Editing Ankylosaurus

{{short description|Ankylosaurid dinosaur genus from the Late Cretaceous Period}}
{{Distinguish|Ankylosuchus}}
{{featured article}}
{{Use American English|date=January 2023}}
{{Use mdy dates|date=January 2024}}
{{speciesbox
| fossil_range = [[Late Cretaceous]] ([[Maastrichtian]]), {{fossilrange|68|66}}
| image = Ankylosaur head - cast - Custer County Montana - Museum of the Rockies - 2013-07-08.jpg
| image_upright = 1.1
| image_alt = Brown skull cast on a pedestal
| image_caption = Cast of ''Ankylosaurus'' skull (AMNH 5214) in front view, [[Museum of the Rockies]]
| genus = Ankylosaurus
| parent_authority = [[Barnum Brown|Brown]], 1908
| species = magniventris
| authority = Brown, 1908
}}

'''''Ankylosaurus'''''{{refn|Pronounced {{IPAc-en|ˌ|æ|ŋ|k|ə|l|oʊ|ˈ|s|ɔːr|ə|s}} {{respell|ANG|kə|loh|SOR|əs}}<ref>{{MerriamWebsterDictionary|Ankylosaurus}}</ref>|group=nb}} is a [[genus]] of [[Thyreophora|armored dinosaur]]. Its [[fossil]]s have been found in [[geological formation]]s dating to the very end of the [[Cretaceous]] [[Period (geology)|Period]], about 68–66 million years ago, in western North America, making it among the last of the non-avian [[dinosaur]]s. It was named by [[Barnum Brown]] in 1908; it is [[Monotypic taxon|monotypic]], containing only '''''A. magniventris'''''. The [[Binomial nomenclature|generic name]] 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 [[Ankylosauria]] are represented by more extensive fossil material, ''Ankylosaurus'' is often considered the [[archetype|archetypal]] member of its group, despite having some unusual features.

Possibly the largest known [[Ankylosauridae|ankylosaurid]], ''Ankylosaurus'' is estimated to have been between {{cvt|6|and|8|m|ft}} long and to have weighed between {{cvt|4.8|and|8|MT|ST}}. It was [[Quadrupedalism|quadrupedal]], 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 [[Club (anatomy)|club]] 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.

''Ankylosaurus'' is a member of the family Ankylosauridae, and its closest relatives appear to be ''[[Anodontosaurus]]'' and ''[[Euoplocephalus]]''. ''Ankylosaurus'' 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 [[Browsing (herbivory)|browser]]. [[Paranasal sinuses|Sinuses]] 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 [[Intraspecific competition|intraspecific combat]]. Specimens of ''Ankylosaurus'' have been found in the [[Hell Creek Formation|Hell Creek]], [[Lance Formation|Lance]], [[Scollard Formation|Scollard]], [[Frenchman Formation|Frenchman]], and [[Ferris Formation|Ferris]] formations, but it appears to have been rare in its environment. Although it lived alongside ''[[Denversaurus]]'', a [[Nodosauridae|nodosaurid]] ankylosaur, their ranges and [[ecological niche]]s do not appear to have overlapped, and ''Ankylosaurus'' may have inhabited upland areas. ''Ankylosaurus'' also lived alongside dinosaurs such as ''[[Tyrannosaurus]]'', ''[[Triceratops]]'', and ''[[Edmontosaurus]]''.

==History of discovery==
[[File:Skull of Ankylosaurus.jpg|thumb|left|alt=Photograph of dorsal view of fossilized skull next to sketch of the same|Skull of [[holotype]] AMNH 5895 and reconstruction diagram of same]]
In 1906, an [[American Museum of Natural History]] expedition led by American [[Paleontology|paleontologist]] [[Barnum Brown]] discovered the [[Type (biology)#Type specimen|type specimen]] of ''Ankylosaurus magniventris'' (AMNH 5895) in the [[Hell Creek Formation]], near Gilbert Creek, [[Montana]]. 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 [[osteoderms]] (armor plates). Brown scientifically [[species description|described]] the animal in 1908; the generic name is derived from the [[Ancient Greek|Greek]] words {{lang|grc|αγκυλος}} ''{{lang|grc-Latn|ankulos}}'' ('bent' or 'crooked'), referring to the medical term [[ankylosis]], the stiffness produced by the fusion of bones in the skull and body, and {{lang|grc|σαυρος}} ''{{lang|grc-Latn|sauros}}'' ('lizard'). The name can be translated as "fused lizard", "stiff lizard", or "curved lizard". The [[type species]] name, ''magniventris,'' is derived from the {{langx|la|magnus}} ('great') and {{langx|la|venter}} ('belly'), referring to the great width of the animal's body.<ref name="brown1908"/><ref name=pronunciation/><ref name=Oxford/>
[[File:Ankylosaurus.jpg|thumb|alt=Images of skeleton: side view facing left, dorsal view, and dorsal view of dorsal plates|1908 skeletal reconstruction of the holotype, with missing parts restored after ''[[Stegosaurus]]'']]
The skeletal reconstruction accompanying the 1908 description restored the missing parts in a fashion similar to ''[[Stegosaurus]]'', and Brown likened the result to the extinct armored mammal ''[[Glyptodon]]''.<ref name="brown1908"/> 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 [[Club (anatomy)|tail club]], 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.<ref name="glut1997"/><ref name="Coombs1978"/><ref>{{cite book
| last1 = Naish
| first1 = D.
|year= 2009
| pages =58–59
|title= The Great Dinosaur Discoveries
|publisher=A & C Black Publishers LTD
|isbn=978-1408119068
|location=London}}</ref> In a 1908 review of Brown's ''Ankylosaurus'' description, the American paleontologist [[Samuel Wendell Williston]] criticized the skeletal reconstruction as being based on too few remains, and claimed that ''Ankylosaurus'' was merely a synonym of the genus ''[[Stegopelta]]'', which Williston had named in 1905. Williston also stated that a skeletal reconstruction of the related ''[[Polacanthus]]'' by Hungarian paleontologist [[Franz Nopcsa von Felső-Szilvás|Franz Nopcsa]] was a better example of how ankylosaurs would have appeared in life.<ref name="Williston1908"/> The claim of synonymy was not accepted by other researchers, and the two genera are now considered distinct.<ref name="carpenter2001"/>

Brown had collected 77 osteoderms while excavating a ''[[Tyrannosaurus]]'' specimen in the [[Lance Formation]] of Wyoming in 1900. He mentioned these osteoderms (specimen AMNH 5866) in his description of ''Ankylosaurus'' but thought they belonged to the ''Tyrannosaurus'' instead. Paleontologist [[Henry Fairfield Osborn]] also expressed this view when he described the ''Tyrannosaurus'' specimen as the now [[junior synonym|synonymous]] genus ''Dynamosaurus'' in 1905. More recent examination has shown them to be similar to those of ''Ankylosaurus''; it seems that Brown had compared them with some ''[[Euoplocephalus]]'' osteoderms, which had been erroneously cataloged as belonging to ''Ankylosaurus'' at the AMNH.<ref name="carpenter2004"/><ref name="osborn1905"/>

