Anodontosaurus is an extinct genus of ankylosaurid dinosaurs within the subfamily Ankylosaurinae. It is known from the entire span of the Late Cretaceous Horseshoe Canyon Formation (mid Late Campanian to "middle" Maastrichtian stage, about 72.8-67 Ma ago) of southern Alberta, Canada. It contains two species, A. lambei and A. inceptus.
|A. lambei holotype skull CMN 8530|
Anodontosaurus was named by Charles Mortram Sternberg in 1928, based on holotype CMN 8530, a partially preserved skeleton including the skull, half ring, armor and other postcranial remains. The badly crushed skeleton was collected by Sternberg in 1916 from a Canadian Museum of Nature quarry, 8 miles southwest of Morrin. It was collected from the upper part of the Lower Horseshoe Canyon Formation (unit 2), dating to the latest Campanian to the earliest Maastrichtian stage of the Late Cretaceous period, about 71-70.2 million years ago. The generic name means "toothless lizard" in Ancient Greek. It was inspired by the fact that compression damage to the specimen had removed the teeth, at the same time shifting various flat round elements below the skull and on top of the left lower jaw, misleading Sternberg into assuming that large "trituration plates" had replaced the normal dentition. The specific name, lambei, honours Lawrence Morris Lambe, the Canadian geologist and palaeontologist from the Geological Survey of Canada where the holotype was reposited.
In 1986 Coombs examined specimen AMNH 5266, at the time by him referred to Euoplocephalus, and determined that it was a juvenile. Its consists of five vertebral centra, a neural arch, one dorsal and two sacral ribs, the right ischium, the complete right hindlimb, the right pes, an incomplete left pes, and various other fragments. AMNH 5266 was discovered in 1912 at Red Deer River and was collected by Barnum Brown with assistance from Peter Kaisen, George Olsen, and Charles M Sternberg in the sediments from the Horseshoe Canyon Formation.
Anodontosaurus was a medium-sized ankylosaurid that was quadrupedal, ground-dwelling and herbivorous. Like other ankylosaurs, Anodontosaurus has armor on a majority of the dorsal surfaces of its body. It has a wide, pointed tail club and the end of its armored tail. The skull features postocular caputegulae, which are small polygonal plates of bone that are present on the cranium and are situated to the immediate rear of the eye.
Coombs supported the assertion that specimen AMNH 5266 represented a juvenile by citing that the vertebral centra were not fused to their neural arches, and that sacral ribs were likewise not fused to vertebrae and to the ilium. Other morphological characters supporting that this is a juvenile specimen include (a) long bones that feature smooth surfaces, which are not marked by the rugosities characteristic of adult bone; (b) the head of the femur is less spherical in shape and is clearly delimited from the adjacent part of the femoral shaft; (c) the distal ends of the tibia and the fibula are not fused to the astragalus and the calcaneum; and (d) the ungual phalanx of the manus is not widest at the proximal articular end as is observed in adults.
In 1971, Walter Coombs concluded that there was only one species of ankylosaurid during the Campanian age of the Late Cretaceous of North America. He synonymised the species Anodontosaurus lambei, Dyoplosaurus acutosquameus, and Scolosaurus cutleri with Euoplocephalus tutus. The synonymisation of Anodontosaurus lambei and Euoplocephalus tutus was generally accepted and thus CMN 8530 was assigned to E. tutus. However, following the redescription of Dyoplosaurus as a valid genus by Arbour et al. (2009), In an abstract presented at the SVP 2010 conference, Victoria Arbour considered Anodontosaurus distinct from Euoplocephalus in distinctive skull and cervical half ring ornamentation, as well as tail club morphology, including the presence of pointed, triangular knob osteoderms in Anodontosaurus. She therefore reassigned all Horseshoe Canyon Formation ankylosaurine specimens previously referred to Euoplocephalus to Anodontosaurus.
