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Edmontonia

Edmontonia is a genus of panoplosaurin nodosaurid dinosaur from the Late Cretaceous Period of what is now western North America. It is named after the Edmonton Formation, the unit of rocks where it was first found.

Description
Size and general build Edmontonia was bulky, broad and tank-like. Its length has been estimated at 6.6 m (22 ft). Edmontonia had small, oval ridged bony plates on its back and head and many sharp spikes along its sides. The four largest spikes jutted out from the shoulders on each side, the second of which was split into subspines in E. rugosidens specimens. Its skull had a pear-like shape when viewed from above. Another similarity with Ankylosauridae is the presence of a secondary bone palate, a possible case of parallel evolution. This has been shown too for Panoplosaurus. Osteoderms Apart from the head armour, the body was covered with osteoderms, skin ossifications. The configuration of the armour of Edmontonia is relatively well known, much of it having been discovered in articulation. The neck and shoulder region was protected by three cervical halfrings, each consisting of fused rounded rectangular, asymmetrically keeled, bone plates. These halfrings did not have a continuous underlying bone band. The first and second halfrings each had three pairs of segments. Below each lower end of the second halfring a side spike was present, a separate triangular osteoderm pointing obliquely forward. In the third halfring over the shoulders, the two pairs of central segments are bordered on each side by a very large forward-pointing spike that is bifurcated, featuring a secondary point above the main one. A third large spike behind it points more sideways; a smaller fourth one, often connected to the third at the base, is directed obliquely to behind. The row of side spikes is continued to the rear but there the osteoderms are much lower, curving strongly to behind, with the point overhanging the rear edge. Gilmore had trouble believing that the shoulder spikes really pointed to the front as this would have greatly hampered the animal while moving through vegetation. He suggested that the points had shifted during the burial of the carcass. However, Carpenter and G.S. Paul, trying to reposition the spikes, found that it was impossible to rotate them without losing conformity with the remainder of the armour. The side spikes have solid, not hollow, bases. The spikes differ in size between E. rugosidens individuals; those of the E. longiceps holotype are relatively small. Behind the third halfring the back and hip are covered by numerous transverse rows of much smaller oval keeled osteoderms. These are not ordered in longitudinal rows. The front rows have plates oriented along the length of the body, but to the rear the long axis of these osteoderms gradually rotates sideways, their keels ultimately running transversely. Rosettes are lacking. The configuration of the tail armour is unknown. The larger plates of all body parts were connected by small ossicles. Such small round scutes also covered the throat. ==Discovery and species==
Discovery and species
In 1915, the American Museum of Natural History obtained the nearly complete, articulated front half of an armoured dinosaur, found the same year by Barnum Brown in Alberta, Canada. In 1922, William Diller Matthew referred this specimen, AMNH 5381, to Palaeoscincus in a popular-science article, not indicating any particular species. It had been intended to name a new Palaeoscincus species in cooperation with Brown but their article was never published. This species was based on type specimen USNM 11868, a skeleton found by George Fryer Sternberg in June 1928. The specific name is derived from Latin rugosus, "rough", and dens, "tooth". In 1940, Loris Shano Russell referred all three specimens to Edmontonia, as an Edmontonia rugosidens. Meanwhile, the type species of Edmontonia, Edmontonia longiceps, had been named by Charles Mortram Sternberg in 1928. The generic name Edmontonia refers to Edmonton or the Edmonton Formation. The specific name longiceps means "long-headed" in Latin. Its holotype is specimen NMC 8531, consisting of a skull, right lower jaw and much of the postcranial skeleton, including the armour. It was discovered near Morrin in 1924 by George Paterson, the teamster of the expedition led by C.M. Sternberg. Edmontonia species include: • E. longiceps, the type, known from a complete skull, is known from the middle Horseshoe Canyon Formation (Unit 2) which used to be dated to 71.5-71 million years ago. This unit, which straddles the Campanian-Maastrichtian boundary, has since been recalibrated to an age of about 72 million years. Isolated bones and shed teeth from E. longiceps are also known from the upper Judith River