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Edmontosaurus

Edmontosaurus, often colloquially and historically known as Anatosaurus or Anatotitan, is a genus of hadrosaurid (duck-billed) dinosaur. It contains two known species: Edmontosaurus regalis and Edmontosaurus annectens. Fossils of E. regalis have been found in rocks of western North America that date from the late Campanian age of the Cretaceous period 73 million years ago, while those of E. annectens were found in the same geographic region from rocks dated to the end of the Maastrichtian age, 66 million years ago. Edmontosaurus was one of the last non-avian dinosaurs ever to exist, and lived alongside dinosaurs like Triceratops, Tyrannosaurus, Ankylosaurus, and Pachycephalosaurus shortly before the Cretaceous–Paleogene extinction event.

Discovery and history
Claosaurus annectens Edmontosaurus has had a very long and complicated history in paleontology, having spent decades with various species classified in other genera. Its taxonomic history intertwines at various points with the genera Agathaumas, Anatosaurus, Anatotitan, Claosaurus, Hadrosaurus, Thespesius, and Trachodon, with references predating the 1980s typically using Anatosaurus, Claosaurus, Thespesius, or Trachodon for edmontosaur fossils (excluding those assigned to E. regalis) depending on the author and the date. Although Edmontosaurus was only named in 1917, its oldest well-supported species (E. annectens) was named in 1892 as a species of Claosaurus. The first well-supported species of Edmontosaurus was named in 1892 as Claosaurus annectens by Othniel Charles Marsh. This species is based on USNM 2414, which is a partial skull-roof and skeleton, with a second skull and skeleton, YPM 2182, designated as the paratype. Both were collected in 1891 by John Bell Hatcher from the late Maastrichtian-age Upper Cretaceous Lance Formation of Niobrara County (then part of Converse County), Wyoming. This species has some historical footnotes attached, as it is among the first dinosaurs to receive a skeletal restoration and is the first hadrosaurid so restored. YPM 2182 and UNSM 2414 are, respectively, the first and second essentially complete mounted dinosaur skeletons in the United States. Hatcher's work led to a brief consensus until post-1910, when new material from Canada and Montana showed a greater diversity of hadrosaurids than previously suspected. His reinstatement of Thespesius for Lance-age hadrosaurids would have other consequences for the taxonomy of Edmontosaurus in the following decades. During this time frame (1902–1915), two additional important specimens of C. annectens were recovered. The first, the "mummified" specimen AMNH 5060, was discovered in 1908 by Charles Hazelius Sternberg and his sons in Lance Formation rocks near Lusk, Wyoming. Sternberg was working for the British Museum of Natural History, but Henry Fairfield Osborn of the American Museum of Natural History was able to purchase the specimen for $2,000. The Sternbergs recovered a second similar specimen from the same area in 1910, which was not as well preserved. However, it was also found with skin impressions. They sold the specimen, SM 4036, to the Senckenberg Museum in Germany. was erroneously described by Glut (1997) as having been assigned to Edmontosaurus regalis by Lull and Wright. More recent reviews of hadrosaurids have concurred. These rocks are older than the rocks in which Claosaurus annectens was found. but then changed his mind. The specimen became the first dinosaur skeleton to be mounted for exhibition in a Canadian museum. Gilmore found that his new species compared closely to what he called Thespesius annectens, but left the two apart because of details of the arms and hands. He also noted that his species had more vertebrae than Marsh's in the back and neck, but proposed that Marsh was mistaken in assuming that the annectens specimens were complete in those regions. This state of affairs persisted for several decades until Michael K. Brett-Surman reexamined the pertinent material for his graduate studies in the 1970s and 1980s. He concluded that the type species of Anatosaurus, A. annectens, was actually a species of Edmontosaurus and that A. copei was different enough to warrant its own genus. Although theses and dissertations are not regarded as official publications by the International Commission on Zoological Nomenclature, which regulates the naming of organisms, his conclusions were known to other paleontologists and were adopted by several popular works of the time. Brett-Surman and Ralph Chapman designated a new genus for A. copei (Anatotitan) in 1990. Of the remaining species, A. saskatchewanensis and A. edmontoni were assigned to Edmontosaurus as well or as a synonym of A. copei. ==Species and distribution==
