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Edmontosaurus annectens

Edmontosaurus annectens, often colloquially and historically known as Anatosaurus, is a species of flat-headed saurolophine hadrosaurid dinosaur from the late Maastrichtian age at the very end of the Cretaceous period, in what is now western North America. Remains of E. annectens have been preserved in the Frenchman, Hell Creek, and Lance Formations. All of these formations are dated to the late Maastrichtian age of the Late Cretaceous period, which represents the last three million years before the extinction of the non-avian dinosaurs. E. annectens is also found in the Laramie Formation, and magnetostratigraphy suggests an age of 69–68 Ma for the Laramie Formation.

Discovery and history
E. annectens has a complicated taxonomic history, with various specimens having been classified in a variety of genera. Its history involves Anatosaurus, Anatotitan, Claosaurus, Diclonius, Hadrosaurus, Thespesius, and Trachodon, as well as Edmontosaurus. References predating the 1980s typically use Anatosaurus, Claosaurus, Diclonius, Thespesius, or Trachodon for E. annectens fossils, depending on the author and date. Cope's Diclonius mirabilis in 1908 The history of E. annectens predates the naming of both the genus Edmontosaurus and the species annectens. The first quality specimen, the former holotype of Anatosaurus copei (Anatotitan), was a complete skull and most of a skeleton collected in 1882 by Dr. J. L. Wortman and R. S. Hill came from northeast of the Black Hills of South Dakota, and originally had extensive skin impressions. It was missing most of its pelvis and part of its torso due to a stream cutting through it. The bill had impressions of a horn-like sheath with a tooth-like series of interlocking points on the upper and lower jaws. His reasoning was that the teeth of the lower jaw were weakly connected to the bone, and liable to break off if used to eat terrestrial food; he described the beak as weak, too. by chance, the lower jaws were missing the walls supporting the teeth from the inside, and the teeth were actually very well-supported. Cope intended to describe the skeleton and skull, but his promised paper never appeared. and it is cataloged as YPM 616. As noted by Lull and Wright, this long, slender partial jaw shares with Cope's specimen a prominent ridge running on its side. However, it is much larger: Cope's specimen had a dentary that is long, whereas Marsh's dentary is estimated at long. 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. The Sternbergs recovered a second similar specimen from the same area in 1910. It was not as well-preserved, but also found with skin impressions. They sold this specimen, SM 4036, to the Senckenberg Museum in Germany. The Horseshoe Canyon Formation is older than the rocks in which Claosaurus annectens was found. now the Frenchman Formation. A multiplicity of names resumed, with the AMNH duckbills being known as Diclonius mirabilis, Trachodon mirabilis, Trachodon annectens, Claosaurus, or Thespesius. Lull and Wright decided to remove the AMNH specimens from Diclonius (or Trachodon), because they found no convincing reason to assign the specimens to either. Because this left the skeletons without a species name, Lull and Wright gave them their own species: Anatosaurus copei, in honor of Cope. Cope's original specimen, AMNH 5730, was made the holotype of the species, with Brown's AMNH 5886 as the plesiotype. 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. The name Edmontosaurus annectens was first coined some time in the 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. His replacement name, Anatotitan (the Latin word anas ("duck"), and the Greek word Titan, meaning large), was known and published as such in the popular literature by 1990. Formal publication of the name Anatotitan copei took place the same year in an article co-written by Brett-Surman with Ralph Chapman (although the name is sometimes credited as Brett-Surman vide Chapman and Brett-Surman, because it came out of Brett-Surman's work). and A. longiceps went to Anatotitan, as either a second species or as a synonym of A. copei. The conception of Edmontosaurus that emerged included three valid species: the type species E. regalis; E. annectens (including Anatosaurus edmontoni, emended to edmontonensis); and E. saskatchewanensis. In a 2011 study by Nicolás Campione and David Evans, the authors conducted the first-ever morphometric analysis of the various specimens assigned to Edmontosaurus. They concluded that only two species are valid: E. regalis, from the late Campanian; and E. annectens, from the late Maastrichtian. Their study provided further evidence that Anatotitan copei is a synonym of E. annectens (specifically, that the long, low skull of A. copei is the result of ontogenetic change, and represents mature E. annectens individuals). E. saskatechwanensis represents young E. annectens, and Anatosaurus edmontoni specimens belong to E. regalis—not E. annectens. The reassessment of Edmontosaurus assigns twenty skulls to E. annectens. Adult skulls of E. annectens can be distinguished from skulls of E. regalis by the elongate snout and other details of skull anatomy, such as the small comb on top of the latter's skull. ==Description==
Description
