Scolosaurus is an extinct genus of ankylosaurid dinosaurs within the subfamily Ankylosaurinae. It is known from the lower levels of the Dinosaur Park Formation and upper levels of the Oldman Formation in the Late Cretaceous (latest middle Campanian stage, about 76.5 Ma ago) of Alberta, Canada. The genus contains two species, S. cutleri and S. thronus. The type species, S. cutleri, measured up to 5.6 metres (18 ft) in length and 2.2 metric tons (2.4 short tons) in body mass. The genus was once believed to represent a synonym of the coeval Euoplocephalus, but this is no longer thought to be the case, and Scolosaurus is now accepted as a valid taxon. Some researchers have considered the genus Oohkotokia, known from the also Campanian-aged Two Medicine Formation of Montana, United States, to be synonymous with Scolosaurus.
Discovery
Scolosaurus was named by Franz Nopcsa von Felső-Szilvás in 1928, based on holotype NHMUK PV 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.
Classification 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:
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