 3630: 2368:Skull of specimen CMN 8880, the largest-known 4953: 4736: 4499: 4460: 4454: 4126: 4117: 4031: 3844: 3758: 3501: 3462: 3397: 3383:. Indiana University Press. pp. 454–83. 3376: 3370: 1449:, and may have been distributed similarly on 1254:, which helped support the massive head. The 1059:Tooth of the holotype in inner and outer view 4617: 4611: 3904: 3795: 3600: 3569: 3565: 3563: 3561: 3559: 3557: 3555: 3553: 3551: 3549: 3547: 3545: 3329: 3265: 3263: 3166: 3160: 544:AMNH 5895 and reconstruction diagram of same 5139: 4755: 4680: 4259: 4257: 4129:"Ankylosauridae (Dinosauria) from Mongolia" 3543: 3541: 3539: 3537: 3535: 3533: 3531: 3529: 3527: 3525: 3139: 3137: 3135: 2950:and other later popular depictions, showed 2936:in New York City. Arbour and Mallon called 2781:. These formations are composed largely of 2559:Shoulder blade and coracoid of the holotype 1176:, tail, and feet, is still unknown. It was 910: in) long and 64.5 cm (2 ft 882: in) long and 74.5 cm (2 ft 6181:Late Cretaceous dinosaurs of North America 4960: 4946: 4730: 4656: 4654: 4422: 4331: 4263: 4233: 3838: 3609: 3495: 3269: 3227: 3225: 2408:The retracted position of the nostrils of 1556:but unlike the pointed club osteoderms of 1372:Left: Cervical half ring from the neck of 1081:bone. The nasal cavities (or chambers) of 1042:of the skull itself. This obliterated the 1008:horns (their shape has been described as " 127: 4594: 4584: 4535: 4525: 4428: 4362: 4203: 4166: 4164: 4100: 3999: 3988:Zoological Journal of the Linnean Society 3913:Zoological Journal of the Linnean Society 3821: 3754: 3752: 3732: 3664: 3654: 3591: 3353: 3260: 3143: 2957:Many traditional popular depictions show 2743:, whether their range overlapped or not. 2372:, including lower jaw (E–F) and tooth (G) 2278:. Though ankylosaurs may not have fed on 1168:The structure of much of the skeleton of 976:Skull (AMNH 5214) from the side and above 677:description, the American paleontologist 4879:franchise gets wrong about paleontology" 4845:"The real scientific history behind the 4815:"We asked a paleontologist how accurate 4254: 4014: 4008: 3861: 3522: 3463:Carpenter, K. (2004). "Redescription of 3132: 2901: 2897: 2690: 2647: 2600: 2554: 2455: 2363: 2249: 1661: 1467: 1297: 1157: 1054: 971: 852: 754: 643: 535: 4786: 4651: 4379: 3222: 2451: 2217:ages of the Late Cretaceous, and there 1153: 531: 510:also lived alongside dinosaurs such as 6168: 4872: 4866: 4842: 4795:from the original on December 26, 2009 4620:Records of the South Australian Museum 4493: 4429:Coombs W. (1972). "The Bony Eyelid of 4227: 4161: 3975: 3938: 3936: 3934: 3749: 2970:has often been conflated with that of 2491:A 2011 study of the nasal passages of 1723:appears to be most closely related to 1194: in) long and was fused with the 502:do not appear to have overlapped, and 6047: 6046: 4979: 4941: 4807: 3796:Arbour, V. M.; Currie, P. J. (2015). 3310: 3187: 2716:Cretaceous–Paleogene extinction event 1656:from the tail (right) of the holotype 1302:Arrangement of armor as suggested by 579:(armor plates). Brown scientifically 4836: 4780: 4741:. Academic Press. pp. 199–204. 3231: 1711:(these families lacked tail clubs). 4766:. Academic Press. pp. 300–02. 4716:. Academic Press. pp. 394–95. 4691:. Academic Press. pp. 302–03. 3931: 3870:Journal of Systematic Palaeontology 3697:Journal of Systematic Palaeontology 3570:Arbour, V.M.; Mallon, J.C. (2017). 3194:Liddell, H. G.; Scott, R. (1980) . 2686: 2576:The squamosal horns of the largest 1699:. This group first appeared in the 824:accepted the teeth as belonging to 13: 4467:Canadian Journal of Earth Sciences 4388:Journal of Vertebrate Paleontology 3469:Canadian Journal of Earth Sciences 2735:it had a more selective diet than 1478:American Museum of Natural History 685:was merely a synonym of the genus 550:American Museum of Natural History 14: 6257: 4899: 2913:(note spikes and dragging tail), 2863:, an indeterminate nodosaur, the 2550: 2221:again, leading to genera such as 1619:. In 1923 Osborn coined the name 1579: 1560:or the narrow, elongated club of 1544: in) wide. The tail club of 803:In 1990, American paleontologist 718:but thought they belonged to the 506:may have inhabited upland areas. 140:skull (AMNH 5214) in front view, 6027: 6026: 6003: 5998: 4967: 4925: 4913: 4196:10.1111/j.1469-7580.2011.01427.x 3043: 2793:environments. The regions where 2789:, which have been attributed to 2746:With its low center of gravity, 2613:fibers. Structurally similar to 2480:animals make sounds through the 1639: 1630: 1588:so distinct that he made it the 1516: in) wide, and 19 cm ( 1483:The tail club (or tail knob) of 1362: 1353: 1104:(opening) near its side where a 152: 52: 4552: 4340:"Hands, feet, and behaviour in 3789: 3762:The Quarterly Review of Biology 3018:Timeline of ankylosaur research 2990:has also been depicted with an 2664:clubs would have had a lowered 2240: 6241:Thyreophorans of North America 3313:The Great Dinosaur Discoveries 3304: 3125:Merriam-Webster.com Dictionary 3112: 3035: 2762:may instead have had such an " 1382:(holotype), and possible back 848: 376:dating to the very end of the 1: 6246:Late Cretaceous thyreophorans 6186:Fossil taxa described in 1908 4764:The Encyclopedia of Dinosaurs 4739:The Encyclopedia of Dinosaurs 4714:The Encyclopedia of Dinosaurs 4689:The Encyclopedia of Dinosaurs 4351:Acta Palaeontologica Polonica 4344:(Dinosauria: Ankylosauridae)" 3961:10.1080/08912963.2016.1208194 3709:10.1080/14772019.2015.1059985 3167:Creisler, B. (July 7, 2003). 