Edmontosaurus annectens

{{short description|Hadrosaurid species from the Late Cretaceous Period}} {{speciesbox | fossil_range = Late Cretaceous (Maastrichtian),<ref name="Holtz2008"/> {{fossilrange|earliest=69|68|66}} {{period fossil range|Cretaceous|68|66}} | image = Oxford Edmontosaurus.jpg | image_caption = Mounted cast of a fossil ''E. annectens'' skeleton, Oxford University Museum of Natural History | genus = Edmontosaurus | species = annectens | authority = (Marsh, 1892) | synonyms = {{Collapsible list| *''Trachodon longiceps'' Marsh, 1890 *''Hadrosaurus longiceps'' (Marsh, 1890) Nopcsa, 1900 *''Claosaurus annectens'' Marsh, 1892 *''Trachodon annectens'' (Marsh, 1892) Hay, 1902 *''Thespesius annectens'' (Marsh, 1892) Sternberg, 1925 *''Thespesius saskatchewanensis'' Sternberg, 1926 *''Trachodon saskatchewanensis'' (Sternberg, 1926) Kuhn, 1936 *''Thespesius longiceps'' (Marsh, 1890) Russell, 1930 *''Anatosaurus annectens'' (Marsh, 1892) Lull & Wright, 1942 *''Anatosaurus longiceps'' (Marsh, 1890) Lull & Wright, 1942 *''Anatosaurus saskatchewanensis'' (Sternberg, 1926) Lull & Wright, 1942 *''Anatosaurus copei'' Lull & Wright, 1942 *''Edmontosaurus copei'' (Lull & Wright, 1942) Brett-Surman, 1975 *''Anatotitan copei'' (Lull & Wright, 1942) Brett-Surman vide Chapman & Brett-Surman, 1990 *''Anatotitan longiceps'' (Marsh, 1890) Olshevsky, 1991 *''Edmontosaurus saskatchewanensis'' (Sternberg, 1926) Horner, Weishampel & Forster, 2004 }} }}

'''''Edmontosaurus annectens''''' (meaning "connected lizard from Edmonton"), often colloquially and historically known as '''''Anatosaurus''''' (meaning "duck lizard"), 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 (between 68 and 66 million years ago<ref name="Holtz2008">Holtz, Thomas R. Jr. (2012) ''Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages,'' [http://www.geol.umd.edu/~tholtz/dinoappendix/HoltzappendixWinter2011.pdf Winter 2011 Appendix.]</ref>).<ref>{{cite journal |last1=Rohrer |first1=Willis L. |last2=Konizeski |first2=Richard L. |title=On the Occurrence of Edmontosaurus in the Hell Creek Formation of Montana |journal=Journal of Paleontology |date=1 May 1960 |volume=34 |issue=3 |pages=464–466 |jstor=1300943 }}</ref> ''E. annectens'' is also found in the Laramie Formation, and magnetostratigraphy suggests an age of 69–68 Ma for the Laramie Formation.<ref name="hicksetal2003">*Hicks, J.F., Johnson, K.R., Obradovich, J. D., Miggins, D.P., and Tauxe, L. 2003. Magnetostratigraphyof Upper Cretaceous (Maastrichtian) to lower Eocene strata of the Denver Basin, Colorado. In K.R. Johnson, R.G. Raynolds and M.L. Reynolds (eds), Paleontology and Stratigraphy of Laramide Strata in the Denver Basin, Pt. II., Rocky Mountain Geology 38: 1-27.</ref>

''Edmontosaurus annectens'' is known from numerous specimens, including at least twenty partial-to-complete skulls, discovered in the U.S. states of Montana, South Dakota, North Dakota, Wyoming, and Colorado, as well as the Canadian province of Saskatchewan. It had an extremely long and low skull, and was quite a large animal, growing up to approximately {{convert|12|m}} in length and {{convert|5.6|MT|ST}} in average asymptotic body mass,<ref name="Wosik2022"/> although it could have been even larger.<ref name="census"/> ''E. annectens'' exhibits one of the most striking examples of the "duckbill" snout that is common to hadrosaurs. It has a long taxonomic history, and specimens have at times been classified as ''Diclonius'', ''Trachodon'', ''Hadrosaurus'', ''Claosaurus'', ''Thespesius'', ''Anatosaurus'', and '''''Anatotitan''''' before all being grouped together in ''Edmontosaurus''.

