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Ankylosaurus

Ankylosaurus is a genus of ankylosaurid dinosaur known for its heavily armored body and distinctive tail club, representing one of the last non-avian dinosaurs before the Cretaceous–Paleogene extinction event.^[1] The type and only recognized species, Ankylosaurus magniventris, was a quadrupedal herbivore that inhabited western North America during the Maastrichtian stage of the Late Cretaceous period, approximately 68 to 66 million years ago.^[1] Fossils of this dinosaur have been primarily discovered in the Hell Creek Formation of Montana, with additional remains from formations in Alberta (Canada), Saskatchewan (Canada), and South Dakota (USA).^[1] Characterized by its tank-like build, Ankylosaurus measured up to 8 meters (26 feet) in length and weighed around 8,000 kilograms (18,000 pounds), making it one of the largest members of the Ankylosauridae family.^[1] Its body was encased in extensive dermal armor, including polygonal osteoderms and bony plates that formed a mosaic-like covering, providing robust protection against predators such as tyrannosaurids.^[2] The dinosaur's skull was broad and low, equipped with a horny beak for cropping vegetation and leaf-shaped teeth suited for grinding tough plant matter, including ferns, cycads, and low-lying shrubs.^[1] At the end of its tail, Ankylosaurus possessed a unique osteoderm-reinforced club, which likely served as a defensive weapon capable of delivering powerful blows to attackers or rivals, as evidenced by the stepwise evolution of this feature in ankylosaurids.^[2]^[3] The genus was first described and named by paleontologist Barnum Brown in 1908, based on fragmentary remains including vertebrae, ribs, and armor collected from the Hell Creek Formation in 1906, marking it as the type genus of the Ankylosauridae family.^[1] Despite its iconic status in popular culture, Ankylosaurus is known from relatively few complete specimens, with no fully articulated skeletons preserving both the skull and tail club together, leading to ongoing research into its precise anatomy and behavior.^[4] As part of the Thyreophora clade within Ornithischia, Ankylosaurus exemplifies the diverse defensive adaptations of armored dinosaurs that dominated Late Cretaceous ecosystems.^[2]

Discovery and Naming

Initial Discovery

The American Museum of Natural History (AMNH) initiated a series of expeditions to the Cretaceous deposits of Montana starting in 1902, led by paleontologist Barnum Brown, with the goal of uncovering the rich Late Cretaceous vertebrate faunas of the region, including dinosaurs from the uppermost Maastrichtian stage.^[5] These efforts built on earlier explorations and focused on productive sites like the Hell Creek Formation, known for yielding fossils of large herbivores and carnivores near the end of the dinosaur era.^[1] In 1906, during one such expedition, Brown discovered the type specimen of Ankylosaurus (AMNH 5895) in the Hell Creek Formation near Gilbert Creek, Montana.^[1] This partial skeleton represented the first definitive evidence of a heavily armored dinosaur from this formation and included the dorsal portion of the skull, five cervical vertebrae, eleven dorsal vertebrae, three caudal vertebrae, the right scapulocoracoid, several ribs, and scattered osteoderms (armor plates).^[6] The find highlighted the diversity of ankylosaurid dinosaurs in the latest Cretaceous ecosystems of western North America. Brown formally named the species Ankylosaurus magniventris in 1908, establishing the new family Ankylosauridae based primarily on this specimen.^[7] The generic name derives from the Greek words ankylos (bent, crooked, or fused, referring to the interlocking armor) and sauros (lizard), while the specific epithet magniventris means "large-bellied," alluding to the wide, robust ribcage evident in the preserved elements.^[6] This initial description emphasized the animal's distinctive fused osteoderms and low-slung body, setting the stage for recognizing ankylosaurids as a specialized group of ornithischian dinosaurs.^[7]

Subsequent Specimens

Following the discovery of the type specimen in 1908, additional Ankylosaurus fossils were recovered starting in 1910. That year, Barnum Brown of the American Museum of Natural History (AMNH) unearthed specimen AMNH 5214 along the Red Deer River in Alberta, Canada, from the Scollard Formation.^[8] This specimen includes a complete skull, mandibles, ribs, caudal vertebrae, limb elements, and the only well-preserved tail club known for the genus, measuring 60 cm long, 49 cm wide, and 19 cm high.^[8] In 1947, Charles M. Sternberg and T. Potter Chamney collected specimen CMN 8880 from the right bank of the Red Deer River in Alberta, Canada, also from the Scollard Formation.^[8] This material consists of a large skull and left mandible, representing one of the most complete cranial specimens and providing key insights into variation in skull size among individuals.^[8] Numerous fragmentary remains of Ankylosaurus have been reported from Late Cretaceous formations across western North America, including the Hell Creek Formation of Montana, the Lance Formation of Wyoming, the Scollard Formation of Alberta, and the Frenchman Formation of Saskatchewan.^[8] These consist primarily of isolated osteoderms, vertebrae, teeth, and partial limb bones, often preserving less than 10% of any single individual's skeleton.^[8] No complete Ankylosaurus skeletons have been found, with only a handful of specimens known overall.^[8] Despite ongoing paleontological work in these regions, no major new Ankylosaurus specimens have been described since CMN 8880, in contrast to the steady stream of discoveries for other ankylosaur genera in recent decades.^[8]