Species and distribution
Edmontosaurus is currently regarded as having two valid species: the type species E. regalis and E. annectens. E. annectens is known from the Frenchman Formation of Saskatchewan, the Hell Creek Formation of Montana, and the Lance Formation of South Dakota and Wyoming. It is limited to late Maastrichtian rocks and is represented by at least twenty skulls, some with postcranial remains. Anatosaurus copei and E. saskatchewanensis are now thought to be growth stages of E. annectens, with A. copei as adults and E. saskatchewanensis as juveniles. However, the identification of Ugrunaaluk as a separate genus was questioned by a 2017 study from Hai Xing and colleagues, who regarded it as a nomen dubium that was indistinguishable from other Edmontosaurus. In 2020, Ryuji Takasaki and colleagues agreed that the Prince Creek remains should be classified as Edmontosaurus, though species designation is unclear because the specimens are juveniles. Another study found the Alaskan material to be referable to Edmontosaurus cf. regalis based on craniomandibular anatomy. Edmontosaurus was also reported from the Javelina Formation of Big Bend National Park, western Texas based on TMM 41442-1, but was later referred to Kritosaurus cf. navajovius by Wagner (2001), before being assigned to Kritosaurus sp. by Lehman et al. (2016). ==Description==
Description
Edmontosaurus has been described in detail from numerous specimens. Traditionally, E. regalis has been regarded as the largest species, though this was challenged by the hypothesis that the larger hadrosaurid Anatotitan copei is a synonym of Edmontosaurus annectens, as put forward by Jack Horner and colleagues in 2004, with a body mass on the order of . E. annectens is often seen as smaller. Two mounted skeletons, USNM 2414 and YPM 2182, measure long and long, respectively. However, these are probably subadult individuals. Two specimens still under study in the collection of the Museum of the Rockies - a tail labelled as MOR 1142 and another labelled as MOR 1609 - indicate that Edmontosaurus annectens could have grown to much larger sizes and reach nearly in length, and weighed up to , but such large individuals were likely very rare. A 2022 study on the osteohistology and growth of E. annectens suggested that previous estimates might have underestimated or overestimated the size of this dinosaur and proposed that a fully grown adult E. annectens would have measured up to in length and approximately in average asymptotic body mass, while the largest individuals measured more than and even up to when based on the comparison between various specimens of different sizes from the Ruth Mason Dinosaur Quarry and other specimens from different localities. According to this analysis, E. regalis may have been heavier, but not enough samples exist to provide a valid estimate and examination on its osteohistology and growth, so the results for E. regalis aren't statistically significant. Skull The skull of a fully grown Edmontosaurus could be over a metre long. One skull of E. annectens (formerly Anatotitan) measures long. The skull was roughly triangular in profile, Viewed from above, the front and rear of the skull were expanded, with the broad front forming a duck-bill or spoon-bill shape. The beak was toothless, and both the upper and lower beaks were extended by keratinous material. The nasal openings of Edmontosaurus were elongate and housed in deep depressions surrounded by distinct bony rims above, behind, and below. In at least one case (the Senckenberg specimen), rarely preserved scleral rings were preserved in the eye sockets. Another rarely seen bone, the stapes (the reptilian ear bone), has also been seen in a specimen of Edmontosaurus. Teeth were present only in the maxillae (upper cheeks) and dentaries (main bone of the lower jaw). The teeth were continually replaced, taking about half a year to form. They were composed of six types of tissues, rivaling the complexity of mammal teeth. They grew in columns, with an observed maximum of six in each, and the number of columns varied based on the animal's size. The ossified tendons are interpreted as having strengthened the vertebral column against gravitational stress, incurred through being a large animal with a horizontal vertebral column otherwise supported mostly by the hind legs and hips. Each foot had three toes, with no big toe or little toe. The toes had hoof-like tips. and the specimen nicknamed "Dakota", the latter apparently including remnant organic compounds from the skin. but a re-examination by Henry Sharpe and colleagues in 2025 suggested that this specimen cannot be confidently referred to any known species of Edmontosaurus, and thus it probably represents a distinct taxon belonging to the Edmontosaurini, leading to a tentative assignment to as E. sp. A preserved rhamphotheca present in the E. annectens specimen LACM 23502, housed in the Los Angeles County Museum, indicates the beak of Edmontosaurus was more hook-shaped and extensive than many illustrations in scientific and public media have previously depicted. Whether or not the specimen in question preserved the true rhamphotheca or just a cast of the inner structure attached to the bone is not known at present. ==Classification==