3023: 2920:Carpenter noted in 2004 that 1600:(only known from teeth), and 1096:Specimen AMNH 5214 has 34–35 498:ankylosaur, their ranges and 4527:10.1371/journal.pone.0006738 3890:10.1080/14772019.2011.569091 3656:10.1371/journal.pbio.1001853 3338:"Review: The Ankylosauridae" 3285:(1): 143–170. Archived from 2986:being depicted with spikes, 2472:, or acted as a chamber for 1502: in) long, 49 cm ( 767:specimen (AMNH 5214) in the 697:by Hungarian paleontologist 612: 595: 7: 4571:(1): Article number 22928. 4433:(Reptilia, Ornithischia)". 3239:Dinosaurs, the encyclopedia 3011: 2809:, had occasional rainfall, 2503:and the looping trachea of 2262:Like other ornithischians, 552:expedition led by American 420:Possibly the largest-known 10: 6262: 6191:Taxa named by Barnum Brown 4843:Waxman, Olivia B. (2018). 4586:10.1038/s41598-021-02273-4 4093:10.1038/s41598-023-45444-1 2596: 2412:were compared to those of 2245: 1715:is considered part of the 1392:Like other ankylosaurids, 1038:", and were the result of 606: 589: 18: 6236:Monotypic dinosaur genera 6055: 6016: 5996: 5885: 5784: 5775: 5740: 5680: 5671: 5615: 5567: 5487: 5478: 5419: 5300: 5291: 5280: 5241: 5154: 5150: 5135: 5116: 5088: 5070: 5061: 5043: 5030: 4975: 4017:American Museum Novitates 3847:American Museum Novitates 3574:Ankylosaurus magniventris 3465:Ankylosaurus magniventris 3336:Williston, S. W. (1908). 3002:has been featured in the 2584:, and this may represent 2571: 2290:probably fed on abundant 2104: 2087: 2080: 2063: 2037: 2020: 2013: 2006: 1980: 1963: 1956: 1949: 1932: 1925: 1887: 1870: 1863: 1846: 1839: 1832: 1815: 1798: 1781: 1774: 1757: 1750: 1652:from the neck (left) and 1488:AMNH 5214 is 60 cm ( 1022:. Below the upper horns, 565:Ankylosaurus magniventris 342:Ankylosaurus magniventris 335: 328: 149:Scientific classification 147: 135: 126: 30: 4142:: 85–151. Archived from 3593:10.1139/facets-2017-0063 2976:(earlier referred to as 2656:displaying its tail club 2464:In 1977, paleontologist 2393:in different ankylosaur 2376:In 1969, paleontologist 1293: 1172:, including most of the 967: 722:instead. Paleontologist 679:Samuel Wendell Williston 413:is often considered the 19:Not to be confused with 6221:Paleontology in Alberta 6216:Paleontology in Wyoming 6211:Paleontology in Montana 4435:Journal of Paleontology 4297:Journal of Paleontology 4266:Journal of Paleontology 4236:Journal of Paleontology 4136:Palaeontologia Polonica 3342:The American Naturalist 3279:Journal of Paleontology 3202:Oxford University Press 3197:A Greek-English Lexicon 2775:Western Interior Seaway 1691:(cranial ornamentation) 1310:A prominent feature of 941: in to 32 ft 474:have been found in the 4500:Arbour, V. M. (2009). 4223:on September 24, 2015. 4127:Maryanska, T. (1977). 3739:Rocky Mountain Geology 3733:Carpenter, K. (1982). 2917: 2704: 2657: 2606: 2560: 2461: 2373: 2259: 1692: 1480: 1307: 1165: 1060: 977: 864:was the largest-known 858: 760: 724:Henry Fairfield Osborn 654: 638: 630: 571:, near Gilbert Creek, 545: 6148:Paleobiology Database 4873:Dorsch, Rita (2022). 4364:10.4202/app.2010.0055 4293:Pinacosaurus grangeri 4044:The Armored Dinosaurs 3380:The Armored Dinosaurs 3200:(abridged ed.). 2966:itself. The armor of 2905: 2898:Cultural significance 2769:The formations where 2741:ecological separation 2739:, further indicating 2694: 2651: 2604: 2558: 2459: 2391:evolved independently 2367: 2253: 1737:phylogenetic analysis 1665: 1471: 1301: 1161: 1058: 975: 856: 758: 647: 539: 374:geological formations 142:Museum of the Rockies 6231:Ornithischian genera 6226:Maastrichtian genera 4664:The Dinosauria (2nd) 4463:Euoplocephalus tutus 4040:Euoplocephalus tutus 3504:Bulletin of the AMNH 3232:Glut, D. F. (1997). 2915:Royal Alberta Museum 2872:, and the theropods 2671:intraspecific combat 2452:Airspaces and senses 2355:hindgut fermentation 1684: 1397: 1333:, the osteoderms of 1279: 1271: 1220: in) long. The 1154:Postcranial skeleton 1063:The snout region of 1031: 990:individual variation 780:Charles M. Sternberg 730:specimen as the now 625:is derived from the 569:Hell Creek Formation 532:History of discovery 468:intraspecific combat 321:A. magniventris 4577:2021NatSR..1122928P 4518:2009PLoSO...4.6738A 4479:1982CaJES..19..689C 4317:10.1017/jpa.2014.14 4309:2015JPal...89..168B 4085:2023NatSR..1318242B 3882:2012JSPal..10..301T 3481:2004CaJES..41..961C 3270:Coombs, W. (1978). 2947:The Age of Reptiles 2865:pachycephalosaurian 2821:, with less common 2545:stereoscopic vision 2205:ages of the Middle 1735:is based on a 2015 1400:cervical half-rings 567:(AMNH 5895) in the 372:have been found in 4565:Scientific Reports 4184:Journal of Anatomy 4073:Scientific Reports 3949:Historical Biology 3802:Journal of Anatomy 3311:Naish, D. (2009). 3175:on August 18, 2010 3128:. Merriam-Webster. 