Historical Reconstructions

The initial scientific reconstruction of Ankylosaurus magniventris was presented by Barnum Brown in 1908, shortly after the discovery of the type specimen, portraying the dinosaur in a semi-upright posture reminiscent of Stegosaurus, with a speculative arrangement of armor featuring closely packed thoracic osteoderms and coossified bands around the pelvis.^[7] At that time, the limited fossil material—primarily consisting of the holotype skull, vertebrae, ribs, and isolated osteoderms—led to an interpretation of the tail as flexible and lacking a club, influencing early artistic depictions that emphasized a more lizard-like, sprawling form rather than the rigid structure later recognized.^[8] In the 1910s, the discovery of additional specimens, such as AMNH 5214, revealed the presence of a tail club, prompting revisions in subsequent reconstructions. By the 1920s and 1940s, museum displays at the American Museum of Natural History (AMNH) and the Canadian Museum of Nature (CMN) featured partial mounts and restorations emphasizing a broad-bodied form with the tail club prominently raised, often incorporating osteoderms borrowed from related taxa like Edmontonia (formerly "Palaeoscincus"), which resulted in exaggerated features such as pectoral spikes in artistic renderings like Charles Knight's 1930 mural.^[8] These depictions, while advancing the understanding of the animal's armored profile, still reflected the constraints of fragmentary remains, perpetuating overly flexible tail interpretations in some illustrations until better comparative material from other ankylosaurids became available.^[9] Significant shifts occurred in the 1970s and 1990s, as paleontologists like Walter Coombs analyzed forelimb musculature and pelvic structure, establishing Ankylosaurus as a fully quadrupedal animal with a low-slung, barrel-shaped body supported primarily by robust forelimbs, rather than the semi-erect stance of earlier models.^[9] This revision was driven by comparisons with more complete ankylosaurid specimens, such as those of Euoplocephalus, which highlighted the dinosaur's stiff tail and grounded posture adapted for stability amid heavy armor.^[8] Post-2000 reconstructions have increasingly relied on digital modeling and computed tomography (CT) scans of skulls and osteoderms, enabling precise visualizations of internal cranial features and armor patterning, as seen in updated restorations that refine the arrangement of cervical half-rings and thoracic plates based on specimens like AMNH 5214.^[8] These modern approaches, exemplified by three-dimensional analyses in studies from the 2010s, correct prior conflations with other genera and portray Ankylosaurus as a heavily built quadruped with a defensive tail club wielded via a stiffened handle.^[10]

Description

Skull

The skull of Ankylosaurus magniventris is low-profile and broad, measuring up to 67 cm in basal length in the largest known specimens, such as CMN 8880. It features a robust construction with fused nasal bones forming a median nasal caputegulum, and the external nares are oriented ventrolaterally, positioned posteriorly and not visible from an anterior view. In Ankylosaurus, the loreal caputegulum roofs the nasal vestibule entirely, differing from other ankylosaurids such as Euoplocephalus, where supranarial caputegulae roof it, potentially influencing airflow dynamics.^[8] The skull bears prominent horns on the squamosals, quadratojugals, and nasals, with pyramidal shapes on the squamosals and quadratojugals that are proportionally longer than in relatives like Euoplocephalus. The bone surfaces exhibit rough textures indicative of keratin sheaths covering these horns, enhancing their defensive role, as inferred from vascular patterns common in ornithischian cranial ornamentation. Additional caputegulae, including rectangular to hexagonal frontonasals and a bulbous, laterally expanded loreal, contribute to the armored cranial profile.^[8] Dentition consists of leaf-shaped, asymmetrical teeth adapted for grinding vegetation, with crowns up to 9.1 mm high in specimen AMNH 5214. The maxilla accommodates 34–36 such teeth, more than in Anodontosaurus or Euoplocephalus, reflecting a capacity for processing tough plant matter through occlusion. Tooth size varies, but overall they are proportionally small relative to skull length.^[8] The braincase is short and robust, consistent with the reduced encephalization typical of ankylosaurids, though direct endocranial volumes remain undocumented for Ankylosaurus. Internal nasal passages form a looping airway system, as partially preserved in AMNH 5895, suggesting complex respiratory or sensory adaptations.^[8]

Postcranial Skeleton

The postcranial skeleton of Ankylosaurus magniventris features a robust, barrel-shaped torso composed of approximately 75–80 vertebrae, though the exact count remains uncertain due to the fragmentary nature of known fossils. Preserved specimens, such as the holotype AMNH 5895, include five cervical vertebrae and eleven dorsal vertebrae, contributing to a broad, stable axial structure. The sacral region is fused, with multiple sacral vertebrae co-ossified to the pelvis for enhanced rigidity and support of the heavy body.^[8]^[11] The appendicular skeleton consists of sturdy, pillar-like limbs suited to quadrupedal weight-bearing. The forelimbs are slightly shorter than the hindlimbs, as evidenced by a humerus length of 54 cm compared to a femur of 67 cm in specimen AMNH 5214; the humerus and radius are particularly robust, with expanded proximal and distal ends. Foot elements are incompletely preserved, but available phalanges from related ankylosaurids suggest a hoof-like configuration with reduced digits.^[8]^[7] The pelvis exhibits broad, flaring ilia that accommodate the wide torso, although preservation is poor across specimens, limiting detailed analysis. The tail includes an estimated 8–13 proximal free caudal vertebrae based on partial sequences in AMNH 5895 and AMNH 5214, transitioning to stiffened distal segments of seven co-ossified vertebrae that form the handle supporting the tail club.^[8]^[11] Broad ribs, with six preserved in AMNH 5214 and some co-ossified to the posterior dorsals in AMNH 5895, along with gastralia, enclose the torso in a basket-like framework that reinforces the body's defensive posture. Overall body length is estimated at 6–8 m using scaling from specimens like AMNH 5214 and CMN 8880, with masses of 4.8–8 tonnes derived from volumetric reconstructions. This skeletal framework was enveloped by osteoderms that provided additional protection to the underlying bones.^[8]^[7]