Classification
Edmontosaurus was a hadrosaurid (a duck-billed dinosaur), a member of a family of dinosaurs which to date are known only from the Late Cretaceous. It is classified within the Saurolophinae (alternately Hadrosaurinae), a clade of hadrosaurids which lacked hollow crests. Other members of the group include Brachylophosaurus, Gryposaurus, Lophorhothon, Maiasaura, Naashoibitosaurus, Prosaurolophus, and Saurolophus. or includes the species Anatosaurus annectens (alternately Edmontosaurus annectens), While the status of Edmontosaurus as a saurolophine has not been challenged, its exact placement within the clade is uncertain. Early phylogenies, such as that presented in R. S. Lull and Nelda Wright's influential 1942 monograph, had Edmontosaurus and various species of Anatosaurus (most of which would be later considered as additional species or specimens of Edmontosaurus) as one lineage among several lineages of "flat-headed" hadrosaurs. One of the first analyses using cladistic methods found it to be linked with Anatosaurus (=Anatotitan) and Shantungosaurus in an informal "edmontosaur" clade, which was paired with the spike-crested "saurolophs" and more distantly related to the "brachylophosaurs" and arch-snouted "gryposaurs". However, the most recent review of Hadrosauridae, by Jack Horner and colleagues (2004), came to a noticeably different result: Edmontosaurus was nested between Gryposaurus and the "brachylophosaurs", and distant from Saurolophus. }} ==Paleobiology==
Paleobiology
Diet and feeding of E. annectens as semi-aquatic animals that could only chew soft water plants, a popular idea at the time which is now outdated As a hadrosaurid, Edmontosaurus was a large terrestrial herbivore. Its teeth were continually replaced and packed into dental batteries that contained hundreds of teeth, only a relative handful of which were in use at any time. The animal's feeding range would have been from ground level to around above. An example of this is William Morris's 1970 interpretation of an edmontosaur skull with nonbony beak remnants. He proposed that the animal had a diet much like that of some modern ducks, filtering plants and aquatic invertebrates like mollusks and crustaceans from the water and discharging water via V-shaped furrows along the inner face of the upper beak. In high latitude regions like Alaska, conifers would have been the dominant food source, although the diet of Edmontosaurus would have been seasonally variable. The tooth structure indicates combined slicing and grinding capabilities. but this material has not been described. The plant remains in the Senckenberg Museum specimen have been described, but have proven difficult to interpret. The plants found in the carcass included needles of the conifer Cunninghamites elegans, twigs from conifer and broadleaf trees, and numerous small seeds or fruits. Upon their description in 1922, they were the subject of a debate in the German-language journal Paläontologische Zeitschrift. Kräusel, who described the material, interpreted it as the gut contents of the animal, At the time, hadrosaurids were thought to have been aquatic animals, and Kräusel made a point of stating that the specimen did not rule out hadrosaurids eating water plants. until it was brought up by John Ostrom in the course of an article reassessing the old interpretation of hadrosaurids as water-bound. Instead of trying to adapt the discovery to the aquatic model, he used it as a line of evidence that hadrosaurids were terrestrial herbivores. Other authors have noted that because the plant fossils were removed from their original context in the specimen and were heavily prepared, it is no longer possible to follow up on the original work, leaving open the possibility that the plants were washed-in debris. Isotopic studies The diet and physiology of Edmontosaurus have been probed by using stable isotopes of carbon and oxygen as recorded in tooth enamel. When feeding, drinking, and breathing, animals take in carbon and oxygen, which become incorporated into bone. The isotopes of these two elements are determined by various internal and external factors, such as the type of plants being eaten, the physiology of the animal, salinity, and climate. If isotope ratios in fossils are not altered by fossilization and later changes, they can be studied for information