Ankylosaurus
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 68–66 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
In 1906, an American Museum of Natural History expedition led by American paleontologist Barnum Brown discovered the type specimen of Ankylosaurus magniventris in the Hell Creek Formation, near Gilbert Creek, Montana. The specimen 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. Brown scientifically described the animal in 1908; the generic name is derived from the Greek words αγκυλος ankulos, referring to the medical term ankylosis, the stiffness produced by the fusion of bones in the skull and body, and σαυρος sauros. The name can be translated as "fused lizard", "stiff lizard", or "curved lizard". The type species name, magniventris, is derived from the and , referring to the great width of the animal's body.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. 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 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. 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. The claim of synonymy was not accepted by other researchers, and the two genera are now considered distinct.
Brown had collected 77 osteoderms while excavating a Tyrannosaurus specimen in the Lance Formation of Wyoming in 1900. He mentioned these osteoderms 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 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.
In 1910, another AMNH expedition led by Brown discovered an Ankylosaurus specimen 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 collectors Charles M. Sternberg and T. Potter Chamney collected a skull and mandible, 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 was discovered in the 1960s in the 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.
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 within the nasal chambers. 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.
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. 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. 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.
Description
Ankylosaurus was the largest-known ankylosaurine dinosaur and possibly the largest ankylosaurid. In 2004 Carpenter estimated that the individual with the largest-known skull, which is long and wide, was about long and had a hip height of about. The smallest-known skull is long and wide, and Carpenter estimated that it measured about long and about tall at the hips. The English paleontologist Roger B. J. Benson and colleagues estimated the weight for AMNH 5214 at in 2014.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
The three known Ankylosaurus skulls differ in various details; this is thought to be the result of taphonomy 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 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, 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 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.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 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 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 choanae, and the air passage was looped. The maxillae expanded to the sides, giving the impression of a bulge, which may have been due to the sinuses inside. The maxillae had a ridge that may have been the attachment site for fleshy cheeks; the presence of cheeks in ornithischians is controversial, but some nodosaurs had armor plates that covered the cheek region, which may have been embedded in the flesh.
Specimen AMNH 5214 has 34–35 dental alveoli in the maxilla. The tooth rows in the maxillae of this specimen are about long. Each alveolus had a foramen near its side where a 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 and are about long. The incomplete mandible of the largest specimen 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. Like other ankylosaurs, Ankylosaurus had small, phylliform teeth, which were compressed sideways. The teeth were mostly taller than they were wide, and were very small; their size in proportion to the skull meant that the jaws of 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 tooth crowns were usually flatter on one side than the other. Ankylosaurus teeth are diagnostic and can be distinguished from the teeth of other ankylosaurids based on their smooth sides. The denticles were large, their number ranging from six to eight on the front part of the tooth, and five to seven behind.