Armor

The dermal armor of Ankylosaurus consisted primarily of osteoderms, which are bony deposits embedded within the skin that formed a protective covering over much of the body. These included large polygonal scutes on the back, some measuring up to approximately 30 cm in length, arranged in irregular rows along the dorsal surface. Along the sides, smaller, keeled plates provided additional coverage, typically more elongate and pointed to accommodate the body's contours. The neck was safeguarded by cervical half-rings, comprising three pairs of fused osteoderms that encircled the region while permitting limited mobility. Each half-ring featured multiple keeled, oval-based elements, with the medial ones often flatter and less pronounced. These structures were positioned to shield the vulnerable cervical area without overly restricting head movement. A distinctive feature was the tail club, formed by a series of stiffened caudal vertebrae creating a pyramidal handle, topped by a bulbous knob assembled from fused osteoderms. The knob varied in size among specimens, typically ranging from 30 to 60 cm in width, with one well-preserved example measuring 49 cm wide, 60 cm long, and 19 cm high.^[12] Osteoderms were composed of dense, thick cortical bone surrounding a core of cancellous tissue, often exhibiting prominent vascular grooves on their surfaces that suggest nutrient supply during growth. These bony elements were likely sheathed in a layer of keratin, enhancing their durability and possibly their coloration or texture. The osteoderms were loosely attached to the underlying endoskeleton via connective tissue, allowing some flexibility. Overall, the distribution of osteoderms was densest on the dorsal surface and along the tail, with an estimated total exceeding 200 per individual, varying in size from small nodules under 1 cm to larger plates over 30 cm, ensuring comprehensive coverage while adapting to different body regions.

Classification

Taxonomic History

Ankylosaurus was established as a genus by Barnum Brown in 1908, with the type species A. magniventris based on fragmentary remains including a partial skull, vertebrae, ribs, and armor ossicles from the Hell Creek Formation of Montana. Brown assigned the new family Ankylosauridae to the suborder Stegosauria, reflecting the limited comparative material available at the time, which led to its initial grouping with plated dinosaurs like Stegosaurus. In 1923, Henry Fairfield Osborn formally named the suborder Ankylosauria to distinguish ankylosaurids from stegosaurs, recognizing their unique osteoderm arrangements and body form as warranting separation within Thyreophora. Franz Nopcsa further advanced the taxonomy in 1928 by elevating Ankylosauridae to family status and providing detailed diagnoses that emphasized the fused armor and tail club as diagnostic traits, solidifying Ankylosaurus as a core member. Early synonymy debates arose with related North American ankylosaurids, particularly regarding potential mergers with Euoplocephalus tutus from the Dinosaur Park Formation; some mid-20th-century workers, including L.S. Russell in the 1940s, initially questioned distinctions based on overlapping armor patterns but ultimately affirmed A. magniventris as a valid, monotypic species due to its larger size, more robust tail club, and later stratigraphic position in the Maastrichtian. Walter Coombs's revisions in the 1970s and 1980s reinforced this by analyzing cranial and postcranial metrics, excluding nodosaurid referrals (such as misidentified specimens once tentatively linked to Ankylosaurus but later assigned to genera like Edmontonia) and confirming Ankylosauridae's internal divisions. A comprehensive redescription by Victoria M. Arbour and Jordan C. Mallon in 2017 utilized computed tomography and comparative morphometrics on the holotype (AMNH 5895) and referred specimens, distinguishing Ankylosaurus from close relatives like Anodontosaurus lambei (previously synonymized with Euoplocephalus but revived as distinct) through differences in skull length-to-width ratios, narial aperture position, and quadrate orientation. This work upheld A. magniventris as the sole valid species, attributing prior uncertainties to the genus's fragmentary fossil record. No additional species have been proposed since 2020, owing to the paucity of complete skeletons and reliance on diagnostic cranial elements for new ankylosaurid referrals.^[8]

Phylogenetic Relationships

Ankylosaurus belongs to the family Ankylosauridae, a derived clade within the larger suborder Ankylosauria of ornithischian dinosaurs. Cladistic analyses consistently position it as a highly derived ankylosaurid from the Late Maastrichtian stage of the Late Cretaceous, approximately 68–66 million years ago.^[13] It shares key synapomorphies with other ankylosaurids, including fused cervical half-rings that form protective bony collars around the neck.^[14] Phylogenetic studies incorporating cranial and postcranial data have refined its relationships within Ankylosauridae. In North American taxa, Ankylosaurus forms a clade with sister taxa Anodontosaurus and Euoplocephalus, characterized by shared features such as robust tail clubs and extensive dermal armor patterning.^[13] The evolutionary timeline of Ankylosauria traces its origins to the Middle Jurassic, around 160 million years ago, with early forms appearing in Laurasian deposits.^[15] Ankylosaurus stands as one of the final surviving genera in this lineage, enduring until the Cretaceous-Paleogene extinction event. Ankylosaurus exhibits no close relatives beyond the Laramidian continent of western North America, underscoring its endemism in Late Cretaceous ecosystems.^[16] This contrasts with Gondwanan ankylosaurs, such as the basal form Spicomellus from Middle Jurassic Africa, whose 2025 osteological redescription reveals unique spike-like armor and challenges prior models of early ankylosaur distribution.