[[File:Ankylosaurus excavation.jpg|thumb|left|alt=Photograph in black and white of rugged, fissured cliff face|Excavation of AMNH 5214 (center, above the pick), 1910]]
In 1910, another AMNH expedition led by Brown discovered an ''Ankylosaurus'' specimen (AMNH 5214) in the [[Scollard Formation]] by the [[Red Deer River]] 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 [[Fossil collecting|fossil collectors]] [[Charles Mortram Sternberg|Charles M. Sternberg]] and T. Potter Chamney collected a skull and mandible (specimen CMN 8880, formerly NMC 8880), {{convert|1|km|mi|frac=8|sp=us|abbr=off}} north of where the 1910 specimen was found. This is the largest-known ''Ankylosaurus'' skull, but it is damaged in places. A section of caudal vertebrae (specimen CCM V03) was discovered in the 1960s in the [[Powder River (Wyoming and Montana)|Powder River]] drainage, Montana, part of the Hell Creek Formation. In addition to these five incomplete specimens, many other isolated osteoderms and teeth have been found.<ref name="2017desc"/><ref name="carpenter2004"/>

In 1990, American paleontologist [[Walter P. Coombs]] pointed out that the teeth of two skulls assigned to ''A. magniventris'' differed from those of the holotype specimen in some details, and though he expressed a "considerate temptation" to name a new species of ''Ankylosaurus'' 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 [[Matrix (geology)|matrix]] within the nasal chambers.<ref name="Coombs teeth"/> The American paleontologist [[Kenneth Carpenter]] accepted the teeth as belonging to ''A. magniventris'' in 2004, and that all the specimens belonged to the same species, noting that the teeth of other ankylosaurs are highly variable.<ref name="carpenter2004"/>

Most of the known ''Ankylosaurus'' specimens were not scientifically described at length, though several paleontologists planned to do so until Carpenter redescribed the genus in 2004.<ref name="carpenter2004"/> In 2017 the Canadian paleontologists [[Victoria M. Arbour]] 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 ''Ankylosaurus'' 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.<ref name="2017desc"/>

==Description==
[[File:Ankylosaurus Scale V2.svg|thumb|right|alt=Outline of human superimposed on outline of ''Ankylosaurus''|Size of the smallest (AMNH 5214) and largest (CMN 8880) known specimens, compared to a human]]
''Ankylosaurus'' was the largest-known [[Ankylosauridae|ankylosaurine]] dinosaur and possibly the largest ankylosaurid.<ref name="2017desc"/> In 2004 Carpenter estimated that the individual with the largest-known skull (specimen CMN 8880), which is {{cvt|64.5|cm|ftin}} long and {{cvt|74.5|cm|ftin}} wide, was about {{cvt|6.25|m|ftin}} long and had a hip height of about {{cvt|1.7|m|ftin}}. The smallest-known skull (specimen AMNH 5214) is {{cvt|55.5|cm|ftin}} long and {{cvt|64.5|cm|ftin}} wide, and Carpenter estimated that it measured about {{cvt|5.4|m|ftin}} long and about {{cvt|1.4|m|ftin}} tall at the hips.<ref name="carpenter2004"/> The English paleontologist Roger B. J. Benson and colleagues estimated the weight for AMNH 5214 at {{cvt|4.78|MT|ST}} in 2014.<ref name="Bensonetal2014">{{cite journal | last1 = Benson | first1 = R. B. J. | last2 = Campione | first2 = N. E. | last3 = Carrano | first3 = M. T. | last4 = Mannion | first4 = P. D. | last5 = Sullivan | first5 = C. | year = 2014 | title = Rates of Dinosaur Body Mass Evolution Indicate 170 Million Years of Sustained Ecological Innovation on the Avian Stem Lineage | journal = PLOS Biol | volume = 12 | issue = 5| page = e1001853 | doi = 10.1371/journal.pbio.1001853 | last6 = Upchurch |first6 = P. | last7 = Evans | first7 = D. C. | display-authors= 5 | pmid=24802911 | pmc=4011683 | doi-access = free }}</ref>

In 2017, based on comparisons with more complete ankylosaurines, Arbour and Mallon estimated a length of {{convert|7.56|to|9.99|m|ftin|frac=2|abbr=on}} for CMN 8880, and {{convert|6.02|to|7.95|m|ftin|frac=2|abbr=on}} for AMNH 5214. Though the latter is the smallest specimen of ''Ankylosaurus'', its skull is still larger than those of any other ankylosaurins. A few other ankylosaurs reached about {{convert|6|m|ft|0|abbr=on}} 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 {{convert|10|m|ft|sp=us}} for large ''Ankylosaurus'' too long, and suggested a length of {{convert|8|m|ft|abbr=on}} instead. Arbour and Mallon estimated a weight of {{convert|4.78|MT|ST|abbr=on}} for AMNH 5214, and tentatively estimated the weight of CMN 8880 at {{convert|7.95|MT|ST|abbr=on}}.<ref name="2017desc">{{cite journal |first1=V.M. |last1=Arbour |first2=J.C. |last2=Mallon |year=2017 |title=Unusual cranial and postcranial anatomy in the archetypal ankylosaur ''Ankylosaurus magniventris'' |journal=FACETS |volume=2 |issue=2 |pages=764–794 |doi=10.1139/facets-2017-0063 |doi-access=free |bibcode=2017Facet...2..764A }}</ref>

===Skull===
[[File:Ankylosaurus skull AMNH.jpg|thumb|upright|alt=Two views of ''Ankylosaurus'' skull, from above and from the left|Skull (AMNH 5214) from the side and above]]
The three known ''Ankylosaurus'' skulls differ in various details; this is thought to be the result of [[taphonomy]] (changes happening during decay and fossilization of the remains) and [[Polymorphism (biology)|individual variation]]. 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 [[beak]] on the [[premaxilla]]e. The [[orbit (anatomy)|orbit]]s (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 [[Squamosal bone|squamosal]] horns (their shape has been described as "[[pyramid]]al"), 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 ''[[Pinacosaurus]]'' and ''Euoplocephalus''. Below the upper horns, [[Jugal bone|jugal]] horns were present, which pointed backward and down. The horns may have originally been osteoderms that fused to the skull. The [[Scale (anatomy)|scale]]-like cranial ornamentation on the surfaces of ankylosaurs skulls is called "{{Dinogloss|caputegulae}}", and were the result of [[Bone remodeling|remodeling]] of the skull itself. This obliterated the [[Suture (anatomy)|sutures]] 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 ''Ankylosaurus'' 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.<ref name="carpenter2004"/><ref name="2017desc"/><ref name="systematics ankylosaurid"/>

[[File:Ankylosaurus tooth.jpg|thumb|left|upright|Tooth of the holotype in inner and outer view]]
The snout region of ''Ankylosaurus'' 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 or upward. Additionally, the nostrils were not visible from the front because the [[sinuses]] 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 [[Nasal septum|septum]], which separated the nasal passage from the sinus. Each side of the snout had five sinuses, four of which expanded into the [[maxilla]] bone. The nasal cavities (or chambers) of ''Ankylosaurus'' 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 [[Posterior nasal apertures|choanae]] (internal nostrils), and the air passage was looped.<ref name="carpenter2004"/><ref name="2017desc"/> 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 [[nodosaurs]] had armor plates that covered the cheek region, which may have been embedded in the flesh.<ref name="carpenter2004"/>