about the original factors; warmblooded animals will have certain isotopic compositions compared to their surroundings, animals that eat certain types of plants or use certain digestive processes will have distinct isotopic compositions, and so on. Enamel is typically used because the structure of the mineral that forms enamel makes it the most resistant material to chemical change in the skeleton. Such a motion would parallel the effects of mastication in mammals, although accomplishing the effects in a completely different way. Work in the early 2000s has challenged the Weishampel model. A study published in 2008 by Casey Holliday and Lawrence Witmer found that ornithopods like Edmontosaurus lacked the types of skull joints seen in those modern animals that are known to have kinetic skulls (skulls that permit motion between their constituent bones), such as squamates and birds. They proposed that joints that had been interpreted as permitting movement in dinosaur skulls were actually cartilaginous growth zones. An important piece of evidence for Weishampel's model is the orientation of scratches on the teeth, showing the direction of jaw action. Other movements could produce similar scratches though, such as movement of the bones of the two halves of the lower jaw. Not all models have been scrutinized under present techniques. Weishampel developed his model with the aid of a computer simulation. Natalia Rybczynski and colleagues have updated this work with a much more sophisticated three-dimensional animation model, scanning a skull of E. regalis with lasers. They were able to replicate the proposed motion with their model, although they found that additional secondary movements between other bones were required, with maximum separations of between some bones during the chewing cycle. Rybczynski and colleagues were not convinced that the Weishampel model is viable, but noted that they have several improvements to implement to their animation. Planned improvements include incorporating soft tissue and tooth wear marks and scratches, which should better constrain movements. They note that there are several other hypotheses to test as well. Growth In a 2011 study, Campione and Evans recorded data from all known "edmontosaur" skulls from the Campanian and Maastrichtian and used it to plot a morphometric graph, comparing variable features of the skull with skull size. Their results showed that within both recognized Edmontosaurus species, many features previously used to classify additional species or genera were directly correlated with skull size. Campione and Evans interpreted these results as strongly suggesting that the shape of Edmontosaurus skulls changed dramatically as they grew. This has led to several apparent mistakes in classification in the past. The Campanian species Thespesius edmontoni, previously considered a synonym of E. annectens due to its small size and skull shape, is more likely a subadult specimen of the contemporary E. regalis. Similarly, the three previously recognized Maastrichtian edmontosaur species likely represent growth stages of a single species, with E. saskatchewanensis representing juveniles, E. annectens subadults, and Anatotitan copei fully mature adults. The skulls became longer and flatter as the animals grew. In a 2022 study, Wosik and Evans proposed that E. annectens reached maturity in 9 years of age based on their analysis for various specimens from different localities. They found the result to be similar to that of other hadrosaurs. and E. regalis, have been studied in this way. The brain was not particularly large for an animal the size of Edmontosaurus. The space holding it was only about a quarter of the length of the skull, Osteochondrosis, or surficial pits in bone at places where bones articulate, is also known in Edmontosaurus. This condition, resulting from cartilage failing to be replaced by bone during growth, was found to be present in 2.2% of 224 edmontosaur toe bones. The underlying cause of the condition is unknown. Genetic predisposition, trauma, feeding intensity, alterations in blood supply, excess thyroid hormones, and deficiencies in various growth factors have been suggested. Among dinosaurs, osteochondrosis (like tumors) is most commonly found in hadrosaurids. Locomotion Like other hadrosaurids, Edmontosaurus is thought to have been a facultative biped, meaning that it mostly moved on four legs, but could adopt a bipedal stance when needed. It probably went on all fours when standing still or moving slowly, and switched to using the hind legs alone when moving more rapidly. While long thought to have been aquatic or semiaquatic, hadrosaurids were not as well-suited for swimming as other dinosaurs (particularly theropods, who were once thought to have been unable to pursue hadrosaurids