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.
|Skeletal mount of the S. thronus holotype at the Royal Tyrell Museum|
Scolosaurus was named by Franz Nopcsa von Felső-Szilvás in 1928, based on holotype NHMUK R.5161, a nearly complete specimen that preserves the entire skeleton except for the distal end of the tail, the right forelimb, the right hindlimb, and the skull. The rare preservation of osteoderms and skin impression are also present. The fossil skeleton was discovered by William Edmund Cutler, an independent fossil collector in 1914 at Quarry 80 of the Deadlodge Canyon locality. It was collected from the bottom of the Dinosaur Park Formation in fine-grained sandstone and fine-grained claystone sediments that were deposited during the Campanian stage of the Late Cretaceous period, approximately 76.5 million years ago. The holotype specimen is housed in the collection of the Natural History Museum in London, England.
In 2013, Arbour and Currie reassigned specimen MOR 433, upon which the genus Oohkotokia was based, to Scolosaurus. This specimen consists of a partial skull, both humeri, a caudal vertebra and several osteoderms and was recovered in the Upper Member of the Two Medicine Formation, in Montana, which has been dated at approximately 74 million years. The remains were collected in 1986-1987 in grey siltstone that was deposited during the Campanian stage of the Cretaceous period. The specimen is housed in the collection of the Museum of the Rockies in Bozeman, Montana.
The generic name Scolosaurus means "pointed stake lizard" and is derived from the Greek words skolos (σκῶλος) meaning "pointed stake", and saûros (σαύρα) meaning "lizard". The specific name, cutleri, honours its discoverer and the collector of the holotype, W. E. Cutler, who was seriously injured when the specimen fell on him as he was excavating it.
In 1928, Nopcsa assigned the specimen to the family Ankylosauridae, and drew morphological comparisons with the fossil material known from Dyoplosaurus. 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 synonymized the species Anodontosaurus lambei, Dyoplosaurus acutosquameus, and Scolosaurus cutleri with Euoplocephalus tutus but did not provide any justification for these synonymies. The synonymization of Scolosaurus cutleri and Euoplocephalus tutus was generally accepted and thus NHMUK R.5161 was assigned to E. tutus. However, a redescription of Scolosaurus published in 2013 in the Canadian Journal of Earth Sciences by Paul Penkalski and William T. Blows suggested that the genus is a valid taxon. They concluded that Scolosaurus can be distinguished from Euoplocephalus by the form of their cervical armour, the details of other armour and the structure of the forelimb. They also concluded that Scolosaurus and Dyoplosaurus are distinct, due to differences noted in the pelvis and armour. Due to its completeness, the holotype of Scolosaurus has formed the basis for most Euoplocephalus reconstructions since 1971; therefore, most images of Euoplocephalus actually depict Scolosaurus instead.
A 2013 study found that the ankylosaurine Oohkotokia, was indistinguishable from Scolosaurus, and was therefore considered a junior synonym. However, this synonymization is contentious as Oohkotokia was subsequently recognized as valid. Thus, much of the material illustrated as belonging to Scolosaurus may actually pertain to Oohkotokia.
The following cladogram is based on a 2015 phylogenetic analysis of the Ankylosaurinae conducted by Victoria Arbour and Phillip J. Currie. The cladogram follows the biogeographical family tree provided by that study, which is a fusion of the study's 50% majority rule tree as well as the maximum agreement subtree. The study's 50% majority rule tree was a cladogram formed by a collection of clades, although it only included clades that appear in more than 50% of the family trees found during the analysis. The maximum agreement subtree is the cladogram that results from an algorithm which attempts to maximize the amount of taxa included in the result while also retaining the fundamental shape of all other trees in the sample. Some controversial taxa thus had to be omitted by the subtree in order for the resulting cladogram to fulfill the second requirement. The biogeographical tree (i.e. the following cladogram) is basically the 50% majority rule tree, except with some of the polytomies resolved according to the results of the maximum agreement subtree:
The following cladogram is based on a 2017 phylogenetic analysis of the Ankylosaurinae conducted by Victoria Arbour and David Evans. The cladogram depicts the majority rule (average result) of 10 most parsimonious trees, which each are considered to have the fewest evolutionary steps, thus being the most accurate under the principle of Occam's razor:
In 1874, G. M. Dawson excavated specimen USNM 7943 at the Milk River locality of the Frenchman Formation in Alberta. It was collected from terrestrial sediments that are considered to be from the Maastrichtian stage of the Late Cretaceous, approximately 70.6 to 66 million years old. The specimen consisted of a partial first cervical ring, which is part of the dinosaur's neck. In 2013, this material was assigned to Scolosaurus by Arbour and Curry who conducted a detailed phylogenetic analysis of the ankylosauridae. It is currently housed at the Smithsonian Institution in Washington, DC.
In 1928, George F. Sternberg, collected specimen USNM 11892, from the Montanazhdarcho holotype locality, high up in the Two Medicine Formation in Glacier County, Montana. The material, a partial skull, was recovered from channel sandstone sediments that were deposited during the Campanian stage, approximately 74 million years ago. This is also housed at the Smithsonian Institution.