Specimen AMNH 5214 has 34–35 [[Dental alveolus|dental alveoli]] (tooth sockets) in the maxilla. The tooth rows in the maxillae of this specimen are about {{convert|20|cm|in|sp=us}} long. Each alveolus had a [[foramen]] (opening) near its side where a [[Polyphyodont|replacement tooth]] could be seen. Compared to other ankylosaurs, the [[mandible]] of ''Ankylosaurus'' 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 {{convert|41|cm|in|frac=2|sp=us}} long. The incomplete mandible of the largest specimen (CMN 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 [[predentary]] bone of the tip of the mandibles has not yet been found.<ref name="carpenter2004"/> Like other ankylosaurs, ''Ankylosaurus'' had small, phylliform (leaf-shaped) teeth, which were compressed sideways.<ref name="Coombs teeth"/> 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 ''Ankylosaurus'' could accommodate more teeth than other ankylosaurines. The teeth of the largest ''Ankylosaurus'' skull are smaller than those of the smallest skull in the absolute sense. Some teeth from behind in the tooth row curved backwards, and [[Crown (tooth)|tooth crowns]] were usually flatter on one side than the other.<ref name="carpenter2004"/> ''Ankylosaurus'' teeth are [[Taxonomy (biology)#Taxonomic descriptions|diagnostic]] and can be distinguished from the teeth of other ankylosaurids based on their smooth sides. The [[Denticle (tooth feature)|denticles]] were large, their number ranging from six to eight on the front part of the tooth, and five to seven behind.<ref name="carpenter2004"/><ref name="carpenter1982b"/>

===Postcranial skeleton===

[[File:Ankylosaurus magniventris by sphenaphinae.png|thumb|alt=Image of restored specimen of a squat quadruped with knobby back|[[paleoart|Life restoration]]]]
The structure of much of the skeleton of ''Ankylosaurus'', including most of the [[pelvis]], tail, and feet, is still unknown.<ref name="carpenter2004"/> It was [[quadruped]]al, and its hind limbs were longer than its forelimbs.<ref name="coombs"/> In the holotype specimen, the [[scapula]] (shoulder blade) measures {{convert|61.5|cm|ftin|frac=4|abbr=on}} long and was fused with the [[coracoid]] (a rectangular bone connected to the lower end of the scapula). It also had [[Enthesis|entheses]] (connective tissue) for various muscle attachments. The [[humerus]] (upper arm bone) of AMNH 5214 was short, very broad and about {{convert|54|cm|ftin|frac=2|abbr=on}} long. The [[femur]] (thigh bone), also from AMNH 5214, was {{convert|67|cm|ftin|frac=2|abbr=on}} long and very robust. While the feet of ''Ankylosaurus'' are incompletely known, the hindfeet probably had three toes, as is the case in advanced ankylosaurids.<ref name="carpenter2004"/>

The [[cervical vertebrae]] 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 [[ligament]]s, which helped support the massive head. The [[Anatomical terms of location|dorsal]] [[vertebra]]e 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 [[Ossification|ossified]] (turned to bone) [[tendon]]s, which also overlapped some of the vertebrae. The ribs of the last four back vertebrae were fused to the {{Dinogloss|diapophyses}} and {{Dinogloss|parapophyses}} (the structures that articulated the ribs with the vertebrae), and the [[ribcage]] 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.<ref name="carpenter2004"/>

===Armor===
[[File:Ankylosaurus armor.png|thumb|Arrangement of armor as suggested by [[Victoria M. Arbour|Arbour]] and Mallon, 2017]] 
A prominent feature of ''Ankylosaurus'' 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 proposed. The osteoderms ranged from {{convert|1|cm|in|frac=4|sp=us}} in diameter to {{convert|35.5|cm|in|frac=4|abbr=on}} in length, and varied in shape. The osteoderms of ''Ankylosaurus'' were generally thin walled and hollowed on the underside. Compared to ''Euoplocephalus'', the osteoderms of ''Ankylosaurus'' 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 ''Edmontonia'' had high keels stretching from one margin to the other on the midline of its osteoderms. ''Ankylosaurus'' had some smaller osteoderms with a keel across the midline.<ref name="2017desc"/><ref name="carpenter2004"/> 

{{multiple image
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|image1 = Ankylosaurus cervical half rings.gif
|alt1 = Armor

|image2 = Newly identified elements the holotype of Ankylosaurus.gif
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|footer = Left: Cervical half ring from the neck of ''[[Euoplocephalus]]'' (A–B) compared with half-ring fragments (C–J) of ''Ankylosaurus'' (holotype), and possible back [[osteoderm]]s (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)
}}

Like other ankylosaurids, ''Ankylosaurus'' had {{Dinogloss|cervical half-rings}} (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 [[Philip J. Currie]] 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 cervical half-rings, which formed two semi-circular plates of armor around the upper part of the neck, as in the closely related ''[[Anodontosaurus]]'' and ''Euoplocephalus''.<ref name="carpenter2004"/><ref name="systematics ankylosaurid"/> 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 ''Ankylosaurus'' had six keeled osteoderms with oval bases on each half-ring.<ref name="2017desc"/>

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 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 ''Ankylosaurus'', as in ''Euoplocephalus''. ''Ankylosaurus'' 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 ''[[Scolosaurus]]''. 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 ''Saichania'', and may have been distributed similarly on ''Ankylosaurus''. 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 ''Ankylosaurus'' specimens may have been placed on the sides of the pelvis or the tail. Ovoid, keeled, and teardrop-shaped osteoderms are known from ''Ankylosaurus'', and may have been placed on the forelimbs, like those known from ''Pinacosaurus'', but it is unknown whether the hindlimbs bore osteoderms.<ref name="carpenter2004"/><ref name="2017desc"/> 

[[File:Ankylosaurus tail club.jpg|thumb|alt=Fossilized tail club, black in coloring|Only known [[tail club]] (AMNH 5214), [[American Museum of Natural History]]]]
The tail club (or tail knob) of ''Ankylosaurus'' 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 AMNH 5214 is {{convert|60|cm|in|frac=2|abbr=on}} long, {{convert|49|cm|in|frac=2|abbr=on}} wide, and {{convert|19|cm|in|frac=2|abbr=on}} tall. The club of the largest specimen may have been {{convert|57|cm|in|frac=2|abbr=on}} wide. The tail club of ''Ankylosaurus'' was semicircular when seen from above, similar to those of ''Euoplocephalus'' and ''Scolosaurus'' but unlike the pointed club osteoderms of ''Anodontosaurus'' or the narrow, elongated club of ''Dyoplosaurus''. 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 [[Articular processes|zygapophyses]] (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 ''Ankylosaurus'' being wider. The larger width may indicate that the tail of ''Ankylosaurus'' 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.<ref name="2017desc"/><ref name="carpenter2004"/><ref>{{Cite journal|doi=10.1111/joa.12363|pmid=26332595|title=Ankylosaurid dinosaur tail clubs evolved through stepwise acquisition of key features|journal=Journal of Anatomy|volume=227|issue=4|pages=514–23|year=2015|last1=Arbour|first1=V. M.|last2=Currie|first2=P. J.|pmc=4580109}}</ref>