==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''.<ref name=HWF04/><ref name="BSC07">{{cite book |last=Creisler |first=Benjamin S. |title=Horns and Beaks: Ceratopsian and Ornithopod Dinosaurs |publisher=Indiana University Press |year=2007 |isbn=978-0-253-34817-3 |editor-last=Carpenter |editor-first=Kenneth |editor-link=Kenneth Carpenter |location=Bloomington and Indianapolis |pages=185–210 |chapter=Deciphering duckbills: a history in nomenclature}}</ref> 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''=== [[File:Anatotitan copei.jpg|thumb|left|upright|Skeletons (AMNH 5730, left, and AMNH 5886, right), first mounted in the American Museum of Natural History in 1908<ref name="The-Trachodon-Story">"[https://dino.lindahall.org/name.shtml 26b. What's In a Name: The Trachodon Story]"; ''Paper Dinosaurs (1824-1969): An Exhibition of Original Publications from the Collections of the Linda Hall Library''.</ref>]] 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<ref name="DFG97"/> for American paleontologist Edward Drinker Cope. This specimen, found in Hell Creek Formation rocks,<ref name="NGD95">{{cite book |last=Norell |first=M. A. |author2=Gaffney, E. S. |author3=Dingus, L. |title=Discovering Dinosaurs in the American Museum of Natural History |publisher=Knopf |location=New York |year=1995 |pages=[https://archive.org/details/discoveringdinos00nore_0/page/156 156–158] |isbn=0-679-43386-4 |url=https://archive.org/details/discoveringdinos00nore_0/page/156 }}</ref> 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.<ref name=HFO09a/> When describing this specimen, AMNH 5730, Cope assigned it to the species ''Diclonius mirabilis''. This species name was created by combining ''Diclonius'', a hadrosaurid genus Cope had named earlier from teeth, with ''Trachodon mirabilis'', an older name based on teeth that was published by Joseph Leidy. Cope believed that Leidy had failed to properly characterize the genus ''Trachodon'' and later abandoned its use, so he assigned the old species to his newer genus.<ref name=EDC83/> Leidy had come to recognize that his ''Trachodon'' was based on the remains of multiple kinds of dinosaurs, and although he had made some attempts to revise the genus, he had not yet made any formal declaration of his intentions.<ref name=BSC07/>

Cope's description promoted hadrosaurids as amphibious animals, contributing to this long-time image.<ref name=JHO64>{{cite journal |doi=10.2475/ajs.262.8.975 |last=Ostrom |first=John H. |author-link=John Ostrom |title=A reconsideration of the paleoecology of the hadrosaurian dinosaurs |journal=American Journal of Science |volume=262 |pages=975–997 |year=1964 |issue=8|bibcode=1964AmJS..262..975O |doi-access=free }}</ref> 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.<ref name=EDC83/> However, aside from misidentifying several of the skull bones,<ref name=OCM93>{{cite journal |last=Marsh |first=Othniel C. |author-link=Othniel Charles Marsh |year=1893 |title=The skull and brain of ''Claosaurus'' |journal=American Journal of Science |series=3rd Series |volume=45 |issue=265 |pages=83–86|url=https://zenodo.org/record/2198923 |doi=10.2475/ajs.s3-45.265.83 |bibcode=1893AmJS...45...83M |s2cid=131740074 }}</ref> by chance, the lower jaws were missing the walls supporting the teeth from the inside, and the teeth were actually very well-supported.<ref name=JHO64/><ref name=LW42d>Lull and Wright, ''Hadrosaurian Dinosaurs of North America'', pp. 43.</ref> Cope intended to describe the skeleton and skull, but his promised paper never appeared.<ref name=BSC07/> It was purchased for the American Museum of Natural History in 1899, where it acquired its present designation: AMNH 5730.<ref name=LW42b/> [[File:Edmontosaurus annectens, by Charles R. Knight.jpg|thumb|Outdated 1909 life restoration of ''Trachodon'' by Charles R. Knight, based on the two specimens (now classified as ''E. annectens'') mounted in 1908 at the AMNH, New York.<ref name="The-Trachodon-Story"/>]] Several years after Cope's description, his arch-rival, Othniel Charles Marsh, published a paper on a sizable lower jaw recovered by John Bell Hatcher in 1889 from the Lance Formation rocks in Niobrara County, Wyoming.<ref name=LW42c/> Marsh named this partial jaw ''Trachodon longiceps'',<ref name=OCM90>{{cite journal |last=Marsh |first=Othniel C. |author-link=Othniel Charles Marsh |year=1890 |title=Additional characteristics of the Ceratopsidae, with notice of new Cretaceous dinosaurs |journal=American Journal of Science |series=3rd Series |volume=39 |issue=233 |pages=418–426|doi=10.2475/ajs.s3-39.233.418 |bibcode=1890AmJS...39..418M |s2cid=130812960 |url=https://zenodo.org/record/2513377 }}</ref> 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 {{convert|92.0|cm|in}} long, whereas Marsh's dentary is estimated at {{convert|110.0|cm}} long.<ref name=LW42c/>

