Ankylosaurus

Taxobox
name = "Ankylosaurus"
fossil_range = Late Cretaceous

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image_width = 250px
image_caption = An "Ankylosaurus" skull, profile view. This specimen is from the Edmonton Formation of Alberta.
regnum = Animalia
phylum = Chordata
classis = Sauropsida
superordo = Dinosauria
ordo = †Ornithischia
subordo = †Thyreophora
infraordo = †Ankylosauria
familia = †Ankylosauridae
genus = †"Ankylosaurus"
genus_authority = Brown, 1908
subdivision_ranks=Species
subdivision=
*†"A. magniventris" Brown, 1908 (type)

"Ankylosaurus" (pronEng|ˌæŋkɪloʊˈsɔrəs, which means 'rigid lizard') is a genus of ankylosaurid dinosaur, containing one species, "A. magniventris". Fossils of "Ankylosaurus" are found in geologic formations dating to the very end of the Cretaceous Period in western North America.

Although a complete skeleton has not been discovered and several other dinosaurs are represented by more extensive fossil material, "Ankylosaurus" is often considered the archetypal armored dinosaur. Other ankylosaurids shared its well-known features, like the heavily-armored body and massive bony tail club, but "Ankylosaurus" was the largest member of its family.

Paleobiology

In comparison with modern land animals the adult "Ankylosaurus" was very large. Some scientists have estimated a length of 9 meters (30 ft).Vickaryous, M.K., Maryanska, T., & Weishampel, D.B. 2004. Ankylosauria. In: Weishampel, D.B., Dodson, P., & Osmólska, H. (Eds.). "The Dinosauria" (2nd edition). Berkeley: University of California Press. Pp. 363-392.] Another reconstruction suggests a significantly smaller size, at 6.25 m (20.5 ft) long, up to 1.5 m (5 ft) wide and about 1.7 m (5.5 ft) high at the hip. The body shape was low-slung and quite wide. "Ankylosaurus" was quadrupedal, with the hind limbs longer than the forelimbs. Although its feet are still unknown to science, comparisons with other ankylosaurids suggest "Ankylosaurus" probably had five toes on each foot. The skull was low and triangular in shape, wider than it was long. The largest known skull measures 64.5 centimeters (25 in) long and 74.5 cm (29 in) wide. Like other ankylosaurs, "Ankylosaurus" was herbivorous, with small, leaf-shaped teeth suitable for cropping vegetation. "Ankylosaurus" did not share the grinding tooth batteries of the contemporaneous ceratopsid and hadrosaurid dinosaurs, indicating that very little chewing occurred. Bones in the skull and other parts of the body were fused to increase their strength.Carpenter, K. 2004. Redescription of "Ankylosaurus magniventris" Brown 1908 (Ankylosauridae) from the Upper Cretaceous of the Western Interior of North America. "Canadian Journal of Earth Sciences" 41: 961–986.]

Armor

The most obvious feature of "Ankylosaurus" is its armor, consisting of massive knobs and plates of bone, known as osteoderms, embedded in the skin. Osteoderms are also found in the skin of crocodiles, armadillos and some lizards. The bone was probably overlain by a tough, horny layer of keratin. These osteoderms ranged greatly in size, from wide, flat plates to small, round nodules. The plates were aligned in regular horizontal rows down the animal's neck, back, and hips, with the many smaller nodules protecting the areas between the large plates. Smaller plates may have been arranged on the limbs and tail. Compared to the slightly more ancient ankylosaurid "Euoplocephalus", the plates of "Ankylosaurus" were smooth in texture, without the high keels found on the armor of the contemporaneous nodosaurid "Edmontonia". A row of flat, triangular spikes may have protruded laterally along each side of the tail. Tough, rounded scales protected the top of the skull, while four large pyramidal horns projected outwards from its rear corners.

Tail club

The famous tail club of "Ankylosaurus" was also composed of several large osteoderms, which were fused to the last few tail vertebrae. It was heavy and supported by the last seven tail vertebrae, which interlocked to form a stiff rod at the base of the club. Thick tendons have been preserved, which attached to these vertebrae. These tendons were partially ossified (or bony) and were not very elastic, allowing great force to be transmitted to the end of the tail when it was swung. It seems to have been an active defensive weapon, capable of producing enough of a devastating impact to break the bones of an assailant. It has also been proposed that the tail club acted as a decoy for the head, although this idea is now largely discredited.Thulborn, T. 1993. Mimicry in ankylosaurid dinosaurs. "Record of the South Australian Museum" 27: 151–158.]

Environment

"Ankylosaurus magniventris" existed between 68 to 65.5 million years ago, in the final Maastrichtian stage of the Late Cretaceous Period, and was one of the dinosaurs to survive until the Cretaceous-Tertiary extinction event. The type specimen is from the Hell Creek Formation of Montana, while other specimens have been found in the Lance Formation of Wyoming and the Scollard Formation in Alberta, Canada, all of which date to the end of the Cretaceous.Vickaryous, M.K., Maryanska, T., & Weishampel, D.B. 2004. Ankylosauria. In: Weishampel, D.B., Dodson, P., & Osmólska, H. (Eds.). "The Dinosauria" (2nd edition). Berkeley: University of California Press. Pp. 363-392.]