2982:); in addition to 2918: 2869:Pachycephalosaurus 2857:, the hadrosaurid 2764:ecosystem engineer 2705: 2695:Map showing where 2658: 2607: 2561: 2462: 2374: 2298:. Assuming it was 2260: 2193:the authors named 1693: 1481: 1308: 1247:cervical vertebrae 1166: 1061: 978: 859: 837:Victoria M. Arbour 831:Most of the known 769:Scollard Formation 761: 655: 546: 395:, containing only 387:. It was named by 6163: 6162: 6135:Open Tree of Life 6049:Taxon identifiers 6040: 6039: 6012: 6011: 5994: 5993: 5990: 5989: 5986: 5985: 5982: 5981: 5931:Nodocephalosaurus 5667: 5666: 5663: 5662: 5276: 5275: 5195:Serendipaceratops 5131: 5130: 5127: 5126: 5112: 5111: 5108: 5107: 4773:978-0-12-226810-6 4748:978-0-12-226810-6 4723:978-0-12-226810-6 4698:978-0-12-226810-6 4673:978-0-520-24209-8 4644:978-0-520-24209-8 4053:978-0-253-33964-5 4001:10.1111/zoj.12293 3925:10.1111/zoj.12185 3814:10.1111/joa.12363 3623:978-0-521-43810-0 3390:978-0-253-33964-5 3253:978-0-375-82419-7 3215:978-0-19-910207-5 2996:Charles R. Knight 2942:Rudolph Zallinger 2934:1964 World's Fair 2908:1964 World's Fair 2803:temperate climate 2666:moment of inertia 2632:The tail club of 2486:specialized bones 2182: 2181: 2173: 2172: 2164: 2163: 2155: 2154: 2146: 2145: 2137: 2136: 2128: 2127: 2119: 2118: 2052: 2051: 1995: 1994: 1984:Nodocephalosaurus 1911: 1910: 1902: 1901: 1683:with color coded 1680:Nodocephalosaurus 1650:Cervical vertebra 1584:Brown considered 1266:(turned to bone) 1106:replacement tooth 822:Kenneth Carpenter 777:fossil collectors 500:ecological niches 353: 352: 309: 6253: 6201:Hell Creek fauna 6156: 6155: 6143: 6142: 6130: 6129: 6117: 6116: 6104: 6103: 6091: 6090: 6089: 6076: 6075: 6074: 6044: 6043: 6030: 6029: 6007: 6002: 5782: 5781: 5678: 5677: 5485: 5484: 5396:Propanoplosaurus 5343:Glyptodontopelta 5298: 5297: 5289: 5288: 5181:Priodontognathus 5152: 5151: 5148: 5147: 5137: 5136: 5068: 5067: 5041: 5040: 5032: 5031: 5025: 5024: 4977: 4976: 4962: 4955: 4948: 4939: 4938: 4930: 4929: 4928: 4918: 4917: 4916: 4909: 4894: 4893: 4891: 4889: 4875:"Everything the 4870: 4864: 4863: 4861: 4859: 4840: 4834: 4833: 4831: 4829: 4811: 4805: 4804: 4802: 4800: 4784: 4778: 4777: 4759: 4753: 4752: 4734: 4728: 4727: 4709: 4703: 4702: 4684: 4678: 4677: 4658: 4649: 4648: 4630: 4624: 4623: 4615: 4609: 4608: 4598: 4588: 4556: 4550: 4549: 4539: 4529: 4497: 4491: 4490: 4458: 4452: 4450: 4426: 4420: 4419: 4383: 4377: 4376: 4366: 4348: 4335: 4329: 4328: 4288: 4282: 4281: 4261: 4252: 4251: 4231: 4225: 4224: 4222: 4216:. Archived from 4207: 4181: 4168: 4159: 4158: 4156: 4154: 4149:on July 12, 2020 4148: 4133: 4124: 4115: 4114: 4104: 4064: 4058: 4057: 4035: 4029: 4028: 4012: 4006: 4005: 4003: 3979: 3973: 3972: 3940: 3929: 3928: 3908: 3902: 3901: 3865: 3859: 3858: 3842: 3836: 3835: 3825: 3793: 3787: 3786: 3756: 3747: 3746: 3730: 3721: 3720: 3692: 3679: 3678: 3668: 3658: 3634: 3628: 3627: 3607: 3598: 3597: 3595: 3567: 3520: 3519: 3499: 3493: 3492: 3460: 3395: 3394: 3374: 3368: 3367: 3357: 3333: 3327: 3326: 3308: 3302: 3301: 3299: 3297: 3291: 3276: 3267: 3258: 3257: 3229: 3220: 3219: 3191: 3185: 3184: 3182: 3180: 3171:. Archived from 3164: 3158: 3157: 3141: 3130: 3129: 3116: 3100: 3096: 3092: 3087: 3086: 3083: 3082: 3079: 3076: 3073: 3070: 3067: 3064: 3061: 3058: 3055: 3052: 3049: 3039: 2725:are rare in the 2702: 2687:Paleoenvironment 2639:metatarsal bones 2623:thermoregulation 2615:Sharpey's fibres 2466:Teresa Maryańska 2387:dental occlusion 2348: 2347: 2343: 2338: 2337: 2333: 2294:and low-growing 2256:digestion system 2083: 2082: 2016: 2015: 2009: 2008: 1959: 1958: 1952: 1951: 1928: 1927: 1866: 1865: 1842: 1841: 1835: 1834: 1777: 1776: 1753: 1752: 1743: 1742: 1731:. The following 1688: 1643: 1634: 1543: 1542: 1538: 1535: 1529: 1528: 1524: 1521: 1515: 1514: 1510: 1507: 1501: 1500: 1496: 1493: 1406:Philip J. Currie 1401: 1366: 1357: 1324: 1323: 1319: 1306:and Mallon, 2017 1283: 1275: 1237: 1236: 1232: 1229: 1219: 1218: 1214: 1211: 1193: 1192: 1188: 1163:Life restoration 1035: 980:The three known 954: 953: 949: 946: 940: 939: 935: 932: 923: 922: 918: 915: 909: 908: 904: 901: 895: 894: 890: 887: 881: 880: 876: 873: 805:Walter P. Coombs 791: 790: 786: 710:specimen in the 615: 609: 608: 598: 592: 591: 366:armored dinosaur 344: 340: 304: 297: 284: 271: 258: 245: 229: 213: 157: 156: 131: 121: 51: 36:Temporal range: 28: 27: 6261: 6260: 6256: 6255: 6254: 6252: 6251: 6250: 6166: 6165: 6164: 6159: 6151: 6146: 6138: 6133: 6125: 6120: 6112: 6107: 6099: 6094: 6085: 6084: 6079: 6070: 6069: 6064: 6051: 6041: 6036: 6008: 5978: 5881: 5861:Tianzhenosaurus 5825:Minotaurasaurus 5771: 5765:Zhongyuansaurus 5736: 5731:Zhejiangosaurus 5724:Liaoningosaurus 5659: 5617:Struthiosaurini 5611: 5563: 5558:Tatankacephalus 5474: 5430:Gargoyleosaurus 5415: 5327:Gargoyleosaurus 5285: 5272: 5243:Parankylosauria 5237: 5203:Sinankylosaurus 5144: 5123: 5104: 5084: 5057: 5051:Avemetatarsalia 5036:Avemetatarsalia 5026: 4982: 4981: 4971: 4966: 4936: 4926: 4924: 4914: 4912: 4904: 4902: 4897: 4887: 4885: 4871: 4867: 4857: 4855: 4841: 4837: 4827: 4825: 4813: 4812: 4808: 4798: 4796: 4785: 4781: 4774: 4760: 4756: 4749: 4735: 4731: 4724: 4710: 4706: 4699: 4685: 4681: 4674: 4659: 4652: 4645: 4631: 4627: 4616: 4612: 4557: 4553: 4498: 4494: 4487:10.1139/e82-058 4459: 4455: 4427: 4423: 4384: 4380: 4346: 4336: 4332: 4289: 4285: 4262: 4255: 4232: 4228: 4220: 4179: 4169: 4162: 4152: 4150: 4146: 4131: 4125: 4118: 4065: 4061: 4054: 4036: 4032: 4013: 4009: 3980: 3976: 3941: 3932: 3909: 3905: 3866: 3862: 3843: 3839: 3794: 3790: 3757: 3750: 3731: 3724: 3693: 3682: 3649:(5): e1001853. 3635: 3631: 3624: 3608: 3601: 3568: 3523: 3510:(14): 259–265. 