A second mostly complete skeleton, AMNH 5886, was found in 1904 in the Hell Creek Formation rocks at Crooked Creek in central Montana by a local rancher named Oscar Hunter. Upon finding the partially exposed specimen, he and a companion argued about whether or not the remains were recent or fossil. Hunter demonstrated that they were brittle and thus stone by kicking the tops off the vertebrae, an act later lamented by the eventual collector Barnum Brown. Another cowboy, Alfred Sensiba, bought the specimen from Hunter for a pistol and later sold it to Brown, who excavated it for the American Museum of Natural History in 1906.<ref name=NGD95/> This specimen had a nearly complete vertebral column, permitting the restoration of Cope's specimen. In 1908, these two specimens were mounted side by side in the American Museum of Natural History under the name ''Trachodon mirabilis''.<ref name="The-Trachodon-Story"/> Cope's specimen is positioned on all fours with its head down, as if feeding, because it has the better skull, while Brown's specimen, with a less perfect skull, is posed bipedally with the head less accessible. Henry Fairfield Osborn described the tableau as representing the two animals feeding alongside a marsh, the standing individual having been startled by the approach of a ''Tyrannosaurus''. Impressions of appropriate plant remains and shells based on associated fossils were included on the base of the group, including ginkgo leaves, ''Sequoia'' cones, and horsetail rushes.<ref name=HFO09a/>

===Marsh's ''Claosaurus annectens''=== [[File:Large marsh claosaurus.jpg|thumb|Skeletal restoration of the ''E. annectens'' (then ''Claosaurus'') holotype, by Othniel Charles Marsh.]] [[File:Transactions of the Connecticut Academy of Arts and Sciences (1901) (14783568452).jpg|thumb|''E. annectens'' paratype YPM 2182 at the Yale University Museum, the first nearly complete dinosaur skeleton mounted in the United States.<ref name=FAL04/>]] The species now known as ''Edmontosaurus annectens'' was named in 1892 as ''Claosaurus annectens'' by Othniel Charles Marsh. This species is based on USNM 2414, a partial skull-roof and skeleton, with a second skull and skeleton, YPM 2182, being 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.<ref name=OCM92a>{{cite journal |last=Marsh |first=Othniel Charles |author-link=Othniel Charles Marsh |year=1892 |title=Notice of new reptiles from the Laramie Formation |journal=American Journal of Science |volume=43 |issue=257 |pages=449–453|doi=10.2475/ajs.s3-43.257.449 |bibcode=1892AmJS...43..449M |s2cid=131291138 |url=https://zenodo.org/record/2238442 }}</ref> 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.<ref name=BSC07/><ref name=OCM92b>{{cite journal |last=Marsh |first=Othniel Charles |author-link=Othniel Charles Marsh |year=1892 |title=Restorations of ''Claosaurus'' and ''Ceratosaurus'' |journal=American Journal of Science |volume=44 |issue=262 |pages=343–349 |doi=10.2475/ajs.s3-44.262.343|bibcode=1892AmJS...44..343M |hdl=2027/hvd.32044107356040 |s2cid=130216318 |hdl-access=free }}</ref> YPM 2182 and UNSM 2414 are, respectively, the first and second essentially complete mounted dinosaur skeletons in the United States.<ref name="FAL04"/> YPM 2182 was put on display in 1901,<ref name=BSC07/> and USNM 2414 was put on display in 1904.<ref name=FAL04/>