The validity of Anodontosaurus was formalized in three studies. The first, published by Paul Penkalski and William T. Blows in 2013, re-validated Scolosaurus as well. The second study, by Penkalski (2013), named and described Oohkotokia from Montana on the basis of remains that were originally thought to be referable to Euoplocephalus. Penkalski (2013) performed a small phylogenetic analysis of some ankylosaurine specimens. The only Anodontosaurus specimen that was included in this analysis was its holotype. Anodontosaurus was placed in a polytomy with the holotype of Euoplocephalus and some specimens that are referred to it, while Oohkotokia was placed in a clade with Dyoplosaurus, and specimens that are thought to represent either Dyoplosaurus or Scolosaurus. In a study based on the results of her 2010 SVP abstract, Arbour along with Philip Currie formalized the revalidation of Anodonotsaurus, and one specimen from the Dinosaur Park Formation, TMP 1997.132.1, was referred to Anodontosaurus extending the stratigraphic range of the genus back a few million years. The DPF specimen, however, was later made the holotype of a new species, A. inceptus. The following cladogram is based on a 2015 phylogenetic analysis of the Ankylosaurinae conducted by Arbour and Currie:
Ankylosauridae () are a family of the armored dinosaurs within Ankylosauria, and sister group to Nodosauridae. Ankylosaurids appeared 122 million years ago and went extinct 66 million years ago during the Cretaceous–Paleogene extinction event. These animals were mainly herbivorous and were obligate quadrupeds, with leaf-shaped teeth and robust, scute-covered bodies. Ankylosaurids possess a distinctly domed and short snout, wedge-shaped osteoderms on their skull, scutes along their torso, and a tail club.Ankylosauridae is exclusively known from the northern hemisphere, with specimens found in western North America, Europe, and East Asia. The first discoveries within this family were of the genus Ankylosaurus, by Peter Kaiser and Barnum Brown in Montana in 1906. Brown went on to name Ankylosauridae and the subfamily Ankylosaurinae in 1908.Ankylosaurinae
Ankylosaurinae is a subfamily of ankylosaurid dinosaurs, existing from the Early Cretaceous about 105 million years ago until the end of the Late Cretaceous, about 66 mya. Many genera are included in the clade, such as Ankylosaurus, Pinacosaurus, Euoplocephalus, and Saichania.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; the only species in the genus is A. magniventris. The genus name means "fused 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 6 and 8 metres (20 and 26 ft) long and to have weighed between 4.8 and 8 tonnes (4.7 and 7.9 long tons). 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. Ankylosaurus has been found in the Hell Creek, Lance, Scollard, Frenchman, and Ferris formations, but appears to have been rare in its environment. Although it lived alongside 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.Bienosaurus
Bienosaurus (meaning "Bien's lizard") was a dinosaur from the Early Jurassic (probably Sinemurian). The genus was an armoured dinosaur from the Lower Lufeng Formation in Yunnan Province in China.Dongyangopelta
Dongyangopelta is an extinct genus of nodosaurid ankylosaurian dinosaur known from the "middle" Cretaceous Chaochuan Formation (Albian or Cenomanian stage) of Dongyang, Zhejiang Province, China. Dongyangopelta was first named by Rongjun Chen, Wenjie Zheng, Yoichi Azuma, Masateru Shibata, Tianliang Lou, Qiang Jin and Xingsheng Jin in 2013 and the type species is Dongyangopelta yangyanensis. It differs from Zhejiangosaurus, the second nodosaurid from southeast China, in the characters of presacral rod, ilium, and femur. Donyangopelta is distinguishable from Zhejiangosaurus only on the basis of the morphology of its pelvic shield.Euoplocephalus
Euoplocephalus ( yoo-OP-loh-SEF-ə-ləs) is a genus of very large, herbivorous ankylosaurian dinosaurs, living during the Late Cretaceous of Canada. It has only one named species, Euoplocephalus tutus.
The first fossil of Euoplocephalus was found in 1897 in Alberta. In 1902, it was named Stereocephalus, but that name had already been given to an insect, so it was changed in 1910. Later, many more ankylosaurid remains were found from the Campanian of North America and often made separate genera. In 1971, Walter Coombs concluded that they all belonged to Euoplocephalus which then would be one of the best-known dinosaurs. Recently however, experts have come to the opposite conclusion, limiting the authentic finds of Euoplocephalus to about a dozen specimens. These include a number of almost complete skeletons, so much is nevertheless known about the build of the animal.
Euoplocephalus was about five and a half meters long and weighed about two and a half tonnes. Its body was low-slung and very flat and wide, standing on four sturdy legs. Its head had a short drooping snout with a horny beak to bite off plants that were digested in the large gut. Like other ankylosaurids, Euoplocephalus was largely covered by bony armor plates, among them rows of large high-ridged oval scutes. The neck was protected by two bone rings. It could also actively defend itself against predators like Gorgosaurus using a heavy club-like tail end.Horseshoe Canyon Formation
The Horseshoe Canyon Formation is a stratigraphic unit of the Western Canada Sedimentary Basin in southwestern Alberta. It takes its name from Horseshoe Canyon, an area of badlands near Drumheller.
The Horseshoe Canyon Formation is part of the Edmonton Group and is up to 230 metres (750 ft) thick. It is of Late Cretaceous age, Campanian to early Maastrichtian stage (Edmontonian Land-Mammal Age), and is composed of mudstone, sandstone, carbonaceous shales, and coal seams. A variety of depositional environments are represented in the succession, including floodplains, estuarine channels, and coal swamps, which have yielded a diversity of fossil material. Tidally-influenced estuarine point bar deposits are easily recognizable as Inclined Heterolithic Stratification (IHS). Brackish-water trace fossil assemblages occur within these bar deposits and demonstrate periodic incursion of marine waters into the estuaries.