Paleobiology

Locomotion and Posture

Ankylosaurus was an obligate quadruped with a broad, low-slung body supported by short, robust limbs that inferred a wide-track gait, as evidenced by the divergent ilia and pillar-like femoral structure accommodating its armored mass.^[17] The forelimbs exhibited a semi-erect posture, with a stout humerus and prominent deltopectoral crest enabling weight-bearing stability, while the hindlimbs were more sprawling, featuring a massive, straight femur and reduced tibia that positioned the body close to the ground for a stable, ponderous stance. This configuration prioritized slow foraging over agility, with biomechanical analyses of limb scaling indicating the dinosaur could achieve estimated maximum speeds of 6-8 km/h in a walking gait.^[18] The tail of Ankylosaurus demonstrated proximal flexibility for powerful swinging motions, contrasted by distal rigidity in the fused handle vertebrae and osteoderm knob forming the club, allowing effective lateral arcs during locomotion or defense.^[19] Recent 2020s biomechanical models, informed by finite element analyses and pathology evidence from related ankylosaurids, suggest the tail club could achieve near-360-degree rotational mobility, facilitating intraspecific combat or predator deterrence through high-impact swings.^[20] Limb bones, including the robust 67 cm femur, were adapted for heavy weight-bearing to support the 4.8-8 tonne body during extended low-speed traversal of forested environments, rather than rapid evasion.^[17] Evidence for ankylosaurid locomotion, informing inferences for Ankylosaurus, comes from rare trackways of related species from earlier Cretaceous formations. These include 2025 discoveries in the Canadian Rockies (~100 million years old) revealing tridactyl pes prints (~30 cm long) paired with five-digited manus impressions in a narrow-gauge, bird-like quadrupedal pattern indicative of lumbering, coordinated steps among multiple individuals.^[21] Additional 2025 reports from South Korea's Late Cretaceous Jindong Formation (Cenomanian, ~95 million years old) describe a wide-gauge trackway with 35 manus-pes sets, showing differential pes depth and blunt digits that confirm a slow, stable gait with pressure concentrated medially, consistent with the inferred posture and low-speed movement of armored thyreophorans.

Senses and Physiology

Ankylosaurids, including Ankylosaurus, possessed relatively small brains compared to their body size, with endocasts indicating volumes on the order of 80–100 cm³ in large specimens, roughly equivalent to the size of a walnut.^[22] These brains featured expanded olfactory bulbs, which diverged widely at angles of 80–100° and occupied a significant portion of the cranial cavity, suggesting a well-developed sense of smell for detecting food or predators.^[10] In contrast, the size and positioning of the orbits—nearly round to slightly oval and directed somewhat forward due to the tapering skull—imply limited visual acuity and a narrower field of view, consistent with a reliance on other senses in forested or low-light environments.^[23] CT scans of ankylosaurid skulls from the 2010s revealed extensive paranasal air sinuses and highly convoluted nasal passages, which lacked the typical ossified sinuses of theropods but formed looping airways up to 790 mm long in species like Euoplocephalus tutus.^[23] These structures likely served multiple functions, including reducing skull weight for better head support during quadrupedal locomotion and potentially aiding vocalization through resonance, though direct evidence for the latter remains speculative.^[24] Computational fluid dynamics analyses of these airways demonstrated high efficiency in heat exchange, recovering 65–84% of thermal energy from exhaled air to precondition inspired air, thereby supporting thermoregulation in variable climates.^[24] Bone histology in ankylosaurids reveals a fibrolamellar matrix with predominantly parallel-fibered tissue and poor vascularization, indicative of slower growth rates than in theropods or ornithopods and suggesting a metabolic physiology intermediate between ectothermy and full endothermy—possibly an inertial homeotherm maintaining stable body temperature through mass alone.^[25] Extensive remodeling, including multiple generations of secondary osteons, points to low metabolic demands, with cyclical growth lines in some long bones implying seasonal pauses consistent with ectothermic influences.^[25] The respiratory system of Ankylosaurus likely incorporated air sacs, inferred from pneumatic features in the vertebrae and extensive cranial airspaces that connected to the nasal passages, enhancing ventilatory efficiency similar to that in other ornithischians. These adaptations would have lightened the torso and improved gas exchange, though pneumatization was less pervasive than in saurischians. Recent analyses of cranial airspaces in ankylosaurids highlight their role in supporting balance, with complex inner ear structures like the vestibular apparatus potentially aiding postural stability during slow, deliberate movements.^[26]

Feeding and Diet

Ankylosaurus was an obligate herbivore that primarily browsed low-lying vegetation, including ferns, cycads, and horsetails, using its broad beak to crop plants close to the ground.^[27] Its dentition consisted of small, leaf-shaped cheek teeth arranged in a single row, adapted for shearing and initial breakdown of tough, fibrous plant matter rather than grinding large volumes.^[26] This feeding strategy aligned with its low browsing height, typically under 1 meter, allowing access to herbaceous understory plants in its Late Cretaceous habitat.^[27] The jaw mechanics of Ankylosaurus featured a low mechanical advantage, as detailed in a 2023 study by Ballell et al., with both anterior and posterior mechanical advantages lower than those of contemporaneous nodosaurids.^[26] This configuration, combined with a biphasal power stroke involving orthal and palinal movements, enabled precise tooth occlusion for processing vegetation, but resulted in relatively weak bite forces estimated at 166–444 N.^[26] Such adaptations suggest specialization in softer or less fibrous plants, contrasting with the higher bite forces and mechanical advantages of nodosaurids, which likely tackled tougher foliage and contributed to dietary niche partitioning among ankylosaurs.^[26] The asymmetrical arrangement of its dentition further facilitated grinding motions during mastication.^[26] Although primarily herbivorous, some hypotheses propose possible omnivory in Ankylosaurus, with incidental insect consumption inferred from gut content analogs in related ankylosaurids, though direct evidence remains unconfirmed.^[28] Foraging behavior likely involved slow, selective feeding, where the dinosaur methodically cropped and processed vegetation in forested floodplain settings to meet its nutritional needs.^[27]