In the first decade of the twentieth century, two additional important specimens of ''C. annectens'' were recovered. The first, the "Trachodon mummy," AMNH 5060, was discovered in 1908 by Charles Hazelius Sternberg and his sons in the 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.<ref name=NGD95b>{{cite book |last=Norell |first=M. A. |author2=Gaffney, E. S. |author3=Dingus, L. |title=Discovering Dinosaurs in the American Museum of Natural History |publisher=Knopf |location=New York |year=1995 |pages=[https://archive.org/details/discoveringdinos00nore_0/page/154 154–155] |isbn=0-679-43386-4 |url=https://archive.org/details/discoveringdinos00nore_0/page/154 }}</ref> The Sternbergs recovered a second similar specimen from the same area in 1910.<ref name=CB04>{{cite book |last=Dal Sasso |first=Cristiano |author2=Brillante, Giuseppe |year=2004 |title=Dinosaurs of Italy |publisher=Indiana University Press |location=Bloomington and Indianapolis |page=112 |isbn=0-253-34514-6}}</ref> It was not as well-preserved, but also found with skin impressions. They sold this specimen, SM 4036, to the Senckenberg Museum in Germany.<ref name=NGD95b/>

===Canadian discoveries=== ''Edmontosaurus'' itself was coined in 1917 by Lawrence Lambe for two partial skeletons found in the Horseshoe Canyon Formation (formerly the lower Edmonton Formation), along the Red Deer River of southern Alberta.<ref name=LML17>{{cite journal |last=Lambe |first=Lawrence M. |author-link=Lawrence Lambe |year=1917 |title=A new genus and species of crestless hadrosaur from the Edmonton Formation of Alberta |journal=The Ottawa Naturalist |volume=31 |issue=7 |pages=[https://archive.org/details/ottawanaturalist31otta/page/65 65]–73 |url=https://archive.org/details/ottawanaturalist31otta |format=pdf (entire volume, 18 mb) |access-date=2009-03-08}}</ref> The Horseshoe Canyon Formation is older than the rocks in which ''Claosaurus annectens'' was found.<ref name=NCDE11/> Lambe found that his new dinosaur compared best to Cope's ''Diclonius mirabilis''.<ref name=LML17/>

In 1926, Charles Mortram Sternberg named ''Thespesius saskatchewanensis'' for NMC 8509, a skull and partial skeleton from the Wood Mountain plateau of southern Saskatchewan. He had collected this specimen in 1921 from rocks that were assigned to the Lance Formation,<ref name=CMS26>{{cite book|last=Sternberg |first=Charles M. |author-link=Charles Mortram Sternberg |year=1926 |title=A new species of ''Thespesius'' from the Lance Formation of Saskatchewan |publisher=Department of Mines, Geological Survey of Canada |series=Bulletin |volume=44 |pages=77–84}}</ref> now the Frenchman Formation.<ref name=HWF04/> NMC 8509 included an almost complete skull, numerous vertebrae, partial shoulder and hip girdles, and partial back legs, representing the first substantial dinosaur specimen recovered from Saskatchewan. Sternberg opted to assign it to ''Thespesius'' because that was the only hadrosaurid genus known from the Lance Formation at the time.<ref name=CMS26/> At the time, ''T. saskatchewanensis'' was unusual because of its small size, estimated at {{convert|7|to|7.3|m|ft|sp=us}} in length.<ref name=LW42a/>