The Horseshoe Canyon Formation crops out extensively in the area around Drumheller, as well as farther north along the Red Deer River near Trochu and along the North Saskatchewan River in Edmonton. It is overlain by the Battle, Whitemud, and Scollard formations. The Drumheller Coal Zone, located in the lower part of the Horseshoe Canyon Formation, was mined for sub-bituminous coal in the Drumheller area from 1911 to 1979, and the Atlas Coal Mine in Drumheller has been preserved as a National Historic Site. In more recent times, the Horseshoe Canyon Formation has become a major target for coalbed methane (CBM) production.
Dinosaurs found in the Horseshoe Canyon Formation include Albertavenator, Albertosaurus, Anchiceratops, Anodontosaurus, Arrhinoceratops, Atrociraptor, Epichirostenotes, Edmontonia, Edmontosaurus, Hypacrosaurus, Ornithomimus, Pachyrhinosaurus, Parksosaurus, Saurolophus, and Struthiomimus. Other finds have included mammals such as Didelphodon coyi, non-dinosaur reptiles, amphibians, fish, marine and terrestrial invertebrates and plant fossils. Reptiles such as turtles and crocodilians are rare in the Horseshoe Canyon Formation, and this was thought to reflect the relatively cool climate which prevailed at the time. A study by Quinney et al. (2013) however, showed that the decline in turtle diversity, which was previously attributed to climate, coincided instead with changes in soil drainage conditions, and was limited by aridity, landscape instability, and migratory barriers.Invictarx
Invictarx is a genus of herbivorous nodosaurid dinosaur from New Mexico dating from the early Campanian epoch of the Late Cretaceous.Mongolostegus
Mongolostegus is a genus of stegosaur from the Early Cretaceous (Aptian-Albian) of Mongolia. The type and only species is M. exspectabilis, known from a single specimen previously under the nomen nudum "Wuerhosaurus mongoliensis".Nodosaurinae
Nodosaurinae is a group of ankylosaurian dinosaurs named in 1919 by Othenio Abel.Nodosaurus
Nodosaurus (meaning "knobbed lizard") is a genus of herbivorous ankylosaurian dinosaur from the Late Cretaceous, the fossils of which are found in North America.Scolosaurus
Scolosaurus is an extinct genus of ankylosaurid dinosaurs within the subfamily Ankylosaurinae. It is known from either the lower levels of the Dinosaur Park Formation or upper levels of the Oldman Formation (the location of the type specimen's quarry is uncertain) in the Late Cretaceous (latest middle Campanian stage, about 76.5 Ma ago) of Alberta, Canada. It contains two species, S. cutleri and S. thronus.Silvisaurus
Silvisaurus, from the Latin silva "woodland" and Greek sauros "lizard", is a nodosaurid ankylosaur from the middle Cretaceous period.Tarchia
Tarchia (meaning "brainy one") is a genus of herbivorous ankylosaurid dinosaur from the late Cretaceous of Mongolia.Tatisaurus
Tatisaurus is a genus of ornithischian dinosaur from the Early Jurassic from the Lower Lufeng Formation in Yunnan Province in China. Little is known as the remains are fragmentary.Texasetes
Texasetes (meaning "Texas resident") is a genus of ankylosaurian dinosaur from the late Lower Cretaceous of North America. This poorly known genus has been recovered from the Paw Paw Formation (late Albian) near Haslet, Tarrant County Texas, which has also produced the nodosaurid ankylosaur Pawpawsaurus. Texasetes is estimated to have been 2.5–3 meters (8–10 ft) in length. It was named by Coombs in 1995.Tsagantegia
Tsagantegia (; meaning "of Tsagan-Teg"; Tumanova, 1993) is a genus of medium-sized ankylosaurid dinosaur from the Upper Cretaceous Mongolia, during the Cenomanian stage.
The holotype specimen (GI SPS N 700/17), a complete skull, was recovered from the Bayan Shireh Formation (Cenomanian-Santonian), at the Tsagan-Teg ("White Mountain") locality, Dzun-Bayan, in the southeastern Gobi Desert, Mongolia. The genus is monotypic, including only the type species, T. longicranialis.Ziapelta
Ziapelta sanjuanensis is an extinct species of ankylosaurid from Upper Cretaceous (Campanian) formations in New Mexico.Zuul
Zuul is a genus of herbivorous ankylosaurine dinosaur from the Campanian Judith River Formation of Montana. The type species is Zuul crurivastator. It is known from a complete skull and tail, which represents the first ankylosaurin known from a complete skull and tail club, as well as the most complete ankylosaurid specimen thus far recovered from North America. The specimen also preserved in situ osteoderms, keratin, and skin remains.