Ankylosaurus is a genus of armored dinosaur. Its fossils have been found in geological formations dating to the very end of the Cretaceous Period, about 70-66 million years ago, in western North America, making it among the last of the non-avian dinosaurs. It was named by Barnum Brown in 1908; it is monotypic, containing only A. magniventris. The 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 archetypal member of its group, despite having some unusual features.

Possibly the largest known ankylosaurid, Ankylosaurus is estimated to have been between long and to have weighed between . It was 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 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 browser. 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 combat. Specimens of Ankylosaurus have been found in the Hell Creek, Lance, Scollard, Frenchman, and Ferris formations, but it appears to have been rare in its environment. Although it lived alongside Denversaurus, a nodosaurid ankylosaur, their ranges and ecological niches 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

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 paleontologist Barnum Brown discovered the 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 described the animal in 1908; the generic name is derived from the Greek words ' ('bent' or 'crooked'), referring to the medical term ankylosis, the stiffness produced by the fusion of bones in the skull and body, and ' ('lizard'). The name can be translated as "fused lizard", "stiff lizard", or "curved lizard". The type species name, magniventris, is derived from the ('great') and ('belly'), referring to the great width of the animal's body. 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 was a better example of how ankylosaurs would have appeared in life.

In 2017, based on comparisons with more complete ankylosaurines, Arbour and Mallon estimated a length of for CMN 8880, and 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 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 for large Ankylosaurus too long, and suggested a length of instead. Arbour and Mallon estimated a weight of for AMNH 5214, and tentatively estimated the weight of CMN 8880 at .

Skull

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 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 premaxillae. The orbits (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 horns (their shape has been described as "pyramidal"), 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 horns were present, which pointed backward and down. The horns may have originally been osteoderms that fused to the skull. The scale-like cranial ornamentation on the surfaces of ankylosaurs skulls is called "", and were the result of remodeling of the skull itself. This obliterated the 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.

Classification

Brown considered Ankylosaurus so distinct that he made it the type genus of a new 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 suborder Stegosauria.

The retracted position of the nostrils of Ankylosaurus were compared to those of fossorial (digging) worm lizards and 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 omnivorous (eating both plant and animal matter). It may also (or alternatively) have dug in the ground for roots and tubers.

Airspaces and senses

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 resonance.

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.

See also

  • Timeline of ankylosaur research

References

Notes

Citations