3500: 3496: 3489:10.1139/e04-043 3461: 3398: 3391: 3375: 3371: 3348:(501): 629–30. 3334: 3330: 3323: 3309: 3305: 3295: 3293: 3292:on July 8, 2015 3289: 3274: 3268: 3261: 3254: 3230: 3223: 3216: 3192: 3188: 3178: 3176: 3165: 3161: 3148:. 24: 187–201. 3142: 3133: 3118: 3117: 3113: 3104: 3103: 3094: 3090: 3046: 3042: 3040: 3036: 3026: 3014: 2930:popular culture 2924:has become the 2906:Replica of the 2900: 2811:tropical storms 2779:Rocky Mountains 2712:Late Cretaceous 2700: 2689: 2652:Restoration of 2619:palpebral bones 2599: 2574: 2553: 2525:olfactory lobes 2497:mucosa membrane 2474:vocal resonance 2454: 2345: 2341: 2340: 2335: 2331: 2330: 2248: 2243: 2183: 2174: 2165: 2156: 2147: 2138: 2129: 2120: 2053: 1996: 1912: 1903: 1802:Zhejiangosaurus 1690: 1660: 1659: 1658: 1657: 1654:caudal vertebra 1646: 1645: 1644: 1636: 1635: 1582: 1540: 1536: 1533: 1531: 1526: 1522: 1519: 1517: 1512: 1508: 1505: 1503: 1498: 1494: 1491: 1489: 1403: 1390: 1389: 1388: 1387: 1369: 1368: 1367: 1359: 1358: 1321: 1317: 1316: 1296: 1285: 1277: 1234: 1230: 1227: 1225: 1216: 1212: 1209: 1207: 1190: 1186: 1185: 1156: 1037: 970: 951: 947: 944: 942: 937: 933: 930: 928: 920: 916: 913: 911: 906: 902: 899: 897: 892: 888: 885: 883: 878: 874: 871: 869: 851: 826:A. magniventris 809:A. magniventris 788: 784: 783: 712:Lance Formation 559:discovered the 534: 470:. Specimens of 398:A. magniventris 391:in 1908; it is 349: 346: 338: 337: 324: 303: 295: 282: 269: 256: 243: 227: 211: 151: 122: 120: 119: 118: 117: 112: 107: 102: 97: 92: 87: 82: 77: 72: 67: 62: 57: 46: 45: 38:Late Cretaceous 34: 24: 17: 12: 11: 5: 6259: 6249: 6248: 6243: 6238: 6233: 6228: 6223: 6218: 6213: 6208: 6206:Scollard fauna 6203: 6198: 6193: 6188: 6183: 6178: 6161: 6160: 6158: 6157: 6144: 6131: 6118: 6105: 6092: 6077: 6061: 6059: 6053: 6052: 6038: 6037: 6035: 6034: 6024: 6017: 6014: 6013: 6010: 6009: 5997: 5995: 5992: 5991: 5988: 5987: 5984: 5983: 5980: 5979: 5977: 5976: 5969: 5962: 5955: 5948: 5941: 5934: 5927: 5924:Euoplocephalus 5920: 5913: 5910:Anodontosaurus 5906: 5899: 5896:Akainacephalus 5891: 5889: 5883: 5882: 5880: 5879: 5872: 5865: 5857: 5850: 5842: 5835: 5828: 5821: 5814: 5807: 5804:Crichtonsaurus 5800: 5793: 5785: 5779: 5777:Ankylosaurinae 5773: 5772: 5770: 5769: 5761: 5754: 5746: 5744: 5738: 5737: 5735: 5734: 5727: 5720: 5712: 5705: 5698: 5690: 5681: 5675: 5673:Ankylosauridae 5669: 5668: 5665: 5664: 5661: 5660: 5658: 5657: 5654:Struthiosaurus 5650: 5643: 5636: 5629: 5621: 5619: 5613: 5612: 5610: 5609: 5602: 5595: 5588: 5581: 5573: 5571: 5569:Panoplosaurini 5565: 5564: 5562: 5561: 5554: 5547: 5540: 5532: 5525: 5518: 5515:Niobrarasaurus 5511: 5504: 5496: 5488: 5482: 5476: 5475: 5473: 5472: 5464: 5457: 5449: 5442: 5434: 5425: 5423: 5417: 5416: 5414: 5413: 5406: 5399: 5392: 5384: 5380:Polacanthoides 5376: 5368: 5361: 5353: 5350:Horshamosaurus 5346: 5339: 5331: 5323: 5320:Dongyangopelta 5316: 5309: 5301: 5295: 5286: 5283:Euankylosauria 5281: 5278: 5277: 5274: 5273: 5271: 5270: 5263: 5259:Kunbarrasaurus 5255: 5247: 5245: 5239: 5238: 5236: 5235: 5228: 5221: 5213: 5206: 5199: 5191: 5184: 5177: 5170: 5163: 5155: 5145: 5140: 5133: 5132: 5129: 5128: 5125: 5124: 5117: 5114: 5113: 5110: 5109: 5106: 5105: 5103: 5102: 5101: 5100: 5094: 5092: 5086: 5085: 5083: 5082: 5081: 5080: 5071: 5065: 5059: 5058: 5056: 5055: 5054: 5053: 5044: 5038: 5028: 5027: 5023: 5022: 5013: 5004: 4995: 4989: 4973: 4972: 4965: 4964: 4957: 4950: 4942: 4935: 4934: 4922: 4901: 4900:External links 4898: 4896: 4895: 4865: 4835: 4817:Jurassic World 4806: 4779: 4772: 4754: 4747: 4729: 4722: 4704: 4697: 4679: 4672: 4650: 4643: 4635:The Dinosauria 4625: 4610: 4551: 4492: 4453: 4431:Euoplocephalus 4421: 4378: 4357:(3): 489–504. 4330: 4303:(1): 168–182. 4283: 4253: 4226: 4175:Euoplocephalus 4160: 4116: 4059: 4052: 4030: 4019:(2399): 1–11. 4007: 3994:(4): 892–909. 3974: 3955:(4): 539–570. 3930: 3919:(3): 631–652. 3903: 3876:(2): 301–312. 3860: 3837: 3788: 3775:10.1086/410790 3769:(4): 393–418. 3748: 3722: 3680: 3629: 3622: 3599: 3586:(2): 764–794. 3521: 3494: 3396: 3389: 3369: 3355:10.1086/278987 3328: 3322:978-1408119068 3321: 3303: 3259: 3252: 3234:"Ankylosaurus" 3221: 3214: 3186: 3159: 3131: 3120:"Ankylosaurus" 3110: 3102: 3101: 3033: 3032: 3025: 3022: 3021: 3020: 3013: 3010: 2944:'s 1947 mural 2899: 2896: 2854:Thescelosaurus 2850:hypsilophodont 2688: 2685: 2669:predators, in 2598: 2595: 2582:Euoplocephalus 2573: 2570: 2552: 2551:Limb movements 2549: 2501:saiga antelope 2493:Euoplocephalus 2453: 2450: 2432:may have been 2403:Euoplocephalus 2383:Euoplocephalus 2316:Euoplocephalus 2276:Euoplocephalus 2247: 2244: 2242: 2239: 2231:Euoplocephalus 2227:Anodontosaurus 2180: 2179: 2176: 2175: 2171: 2170: 2167: 2166: 2162: 2161: 2158: 2157: 2153: 2152: 2149: 2148: 2144: 2143: 2140: 2139: 2135: 2134: 2131: 2130: 2126: 2125: 2122: 2121: 2117: 2116: 2113: 2112: 2103: 2100: 2099: 2096: 2095: 2086: 2081: 2079: 2076: 2075: 2072: 2071: 2067:Euoplocephalus 2062: 2059: 2058: 2055: 2054: 2050: 2049: 2046: 2045: 2041:Anodontosaurus 2036: 2033: 2032: 2029: 2028: 2019: 2014: 2012: 2007: 2005: 2002: 2001: 1998: 1997: 1993: 1992: 1989: 1988: 1979: 1976: 1975: 1972: 1971: 1962: 1957: 1955: 1950: 1948: 1945: 1944: 1941: 1940: 1931: 1926: 1924: 1918: 1917: 1914: 1913: 1909: 1908: 1905: 1904: 1900: 1899: 1896: 1895: 1886: 1883: 1882: 1879: 1878: 1869: 