===Early classifications=== Because of the incomplete understanding of hadrosaurids at the time, following Marsh's death in 1899, ''Claosaurus annectens'' was variously classified as a species of ''Claosaurus'', ''Thespesius'', or ''Trachodon''. Opinions varied greatly, with textbooks and encyclopedias drawing a distinction between the "''Iguanodon''-like" ''Claosaurus annectens'' and the "duck-billed" ''Hadrosaurus'' (based on Cope's ''Diclonius mirabilis''); conversely, Hatcher explicitly identified ''C. annectens'' as synonymous with the hadrosaurid represented by those same duck-billed skulls,<ref name="BSC07"/> the two differentiated only by individual variation or distortion from pressure.<ref name="JBH02"/> Hatcher's revision, published in 1902, was sweeping, as he considered almost all hadrosaurid genera then known as synonyms of ''Trachodon''. This included ''Cionodon'', ''Diclonius'', ''Hadrosaurus'', ''Ornithotarsus'', ''Pteropelyx'', and ''Thespesius'', as well as ''Claorhynchus'' and ''Polyonax'',<ref name="JBH02"/> fragmentary genera now thought to be ceratopsians. Hatcher's work led to a brief consensus until about 1910, when new material from Canada and Montana showed a greater diversity of hadrosaurids than previously suspected.<ref name="BSC07"/> In 1915, Charles W. Gilmore reassessed hadrosaurids, and recommended that ''Thespesius'' should be reintroduced for hadrosaurids from the Lance Formation and rock units of equivalent age, and that ''Trachodon'', based on inadequate material, should be restricted to a hadrosaurid from the older Judith River Formation and its equivalents. In regards to ''Claosaurus annectens'', he recommended that it be considered the same as ''Thespesius occidentalis''.<ref name="CWG15">{{cite journal |last=Gilmore |first=Charles W. |author-link=Charles W. Gilmore |year=1915 |title=On the genus ''Trachodon'' |journal=Science |volume=41 |pages=658–660 |doi=10.1126/science.41.1061.658 |pmid=17747979 |issue=1061|bibcode=1915Sci....41..658G |url=https://zenodo.org/record/1448167 }}</ref> A multiplicity of names resumed, with the AMNH duckbills being known as ''Diclonius mirabilis'', ''Trachodon mirabilis'', ''Trachodon annectens'', ''Claosaurus'', or ''Thespesius''.<ref name=BSC07/>

===''Anatosaurus'' to the present=== thumb|AMNH 5060: a well-preserved specimen of ''E. annectens'' This confusing situation was temporarily resolved in 1942 by Richard Swann Lull and Nelda Wright. In their monograph on hadrosaurian dinosaurs of North America, they opted to settle the questions revolving around the AMNH duckbills, Marsh's ''Claosaurus annectens'', and several other species, by creating a new generic name. They created the new genus ''Anatosaurus'' (meaning "duck lizard", because of its wide, duck-like beak; Latin ''anas'' = duck + Greek ''sauros'' = lizard), and made Marsh's species the type species, calling it ''Anatosaurus annectens''. They also assigned Marsh's ''Trachodon longiceps'' to this genus, a pair of species that had been assigned to ''Thespesius'' under Gilmore's "Lance Formation hadrosaurid" conception (''T. edmontoni'' from Gilmore in 1924 and ''T. saskatchewanensis''), and Cope's ''Diclonius mirabilis''.<ref name=LW42e>Lull and Wright, ''Hadrosaurian Dinosaurs of North America'', pp. 154–164.</ref> 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.<ref name=LW42b/> ''Anatosaurus'' would come to be called the "classic duck-billed dinosaur".<ref name=DFG82>{{cite book |last=Glut |first=Donald F. |title=The New Dinosaur Dictionary |year=1982 |publisher=Citadel Press |location=Secaucus, NJ |isbn=0-8065-0782-9 |page=[https://archive.org/details/newdinosaurdicti00glut/page/57 57] |url=https://archive.org/details/newdinosaurdicti00glut/page/57 }}</ref>

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.<ref name=MKBS75>{{cite book |last=Brett-Surman |first=Michael K. |year=1975 |title=The appendicular anatomy of hadrosaurian dinosaurs |series=M.A. thesis |publisher=University of California |location=Berkeley}}</ref><ref name=MKBS79>{{cite journal |last=Brett-Surman |first=Michael K. |year=1979 |title=Phylogeny and paleobiogeography of hadrosaurian dinosaurs |journal=Nature |volume=277 |issue=5697 |pages=560–562 |doi=10.1038/277560a0|bibcode=1979Natur.277..560B |s2cid=4332144 }}</ref><ref name=MKBS89>{{cite book |last=Brett-Surman |first=Michael K. |year=1989 |title=A revision of the Hadrosauridae (Reptilia: Ornithischia) and their evolution during the Campanian and Maastrichtian |series=Ph.D. dissertation |publisher=George Washington University |location=Washington, D.C. }}</ref> 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.<ref name=DFG82b>{{cite book |last=Glut |first=Donald F. |title=The New Dinosaur Dictionary |year=1982 |publisher=Citadel Press |location=Secaucus, NJ |isbn=0-8065-0782-9 |pages=[https://archive.org/details/newdinosaurdicti00glut/page/49 49, 53] |url=https://archive.org/details/newdinosaurdicti00glut/page/49 }}</ref><ref name=DL83>{{cite book |last=Lambert |first=David |author2=the Diagram Group |title=A Field Guide to Dinosaurs |year=1983 |publisher=Avon Books |location=New York |isbn=0-380-83519-3 |pages=[https://archive.org/details/fieldguidetodino00lamb/page/156 156–161] |url=https://archive.org/details/fieldguidetodino00lamb/page/156 }}</ref> 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.<ref name="DL90">{{cite book |last=Lambert |first=David |author2=the Diagram Group |title=The Dinosaur Data Book |year=1990 |publisher=Avon Books |location=New York |isbn=0-380-75896-2 |pages=[https://archive.org/details/dinosaurdatabook00lamb/page/41 41] |url=https://archive.org/details/dinosaurdatabook00lamb/page/41 }}</ref> 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).<ref name="RCMBS90"/> Because the type species of ''Anatosaurus'' (''A. annectens'') was sunk into ''Edmontosaurus'', the name ''Anatosaurus'' is abandoned as a junior synonym of ''Edmontosaurus''.