Growth and Ontogeny

Ankylosaurus exhibited a growth pattern typical of large ornithischian dinosaurs, characterized by rapid early ontogeny followed by a marked slowdown toward maturity. Fossil evidence from related ankylosaurids, including bone histology analyses, indicates that juveniles grew quickly, potentially reaching lengths of approximately 4 meters by around age 10 through fibrolamellar bone deposition with high vascularization, before transitioning to slower parallel-fibered bone formation in subadulthood.^[25] This early acceleration likely supported the development of body armor and defensive structures, with overall body size scaling proportionally with age as evidenced by limb bone proportions in preserved specimens.^[8] Ontogenetic changes were particularly evident in the tail club and dermal armor. Juvenile Ankylosaurus and closely related ankylosaurids possessed smaller, less developed tail clubs, often with reduced knob sizes under 200 mm in diameter and limited ossification, rendering them ineffective for bone-breaking impacts compared to the robust, enlarged clubs of adults exceeding 500 mm.^[19] Bone histology reveals that armor osteoderms underwent progressive fusion and remodeling with age, starting with loosely integrated plates in juveniles and increasing density through cancellous bone infilling and secondary osteons by maturity, as cyclical growth marks (lines of arrested growth, or LAGs) accumulate in the cortex.^[29] No hatchling fossils of Ankylosaurus have been identified, in contrast to the 2025 discovery of a hatchling Liaoningosaurus paradoxus from the Lower Cretaceous of China, dated to approximately 115 million years ago, which represents the only confirmed ankylosaur hatchling known to date.^[30] Longevity estimates for Ankylosaurus derive from growth ring analyses in ankylosaurid long bones and osteoderms, suggesting lifespans of 20–30 years, with maturity attained between 15 and 20 years as growth rates decline and an external fundamental system forms in the outermost bone layers.^[25] Evidence for sexual dimorphism remains unconfirmed, as protracted growth trajectories in non-avian dinosaurs complicate differentiation of size variation due to sex versus individual growth rates.^[31] A 2017 study on Ankylosaurus specimens highlighted significant size variation among individuals, from estimated lengths of 6 meters in smaller examples to nearly 10 meters in larger ones, attributable to differences in growth rates rather than taxonomic distinctions.^[8]

Defense

The primary defensive adaptation of Ankylosaurus was its tail club, a specialized structure formed by fused caudal vertebrae creating a rigid "handle" and aggregated osteoderms forming a bulbous knob at the distal end. This weapon enabled powerful lateral swings targeted at predators such as Tyrannosaurus rex, potentially striking the legs or body to inflict debilitating injuries. Biomechanical models indicate that large tail clubs in ankylosaurids, including Ankylosaurus, could generate impact forces exceeding 14,000 newtons—equivalent to approximately 1.4 tons—sufficient to fracture bone upon contact.^[19] Complementing the tail club, the extensive dermal armor of Ankylosaurus provided passive protection against predatory attacks. The dorsal surface was covered in thick, interlocking osteoderms, particularly dense over the thoracic region, which deterred deep bites by distributing force and resisting penetration. Anteriorly, large half-rings of fused osteoderms shielded the neck and shoulders, vulnerable areas during close encounters with large theropods. Inferred behaviors suggest Ankylosaurus relied on stationary clubbing or limited charging motions to deploy its defenses, given its low-slung posture and restricted mobility.^[32] Fossil evidence includes healed pathologies on osteoderms and tail clubs in related ankylosaurids, indicating successful repulsion of attacks, though direct predation traces on Ankylosaurus specimens are rare.^[33] Recent finite element analyses confirm the structural integrity of tail clubs under high-impact loads, with peak stresses below failure thresholds for bone, supporting their role in predator deterrence.^[32]

Paleoenvironment

Geological Context

Ankylosaurus fossils are primarily known from the Hell Creek Formation in Montana and South Dakota, USA; the Lance Formation in Wyoming, USA; the Scollard Formation in Alberta, Canada; and the Frenchman Formation in Saskatchewan, Canada, all dating to the Maastrichtian stage of the Late Cretaceous, approximately 68 to 66 million years ago.^[6]^[34] These formations represent the final 1.5 to 1.9 million years of the Cretaceous Period, immediately preceding the Cretaceous-Paleogene (K-Pg) extinction event at 66 Ma.^[34]^[35] The paleoenvironment of these deposits consisted of coastal floodplains characterized by meandering rivers, swamps, marshes, estuaries, and lowland forests.^[34] The region experienced a subtropical climate that was warm and humid, supporting lush vegetation and diverse ecosystems along the western margin of the Western Interior Seaway.^[36]^[37] Sedimentologically, the formations comprise fluvial deposits including sandstones, mudstones, siltstones, shales, and minor coal seams, indicative of seasonal flooding and overbank deposition.^[34] Volcanic ash layers, preserved as bentonites, occur intermittently and provide key radiometric dating points through their radioactive minerals.^[37]^[35] Fossils of Ankylosaurus are restricted to western North America, specifically the landmass known as Laramidia, with no known occurrences outside this continent.^[6]^[38] This distribution reflects the isolation of Laramidia by the Western Interior Seaway during the Late Cretaceous.^[38]

Associated Fauna and Flora

Ankylosaurus inhabited the Maastrichtian-age Hell Creek Formation of western North America, sharing its ecosystem with a diverse assemblage of vertebrates in a high-biodiversity setting dominated by large herbivores and apex predators. The primary predators included the large theropod Tyrannosaurus rex, which comprised about 24% of dinosaur specimens, and smaller carnivores such as Troodon formosus, known primarily from isolated teeth in microvertebrate sites. Among potential competitors for resources, the hadrosaur Edmontosaurus annectens (20% of specimens) and ceratopsian Triceratops horridus (40% of specimens) were the most abundant herbivores, likely occupying overlapping niches in floodplain habitats. Other herbivores included the nodosaurid Denversaurus schlessmani, a close relative that coexisted with Ankylosaurus as part of the rare armored dinosaur component (collectively ~1% of fauna), alongside smaller ornithischians like Thescelosaurus neglectus.^[39] The community also featured small mammals, such as multituberculates, and birds, including enantiornithines, contributing to the overall vertebrate diversity. The flora of the Hell Creek Formation was predominantly angiosperm-dominated, forming a multi-tiered structure in a warm, moist coastal plain environment with meandering rivers.^[40] Flowering plants, including palms, magnolias, and early fruits, were widespread, supported by fossil evidence of leaves, seeds, and pollen that indicate a lush, tropical understory with herbaceous diversity.^[37] Conifers, such as araucarians and cypresses, were common in upland and riparian zones, while ferns and allies occupied shaded or disturbed ground, as revealed by palynological records showing a mix of evergreen conifers and deciduous angiosperms.^[40]^[37] Pollen assemblages further highlight a diverse understory of ferns, mosses, and low-growing angiosperms, reflecting seasonal flooding that preserved these elements in fine sediments.^[37] Within this ecosystem, Ankylosaurus functioned as a low-browser, targeting vegetation below 1 meter in height, such as ferns and low angiosperms, thereby filling a mid-trophic niche as a primary consumer amid taller herbivores.^[41] Its rarity—accounting for less than 0.1% of the fauna—suggests a specialized role, potentially involving interactions like trampling understory plants during movement in forested floodplains.^[37] This positioning allowed coexistence with abundant competitors by exploiting ground-level resources in a vertically stratified plant community.^[41]