==Classification==
Brown considered ''Ankylosaurus'' so distinct that he made it the [[Type (biology)|type genus]] of a new [[Family (biology)|family]], Ankylosauridae, typified by massive, triangular skulls, short necks, stiff backs, broad bodies, and osteoderms. He also classified ''Palaeoscincus'' (only known from teeth), and ''Euoplocephalus'' (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 ''Euoplocephalus'' and ''Ankylosaurus''. Having for comparison only a few, incomplete members of the family, he believed the group was part of the [[Order (biology)|suborder]] [[Stegosauria]].<ref name="brown1908"/> In 1923 Osborn coined the name [[Ankylosauria]], thereby placing the ankylosaurids in their own suborder.<ref name = "osborn1923"/>

{{multiple image
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|image1 = Ankylosaurin skulls.gif
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|image2 = Ankylosaur caputegulae.gif
|alt2 = 

|footer = Skulls of the [[ankylosaurins]] with color coded {{Dinogloss|caputegulae}} (cranial ornamentation)
}}

{{multiple image
|align = right
|total_width = 350
|image1 = Ankylosaurus neck vertebra.jpg
|alt1 = 

|image2 = Ankylosaurus tail vertebra.jpg
|alt2 = 

|footer = [[Cervical vertebra]] from the neck (left) and [[caudal vertebra]] from the tail (right) of the holotype
}}

Ankylosauria and Stegosauria are now grouped together within the clade [[Thyreophora]]. This group first appeared in the [[Sinemurian]] age, and survived for 135 million years until disappearing in the [[Maastrichtian]]. They were widespread and inhabited a broad range of environments.<ref name="carpenter2004"/><ref name="Coombs1978"/> 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 [[Polacanthinae|Polacanthidae]] (these families lacked tail clubs). ''Ankylosaurus'' is considered part of the [[subfamily]] [[Ankylosaurinae]] (members of which are called ankylosaurines) within Ankylosauridae.<ref name="thompson"/> ''Ankylosaurus'' appears to be most closely related to ''Anodontosaurus'' and ''Euoplocephalus''.<ref name="Arbour2014II"/> The following [[cladogram]] is based on a 2015 [[Phylogenetics|phylogenetic analysis]] of the Ankylosaurinae conducted by Arbour and Currie:<ref name="systematics ankylosaurid"/>

{{clade| style=font-size:85%; line-height:85%
|label1=[[Ankylosaurinae]]
|1={{clade
 |1=''[[Crichtonpelta]]''
 |2={{clade
 |1=''[[Tsagantegia]]''
 |2=''[[Zhejiangosaurus]]''
 |3=''[[Pinacosaurus]]''
 |4={{clade
 |1={{clade
 |1=''[[Saichania]]''
 |2={{clade
 |1=''[[Tarchia]]''
 |2=''[[Zaraapelta]]'' }} }}
 |label2=[[Ankylosaurini]]
 |2={{clade
 |1=''[[Dyoplosaurus]]''
 |2={{clade
 |1={{clade
 |1=''[[Talarurus]]''
 |2=''[[Nodocephalosaurus]]'' }}
 |2={{clade
 |1={{clade
 |1='''''Ankylosaurus'''''
 |2=''[[Anodontosaurus]]'' }}
 |2=''[[Euoplocephalus]]''
 |3={{clade
 |1=''[[Scolosaurus]]''
 |2=''[[Ziapelta]]'' }} }} }} }} }} }} }} }}

Because ''Ankylosaurus'' and other Late Cretaceous North American ankylosaurids were grouped with Asian genera (in a [[Tribe (biology)|tribe]] the authors named [[Ankylosaurini]]), Arbour and Currie suggested that earlier North American ankylosaurids had gone extinct by the late [[Albian]] or [[Cenomanian]] ages of the Middle [[Cretaceous]]. Ankylosaurids thereafter recolonized North America from Asia during the [[Campanian]] or [[Turonian]] ages of the Late Cretaceous, and there [[Phylogenetic diversity|diversified]] again, leading to genera such as ''Ankylosaurus'', ''Anodontosaurus'', and ''Euoplocephalus''. The theory explains a 30-million-year gap in the [[fossil]] record of North American ankylosaurids between the ages.<ref name="systematics ankylosaurid"/>

==Paleobiology==
===Feeding===

[[File:Ankylosaurus ribs.jpg|thumb|left|Back vertebra with fused ribs of the holotype. The broad body housed a large [[Digestion|digestion system]].]] 
Like other ornithischians, ''Ankylosaurus'' was [[Herbivore|herbivorous]]. Its wide muzzle was adapted for non-selective low-[[Browsing (herbivory)|browse]] cropping,<ref name="carpenter2004"/> although not to the extent seen in some related genera, especially ''Euoplocephalus''.<ref name="2017desc"/><ref name="Ősi"/> Though ankylosaurs may not have fed on [[Fiber|fibrous]] and [[woody plant]]s, they may have had a varied diet, including tough leaves and pulpy fruits.<ref name="tongue"/> ''Ankylosaurus'' probably fed on abundant [[fern]]s and low-growing [[shrub]]s. Assuming it was [[endotherm]]ic, ''Ankylosaurus'' would have eaten {{convert|60|kg|lb|abbr=off}} of ferns per day, similar to the amount of dry vegetation a large [[elephant]] would consume. The requirements for nutrition could have been more effectively met if ''Ankylosaurus'' ate fruit, which its small, cusp-like teeth and the shape of its beak seem well adapted for, compared to for example ''Euoplocephalus''. Certain invertebrates, which the small teeth may have been adapted for handling, could also have provided supplemental nutrition.<ref name="2017desc"/>

Fossils of ''Ankylosaurus'' teeth exhibit wear on the face of the crown rather than on the tip of the crown, as in nodosaurid ankylosaurs.<ref name="carpenter2004"/> In 1982 Carpenter ascribed to baby ''Ankylosaurus'' two very small teeth that originate from the Lance and Hell Creek Formations and measure {{convert|3.2|to|3.3|mm|in|frac=128|abbr=on}} 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.<ref name="carpenter1982b"/> Since adult ''Ankylosaurus'' did little chewing of its food, it would have spent less time in the day foraging than an elephant.<ref name="2017desc"/> Based on the broadness of the ribcage, the digestion of unchewed food may have been facilitated by [[hindgut fermentation]] like in modern herbivorous lizards, which have several chambers in their enlarged [[Large intestine|colon]].<ref name="carpenter2004"/>

[[File:Ankylosaurus skull CMN 8880.gif|thumb|right|Skull of specimen CMN 8880, the largest-known [[ankylosaurid]], including lower jaw (E–F) and tooth (G)]]
In 1969, paleontologist [[Georg Haas (paleontologist)|Georg Haas]] 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.<ref name="Hans1969"/> Later research on ''Euoplocephalus'' indicates that forward and sideways jaw movement was possible in these animals, the skull being able to withstand considerable forces.<ref name=Rybzynski2001/> A 2016 study of the [[Occlusion (dentistry)|dental occlusion]] (contact between the teeth) of ankylosaur specimens found that the ability for backwards (palinal) jaw movement [[convergent evolution|evolved independently]] in different ankylosaur [[Lineage (evolution)|lineages]], including Late Cretaceous North American ankylosaurids like ''Ankylosaurus'' and ''Euoplocephalus''.<ref name="Ősi">{{cite journal|last1=Ősi|first1=A.|last2=Prondvai|first2=E.|last3=Mallon|first3=J.|last4=Bodor|first4=E. R.|title=Diversity and convergences in the evolution of feeding adaptations in ankylosaurs (Dinosauria: Ornithischia)|journal=Historical Biology|volume=29|issue=4|date=2016|pages=539–570|doi=10.1080/08912963.2016.1208194|s2cid=55372674|url=https://www.researchgate.net/publication/305480013}}</ref>