into water). Hadrosaurids had slim hands with short fingers, making their forelimbs ineffective for propulsion, and the tail was also not useful for propulsion because of the ossified tendons that increased its rigidity, and the poorly developed attachment points for muscles that would have moved the tail from side to side. Social behavior Extensive bone beds are known for Edmontosaurus, and such groupings of hadrosaurids are used to suggest that they were gregarious, living in groups. Unlike many other hadrosaurids, Edmontosaurus lacked a bony crest. It may have had soft-tissue display structures in the skull, though: the bones around the nasal openings had deep indentations surrounding the openings, and this pair of recesses are postulated to have held inflatable air sacs, perhaps allowing for both visual and auditory signaling. Edmontosaurus has been considered a possibly migratory hadrosaurid by some authors. A 2008 review of dinosaur migration studies by Phil R. Bell and Eric Snively proposed that E. regalis was capable of an annual round-trip journey, provided it had the requisite metabolism and fat deposition rates. Such a trip would have required speeds of about , and could have brought it from Alaska to Alberta. In contrast to Bell and Snively, Anusuya Chinsamy and colleagues concluded from a study of bone microstructure that polar Edmontosaurus overwintered. ==Paleoecology==
Paleoecology
Distribution . The dark bands are coal seams. Edmontosaurus was a wide-ranging genus in both time and space. At the southern range of its distribution, the rock units from which it is known can be divided into two groups by age: the older Horseshoe Canyon and St. Mary River formations, and the younger Frenchman, Hell Creek, and Lance formations. The time span covered by the Horseshoe Canyon Formation and equivalents is also known as Edmontonian, and the time span covered by the younger units is also known as Lancian. The Edmontonian and Lancian time intervals had distinct dinosaur faunas. At its northern range, Edmontosaurus is known from a single locality; the Liscomb Bonebed of the Prince Creek Formation. Although sometimes reported as of exclusively early Maastrichtian age, Edmontosaurus regalis is known from the lowest of five units within the Horseshoe Canyon Formation, but is absent from at least the second to the top. As many as three quarters of the dinosaur specimens from badlands near Drumheller, Alberta may pertain to Edmontosaurus. Ecosystem . The Lancian time interval was the last interval before the Cretaceous–Paleogene extinction event that eliminated non-avian dinosaurs. Edmontosaurus was one of the more common dinosaurs of the interval. Robert T. Bakker reported that it made up one-seventh of the large dinosaur sample, with most of the rest (five-sixths) made up of the horned dinosaur Triceratops. The coastal plain TriceratopsEdmontosaurus association, dominated by Triceratops, extended from present-day Colorado to Saskatchewan. The Lance Formation, as typified by exposures approximately north of Fort Laramie in eastern Wyoming, has been interpreted as a bayou setting similar to the Louisiana coastal plain. It was closer to a large delta than the Hell Creek Formation depositional setting to the north and received much more sediment. Tropical araucarian conifers and palm trees dotted the hardwood forests, differentiating the flora from the northern coastal plain. The climate was humid and subtropical, with conifers, palmettos, and ferns in the swamps, and conifers, ash, live oak, and shrubs in the forests. However, more modern studies dispute the idea of an attack but rather other factors unrelated to an attack from a tyrannosaur. Another specimen of E. annectens, pertaining to a long individual from South Dakota, shows evidence of tooth marks from small theropods on its lower jaws. Some of the marks are partially healed. Michael Triebold, informally reporting on the specimen, suggested a scenario where small theropods attacked the throat of the edmontosaur; the animal survived the initial attack but succumbed to its injuries shortly thereafter. Some edmontosaur bone beds were sites of scavenging. Albertosaurus and Saurornitholestes tooth marks are common at one Alberta bone bed, and Daspletosaurus fed on Edmontosaurus and fellow hadrosaurid Saurolophus at another Alberta site. However more recent studies suggest that any and all evidence for Daspletosaurus being present in the Horseshoe Canyon Formation is referrable to Albertosaurus. A 2026 paper reported a speciman of E. annectens with a tyrannosaurid tooth embedded in the nasal around the time of death of the animal's death, interpreted as likely resulting from a bite to the snout of the animal during a predation attempt. ==See also==
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