Cultural Impact

In Popular Media

Ankylosaurus first gained public attention through museum dioramas in the early 20th century, where it was often reconstructed as a heavily armored, low-slung quadruped amid Cretaceous landscapes, emphasizing its bony plates and tail club as defensive features.^[42] These displays, such as those at major natural history museums, portrayed it alongside other armored dinosaurs to highlight its role as a "living tank" in prehistoric ecosystems.^[43] In the 1940s, Ankylosaurus appeared in Disney's animated film Fantasia (1940), specifically in the "Rite of Spring" sequence, where it is depicted as part of a herd of armored herbivores, swinging its tail club in defense against predators like Tyrannosaurus rex.^[44] This portrayal, influenced by contemporary paleontological illustrations, showed it with exaggerated armor and a sluggish gait, contributing to its image as an indomitable defender in early popular media.^[45] The Jurassic Park franchise, beginning with Jurassic Park (1993), has featured Ankylosaurus as a formidable, tank-like herbivore, protected by osteoderm-covered armor and capable of using its tail club to fend off threats like Tyrannosaurus or hybrid dinosaurs.^[46] In films such as Jurassic Park III (2001), Jurassic World: Fallen Kingdom (2018), and Jurassic World Dominion (2022), it serves as a symbol of armored resilience, often in defensive confrontations or herd scenes that underscore its role as a protector among herbivores.^[47]^[48] Similarly, the 1999 BBC series Walking with Dinosaurs depicted Ankylosaurus in its finale episode "Death of a Dynasty," portraying it as a heavily plated, low-slung grazer that employs its tail club effectively against a Tyrannosaurus, blending dramatic encounters with then-current scientific understanding of its physiology.^[49] In video games, Ankylosaurus is prominently featured in ARK: Survival Evolved (2015), where players can tame it as a versatile mount prized for its powerful tail swing that excels at harvesting metal and other resources from rock formations.^[50] Earlier titles, such as those from the 1990s and early 2000s, frequently inaccurately depicted it with an upright, bipedal or semi-erect posture, diverging from its quadrupedal, sprawling build.^[51] Since the 1950s, Ankylosaurus has been a staple in children's dinosaur encyclopedias and illustrated books, often highlighted for its distinctive armor and clubbed tail as iconic traits of Cretaceous herbivores.^[52] Complementary toys from this era onward, including plastic figures and playsets, emphasize its bony armor through molded osteoderms and reinforced plating, positioning it as a durable, battle-ready plaything.^[53]

In Scientific and Educational Contexts

In educational contexts, Ankylosaurus serves as an iconic example of dinosaur diversity and adaptation, featured prominently in museum exhibits and curricula to illustrate Cretaceous ecosystems and extinction events. The Natural History Museum in London displays Ankylosaurus models and fossils in its Dinosaurs Gallery, using them to engage visitors with concepts of herbivory, armor evolution, and predator-prey dynamics, supported by interactive resources like activity kits.^[1] Similarly, the Canadian Museum of Nature incorporates Ankylosaurus specimens in paleontology programs, emphasizing its role in understanding the Maastrichtian stage of the Late Cretaceous, with educational materials linking it to broader themes of biodiversity in western North America.^[54] These institutions, along with sites like the San Diego Natural History Museum, utilize Ankylosaurus to teach about thyreophoran phylogeny and fossil preservation biases, fostering public appreciation for paleontological methods through guided tours and digital content.^[55]