1864: 1862: 1859: 1858: 1855: 1854: 1845: 1840: 1838: 1833: 1831: 1828: 1827: 1824: 1823: 1814: 1811: 1810: 1807: 1806: 1797: 1794: 1793: 1790: 1789: 1780: 1775: 1773: 1770: 1769: 1766: 1765: 1756: 1751: 1749: 1747:Ankylosaurinae 1741: 1729:Euoplocephalus 1725:Anodontosaurus 1675:Euoplocephalus 1666:Skulls of the 1648: 1647: 1638: 1637: 1629: 1628: 1627: 1626: 1625: 1606:Euoplocephalus 1602:Euoplocephalus 1581: 1580:Classification 1578: 1558:Anodontosaurus 1550:Euoplocephalus 1433:Euoplocephalus 1417:Euoplocephalus 1412:Anodontosaurus 1375:Euoplocephalus 1371: 1370: 1361: 1360: 1352: 1351: 1350: 1349: 1348: 1331:Euoplocephalus 1295: 1292: 1155: 1152: 1098:dental alveoli 1020:Euoplocephalus 969: 966: 850: 847: 773:Red Deer River 745:Euoplocephalus 633:('great') and 554:paleontologist 533: 530: 451:Euoplocephalus 445:Anodontosaurus 351: 350: 347: 333: 332: 326: 325: 317: 315: 311: 310: 293: 289: 288: 280: 276: 275: 273:Ankylosaurinae 267: 263: 262: 260:Ankylosauridae 254: 250: 249: 241: 234: 233: 225: 218: 217: 209: 202: 201: 196: 189: 188: 183: 179: 178: 173: 169: 168: 163: 159: 158: 145: 144: 133: 132: 124: 123: 115: 114: 113: 108: 103: 98: 93: 88: 83: 78: 73: 68: 63: 58: 53: 35: 15: 9: 6: 4: 3: 2: 6258: 6247: 6244: 6242: 6239: 6237: 6234: 6232: 6229: 6227: 6224: 6222: 6219: 6217: 6214: 6212: 6209: 6207: 6204: 6202: 6199: 6197: 6194: 6192: 6189: 6187: 6184: 6182: 6179: 6177: 6176:Ankylosaurids 6174: 6173: 6171: 6154: 6149: 6145: 6141: 6136: 6132: 6128: 6123: 6119: 6115: 6110: 6106: 6102: 6097: 6093: 6088: 6082: 6078: 6073: 6067: 6063: 6062: 6060: 6058: 6054: 6050: 6045: 6033: 6025: 6023: 6019: 6018: 6015: 6006: 6001: 5975: 5974: 5970: 5968: 5967: 5963: 5961: 5960: 5956: 5954: 5953: 5949: 5947: 5946: 5942: 5940: 5939: 5935: 5933: 5932: 5928: 5926: 5925: 5921: 5919: 5918: 5914: 5912: 5911: 5907: 5905: 5904: 5900: 5898: 5897: 5893: 5892: 5890: 5888: 5887:Ankylosaurini 5884: 5878: 5877: 5873: 5871: 5870: 5866: 5863: 5862: 5858: 5856: 5855: 5851: 5848: 5847: 5843: 5841: 5840: 5836: 5834: 5833: 5829: 5827: 5826: 5822: 5820: 5819: 5815: 5813: 5812: 5808: 5806: 5805: 5801: 5799: 5798: 5797:Crichtonpelta 5794: 5792: 5791: 5790:Bissektipelta 5787: 5786: 5783: 5780: 5778: 5774: 5767: 5766: 5762: 5760: 5759: 5755: 5753: 5752: 5748: 5747: 5745: 5743: 5742:Shamosaurinae 5739: 5733: 5732: 5728: 5726: 5725: 5721: 5718: 5717: 5713: 5711: 5710: 5706: 5704: 5703: 5699: 5696: 5695: 5691: 5688: 5687: 5686:Ahshislepelta 5683: 5682: 5679: 5676: 5674: 5670: 5656: 5655: 5651: 5649: 5648: 5644: 5642: 5641: 5637: 5635: 5634: 5633:Hungarosaurus 5630: 5628: 5627: 5623: 5622: 5620: 5618: 5614: 5608: 5607: 5603: 5601: 5600: 5599:Panoplosaurus 5596: 5594: 5593: 5589: 5587: 5586: 5582: 5580: 5579: 5575: 5574: 5572: 5570: 5566: 5560: 5559: 5555: 5553: 5552: 5548: 5546: 5545: 5541: 5538: 5537: 5533: 5531: 5530: 5526: 5524: 5523: 5519: 5517: 5516: 5512: 5510: 5509: 5505: 5502: 5501: 5500:Ahshislepelta 5497: 5495: 5494: 5490: 5489: 5486: 5483: 5481: 5477: 5470: 5469: 5465: 5463: 5462: 5458: 5455: 5454: 5450: 5448: 5447: 5446:Hoplitosaurus 5443: 5440: 5439: 5435: 5432: 5431: 5427: 5426: 5424: 5422: 5421:Polacanthinae 5418: 5412: 5411: 5407: 5405: 5404: 5403:Rhadinosaurus 5400: 5398: 5397: 5393: 5390: 5389: 5385: 5382: 5381: 5377: 5374: 5373: 5369: 5367: 5366: 5362: 5359: 5358: 5354: 5352: 5351: 5347: 5345: 5344: 5340: 5337: 5336: 5332: 5329: 5328: 5324: 5322: 5321: 5317: 5315: 5314: 5310: 5308: 5307: 5306:Acanthopholis 5303: 5302: 5299: 5296: 5294: 5290: 5287: 5284: 5279: 5269: 5268: 5264: 5261: 5260: 5256: 5254: 5253: 5252:Antarctopelta 5249: 5248: 5246: 5244: 5240: 5234: 5233: 5229: 5227: 5226: 5225:Tianchisaurus 5222: 5219: 5218: 5217:Stegosaurides 5214: 5212: 5211: 5207: 5205: 5204: 5200: 5197: 5196: 5192: 5190: 5189: 5185: 5183: 5182: 5178: 5176: 5175: 5171: 5169: 5168: 5164: 5162: 5161: 5157: 5156: 5153: 5149: 5146: 5143: 5138: 5134: 5122: 5121: 5115: 5098: 5097: 5096: 5095: 5093: 5091: 5087: 5079: 5075: 5074: 5073: 5072: 5069: 5066: 5064: 5060: 5052: 5048: 5047: 5046: 5045: 5042: 5039: 5037: 5033: 5029: 5021: 5017: 5014: 5012: 5008: 5005: 5003: 4999: 4996: 4994: 4990: 4988: 4984: 4983: 4978: 4974: 4970: 4963: 4958: 4956: 4951: 4949: 4944: 4943: 4940: 4933: 4932:United States 4923: 4921: 4911: 4910: 4907: 4884: 4880: 4878: 4877:Jurassic Park 4869: 4854: 4850: 4848: 4847:Jurassic Park 4839: 4823: 4820: 4819:really is..." 4818: 4810: 4794: 4790: 4783: 4775: 4769: 4765: 4758: 4750: 4744: 4740: 4733: 4725: 4719: 4715: 4708: 4700: 4694: 4690: 4683: 4675: 4669: 4665: 4657: 4655: 4646: 4640: 4636: 4629: 4622:. 27: 151–58. 4621: 4614: 4606: 4602: 4597: 4592: 4587: 4582: 4578: 4574: 4570: 4566: 4562: 4555: 4547: 4543: 4538: 4533: 4528: 4523: 4519: 4515: 4511: 4507: 4503: 4496: 4488: 4484: 4480: 4476: 4473:(4): 689–97. 4472: 4468: 4464: 4457: 4448: 4444: 4441:(5): 637–50. 4440: 4436: 4432: 4425: 4417: 4413: 4409: 4405: 4401: 4397: 4394:(4): 874–93. 4393: 4389: 4382: 4374: 4370: 4365: 4360: 4356: 4352: 4345: 4343: 4334: 4326: 4322: 4318: 4314: 4310: 4306: 4302: 4298: 4294: 4287: 4279: 4275: 4272:(3): 666–84. 4271: 4267: 4260: 4258: 4249: 4245: 4242:(3): 642–57. 4241: 4237: 4230: 4219: 4215: 4211: 4206: 4201: 4197: 4193: 4190:(6): 661–75. 