The retracted position of the nostrils of ''Ankylosaurus'' were compared to those of [[fossorial]] (digging) [[Amphisbaenia|worm lizards]] and [[Scolecophidia|blind snakes]] by Arbour and Mallon in 2017, and though it was probably not a burrowing animal, the snout of ''Ankylosaurus'' may indicate earth-moving behavior. These factors, as well as the low rate of tooth formation in ankylosaurs compared to other ornithischians, indicate that ''Ankylosaurus'' may have been [[Omnivore|omnivorous]] (eating both plant and animal matter). It may also (or alternatively) have dug in the ground for roots and [[tuber]]s.<ref name="2017desc"/> A 2023 study by paleontologist Antonio Ballell and colleagues found that North American ankylosaurids from the latest Cretaceous (including ''Ankylosaurus'') 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 [[bite force]], which suggests diverging jaw mechanics and dietary partitioning between the two.<ref name="Divergent strategies">{{cite journal |last1=Ballell |first1=Antonio |last2=Mai |first2=Bohao |last3=Benton |first3=Michael J. |title=Divergent strategies in cranial biomechanics and feeding ecology of the ankylosaurian dinosaurs |journal=Scientific Reports |date=2023 |volume=13 |issue=1 |page=18242 |doi=10.1038/s41598-023-45444-1|pmid=37880323 |pmc=10600113 |bibcode=2023NatSR..1318242B }}</ref>

===Airspaces and senses===
[[File:Ankylosaurus nasal chambers.jpg|thumb|left|alt=Diagram of internal chambers of a skull|Diagram showing nasal chambers inside the snout (holotype)]]
In 1977, paleontologist [[Teresa Maryańska]] proposed that the complex sinuses and nasal cavities of ankylosaurs may have lightened the weight of the skull, housed a nasal [[gland]], or acted as a chamber for [[Vocal resonation|vocal resonance]].<ref name="carpenter2004"/><ref name=Maryanska1977/> Carpenter rejected these hypotheses, arguing that [[tetrapod]] animals make sounds through the [[larynx]], 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.<ref name="carpenter2004"/> 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 [[nasal concha|specialized bones]].<ref name=Maryanska1977/>

A 2011 study of the nasal passages of ''Euoplocephalus'' 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 [[Mucous membrane of nose|mucosa membrane]] (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 [[saiga antelope]] and the looping trachea of [[Crane (bird)|crane]]s and [[swan]]s. 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 [[olfaction]] (sense of smell) as the olfactory region is pushed to the sides of the main airway.<ref name="Witmer"/>

According to Carpenter, the shape of the nasal chambers of ''Ankylosaurus'' 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 [[Olfactory bulb|olfactory lobes]].<ref name="carpenter2004"/> The enlarged olfactory region of ankylosaurids indicates a well-developed sense of smell.<ref name="Witmer"/> Though hindwards retraction of the nostrils is seen in aquatic animals and animals with a [[proboscis]], it is unlikely either possibility applies to ''Ankylosaurus'', 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 [[moose]], little is known about this feature.<ref name="2017desc"/> The position of the orbits of ''Ankylosaurus'' suggest some [[Stereopsis|stereoscopic vision]].<ref name="carpenter2004"/>

===Limb movements===
[[File:Ankylosaurus scapula.jpg|thumb|Shoulder blade and coracoid of the holotype]]
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 diggers, though the hoof-like structure of the [[Manus (anatomy)|manus]] would have limited fossorial activity. Ankylosaurs were likely to have been slow-moving and sluggish animals,<ref name="digger"/><ref name="hindlimb"/> though they may have been capable of quick movements when necessary.<ref name="coombs"/>

===Growth===
The squamosal horns of the largest ''Ankylosaurus'' specimen are blunter than those of the smallest specimen, which is also the case in ''Euoplocephalus'', and this may represent [[ontogeny|ontogenetic]] variation (related to growth development).<ref name="2017desc"/> Studies of specimens of ''Pinacosaurus'' 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 weight. In the cervical half-rings, the underlying bone band developed outgrowths connecting it with the underlying osteoderms, which simultaneously fused to each other.<ref name="Burns Postcrania"/> 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.<ref name="Currie2011"/>

===Defense===
[[File:Osteoderms of Ankylosaurus.jpg|thumb|left|alt=Two fossilized knobs of bone, black with white streaks|Possible back osteoderm (holotype), in outer and inner view]]
The osteoderms of ankylosaurids were thin in comparison to those of other ankylosaurs, and appear to have been strengthened by randomly distributed cushions of [[collagen]] fibers. Structurally similar to [[Sharpey's fibres]], 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.<ref name="Scheyer"/> The [[Palpebral (bone)|palpebral bones]] over the eyes may have provided additional protection for them.<ref name=Coombs1972/> Carpenter suggested in 1982 that the heavily vascularized armor may also have had a role in [[thermoregulation]] as in modern [[crocodilian]]s.<ref name=Carpenter1982/>

The tail club of ''Ankylosaurus'' 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.<ref name="carpenter2004"/> Coombs suggested in 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 [[metatarsal bones]] of large [[Theropoda|theropods]].<ref name="hindlimb"/>

[[File:Anodontosaurus and (B) Ankylosaurus tail clubs.jpeg|thumb|alt=Diagrams of two tail clubs|Comparison between the tail clubs of ''[[Anodontosaurus]]'' and ''Ankylosaurus'']]
A 2009 study estimated that ankylosaurids could swing their tails at 100 degrees laterally, and the mainly [[cancellous]] clubs would have had a lowered [[moment of inertia]] 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 predators, in [[Intraspecific competition|intraspecific combat]], or both.<ref name=arbour/> Other studies have found evidence of ankylosaurids using their tail clubs for intraspecific combat. One specimen of ''[[Tarchia]]'' 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.<ref>{{cite journal | vauthors = Park JY, Lee YN, Kobayashi Y, Jacobs LL, Barsbold R, Lee HJ, Kim N, Song KY, Polcyn MJ | title = A new ankylosaurid from the Upper Cretaceous Nemegt Formation of Mongolia and implications for paleoecology of armoured dinosaurs | journal = Scientific Reports | volume = 11 | issue = 1 | pages = Article number 22928 | year = 2021 | doi = 10.1038/s41598-021-02273-4 | pmid = 34824329 | pmc = 8616956 | bibcode = 2021NatSR..1122928P | doi-access = free }}</ref>

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.<ref name="thulborn1993"/> Carpenter has rejected this idea, as tail club shape is highly variable among ankylosaurids, even in the same genus.<ref name="carpenter2004"/>

==Paleoenvironment==

[[File:Ankylosaurus map.gif|thumb|left|Map showing where ''Ankylosaurus'' fossils have been discovered; the holotype is shown in red (<span style="color:#9F000F">◆</span>)]] 
''Ankylosaurus'' existed between 68 and 66 million years ago, in the final, or Maastrichtian, stage of the [[Late Cretaceous]] Period. It was among the last dinosaur genera that appeared before the [[Cretaceous–Paleogene extinction event]]. 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.<ref name="vickaryousetal2004"/><ref name=WETAL04/><ref name="2017desc"/>