References

[1]
Ankylosaurus | Natural History Museum
This armoured dinosaur had bony plates covering its body and a heavy club at the end of its tail. Though it's the most famous member of the ankylosaur group, ...
[2]
Ankylosaurid dinosaur tail clubs evolved through stepwise ...
Aug 31, 2015 · Ankylosaurid ankylosaurs were quadrupedal, herbivorous dinosaurs with abundant dermal ossifications. They are best known for their distinctive ...
[3]
Ankylosaurs may have used their tail clubs to fight each other, not ...
Dec 7, 2022 · While stegosaurs often had a spiked structure on the end of their tail, ankylosaurs instead had a hard, bony structure known as a tail club.
[4]
Types of Dinosaurs | American Museum of Natural History
Somewhat reminiscent of mammalian armadillos and their relatives the glyptodonts, Ankylosaurus was one of the armored reptilian tanks of the Cretaceous.
[5]
Brown Expedition to the Cretaceous of Montana (1902-1910)
Most notably, Brown discovered portions of vertebrae found together which Henry Fairfield Osborn later described as the "type of the new genus Tyrannosaurus ...
[6]
Redescription of Ankylosaurus magniventris Brown 1908 ...
Redescription of Ankylosaurus magniventris Brown 1908 (Ankylosauridae) from the Upper Cretaceous of the Western Interior of North America ... PDF. View PDF ...
[7]
The Ankylosauridae, a new family of armored dinosaurs from the ...
The Ankylosauridae, a new family of armored dinosaurs from the Upper Cretaceous. Bulletin of the AMNH ; v. 24, article 12.
[8]
Unusual cranial and postcranial anatomy in the archetypal ...
Oct 12, 2017 · The iconic Maastrichtian dinosaur Ankylosaurus magniventris was the last and largest of the ankylosaurid dinosaurs. A member of the tail-clubbed ...
[9]
Osteology and Myology of the Hindlimb in the Ankylosauria ... - jstor
fused to the ilium. The widely reproduced reconstruction of. Ankylosaurus originally published by Brown. (1908, figs. 19 and 20) as well as ...
[10]
The internal cranial morphology of an armoured dinosaur ...
Sep 29, 2011 · This paper presents corroborating evidence from direct osteological observation and from reconstruction based on computed X-ray tomography (CT) on the internal ...
[11]
An exceptionally preserved armored dinosaur reveals the ...
Nov 29, 2017 · Thickness and relative contribution of the keratinized epiosteodermal scales/sheaths varies greatly by region, ranging from 2% to 6% for ...
[12]
(PDF) Redescription of Ankylosaurus magniventris Brown 1908 ...
Aug 10, 2025 · The armor-plated dinosaur Ankylosaurus magniventris is redescribed based on specimens from the Hell Creek Formation of northeastern Montana, USA.
[13]
https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0068717
[14]
https://pmc.ncbi.nlm.nih.gov/articles/PMC6063217/
[15]
A new southern Laramidian ankylosaurid, Akainacephalus johnsoni ...
Jul 19, 2018 · Cervical half rings. A partial anterior cervical half ring and a nearly complete posteriorcervical half ring is preserved (Figs. 25A–25F) ...
[16]
50% majority rule tree of Ankylosauria, highlighting North American...
After becoming globally distributed in the Early–Middle Jurassic (200–160 Ma), dinosaurs experienced vicariance driven by Pangaean fragmentation. Regional ...
[17]
Paleodiversity of Late Cretaceous Ankylosauria from Mexico and ...
Jun 6, 2018 · Ankylosaurs are characteristic and abundant components of dinosaur assemblages from the Late Cretaceous of western North America (Laramidia).
[18]
(PDF) Ankylosaurs - ResearchGate
Aug 4, 2017 · The armored bodies of these obligatory quadrupeds are barrel shaped (Carpenter 2012; Vickaryous et al. 2004). Ankylosauria are divided into two ...
[19]
Speeds and gaits of dinosaurs - ScienceDirect.com
Most quadrupedal dinosaurs seem to have been restricted to a walking gait. Stegosaurs and ankylosaurs may have had maximum speeds as low as 6–8 km/h. Sauropods ...
[20]
Estimating Impact Forces of Tail Club Strikes by Ankylosaurid ...
CT scans showed that knob osteoderms were predominantly cancellous, which would have lowered the rotational inertia of the tail club and made it easier to ...<|separator|>
[21]
Palaeopathological evidence for intraspecific combat in ... - Journals
Dec 7, 2022 · victoria.arbour@gmail.com. Contribution: Conceptualization, Data ... We found pathological osteoderms (armour plates) in the holotype ...<|control11|><|separator|>
[22]
A new thyreophoran ichnotaxon from British Columbia, Canada confirms the presence of ankylosaurid dinosaurs in the mid Cretaceous of North America
### Key Findings on Ankylosaurid Trackways, Gait, Posture, and Locomotion
[23]
To Think Like a Dinosaur: Paleontologists Created the Most Detailed ...
Jun 5, 2020 · A 3D reconstruction of the endocast of its brain cavity was made using CT. It is important to understand that the digital 'cast' of the ...Missing: Ankylosaurus scan
[24]
The Paranasal Air Sinuses of Predatory and Armored Dinosaurs ...
Oct 24, 2008 · The paranasal air sinuses and nasal cavities were studied along with other cephalic spaces (brain cavity, paratympanic sinuses) in certain dinosaurs.
[25]
Convoluted nasal passages function as efficient heat exchangers in ...
Dec 19, 2018 · Previous research suggested that these convoluted nasal passages functioned as heat exchangers analogous to the respiratory turbinates of mammals and birds.
[26]
Long Bone Histology and Growth Patterns in Ankylosaurs - NIH
Jul 24, 2013 · Ankylosaur long bone histology is characterized by a fibrolamellar bone architecture. The bone matrix type in ankylosaurs is closest to that of Stegosaurus.<|control11|><|separator|>
[27]
Divergent strategies in cranial biomechanics and feeding ecology of ...
Oct 25, 2023 · Three-dimensional digital models of the skulls and mandibles of the ankylosaurid Euoplocephalus tutus (AMNH 5405) and the nodosaurid ...
[28]
Feeding height stratification among the herbivorous dinosaurs from ...
Apr 4, 2013 · Most herbivorous dinosaur species from the Dinosaur Park Formation were restricted to feeding no higher than approximately 1 m above the ground.