4189: 4185: 4178: 4176: 4167: 4165: 4145: 4141: 4137: 4130: 4123: 4121: 4112: 4108: 4103: 4098: 4094: 4090: 4086: 4082: 4078: 4074: 4070: 4063: 4055: 4049: 4045: 4041: 4034: 4026: 4022: 4018: 4011: 4002: 3997: 3993: 3989: 3985: 3978: 3970: 3966: 3962: 3958: 3954: 3950: 3946: 3939: 3937: 3935: 3926: 3922: 3918: 3914: 3907: 3899: 3895: 3891: 3887: 3883: 3879: 3875: 3871: 3864: 3856: 3852: 3848: 3841: 3833: 3829: 3824: 3819: 3815: 3811: 3808:(4): 514–23. 3807: 3803: 3799: 3792: 3784: 3780: 3776: 3772: 3768: 3764: 3763: 3755: 3753: 3745:(2): 123–134. 3744: 3740: 3736: 3729: 3727: 3718: 3714: 3710: 3706: 3702: 3698: 3691: 3689: 3687: 3685: 3676: 3672: 3667: 3662: 3657: 3652: 3648: 3644: 3640: 3633: 3625: 3619: 3615: 3614: 3606: 3604: 3594: 3589: 3585: 3581: 3577: 3575: 3566: 3564: 3562: 3560: 3558: 3556: 3554: 3552: 3550: 3548: 3546: 3544: 3542: 3540: 3538: 3536: 3534: 3532: 3530: 3528: 3526: 3517: 3513: 3509: 3505: 3498: 3490: 3486: 3482: 3478: 3475:(8): 961–86. 3474: 3470: 3466: 3459: 3457: 3455: 3453: 3451: 3449: 3447: 3445: 3443: 3441: 3439: 3437: 3435: 3433: 3431: 3429: 3427: 3425: 3423: 3421: 3419: 3417: 3415: 3413: 3411: 3409: 3407: 3405: 3403: 3401: 3392: 3386: 3382: 3381: 3373: 3365: 3361: 3356: 3351: 3347: 3343: 3339: 3332: 3324: 3318: 3314: 3307: 3288: 3284: 3280: 3273: 3266: 3264: 3255: 3249: 3245: 3241: 3240: 3235: 3228: 3226: 3217: 3211: 3207: 3203: 3199: 3198: 3190: 3174: 3170: 3163: 3155: 3151: 3147: 3140: 3138: 3136: 3127: 3126: 3121: 3115: 3111: 3109: 3108: 3099: 3098: 3085: 3038: 3034: 3031: 3030: 3019: 3016: 3015: 3009: 3007: 3006: 3005:Jurassic Park 3001: 2997: 2993: 2989: 2985: 2981: 2980: 2979:Palaeoscincus 2975: 2974: 2969: 2965: 2960: 2955: 2953: 2949: 2948: 2943: 2939: 2935: 2931: 2927: 2923: 2916: 2912: 2909: 2904: 2895: 2893: 2892:Tyrannosaurus 2889: 2888: 2883: 2882: 2877: 2876: 2875:Struthiomimus 2871: 2870: 2866: 2862: 2861: 2860:Edmontosaurus 2856: 2855: 2851: 2847: 2846: 2841: 2840: 2836: 2832: 2828: 2824: 2820: 2816: 2812: 2808: 2804: 2800: 2796: 2792: 2788: 2784: 2780: 2776: 2772: 2767: 2765: 2761: 2757: 2753: 2749: 2744: 2742: 2738: 2733: 2728: 2724: 2719: 2717: 2713: 2709: 2698: 2693: 2684: 2680: 2678: 2677: 2672: 2667: 2663: 2655: 2650: 2646: 2644: 2640: 2635: 2630: 2628: 2625:as in modern 2624: 2620: 2616: 2612: 2603: 2594: 2591: 2587: 2583: 2579: 2569: 2567: 2557: 2548: 2546: 2543:suggest some 2542: 2538: 2534: 2530: 2526: 2521: 2516: 2514: 2510: 2506: 2502: 2498: 2494: 2489: 2487: 2483: 2479: 2475: 2471: 2467: 2458: 2449: 2447: 2443: 2439: 2435: 2431: 2427: 2423: 2419: 2415: 2411: 2406: 2404: 2400: 2396: 2392: 2388: 2384: 2379: 2371: 2366: 2362: 2360: 2356: 2352: 2328: 2324: 2319: 2317: 2313: 2309: 2305: 2301: 2297: 2293: 2289: 2285: 2281: 2277: 2273: 2269: 2265: 2257: 2252: 2238: 2236: 2232: 2228: 2224: 2220: 2216: 2212: 2208: 2204: 2200: 2196: 2195:Ankylosaurini 2192: 2188: 2178: 2177: 2169: 2168: 2160: 2159: 2151: 2150: 2142: 2141: 2133: 2132: 2124: 2123: 2115: 2114: 2111: 2110: 2109: 2102: 2101: 2098: 2097: 2094: 2093: 2092: 2085: 2084: 2078: 2077: 2074: 2073: 2070: 2069: 2068: 2061: 2060: 2057: 2056: 2048: 2047: 2044: 2043: 2042: 2035: 2034: 2031: 2030: 2027: 2026: 2025: 2018: 2017: 2011: 2010: 2004: 2003: 2000: 1999: 1991: 1990: 1987: 1986: 1985: 1978: 1977: 1974: 1973: 1970: 1969: 1968: 1961: 1960: 1954: 1953: 1947: 1946: 1943: 1942: 1939: 1938: 1937: 1930: 1929: 1923: 1922:Ankylosaurini 1920: 1919: 1916: 1915: 1907: 1906: 1898: 1897: 1894: 1893: 1892: 1885: 1884: 1881: 1880: 1877: 1876: 1875: 1868: 1867: 1861: 1860: 1857: 1856: 1853: 1852: 1851: 1844: 1843: 1837: 1836: 1830: 1829: 1826: 1825: 1822: 1821: 1820: 1813: 1812: 1809: 1808: 1805: 1804: 1803: 1796: 1795: 1792: 1791: 1788: 1787: 1786: 1779: 1778: 1772: 1771: 1768: 1767: 1764: 1763: 1762: 1761:Crichtonpelta 1755: 1754: 1748: 1745: 1744: 1740: 1738: 1734: 1730: 1726: 1722: 1718: 1714: 1710: 1709:Polacanthidae 1706: 1705:Maastrichtian 1702: 1698: 1689: 1682: 1681: 1676: 1672: 1669: 1668:ankylosaurins 1664: 1655: 1651: 1642: 1633: 1624: 1622: 1618: 1615: 1611: 1607: 1603: 1599: 1598:Palaeoscincus 1595: 1591: 1587: 1577: 1575: 1571: 1567: 1563: 1559: 1555: 1551: 1547: 1486: 1479: 1476:(AMNH 5214), 1475: 1470: 1466: 1464: 1460: 1456: 1452: 1448: 1444: 1443: 1438: 1434: 1430: 1424: 1422: 1418: 1414: 1413: 1407: 1402: 1395: 1385: 1381: 1377: 1376: 1365: 1356: 1347: 1345: 1341: 1336: 1332: 1328: 1313: 1305: 1300: 1291: 1289: 1284: 1276: 1269: 1265: 1260: 1257: 1253: 1248: 1243: 1241: 1223: 1205: 1201: 1197: 1183: 1179: 1175: 1171: 1164: 1160: 1151: 1149: 1145: 1141: 1137: 1133: 1129: 1124: 1120: 1115: 1111: 1107: 1103: 1099: 1094: 1092: 1088: 1084: 1080: 1076: 1071: 1066: 1057: 1053: 1050: 1045: 1041: 1036: 1029: 1025: 1021: 1017: 1016: 1011: 1007: 1003: 999: 995: 991: 987: 983: 974: 965: 962: 958: 925: 867: 866:ankylosaurine 863: 855: 846: 843: 838: 834: 829: 827: 823: 819: 814: 810: 806: 801: 799: 795: 781: 778: 774: 770: 766: 757: 753: 752:at the AMNH. 751: 747: 746: 741: 737: 733: 729: 728:Tyrannosaurus 725: 721: 720:Tyrannosaurus 717: 713: 709: 708: 707:Tyrannosaurus 702: 700: 696: 695: 690: 689: 684: 680: 676: 672: 668: 667: 662: 661: 653: 652: 646: 642: 640: 636: 632: 628: 624: 623:magniventris, 620: 616: 614: 603: 599: 597: 586: 582: 578: 574: 570: 566: 562: 561:type specimen 558: 555: 551: 543: 538: 529: 527: 526: 525:Edmontosaurus 521: 520: 515: 514: 513:Tyrannosaurus 509: 505: 501: 497: 493: 489: 485: 481: 477: 473: 