thumb|A well-preserved skin impression of the specimen nicknamed "Dakota," which was found in 1999 Of the remaining species of ''Anatosaurus'', ''A. saskatchewanensis'' and ''A. edmontoni'' were assigned to ''Edmontosaurus'' as well,<ref name="WH90">{{cite book |last=Weishampel |first=David B. |author-link=David B. Weishampel |title=The Dinosauria |last2=Horner |first2=Jack R. |author-link2=Jack Horner (paleontologist) |publisher=University of California Press |year=1990 |isbn=0-520-06727-4 |editor-last=Weishampel |editor-first=David B. |edition=1st |location=Berkeley |pages=534–561 |chapter=Hadrosauridae |editor-last2=Osmólska |editor-first2=Halszka |editor-link2=Halszka Osmólska |editor-last3=Dodson |editor-first3=Peter |editor-link3=Peter Dodson}}</ref> and ''A. longiceps'' went to ''Anatotitan'', as either a second species<ref name="olshevsky1991">{{cite book |author=Olshevsky |first=George |title=A Revision of the Parainfraclass Archosauria Cope, 1869, Excluding the Advanced Crocodylia |publisher=Publications Requiring Research |year=1991 |series=Mesozoic Meanderings No. 2 |location=San Diego}}</ref> or as a synonym of ''A. copei''.<ref name=WH90/> ''A. longiceps'' may be a synonym of ''E. annectens'',<ref name=HWF04/> though it has also been treated as a ''nomen dubium'' by some.<ref name=lundgates2006>Lund, E. & Gates, T. (2006). "A historical and biogeographical examination of hadrosaurian dinosaurs." Pp. 263-276 in Lucas, S.G. and Sullivan, R.M. (eds.), ''Late Cretaceous vertebrates from the Western Interior''. New Mexico Museum of Natural History and Science Bulletin '''35'''.</ref>

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''.<ref name="WH90"/> The debate about the proper taxonomy of the ''A. copei'' specimens continues to the present day. Returning to Hatcher's argument of 1902, Jack Horner, David B. Weishampel, and Catherine Forster regarded ''Anatotitan copei'' as representing specimens of ''Edmontosaurus annectens'' with crushed skulls.<ref name=HWF04/> In 2007, another "mummy" was announced. Nicknamed "Dakota," it was discovered in 1999 by Tyler Lyson, and came from the Hell Creek Formation of North Dakota.<ref name="ng">{{cite news |title= Mummified Dinosaur Unveiled |url= http://news.nationalgeographic.com/news/2007/12/photogalleries/dinosaur-pictures/index.html |archive-url= https://web.archive.org/web/20071204110439/http://news.nationalgeographic.com/news/2007/12/photogalleries/dinosaur-pictures/index.html |archive-date= December 4, 2007 |publisher= National Geographic News |date= 2007-12-03 |access-date= 2007-12-03 }}</ref><ref name="wp">{{cite news |last=Lee |first=Christopher |date=2007-12-03 |title=Scientists Get Rare Look at Dinosaur Soft Tissue |url=https://www.washingtonpost.com/wp-dyn/content/story/2007/12/03/ST2007120300591.html |access-date=2007-12-03 |newspaper=The Washington Post}}</ref>