[29]
Ankylosaurs Slurped Food With Powerful Tongues
Jul 27, 2015 · The little leaf-shaped teeth in ankylosaurian mouths accorded better with an herbivorous diet, and a rare find of ankylosaur gut contents ...
[30]
(PDF) Histology of ankylosaur osteoderms: Implications for ...
Aug 10, 2025 · Ankylosaurids have very thin osteoderms that were greatly strengthened by the incorporation of structural collagen fiber bundles, similar but ...
[31]
Bone histology of Liaoningosaurus paradoxus (Ornithischia: Ankylosauria) from the Lower Cretaceous of Liaoning Province, China
### Summary of Bone Histology of Liaoningosaurus paradoxus
[32]
Protracted growth impedes the detection of sexual dimorphism in ...
Aug 6, 2025 · Hone & Mallon (2017) assessed detection of sexual dimorphism in alligators based on body size, and suggested a minimum of 60 animals might be ...Missing: Ankylosaurus | Show results with:Ankylosaurus
[33]
Finite Element Analyses of Ankylosaurid Dinosaur Tail Club Impacts
Aug 26, 2009 · Four ankylosaurid tail clubs (Tables 1 and 2) were scanned using computed tomography (CT), to derive three-dimensional models for use in FEA ( ...
[34]
Palaeopathological evidence for intraspecific combat in ... - NIH
Dec 7, 2022 · Ankylosaurid dinosaurs were heavily armoured herbivores with tails modified into club-like weapons. These tail clubs have widely been considered ...Missing: Ankylosaurus | Show results with:Ankylosaurus
[35]
[PDF] A Census of Dinosaur Fossils Recovered From the Hell Creek and ...
A census of Hell Creek and Lance Formation dinosaur remains was conducted from April, 2017 through February of 2018. Online databases were reviewed and ...
[36]
First high-precision U–Pb CA–ID–TIMS age for the Battle Formation ...
Jan 25, 2019 · Accordingly, a minimum age of ∼67.20 Ma is assessed for the base of the Hell Creek Formation in its type area. La Formation de Battle (BFm) est ...
[37]
[PDF] The Hell Creek Formation - Montana Department of Transportation
through the coastal plain in a warm and humid climate, depositing sediment which would later become known as the Hell Creek Formation. The tan sandstones ...Missing: paleoclimate | Show results with:paleoclimate
[38]
[PDF] Last of the Dinosaurs: Hell Creek Rocks - USDA Forest Service
Mar 5, 2024 · The climate is warm and humid. The vegetation is tropical and lush. Suddenly and without warning the bright flash of an immense meteor ...
[39]
Distribution of Late Cretaceous Laramidian ankylosaurid...
No ankylosaurids are known from North America prior to this, and Arbour et al. (2016) noted that after Cedarpelta, no definitive ankylosaurid body fossils are ...
[40]
latest cretaceous occurrence of nodosaurid ankylosaurs - jstor
ABSTRACT-The presence of nodosaurid ankylosaurs in the Lance, Hell Creek, and Laramie for- mations of western North America is confirmed, thereby extending ...<|separator|>
[41]
https://doi.org/10.1186/1472-6785-13-14
[42]
https://pseudoplocephalus.com/2011/01/26/ankylosaurus-through-the-ages/
[43]
Ankylosaurus through the Ages | pseudoplocephalus
Jan 26, 2011 · Brown 1908. The Ankylosauridae, a new family of armored dinosaurs from the upper Cretaceous. ... Barnum Brown's 1908 description included ...
[44]
Dinosaurs, dioramas, and the strange world of natural history
Sep 7, 2017 · By the turn of the 20th century, 'modern' dioramas which combine painted backdrops, models and taxidermy, were beginning to take hold. Among ...
[45]
Fantasia [1940] - Ankylosaurus Screen Time - YouTube
Nov 30, 2021 · Fantasia [1940] - Ankylosaurus Screen Time. 14K views · 3 years ... "Hunted by Moonlight" | Dinosauria Series | Animated Short Film (2025).
[46]
Bringing Fantasia's Dinosaurs To Life | - Cartoon Research
Sep 20, 2022 · Walt Disney's fascination with making dinosaurs “real” goes back several decades to the animated feature Fantasia in 1940. Disney transformed ...Missing: Ankylosaurus | Show results with:Ankylosaurus
[47]
Ankylosaurus - Dinosaurs - Jurassic World Evolution 3
Famed for its clubbed tail and the armour plates that cover its back, Ankylosaurus is a genus of Ankylosaurus that lived around 70m years ago during the Late Cr ...
[48]
The History of the Ankylosaurus in the Jurassic Park Franchise
Oct 20, 2018 · To quote Fallen Kingdom: "This is a herbivorous quadruped. Late Cretaceous. This is one of the largest armored dinosaurs. Known by ...
[49]
TRILOGY OF LIFE - Walking with Dinosaurs - "Ankylosaurus"
Aug 11, 2018 · All content in this video is the property of BBC worldwide! I'm simply using it under fair use! Music composed by Benjamin Bartlett.
[50]
Ankylosaurus - ARK Official Community Wiki
Oct 4, 2025 · It is identical to the normal Ankylosaurus except for its glowing robe. It is not Radiation immune in ARK: Survival Evolved, while such passive ...Basic Info · Abilities · Taming · CombatMissing: video | Show results with:video
[51]
Ankylosaurus (Jurassic vs Accurate) - YouTube
Jun 6, 2023 · Taking a look at the armored tank of the Jurassic Franchise, and seeing how it holds up in terms of accuracy! Thumbnail art: (Left) Jurassic ...
[52]
Vintage Dinosaur Art: Rourke's Ankylosaurus
May 7, 2012 · Ankylosaurus herself is pretty respectable for an 80's reconstruction; Bernard Long avoids the old neckless sluggard look of mid-century depictions.
[53]
https://www.rainbowresource.com/016311.html
[54]
FIRST IMAGES For The NEW Walking With Dinosaurs 2025!
Jan 22, 2025 · Share your videos with friends, family, and the world.
[55]
Ankylosaurus | The Canadian Encyclopedia
Jan 31, 2025 · Ankylosaurus was a large, armored, herbivorous dinosaur, one of the last non-avian dinosaurs, reaching up to 8m long and weighing up to 8 ...Missing: CMN 8880
[56]
https://doi.org/10.1371/journal.pone.0068590
[57]
The Nat | Ankylosaur - San Diego Natural History Museum
Ankylosaurs were sturdy, armored dinosaurs, distantly related to the stegosaurs, with their large bony plates and tail spikes. Much of the body of ankylosaurs ...