Fossils of ''Ankylosaurus'' are rare in the [[sediments]] 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 ''Edmontonia'' 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 it had a more selective diet than ''Ankylosaurus'', further indicating [[Niche differentiation|ecological separation]], whether their range overlapped or not.<ref name="2017desc"/><ref name="carpenter2004"/>

With its low center of gravity, ''Ankylosaurus'' 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, ''Ankylosaurus'' does not appear to have congregated in groups (though some ankylosaurs appear to have congregated when young). It is therefore improbable that ''Ankylosaurus'' was able to modify the landscape of its ecosystem in the way elephants do; [[hadrosaurid]]s may instead have had such an "[[ecosystem engineer]]" role.<ref name="2017desc"/>

The formations where ''Ankylosaurus'' fossils have been found represent different sections of the western shore of the [[Western Interior Seaway]] dividing western and eastern North America during the Cretaceous, a broad coastal plain extending westward from the seaway to the newly formed [[Rocky Mountains]]. These formations are composed largely of [[sandstone]] and [[mudstone]], which have been attributed to [[floodplain]] environments.<ref name="lofgren1997"/><ref name="breithaupt1997"/><ref name="eberth1997"/> The regions where ''Ankylosaurus'' and other Late Cretaceous ankylosaurs have been found had a warm [[Subtropics|subtropical]]/[[temperate|temperate climate]], which was [[monsoonal]], had occasional rainfall, [[tropical storms]], and [[forest fires]].<ref name="Ősi"/> In the Hell Creek Formation, many types of plants were supported, primarily [[angiosperm]]s, with less common [[conifer]]s, ferns and [[cycad]]s. An abundance of fossil leaves found at dozens of different sites indicates that the area was largely forested by small trees.<ref name="johnson1997"/> ''Ankylosaurus'' shared its environment with other dinosaurs that included the [[Ceratopsidae|ceratopsids]] ''[[Triceratops]]'' and ''[[Torosaurus]]'', the [[hypsilophodont]] ''[[Thescelosaurus]]'', the hadrosaurid ''[[Edmontosaurus]]'', an indeterminate nodosaur, the [[pachycephalosauria]]n ''[[Pachycephalosaurus]]'', and the theropods ''[[Struthiomimus]]'', ''[[Ornithomimus]]'', ''[[Pectinodon]]'', and ''Tyrannosaurus''.<ref name=WETAL04/><ref name=HCFF/>

==Cultural significance==

[[File: Royal Alberta museum Ankylosaurus.jpg|thumb|alt=Photograph of a dinosaur diorama|Replica of the [[1964 World's Fair]] ''Ankylosaurus'' (note spikes and dragging tail), [[Royal Alberta Museum]]]]
Carpenter noted in 2004 that ''Ankylosaurus'' has become the [[Archetype|archetypal]] member of its group, and the best-known ankylosaur in [[popular culture]], perhaps due to a life-sized reconstruction of the animal being featured at the [[1964 New York World's Fair|1964 World's Fair]] in New York City.<ref name="carpenter2004"/> Arbour and Mallon called ''Ankylosaurus'' an "iconic" dinosaur in 2017, and noted that the World's Fair sculpture, as well as the American artist [[Rudolph Zallinger]]'s 1947 mural ''[[The Age of Reptiles]]'' and other later popular depictions, showed ''Ankylosaurus'' with a tail club, following the first discovery of the feature in 1910.<ref name="2017desc"/>

Many traditional popular depictions show ''Ankylosaurus'' 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 present in many traditional depictions, but are not known from ''Ankylosaurus'' itself.<ref name="glut1997"/> The armor of ''Ankylosaurus'' has often been conflated with that of ''[[Edmontonia]]'' (earlier referred to as ''[[Palaeoscincus]]''); in addition to ''Ankylosaurus'' being depicted with spikes, ''Edmontonia'' has also been depicted with an ''Ankylosaurus''-like tail club (a feature nodosaurids did not have), including in a mural by the American artist [[Charles R. Knight]] from 1930.<ref name="2017desc"/> ''Ankylosaurus'' has been featured in the ''[[Jurassic Park]]'' franchise, where they are depicted as attacking with their tails and running, abilities that have been criticized as unlikely by paleontologists.<ref name="Wired">{{cite magazine |title=We asked a paleontologist how accurate ''Jurassic World'' really is... |url=https://www.wired.co.uk/article/jurassic-world-dinosaur-expert |magazine=Wired UK |access-date=August 29, 2023 |date=2015 |archive-date=September 20, 2020 |archive-url=https://web.archive.org/web/20200920075538/https://www.wired.co.uk/article/jurassic-world-dinosaur-expert |url-status=live }}</ref><ref name="Time">{{cite magazine |last1=Waxman |first1=Olivia B. |title=The real scientific history behind the ''Jurassic Park'' dinosaurs |url=https://time.com/5313949/real-dinosaurs-jurassic-world-fallen-kingdom-jurassic-park-fact-check-checking-history/ |access-date=August 29, 2023 |magazine=Time |date=2018 |archive-date=November 16, 2021 |archive-url=https://web.archive.org/web/20211116232936/https://time.com/5313949/real-dinosaurs-jurassic-world-fallen-kingdom-jurassic-park-fact-check-checking-history/ |url-status=live }}</ref><ref name="Looper">{{cite news |last1=Dorsch |first1=Rita |title=Everything the ''Jurassic Park'' franchise gets wrong about paleontology |url=https://www.looper.com/892011/everything-the-jurassic-park-franchise-gets-wrong-about-paleontology/ |work=Looper |date=2022|access-date=August 29, 2023 }}</ref>

==See also==
* [[Timeline of ankylosaur research]]