The skull and skeleton of E. annectens are very well-known. Edward Drinker Cope estimated the length of one specimen as about long, with a skull long. This body length estimate was later revised down to a length of . To be fair to Cope, a dozen vertebrae, the hips, and thigh bones had been carried away by a stream cutting through the skeleton, and the tip of the tail was incomplete. A second skeleton currently exhibited next to Cope's specimen, but in a standing posture, is estimated at long, with its head above the ground. Other sources have estimated the length of E. annectens as approximately . Most specimens are somewhat shorter, representing individuals that are not fully grown. Two well-known mounted skeletons, USNM 2414 and YPM 2182, measure long and long, respectively. E. annectens may have weighed about when fully grown. potentially making it one of the largest hadrosaurids ever. However, Jack Horner and his colleagues suggested that such large individuals would have been extremely rare. The 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 argued that a fully grown adult E. annectens would have measured up to in length and in average asymptotic body mass, while the largest individuals measured more than and even up to , based on the comparison between various specimens of different sizes from the Ruth Mason Dinosaur Quarry and other specimens from different localities. The skull of E. annectens is known for its long, wide muzzle. Cope compared this feature to that of a goose in side view, and to a short-billed spoonbill in top view. The extreme length and breadth did not appear until an individual reached maturity, so many specimens lack the distinctive shape. The ridge on the lower jaw may have reinforced the long, slender structure. As mounted, the vertebral column of E. annectens includes twelve neck, twelve back, nine sacral, and at least thirty tail vertebrae. Henry Fairfield Osborn used the skeletons in the American Museum of Natural History to portray both quadrupedal and bipedal stances for E. annectens. ==Classification==
Classification
E. annectens was a saurolophine, or "flat-headed", hadrosaurid. This group was historically known as Hadrosaurinae. Species now considered to be synonymous with Edmontosaurus annectens were long recognized as closely related to both the genus and the species. However, the skull of the sub-adult type specimen of E. annectens differs noticeably from fully mature remains, so many researchers had classified the two growth stages as different species, or even different genera. On the other side of the issue, other authors, from John Bell Hatcher in 1902, }} ==Paleobiology==
Paleobiology
As a hadrosaurid, Edmontosaurus annectens was a fairly large herbivore, eating plants with a sophisticated skull that permitted a grinding motion analogous to chewing. Their teeth were continually replaced and packed into dental batteries that contained hundreds of teeth, but only a relative handful of them were in use at any time. Plant material would have been cropped by the broad beak, and held in the jaws by a cheek-like structure. Feeding would have been from the ground up to around above the ground. Like other hadrosaurs, they could have moved both bipedally and quadrupedally. A preserved rhamphotheca present in specimen LACM 23502, housed in the Los Angeles County Museum, also indicates the beak of Edmontosaurus was more hook-shaped and extensive than many illustrations in scientific and public media have previously depicted. Growth , nicknamed Diana and Leon 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 skulls with skull size. Their results showed that, in both recognized Edmontosaurus species, many features previously used to classify additional species or genera were directly correlated to skull size. Campione and Evans interpreted these results as strongly suggesting that the shape of Edmontosaurus skulls changed dramatically as they grew and matured. This has led to several apparent mistakes in past classification. 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 being 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 nine years, based on their analysis for various specimens from different localities. They found the result to be similar to that of other hadrosaurs. ==Paleoecology==
Paleoecology
''. True E. annectens remains are known only from latest Maastrichtian rocks of the Hell Creek and Lance Formations of South Dakota, Montana, and Wyoming, alongside the Frenchman Formation of Saskatchewan. The coastal plain TriceratopsEdmontosaurus association, dominated by Triceratops, extended from Colorado to Saskatchewan. Typical dinosaur faunas of the Lancian formations where Edmontosaurus annectens has been found also included: the hypsilophodont Thescelosaurus; the rare ceratopsid Torosaurus; the pachycephalosaurid Pachycephalosaurus; the ankylosaurid Ankylosaurus; and the theropods Ornithomimus, Pectinodon, Acheroraptor, Dakotaraptor, and Tyrannosaurus rex. The Hell Creek Formation, as typified by exposures in the Fort Peck area of Montana, has been interpreted as a flat, forested floodplain, with a relatively dry subtropical climate supporting a variety of plants that ranged from angiosperm trees to conifers, such as bald cypress, as well as ferns and ginkgos. The coastline was hundreds of kilometres or miles to the east. Stream-dwelling turtles and tree-dwelling multituberculate mammals were diverse, and monitor lizards as large as the modern Komodo dragon hunted on the ground. Triceratops was the most abundant large dinosaur, and Thescelosaurus the most abundant small herbivorous dinosaur. Edmontosaur remains have been collected here from stream channel sands, and include fossils from individuals as young as a metre/yard-long infant. The edmontosaur fossils potentially represented accumulations from groups on the move. 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 consequently 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. Freshwater fish, salamanders, turtles, lizards, snakes, shorebirds, and small mammals lived alongside the dinosaurs. Small dinosaurs are not known in as great of abundance here as in the Hell Creek rocks, but Thescelosaurus once again seems to have been relatively common. Triceratops in this formation is known from many skulls, which tend to be somewhat smaller than those of more northern individuals. The Lance Formation is the setting of two edmontosaur "mummies". ==See also==
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