Other referred specimens include FPDM V-31, NSM PV 20381 and TMP 2001.42.9. FPDM V-31 and TMP 2001.42.9 are both skulls, in various states of preservation. NSM PV 20381 includes a skull, dorsal vertebrae, caudal vertebrae, ribs, both scapulae, both ilia, partial ischia, and both femora, both tibiae and fibulae.
A differential diagnosis is a statement of the anatomical features of an organism (or group) that collectively distinguish it from all other organisms. Some, but not all, of the features in a diagnosis are also autapomorphies. An autapomorphy is a distinctive anatomical feature that is unique to a given organism.
According to Arbour and Currie (2013), Scolosaurus (including the Two Medicine material) can be distinguished from other ankylosaurines based on the following characteristics:
Argon-argon radiometric dating indicates that the Two Medicine Formation was deposited between 83.5 and 70.6 million years ago, during the Campanian stage of the Late Cretaceous period, in what is now northwestern Montana. If Oohkotokia is the same as Scolosaurus it would mean that Scolosaurus existed for around 3 million years. The Two Medicine Formation correlates to the Belly River Group in southwest Alberta, and the Pakowki Formation eastward. The Two Medicine Formation was deposited by rivers and deltas between the western shoreline of the Western Interior Seaway and the eastward advancing margin of the Cordilleran Overthrust Belt. Since the mid-Cretaceous, North America had been divided in half by this seaway, with much of Montana and Alberta below the surface of the water. However, the uplift of the Rocky Mountains forced the seaway to retreat eastwards and southwards. Rivers flowed down from the mountains and drained into the seaway, carrying sediment that formed the Two Medicine Formation and the Judith River Group. About 73 million years ago, the seaway began to advance westwards and northwards again, and the entire region was covered by the Bearpaw Sea, now preserved throughout the Western US and Canada by the massive Bearpaw Shale, which overlies the Two Medicine. Below this formation are the nearshore deposits of the Virgelle Sandstone. Lithologies, invertebrate faunas, and plant and pollen data support that the Two Medicine Formation was deposited in a seasonal, semi-arid climate with possible rainshadows from the Cordilleran highlands. This region experienced a long dry season and warm temperatures. The extensive red beds and caliche horizons of the upper Two Medicine are evidence of at least seasonally arid conditions.
Scolosaurus shared its paleoenvironment with other dinosaurs, such as the duck-billed hadrosaurs Hypacrosaurus, Acristavus, Gryposaurus, Brachylophosaurus, Glishades, Prosaurolophus and Maiasaura, and the ankylosaur Edmontonia. Volcanic eruptions from the west periodically blanketed the region with ash, resulting in large-scale mortality, while simultaneously enriching the soil for future plant growth. Fluctuating sea levels also resulted in a variety of other environments at different times and places within the Judith River Group, including offshore and nearshore marine habitats, coastal wetlands, deltas and lagoons, in addition to the inland floodplains. The Two Medicine Formation was deposited at higher elevations farther inland than the other two formations. Oviraptorosaurs like Caenagnathus and Chirostenotes may have preyed upon the ornithopod Orodromeus. A large variety of ceratopsians coexisted in this region, which included Achelousaurus, Brachyceratops, Cerasinops, Einiosaurus, Prenoceratops and Rubeosaurus. Carnivores included an unnamed troodontid, possibly Stenonychosaurus, the dromaeosaurs Bambiraptor and Saurornitholestes, and the large tyrannosaurids Daspletosaurus and Gorgosaurus.
The excellent vertebrate fossil record of Two Medicine and Judith River rocks resulted from a combination of abundant animal life, periodic natural disasters, and the deposition of large amounts of sediment. Many types of freshwater and estuarine fish are represented, including sharks, rays, sturgeons, gars and others. This region preserves the remains of many aquatic amphibians and reptiles, including bivalves, gastropods, frogs, salamanders, turtles, Champsosaurus and crocodilians. Terrestrial lizards, including whiptails, skinks, monitors and alligator lizards have also been discovered. Pterosaurs like Montanazhdarcho and Piksi as well as birds like Apatornis and Avisaurus flew overhead. Several varieties of mammals, such as the multituberculate Cimexomys coexisted with dinosaurs in the Two Medicine Formation and the various other formations that make up the Judith River wedge. Fossilized eggs belonging to a dromaeosaur have been recovered here. When water was plentiful, the region could support a great deal of plant and animal life, but periodic droughts often resulted in mass mortality.
Ankylosauridae () is a family of armored dinosaurs within Ankylosauria, and is the 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.Anodontosaurus
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.Bienosaurus
Bienosaurus (meaning "Bien's lizard") is a genus of thyreophoran dinosaur from the Lower Jurassic (probably Sinemurian) Lower Lufeng Formation in Yunnan Province in China.Bissektipelta
Bissektipelta is a genus of herbivorous ankylosaurid dinosaur from the Upper Cretaceous of Uzbekistan. Bissektipelta is monospecific, containing only the species B. archibaldi.Corythosaurus
Corythosaurus is a genus of hadrosaurid "duck-billed" dinosaur from the Upper Cretaceous Period, about 77–75.7 million years ago. It lived in what is now North America. Its name means "helmet lizard", derived from Greek κόρυς. It was named and described in 1914 by Barnum Brown. Corythosaurus is now thought to be a lambeosaurine, related to Nipponosaurus, Velafrons, Hypacrosaurus, and Olorotitan. Corythosaurus has an estimated length of 9 metres (30 ft), and has a skull, including the crest, that is 70.8 centimetres (27.9 in) tall.