==References==
'''Notes'''
{{Reflist|group=nb}}
'''Citations'''
{{Reflist|30em|refs=
<ref name=HCFF>{{cite web|author=Bigelow, P. |title=Cretaceous 'Hell Creek Faunal Facies'; Late Maastrichtian |url=http://www.scn.org/~bh162/hellcreek2.html |access-date=March 24, 2014 |archive-url=https://web.archive.org/web/20091226162134/http://www.scn.org/~bh162/hellcreek2.html |archive-date=December 26, 2009 |url-status=live }}</ref>
<ref name=WETAL04>{{cite book|author1=Weishampel, D. B. |author2=Barrett, P. M. |author3=Coria, R. A. |author4=Le Loeuff, J. |author5=Xu X. |author6=Zhao X. |author7=Sahni, A. |author8=Gomani, E. M. P. |author9=Noto, C. R. |year=2004|contribution=Dinosaur Distribution|editor1=Weishampel, D. B. |editor2=Dodson, P. |editor3=Osmolska, H.. |title=The Dinosauria (2nd)|publisher=University of California Press|pages=517–606|isbn=978-0-520-24209-8}}</ref>
<ref name="eberth1997">{{cite book|author =Eberth, D. A.|year=1997|contribution=Edmonton Group |editor1=Currie, P. J. |editor2=Padian, K. |title=The Encyclopedia of Dinosaurs |publisher=Academic Press |pages=199–204|isbn=978-0-12-226810-6}}</ref>
<ref name="breithaupt1997">{{cite book|author =Breithaupt, B. H.|year=1997|contribution=Lance Formation |editor1=Currie, P.J. |editor2=Padian, K.|title=The Encyclopedia of Dinosaurs |publisher=Academic Press |pages=394–95 |isbn=978-0-12-226810-6}}</ref>
<ref name="lofgren1997">{{cite book|author =Lofgren, D. F.|year=1997|contribution=Hell Creek Formation|editor1=Currie, P.J. |editor2=Padian, K. |title=The Encyclopedia of Dinosaurs |publisher=Academic Press |pages=302–03 |isbn=978-0-12-226810-6}}</ref>
<ref name="carpenter2001">{{cite book |author =Carpenter, K. |year=2001 |title=The Armored Dinosaurs |editor=Carpenter, K. |contribution=Chapter 21: Phylogenetic Analysis of the Ankylosauria |pages=454–83 |publisher=Indiana University Press |isbn=978-0-253-33964-5 |url=https://books.google.com/books?id=04tQ5_qJN8MC&q=The%20Armored%20Dinosaurs&pg=PA455}}</ref>
<ref name="brown1908">{{cite journal |author =Brown, B. |year=1908 |title=The Ankylosauridae, a new family of armored dinosaurs from the Upper Cretaceous |journal=Bulletin of the American Museum of Natural History |series=24 |pages=187–201 |hdl=2246/1435 }}</ref>
<ref name="thulborn1993">{{cite journal |author =Thulborn, T. |year=1993 |title=Mimicry in ankylosaurid dinosaurs |journal=Records of the South Australian Museum |series=27 |pages=151–58}}</ref>
<ref name=arbour>{{cite journal|author =Arbour, V. M.|year=2009|title= Estimating impact forces of tail club strikes by ankylosaurid dinosaurs|journal=PLOS ONE |volume=4|issue=8|page=e6738|doi=10.1371/journal.pone.0006738|bibcode=2009PLoSO...4.6738A|pmid=19707581|pmc=2726940|doi-access=free}}</ref>
<ref name="carpenter2004">{{cite journal |author =Carpenter, K. |year=2004 |title=Redescription of ''Ankylosaurus magniventris'' Brown 1908 (Ankylosauridae) from the Upper Cretaceous of the Western Interior of North America |journal=Canadian Journal of Earth Sciences |volume=41 |issue=8 |pages=961–86|doi=10.1139/e04-043|bibcode=2004CaJES..41..961C}}</ref>
<ref name="Burns Postcrania">{{cite journal |last1=Burns|first1=M |last2=Tumanova|first2=T |last3=Currie|first3=P |title=Postcrania of juvenile ''Pinacosaurus grangeri'' (Ornithischia: Ankylosauria) from the Upper Cretaceous Alagteeg Formation, Alag Teeg, Mongolia: implications for ontogenetic allometry in ankylosaurs|journal=Journal of Paleontology |date=2015 |volume=89 |issue=1 |pages=168–182 |doi=10.1017/jpa.2014.14 |bibcode=2015JPal...89..168B |s2cid=130610291 }}</ref>
<ref name="vickaryousetal2004">{{cite book |author=Vickaryous, M. K. |author2=Maryanska, T. |author3=Weishampel, D. B.|contribution=Ankylosauria|editor1=Weishampel, D. B. |editor2=Dodson, P. |editor3=Osmólska, H. |year=2004 |title=The Dinosauria |publisher=University of California Press |pages=363–92 |isbn=978-0-520-24209-8}}</ref>
<ref name="johnson1997">{{cite book|author =Johnson, K. R.|year=1997|contribution=Hell Creek Flora |editor1=Currie, P. J. |editor2=Padian, K. |title=The Encyclopedia of Dinosaurs |publisher=Academic Press |pages=300–02 |isbn=978-0-12-226810-6}}</ref>
<ref name=pronunciation>{{cite web|author=Creisler, B. |title=Dinosauria Translation and Pronunciation Guide A |date=July 7, 2003 |url=http://www.dinosauria.com/dml/names/dinoa.htm |access-date=September 3, 2010 |archive-url=https://web.archive.org/web/20100818013919/http://www.dinosauria.com/dml/names/dinoa.htm |archive-date=August 18, 2010 |url-status=dead }}</ref>
<ref name="hindlimb">{{cite journal|author =Coombs, W.|year=1979 |title=Osteology and myology of the hindlimb in the Ankylosauria (Reptillia, Ornithischia)|journal=Journal of Paleontology |volume=53 |issue=3 |pages=666–84 |jstor=1304004}}</ref>
<ref name="Witmer">{{cite journal |author=Miyashita, T. |author2=Arbour V. M. |author3=Witmer L. M. |author4=Currie, P. J.|year=2011 |title=The internal cranial morphology of an armoured dinosaur ''Euoplocephalus'' corroborated by X-ray computed tomographic reconstruction|journal=Journal of Anatomy |volume=219|issue=6|pages=661–75|doi=10.1111/j.1469-7580.2011.01427.x |pmid=21954840 |pmc=3237876 |url=http://www.oucom.ohiou.edu/dbms-witmer/Downloads/2011_Miyashita_et_al._Euoplocephalus_head_anatomy.pdf |url-status=dead |archive-url=https://web.archive.org/web/20150924062640/http://www.oucom.ohiou.edu/dbms-witmer/Downloads/2011_Miyashita_et_al._Euoplocephalus_head_anatomy.pdf |archive-date=September 24, 2015}}</ref>
<ref name="digger">{{cite journal|author =Coombs, W.|year=1978|title=Forelimb muscles of the Ankylosauria (Reptilia, Ornithischia)|journal=Journal of Paleontology|volume=52|issue=3|pages=642–57|jstor=1303969}}</ref>
<ref name="Currie2011">{{cite journal|author1=Currie, P. J.|author2=Badamgarav, D.|author3=Koppelhus, E. B.|author4=Sissons, R.|author5=Vickaryous, M. K.|year=2011|title=Hands, feet, and behaviour in ''Pinacosaurus'' (Dinosauria: Ankylosauridae)|journal=Acta Palaeontologica Polonica|volume=56|issue=3|pages=489–504|doi=10.4202/app.2010.0055|s2cid=129291148|url=http://www.app.pan.pl/archive/published/app56/app20100055.pdf|doi-access=free|archive-date=July 8, 2021|access-date=September 14, 2019|archive-url=https://web.archive.org/web/20210708144810/http://www.app.pan.pl/archive/published/app56/app20100055.pdf|url-status=live}}</ref>
<ref name="Scheyer">{{cite journal|author1=Scheyer, T. M. |author2=Sander, P. M. |year=2004|title=Histology of ankylosaur osteoderms: implications for systematics and function|journal=Journal of Vertebrate Paleontology|volume=24|issue=4|pages=874–93|jstor=4524782|doi=10.1671/0272-4634(2004)024[0874:hoaoif]2.0.co;2|s2cid=86333501 }}</ref>