469: 465: 461: 457: 453: 452: 447: 446: 441: 437: 435: 431: 427: 423: 418: 416: 412: 408: 404: 400: 399: 394: 390: 386: 382: 379: 375: 371: 367: 363: 359: 358: 345: 343: 334: 331: 330:Binomial name 327: 323: 322: 316: 313: 312: 307: 302: 301: 294: 291: 290: 287: 286:Ankylosaurini 281: 278: 277: 274: 268: 265: 264: 261: 255: 252: 251: 248: 242: 239: 236: 235: 232: 226: 223: 220: 219: 216: 210: 207: 204: 203: 200: 197: 194: 191: 190: 187: 184: 181: 180: 177: 174: 171: 170: 167: 164: 161: 160: 155: 150: 146: 143: 139: 134: 130: 125: 111: 106: 101: 96: 91: 86: 81: 76: 71: 66: 61: 56: 50: 43: 42:Maastrichtian 39: 33: 29: 26: 22: 6087:Ankylosaurus 6057:Ankylosaurus 6056: 5971: 5964: 5957: 5950: 5943: 5936: 5929: 5922: 5917:Dyoplosaurus 5915: 5908: 5903:Ankylosaurus 5902: 5901: 5894: 5874: 5867: 5859: 5852: 5844: 5837: 5832:Pinacosaurus 5830: 5823: 5816: 5809: 5802: 5795: 5788: 5763: 5756: 5749: 5729: 5722: 5716:Hylaeosaurus 5714: 5707: 5700: 5692: 5684: 5652: 5645: 5640:Pawpawsaurus 5638: 5631: 5624: 5604: 5597: 5590: 5585:Denversaurus 5583: 5576: 5556: 5549: 5542: 5536:Peloroplites 5534: 5527: 5520: 5513: 5508:Borealopelta 5506: 5498: 5493:Acantholipan 5491: 5480:Nodosaurinae 5466: 5459: 5453:Peloroplites 5451: 5444: 5436: 5428: 5408: 5401: 5394: 5386: 5378: 5372:Mymoorapelta 5370: 5363: 5357:Hylaeosaurus 5355: 5348: 5341: 5333: 5325: 5318: 5313:Anoplosaurus 5311: 5304: 5293:Nodosauridae 5265: 5257: 5250: 5230: 5223: 5215: 5208: 5201: 5193: 5186: 5179: 5172: 5165: 5160:Cryptosaurus 5158: 5142:Ankylosauria 5120:Ankylosauria 5118: 5090:Ankylosauria 5078:Ornithischia 5063:Ornithischia 5015: 5011:Ornithischia 5006: 4997: 4969:Ankylosauria 4886:. Retrieved 4882: 4876: 4868: 4856:. Retrieved 4852: 4846: 4838: 4826:. Retrieved 4821: 4816: 4809: 4797:. Retrieved 4787:Bigelow, P. 4782: 4763: 4757: 4738: 4732: 4713: 4707: 4688: 4682: 4663: 4634: 4628: 4619: 4613: 4568: 4564: 4554: 4512:(8): e6738. 4509: 4505: 4495: 4470: 4466: 4462: 4456: 4438: 4434: 4430: 4424: 4391: 4387: 4381: 4354: 4350: 4342:Pinacosaurus 4341: 4333: 4300: 4296: 4292: 4286: 4269: 4265: 4239: 4235: 4229: 4218:the original 4187: 4183: 4174: 4151:. Retrieved 4144:the original 4139: 4135: 4079:(1): 18242. 4076: 4072: 4062: 4043: 4039: 4033: 4016: 4010: 3991: 3987: 3977: 3952: 3948: 3916: 3912: 3906: 3873: 3869: 3863: 3849:(95): 1–10. 3846: 3840: 3805: 3801: 3791: 3766: 3760: 3742: 3738: 3700: 3696: 3646: 3642: 3632: 3612: 3583: 3579: 3573: 3507: 3503: 3497: 3472: 3468: 3464: 3379: 3372: 3345: 3341: 3331: 3312: 3306: 3294:. Retrieved 3287:the original 3282: 3278: 3238: 3196: 3189: 3179:September 3, 3177:. Retrieved 3173:the original 3162: 3145: 3123: 3114: 3106: 3105: 3037: 3028: 3027: 3003: 3000:Ankylosaurus 2999: 2992:Ankylosaurus 2991: 2987: 2984:Ankylosaurus 2983: 2977: 2971: 2968:Ankylosaurus 2967: 2964:Ankylosaurus 2963: 2959:Ankylosaurus 2958: 2956: 2952:Ankylosaurus 2951: 2945: 2938:Ankylosaurus 2937: 2922:Ankylosaurus 2921: 2919: 2911:Ankylosaurus 2910: 2891: 2885: 2881:Ornithomimus 2879: 2873: 2867: 2858: 2852: 2843: 2837: 2831:Ankylosaurus 2830: 2825:, ferns and 2815:forest fires 2805:, which was 2795:Ankylosaurus 2794: 2771:Ankylosaurus 2770: 2768: 2760:hadrosaurids 2756:Ankylosaurus 2755: 2752:Ankylosaurus 2751: 2748:Ankylosaurus 2747: 2745: 2737:Ankylosaurus 2736: 2731: 2723:Ankylosaurus 2722: 2720: 2708:Ankylosaurus 2707: 2706: 2697:Ankylosaurus 2696: 2681: 2674: 2659: 2654:Ankylosaurus 2653: 2634:Ankylosaurus 2633: 2631: 2627:crocodilians 2608: 2590:Pinacosaurus 2589: 2581: 2578:Ankylosaurus 2577: 2575: 2562: 2541:Ankylosaurus 2540: 2533:Ankylosaurus 2532: 2520:Ankylosaurus 2519: 2517: 2492: 2490: 2463: 2442:Ankylosaurus 2441: 2430:Ankylosaurus 2429: 2426:Ankylosaurus 2425: 2422:blind snakes 2418:worm lizards 2410:Ankylosaurus 2409: 2407: 2402: 2399:Ankylosaurus 2398: 2382: 2375: 2370:ankylosaurid 2351:Ankylosaurus 2350: 2327:Ankylosaurus 2326: 2323:Ankylosaurus 2322: 2320: 2315: 2312:Ankylosaurus 2311: 2304:Ankylosaurus 2303: 2288:Ankylosaurus 2287: 2284:woody plants 2275: 2264:Ankylosaurus 2263: 2261: 2241:Paleobiology 2230: 2226: 2223:Ankylosaurus 2222: 2187:Ankylosaurus 2186: 2184: 2106: 2105: 2089: 2088: 2065: 2064: 2039: 2038: 2024:Ankylosaurus 2023: 2022: 2021: 1982: 1981: 1965: 1964: 1936:Dyoplosaurus 1934: 1933: 1889: 1888: 1872: 1871: 1848: 1847: 1819:Pinacosaurus 1817: 1816: 1800: 1799: 1783: 1782: 1759: 1758: 1728: 1724: 1721:Ankylosaurus 1720: 1713:Ankylosaurus 1712: 1694: 1678: 1674: 1671:Ankylosaurus 1670: 1621:Ankylosauria 1610:Ankylosaurus 1609: 1605: 1601: 1597: 1586:Ankylosaurus 1585: 1583: 1574:Ankylosaurus 1573: 1570:Ankylosaurus 1569: 1566:zygapophyses 1562:Dyoplosaurus 1561: 1557: 1553: 1549: 1546:Ankylosaurus 1545: 1485:Ankylosaurus 1484: 1482: 1463:Pinacosaurus 1462: 1459:Ankylosaurus 1458: 1455:Ankylosaurus 1454: 1451:Ankylosaurus 1450: 1446: 1440: 1437:Ankylosaurus 1436: 1432: 1429:Ankylosaurus 1428: 1425: 1421:Ankylosaurus 1420: 1416: 1410: 1394:Ankylosaurus 1393: 1391: 1380:Ankylosaurus 1379: 1373: 1344:Ankylosaurus 1343: 1339: 1335:Ankylosaurus 1334: 1330: 1327:Ankylosaurus 1326: 1312:Ankylosaurus 1311: 1309: 1282:parapophyses 1244: 1240:Ankylosaurus 1239: 1170:Ankylosaurus 1169: 1167: 1140:Ankylosaurus 1139: 1136:tooth crowns 1132:Ankylosaurus 1131: 1128:Ankylosaurus 1127: 1123:Ankylosaurus 1122: 1114:Ankylosaurus 