Formation in Montana. • E. rugosidens. This species has been given its own genus, Chassternbergia, first coined as a subgenus by Dr. Robert Thomas Bakker in 1988, as Edmontonia (Chassternbergia) rugosidens and is based on differences in skull proportion from E. longiceps and its earlier time period. It was given its full generic name in 1991 by George Olshevsky. The name Chassternbergia honours Charles, "Chas", M. Sternberg. This subgenus or genus name is rarely applied. E. rugosidens is found in the Campanian lower Dinosaur Park Formation, dating from about 76.5-75 million years ago. RTMP 98.98.01, a skull and right lower jaw; and RTMP 2001.12.158, a skull. This taxon was erected by Bakker in 1988 for a skull from the Late Maastrichtian Upper Cretaceous lower Hell Creek Formation of South Dakota, specimen DMNH 468 which was first described as Edmontonia sp. by Carpenter and Breithaupt (1986). This type specimen of Denversaurus is in the collections of the Denver Museum of Natural History (now the Denver Museum of Nature and Science), Denver, Colorado for which the genus was named. The specific name honours Lee E. Schlessman, whose Schlessman Family Foundation sponsored the museum. Bakker described the skull as being much wider at the rear than Edmontonia specimens. and considered the taxon a junior synonym of Edmontonia longiceps. Recent phylogenetic analyses included Denversaurus as a valid genus closely related to Edmontonia. Edmontonia australis was named by Tracy Lee Ford in 2000 on the basis of cervical scutes, the holotype NMMNH P-25063, a pair of medial keeled neck osteoderms from the Maastrichtian Kirtland Formation of New Mexico and the paratype NMMNH P-27450, a right middle neck plate. The naming history was further complicated in 1971, when Walter Preston Coombs Jr renamed both Edmontonia species, into Panoplosaurus longiceps and Panoplosaurus rugosidens respectively. The latter species, which due to its much more complete material has determined the image of Edmontonia, until 1940 thus appeared under the name of Palaeoscincus, and during the 1970s and 1980s was shown as "Panoplosaurus" until newer research revived the name Edmontonia. In 2010, G.S. Paul suggested that E. rugosidens was the direct ancestor of Edmontonia longiceps and the latter was again the direct ancestor of E. schlessmani. ==Phylogeny==
Phylogeny
C.M. Sternberg originally did not provide a classification of Edmontonia. In 1930, L.S. Russell placed the genus in the Nodosauridae, which has been confirmed by subsequent analyses. Edmontonia was generally shown to be a derived nodosaurid, closely related to Panoplosaurus. Russell in 1940 named a separate Edmontoniinae. In 1988 Bakker proposed that the Edmontoniinae with the Panoplosaurinae should be joined into Edmontoniidae, the presumed sister group of the Nodosauridae within Nodosauroidea which he assumed not be ankylosaurians but the last surviving stegosaurians. including in the 2018 phylogenetic analysis of Rivera-Sylva and colleagues shown below; limited to the relationships within Panoplosaurini. }} ==Paleobiology==
Paleobiology
Function of the armour The large spikes were probably used between males in contests of strength to defend territory or gain mates. The spikes would also have been useful for intimidating predators or rival males, passive protection, or for active self-defense. The large forward pointing shoulder spikes could have been used to run through attacking theropods. Carpenter suggested that the larger spikes of AMNH 5665 indicated this was a male specimen, a case of sexual dimorphism. However, he admitted the possibility of ontogeny, older individuals having longer spikes, as the specimen was relatively large. Traditionally it had been assumed that to protect themselves from predators, nodosaurids like Edmontonia might have crouched down on the ground to minimize the possibility of attack to their defenseless underbelly, trying to prevent being flipped over by a predator. ==Paleoecology==
Paleoecology
Rings in the petrified wood of trees contemporary with Edmontonia show evidence of strong seasonal changes in precipitation and temperature; Edmontonia rugosidens existed in the upper section of the Dinosaur Park Formation, about 76.5–75 million years ago. It lived alongside numerous other giant herbivores, such as the hadrosaurids Gryposaurus, Corythosaurus and Parasaurolophus, the ceratopsids Centrosaurus and Chasmosaurus, and ankylosaurids Scolosaurus The only large predators known from the same levels of the formation as Edmontonia are the tyrannosaurids Gorgosaurus libratus and an unnamed species of Daspletosaurus. Adult Albertosaurus was the apex predator in this environment, with intermediate niches possibly filled by juvenile albertosaurs. ==See also==
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