<ref name=Carpenter1982>{{cite journal|author = Carpenter, K.|year=1982|title=Skeletal and dermal armor reconstruction of ''Euoplocephalus tutus'' (Ornithischia: Ankylosauridae) from the Late Cretaceous Oldman Formation of Alberta|journal=Canadian Journal of Earth Sciences|volume=19|issue=4|pages=689–97|doi=10.1139/e82-058|bibcode=1982CaJES..19..689C}}</ref>
<ref name="Hans1969">{{cite journal|author =Haas, G.|year=1969|title=On the jaw musculature of ankylosaurs|journal=American Museum Novitates|issue=2399|pages=1–11 |hdl=2246/2609 }}</ref>
<ref name=Rybzynski2001>{{cite book|author1=Rybczynski, N. |author2=Vickaryous, M. K. |year=2001|contribution=Chapter 14: Evidence of Complex Jaw Movement in the Late Cretaceous Ankylosaurid, ''Euoplocephalus tutus'' (Dinosauria: Thyreophora)|pages=299–317|editor=K. Carpenter|title=The Armored Dinosaurs|publisher=Indiana University Press|isbn=978-0-253-33964-5}}</ref>
<ref name=Coombs1972>{{cite journal|author=Coombs W.|year=1972|title=The Bony Eyelid of ''Euoplocephalus'' (Reptilia, Ornithischia)|journal=Journal of Paleontology |volume=46 |issue=5 |pages=637–50 |jstor=1303019}}.</ref>
<ref name=Maryanska1977>{{cite journal|author=Maryanska, T.|year=1977|title=Ankylosauridae (Dinosauria) from Mongolia|journal=Palaeontologia Polonica|volume=37|pages=85–151|url=http://palaeontologia.pan.pl/Archive/1977-37_85-151_19-36.pdf|access-date=July 25, 2015|archive-date=July 12, 2020|archive-url=https://web.archive.org/web/20200712160519/http://palaeontologia.pan.pl/Archive/1977-37_85-151_19-36.pdf|url-status=dead}}</ref>
<ref name="coombs">{{cite journal|author =Coombs, W. P.|title=Theoretical aspects of cursorial adaptations in dinosaurs |journal=[[The Quarterly Review of Biology]] |volume=53 |year=1978 |pages=393–418 |doi=10.1086/410790 |issue=4|s2cid=84505681 }}</ref>
<ref name="Oxford">{{cite book | author1=Liddell, H. G. | author2=Scott, R. | edition=abridged | isbn=978-0-19-910207-5 | orig-year=1871 | title=A Greek-English Lexicon | publisher=[[Oxford University Press]] | year=1980 | page=[https://archive.org/details/lexicon00lidd/page/5 5] | title-link=A Greek-English Lexicon }}</ref>
<ref name="osborn1905">{{cite journal|author =Osborn, H. F. |year=1905 |title=Tyrannosaurus and other Cretaceous carnivorous dinosaurs |journal=Bulletin of the AMNH |volume=21 |issue=14 |pages=259–265 |hdl=2246/1464 }}</ref>
<ref name = "osborn1923">{{cite journal|author =Osborn, H. F.|year=1923|title=Two Lower Cretaceous dinosaurs of Mongolia|journal=American Museum Novitates|issue=95|pages=1–10 |hdl=2246/3267 }}</ref>
<ref name="Coombs teeth">{{cite book|author =Coombs, W.|year=1990|contribution=Teeth and taxonomy in ankylosaurs |editor1=Carpenter, K. |editor2=Currie, P. J.|title=Dinosaur systematics: Approaches and perspectives |pages=269–79 |publisher=Cambridge University Press |url=https://books.google.com/books?id=6ZV1KcVNM18C&pg=PA269 |isbn=978-0-521-43810-0}}</ref>
<ref name="Coombs1978">{{cite journal|author=Coombs, W.|year=1978|title=The families of the ornithischian dinosaur order Ankylosauria|journal=Journal of Paleontology|volume=21|issue=1|pages=143–170|url=http://cdn.palass.org/publications/palaeontology/volume_21/pdf/vol21_part1_pp143-170.pdf|access-date=July 8, 2015|archive-url=https://web.archive.org/web/20150708192417/http://cdn.palass.org/publications/palaeontology/volume_21/pdf/vol21_part1_pp143-170.pdf|archive-date=July 8, 2015|url-status=dead}}</ref>
<ref name="Williston1908">{{cite journal|author=Williston, S. W.|year=1908|title=Review: The Ankylosauridae|journal=The American Naturalist|volume=42|issue=501|pages=629–30|doi=10.1086/278987|jstor=2455817|url=https://zenodo.org/record/1431351|doi-access=free|archive-date=November 12, 2019|access-date=November 12, 2019|archive-url=https://web.archive.org/web/20191112110536/https://zenodo.org/record/1431351|url-status=live}}</ref>
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<ref name="thompson">{{Cite journal | last1 = Thompson | first1 = R. S. | last2 = Parish | first2 = J. C. | last3 = Maidment | first3 = S. C. R. | last4 = Barrett | first4 = P. M. | title = Phylogeny of the ankylosaurian dinosaurs (Ornithischia: Thyreophora) | doi = 10.1080/14772019.2011.569091 | journal = Journal of Systematic Palaeontology | volume = 10 | issue = 2 | pages = 301–312 | year = 2012 | bibcode = 2012JSPal..10..301T | s2cid = 86002282 }}</ref>
<ref name="Arbour2014II">{{cite journal|last1=Arbour|first1=V.M.|last2=Currie|first2=P.J.|last3=Badamgarav|first3=D.|year=2014|title=The ankylosaurid dinosaurs of the Upper Cretaceous Baruungoyot and Nemegt formations of Mongolia|journal=Zoological Journal of the Linnean Society|volume=172|issue=3|pages=631–652|doi=10.1111/zoj.12185}}</ref>
<ref name="systematics ankylosaurid">{{cite journal|author1=Arbour, V. M. |author2=Currie, P. J. |year=2015|title=Systematics, phylogeny and palaeobiogeography of the ankylosaurid dinosaurs|journal=Journal of Systematic Palaeontology|volume=14|issue=5|pages=1–60|doi=10.1080/14772019.2015.1059985|s2cid=214625754 }}</ref>
<ref name="glut1997">{{Cite book| publisher = McFarland & Company, Inc. Publishers| isbn = 978-0-375-82419-7| last = Glut| first = D. F.| title = Dinosaurs, the encyclopedia| year = 1997| chapter = Ankylosaurus| pages = [https://archive.org/details/dinosaursmostcom00holt/page/141 141–143]| chapter-url = https://archive.org/details/dinosaursmostcom00holt| url = https://archive.org/details/dinosaursmostcom00holt/page/141}}</ref>
<ref name="carpenter1982b">{{Cite journal| volume = 20| issue = 2| pages = 123–134| last = Carpenter| first = K.| title = Baby dinosaurs from the Late Cretaceous Lance and Hell Creek formations and a description of a new species of theropod| journal = Rocky Mountain Geology| date = 1982| url = http://rmg.geoscienceworld.org/content/20/2/123.short}}</ref>
}}

==External links==
{{Wikispecies|Ankylosaurus}}
{{Commons category}}
{{wiktionary}}
{{Wikibooks|Wikijunior Dinosaurs/Ankylosaurus}}

{{Thyreophora|A.}}
{{Portal bar|Dinosaurs|United States}}
{{Taxonbar|from=Q40621}}
{{authority control}}

[[Category:Ankylosaurinae]]
[[Category:Dinosaur genera]]
[[Category:Maastrichtian dinosaurs]]
[[Category:Hell Creek Formation]]
[[Category:Lance Formation]]
[[Category:Scollard Formation]]
[[Category:Frenchman Formation]]
[[Category:Dinosaurs of Canada]]
[[Category:Fossil taxa described in 1908]]
[[Category:Taxa named by Barnum Brown]]
[[Category:Dinosaurs of the United States]]