Corythosaurus is known from many complete specimens, including the nearly complete holotype found by Brown in 1911. The holotype skeleton is only missing the last section of the tail, and part of the forelimbs, but was preserved with impressions of polygonal scales. Corythosaurus is known from many skulls with tall crests. The crests resemble the crests of the cassowary and a Corinthian helmet. The most likely function of the crest is thought to be vocalization. As in a trombone, sound waves would travel through many chambers in the crest, and then get amplified when Corythosaurus exhaled. A Corythosaurus specimen has been preserved with its last meal in its chest cavity. Inside the cavity were remains of conifer needles, seeds, twigs, and fruits: Corythosaurus probably fed on all of these.
The two species of Corythosaurus are both present in slightly different levels of the Dinosaur Park Formation. Both still co-existed with theropods and other ornithischians, like Daspletosaurus, Brachylophosaurus, Parasaurolophus, Scolosaurus, and Chasmosaurus.Euoplocephalus
Euoplocephalus ( yoo-OP-loh-KEF-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.Invictarx
Invictarx is a genus of herbivorous nodosaurid dinosaur from New Mexico dating from the early Campanian epoch of the Late Cretaceous.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.Oldman Formation
The Oldman Formation is a stratigraphic unit of Late Cretaceous (Campanian stage) age that underlies much of southern Alberta, Canada. It consists primarily of sandstones that were deposited in fluvial channel and floodplain environments. It was named for exposures along the Oldman River between its confluence with the St. Mary River and the city of Lethbridge, and it is known primarily for its dinosaur remains and other fossils.Oohkotokia
Oohkotokia ( OH-oh-kə-TOH-kee-ə) is a genus of ankylosaurid dinosaur within the subfamily Ankylosaurinae. It is known from the upper levels of the Two Medicine Formation (late Campanian stage, about 74 Ma ago) of Montana, United States. The discovery of Oohkotokia supports that Ankylosaurine dinosaurs existed and flourished continuously in Montana and/or Alberta throughout the late Campanian and early Maastrichtian stages in the Late Cretaceous period. It was a large, heavily built, quadrupedal, herbivore, that could grow up to 6 m (19.7 ft) long.Platypelta
Platypelta is an extinct genus of herbivorous ankylosaurid dinosaurs within the subfamily Ankylosaurinae. It is known from the Late Cretaceous Dinosaur Park Formation (early Late Campanian stage, about 77.5-76.5 Ma ago) of southern Alberta, Canada. The type species is Platypelta coombsi.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 dinosaurs 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 m (8–10 ft) in length. It was named by Coombs in 1995.Tianzhenosaurus
Tianzhenosaurus (Tianzhen + Greek sauros="lizard") is a genus of ankylosaurid dinosaurs discovered in Tianzhen County, at Kangdailiang near Zhaojiagou Village, in Shanxi Province, China, in the Late Cretaceous Huiquanpu Formation. Thus far, a virtually complete skull and postcranial skeleton have been assigned to the genus, which is monotypic (T. youngi Pang & Cheng, 1998).
This was a medium-sized ankylosaurian, the skull measuring 28 cm (11 in) in length, with a total body length around 4 m (13 ft).
Vickaryous et al. (2004) placed Tianzhenosaurus within the Ankylosauridae, nested as the sister group to Pinacosaurus. Some authors have suggested that Tianzhenosaurus is actually a junior synonym of Saichania chulsanensis.Tsagantegia
Tsagantegia (; meaning "of Tsagan-Teg"; Tumanova, 1993) is a genus of medium-sized ankylosaurid dinosaur from the Late Cretaceous of 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.Two Medicine Formation
The Two Medicine Formation is a geologic formation, or rock body, that was deposited between 83.5 ± 0.7 Ma and 70.6 ± 3.4 Ma (million years ago), during Campanian (Late Cretaceous) time, and is located in northwestern Montana and southern Alberta. It crops out to the east of the Rocky Mountain Overthrust Belt, and the western portion (about 600 metres (2,000 ft) thick) of this formation is folded and faulted while the eastern part, which thins out into the Sweetgrass Arch, is mostly undeformed plains. Below the formation are the nearshore (beach and tidal zone) deposits of the Virgelle Sandstone, and above it is the marine Bearpaw Shale. Throughout the Campanian, the Two Medicine Fm. was deposited between the western shoreline of the Late Cretaceous Interior Seaway and the eastward advancing margin of the Cordilleran Overthrust Belt. The Two Medicine Fm. is mostly sandstone, deposited by rivers and deltas.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.