The Lance, Hell Creek and Scollard Formations represent different sections of the western shore of the shallow sea that divided western and eastern North America during the Cretaceous. They represent 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.Lofgren, D.F. 1997. Hell Creek Formation. In: Currie, P.J. & Padian, K. (Eds.). "The Encyclopedia of Dinosaurs". San Diego: Academic Press. Pp. 302-303.] Breithaupt, B.H. 1997. Lance Formation. In: Currie, P.J. & Padian, K. (Eds.). "The Encyclopedia of Dinosaurs". San Diego: Academic Press. Pp. 394-395.] Eberth, D.A. 1997. Edmonton Group. In: Currie, P.J. & Padian, K. (Eds.). "The Encyclopedia of Dinosaurs". San Diego: Academic Press. Pp. 199-204.] The Hell Creek is the best studied of these ancient environments. At the time, this region was subtropical, with a warm and humid climate. Many plant species were supported, primarily angiosperms, with less common conifers, ferns and cycads. An abundance of fossil leaves found at dozens of different sites indicates that the area was largely forested by small trees.Johnson, K.R. 1997. Hell Creek Flora. In: Currie, P.J. & Padian, K. (Eds.). "The Encyclopedia of Dinosaurs". San Diego: Academic Press. Pp. 300-302.]

Fossils of "Ankylosaurus" are considerably rare in these sediments, compared to "Edmontosaurus" and the super-abundant "Triceratops", which make up most of the large herbivore fauna. Another ankylosaur, "Edmontonia", is also found in the same formations. However, "Ankylosaurus" and "Edmontonia" seem to have been separated both geographically and ecologically. "Ankylosaurus" had a wide muzzle, perhaps used for non-selective grazing and may have been limited to the upland regions, away from the coast, while "Edmontonia" had a narrower muzzle, indicating a more selective diet, and seems to have lived at lower elevations, closer to the coast.

Classification

"Ankylosaurus" was named as the type genus of the family Ankylosauridae.Brown, B. 1908. The Ankylosauridae, a new family of armored dinosaurs from the Upper Cretaceous. "Bulletin of the American Museum of Natural History" 24: 187–201.] Ankylosaurids are members of the larger taxon Ankylosauria, which also contains the nodosaurids. Ankylosaur phylogeny is a contentious topic, with several mutually exclusive analyses presented in recent years, so the exact position of "Ankylosaurus" within Ankylosauridae is unknown. "Ankylosaurus" and "Euoplocephalus" are often thought to be sister taxa. However, other analyses have found these genera in different positions.Carpenter, K. 2001. Phylogenetic analysis of the Ankylosauria. In: Carpenter, K. (Ed.). "The Armored Dinosaurs". Bloomington: Indiana University Press. Pp. 454–483.] Hill, R.V., Witmer, L.M., & Norell, M.A. 2003. A new specimen of "Pinacosaurus grangeri" (Dinosauria: Ornithischia) from the Late Cretaceous of Mongolia: ontogeny and phylogeny of ankylosaurs. "American Museum Novitates" 3395: 1-29.] Further discoveries or research may clarify the situation.

Discovery

"Ankylosaurus" was named by American paleontologist Barnum Brown, in 1908. The generic name is derived from the Greek words "αγκυλος"/"ankulos" ('curved') and "σαυρος"/"sauros" ('lizard'). Brown intended this name in the same sense as the medical term ankylosis, to refer to the stiffness produced by the fusion of many bones in the skull and body, so the name is often translated as 'stiffened lizard.' The type species is "A. magniventris", from the Latin "magnus" ('great') and "venter" ('belly'), referring to the great width of the animal's body.

A team led by Brown discovered the type specimen of "A. magniventris" (AMNH 5895) in the Hell Creek Formation of Montana, in 1906. This consisted of the top of the skull, as well as vertebrae, ribs, part of the shoulder girdle and armor. Six years earlier, Brown found the skeleton of a large theropod dinosaur (AMNH 5866) in the Lance Formation of Wyoming. This specimen was named "Dynamosaurus imperiosus" in 1905 but is now thought to belong to "Tyrannosaurus rex". Associated with AMNH 5866 were more than 75 osteoderms of various sizes, which were attributed to "Dynamosaurus". However, these osteoderms are nearly identical in form to those of "A. magniventris" and most probably belong to this species. In 1910, while on an expedition to Alberta, Barnum Brown recovered his third specimen of "A. magniventris" (AMNH 5214), from the Scollard Formation. AMNH 5214 includes a complete skull and the first known tail club, as well as ribs, limb bones and armor. All three of the above specimens are now housed at the American Museum of Natural History in New York City. The largest known skull of this animal (NMC 8880) was collected in Alberta by Charles M. Sternberg, in 1947 and is now housed at the Canadian Museum of Nature. Many other isolated bones, armor plates and teeth have been found over the years.

In popular culture

Since its description in 1908, "Ankylosaurus" has been publicized as the archetypal armored dinosaur, and due to its easily recognizable appearance and the intense public interest in dinosaurs, "Ankylosaurus" has been a feature of worldwide popular culture for many years. A life-sized reconstruction of "Ankylosaurus" featured at the 1964 World's Fair in New York City greatly contributed to its popularity.

Several motion picture series have featured ankylosaurs. "Ankylosaurus" has brief cameos in the animated "The Land Before Time" series throughout the late 1980s and 1990s, though never a large part, and is one of several types of dinosaur shown grazing together in the 2001 film "Jurassic Park III" and its video games. HardRock from the animated television program "Extreme Dinosaurs" (1998) is an "Ankylosaurus". "Ankylosaurus" has also been featured on several television documentary miniseries, including "Walking with Dinosaurs" (1999) and "The Truth About Killer Dinosaurs" (2005).

It has also been featured in computer games like "Zoo Tycoon", "" and "several "Jurassic Park" games". While some of these games depict "Ankylosaurus" as a difficult target, others present the genus as slow-moving, weak herbivores.

References