1113: 1095: 1083:Ankylosaurus 1082: 1065:Ankylosaurus 1064: 1062: 1049:Ankylosaurus 1048: 1019: 1015:Pinacosaurus 1013: 982:Ankylosaurus 981: 979: 961:Ankylosaurus 960: 957:Ankylosaurus 956: 926: 862:Ankylosaurus 861: 860: 842:Ankylosaurus 841: 833:Ankylosaurus 832: 830: 825: 813:Ankylosaurus 812: 808: 802: 798:Powder River 794:Ankylosaurus 793: 765:Ankylosaurus 764: 762: 750:Ankylosaurus 749: 743: 740:Ankylosaurus 739: 736:Dynamosaurus 735: 727: 719: 716:Ankylosaurus 715: 705: 703: 699:Franz Nopcsa 692: 686: 683:Ankylosaurus 682: 675:Ankylosaurus 674: 664: 658: 656: 649: 622: 619:type species 610: 593: 564: 557:Barnum Brown 548:In 1906, an 547: 523: 517: 511: 508:Ankylosaurus 507: 504:Ankylosaurus 503: 472:Ankylosaurus 471: 456:Ankylosaurus 455: 449: 443: 440:Ankylosaurus 439: 438: 426:Ankylosaurus 425: 422:ankylosaurid 419: 411:Ankylosaurus 410: 407:Ankylosauria 403:generic name 397: 396: 389:Barnum Brown 357:Ankylosaurus 356: 355: 354: 341: 336: 320: 319: 300:Ankylosaurus 299: 298: 247:Ankylosauria 237: 221: 215:Ornithischia 205: 192: 138:Ankylosaurus 137: 32:Ankylosaurus 31: 25: 21:Ankylosuchus 6196:Lance fauna 6081:Wikispecies 5952:Scolosaurus 5869:Tsagantegia 5818:Jinyunpelta 5758:Shamosaurus 5709:Chuanqilong 5551:Silvisaurus 5529:Patagopelta 5461:Polacanthus 5410:Sauroplites 5210:Spicomellus 5188:Sarcolestes 5020:Thyreophora 3703:(5): 1–60. 3041:Pronounced 2998:from 1930. 2839:Triceratops 2835:ceratopsids 2819:angiosperms 2799:subtropical 2721:Fossils of 2586:ontogenetic 2321:Fossils of 2300:endothermic 2268:herbivorous 2219:diversified 2091:Scolosaurus 1785:Tsagantegia 1697:Thyreophora 1687:caputegulae 1617:Stegosauria 1554:Scolosaurus 1472:Only known 1442:Scolosaurus 1274:diapophyses 1178:quadrupedal 1034:caputegulae 998:premaxillae 849:Description 694:Polacanthus 660:Stegosaurus 651:Stegosaurus 519:Triceratops 430:quadrupedal 348:Brown, 1908 266:Subfamily: 231:Thyreophora 47:68–66 6170:Categories 6020:See also: 5945:Platypelta 5938:Oohkotokia 5876:Zaraapelta 5751:Gobisaurus 5702:Cedarpelta 5694:Aletopelta 5647:Stegopelta 5592:Edmontonia 5578:Animantarx 5544:Sauropelta 5522:Nodosaurus 5388:Priconodon 5232:Vectipelta 5167:Dracopelta 5099:see below↓ 5002:Dinosauria 4888:August 29, 4858:August 29, 4849:dinosaurs" 4828:August 29, 3204:. p. 3024:References 2988:Edmontonia 2973:Edmontonia 2926:archetypal 2887:Pectinodon 2845:Torosaurus 2791:floodplain 2732:Edmontonia 2662:cancellous 2446:bite force 2434:omnivorous 2416:(digging) 2378:Georg Haas 2207:Cretaceous 2203:Cenomanian 1891:Zaraapelta 1701:Sinemurian 1590:type genus 1384:osteoderms 1340:Edmontonia 1144:diagnostic 1142:teeth are 1119:predentary 1040:remodeling 732:synonymous 688:Stegopelta 577:osteoderms 496:nodosaurid 476:Hell Creek 415:archetypal 378:Cretaceous 199:Dinosauria 5959:Talarurus 5839:Saichania 5626:Europelta 5606:Texasetes 5468:Taohelong 5365:Invictarx 5267:Stegouros 4985:Kingdom: 4920:Dinosaurs 4799:March 24, 4373:129291148 4325:130610291 4025:2246/2609 3855:2246/3267 3717:214625754 3643:PLOS Biol 3516:2246/1464 3154:2246/1435 3107:Citations 2807:monsoonal 2783:sandstone 2727:sediments 2643:theropods 2641:of large 2529:proboscis 2513:olfaction 2414:fossorial 2211:Campanian 1967:Talarurus 1850:Saichania 1733:cladogram 1717:subfamily 1592:of a new 1474:tail club 1447:Saichania 1259:vertebrae 1252:ligaments 1148:denticles 1091:nodosaurs 1010:pyramidal 1006:squamosal 986:taphonomy 671:tail club 666:Glyptodon 602:ankylosis 581:described 540:Skull of 488:Frenchman 393:monotypic 385:dinosaurs 314:Species: 172:Kingdom: 166:Eukaryota 6066:Wikidata 6032:Category 6022:Timeline 5966:Ziapelta 5438:Gastonia 5335:Gastonia 4993:Chordata 4991:Phylum: 4987:Animalia 4822:Wired UK 4793:Archived 4605:34824329 4546:19707581 4506:PLOS ONE 4416:86333501 4214:21954840 4153:July 25, 4111:37880323 4102:10600113 3969:55372674 3898:86002282 3832:26332595 3783:84505681 3675:24802911 3093:-kə-loh- 3012:See also 2823:conifers 2787:mudstone 2766:" role. 2611:collagen 2478:tetrapod 2395:lineages 2308:elephant 2215:Turonian 2185:Because 2108:Ziapelta 1614:suborder 1431:, as in 1264:ossified 1200:entheses 1196:coracoid 1110:mandible 542:holotype 484:Scollard 253:Family: 186:Chordata 182:Phylum: 176:Animalia 162:Domain: 136:Cast of 6140:4946546 6127:1003608 6114:4823327 6101:4530828 5854:Tarchia 5846:Shanxia 4906:Portals 4596:8616956 4573:Bibcode 4537:2726940 4514:Bibcode 4475:Bibcode 4447:1303019 4408:4524782 4305:Bibcode 4278:1304004 4248:1303969 4205:3237876 4081:Bibcode 3878:Bibcode 3823:4580109 3666:4011683 3477:Bibcode 3364:2455817 3296:July 8, 3244:141–143 2676:Tarchia 2597:Defense 2344:⁄ 2334:⁄ 2280:fibrous 2246:Feeding 1874:Tarchia 1539:⁄ 1525:⁄ 1511:⁄ 1497:⁄ 1320:⁄ 1288:ribcage 1268:tendons 1233:⁄ 1215:⁄ 1204:humerus 1189:⁄ 1182:scapula 1102:foramen 1087:choanae 1079:maxilla 1070:sinuses 1044:sutures 996:on the 950:⁄ 936:⁄ 919:⁄ 905:⁄ 891:⁄ 877:⁄ 787:⁄ 771:by the 596:ankulos 590:αγκυλος 573:Montana 464:Sinuses 460:browser 370:fossils 339:† 318:† 292:Genus: 279:Tribe: 116:↓ 6072:Q40621 4883:Looper 4824:. 2015 4770: 4745: 4720: 4695: 4670: 4641: 4603: 4593: 4544: 4534: 4445: 4414: 4406: 4371: 4323: 4276: 4246: 4212: 4202: 4109: 4099: 4050: 3967: 3896: 3830: 3820: 3781: 3715: 3673: 3663: 3620: 3580:FACETS 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