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.<ref name=NCDE11/>

==Description== thumb|left|Scale diagram comparing large adult specimens of ''E. regalis'' (gray) and ''E. annectens'' (green) to a human The skull and skeleton of ''E. annectens'' are very well-known. Edward Drinker Cope estimated the length of one specimen as about {{convert|38|ft}} long, with a skull {{convert|3.87|ft}} long.<ref name=EDC83>{{cite journal |last=Cope |first=Edward D. |author-link=Edward Drinker Cope |year=1883 |title=On the characters of the skull in the Hadrosauridae |journal=Proceedings of the Philadelphia Academy of Natural Sciences |volume=35 |pages=97–107}}</ref>{{Ref|taxonomy|*}} This body length estimate was later revised down to a length of {{Convert|29|ft}}.<ref name=LW42a>{{cite book |last=Lull |first=Richard Swann |author-link=Richard Swann Lull |last2=Wright |first2=Nelda E. |title=Hadrosaurian Dinosaurs of North America |year=1942 |publisher=Geological Society of America |series=Geological Society of America Special Paper '''40''' |page=225 }}</ref> 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.<ref name=HFO09a>{{cite journal |last=Osborn |first=Henry Fairfield |author-link=Henry Fairfield Osborn |year=1909 |title=The Upper Cretaceous iguanodont dinosaurs |journal=Nature |volume=81 |issue=2075 |pages=160–162 |doi=10.1038/081160a0|bibcode=1909Natur..81..160H |doi-access=free }}</ref> A second skeleton currently exhibited next to Cope's specimen, but in a standing posture, is estimated at {{convert|30|ft}} long, with its head {{convert|17|ft}} above the ground.<ref name=HFO09a/> The hip height of this specimen is estimated as approximately {{convert|6.9|ft}}.<ref name=DFG97>{{cite book|chapter=Anatotitan |last=Glut |first=Donald F. |author-link=Donald F. Glut |title=Dinosaurs: The Encyclopedia |url=https://archive.org/details/dinosaursencyclo04dfgl_143 |url-access=limited |year=1997 |publisher=McFarland & Co |location=Jefferson, North Carolina |pages=[https://archive.org/details/dinosaursencyclo04dfgl_143/page/n139 132]–134 |isbn=0-89950-917-7}}</ref> Other sources have estimated the length of ''E. annectens'' as approximately {{convert|39|ft}}.<ref name=WJM70>{{cite journal |last=Morris |first=William J. |year=1970 |title=Hadrosaurian dinosaur bills — morphology and function |journal=Contributions in Science (Los Angeles County Museum of Natural History) |volume=193 |pages=1–14}}</ref><ref name="HDS97">{{cite book |last=Sues |first=Hans-Dieter |author-link=Hans-Dieter Sues |url=https://archive.org/details/isbn_9780253333490/page/338 |title=The Complete Dinosaur |publisher=Indiana University Press |year=1997 |isbn=0-253-33349-0 |editor=Farlow, James O. |location=Bloomington |pages=[https://archive.org/details/isbn_9780253333490/page/338 338] |chapter=Ornithopods |editor2=Brett-Surman, Michael K.}}</ref> Most specimens are somewhat shorter, representing individuals that are not fully grown.<ref name=NCDE11>{{cite journal |last=Campione |first=Nicolás E. |author2=Evans, David C. |year=2011 |title=Cranial growth and variation in ''Edmontosaurs'' (Dinosauria: Hadrosauridae): implications for latest Cretaceous megaherbivore diversity in North America |journal=PLOS ONE |volume=6 |issue=9 |article-number=e25186 |doi=10.1371/journal.pone.0025186 |pmid=21969872 |pmc=3182183|bibcode=2011PLoSO...625186C |doi-access=free }}</ref> Two well-known mounted skeletons, USNM 2414 and YPM 2182, measure {{convert|26.25|ft}} long and {{convert|29.3|ft}} long, respectively.<ref name=LW42a/><ref name=FAL04>{{cite journal |last=Lucas |first=Frederic A. |year=1904 |title=The dinosaur ''Trachodon annectens'' |journal=Smithsonian Miscellaneous Collections |volume=45 |pages=317–320}}</ref> ''E. annectens'' may have weighed about {{Convert|7.3|t|MT|lk=on}} when fully grown.<ref name=DFG97/> thumb|Life reconstruction of ''E. annectens'' based on preserved soft tissues Recently-found specimens that are still under study at the Museum of the Rockies, namely MOR 1142 ("X-rex") and MOR 1609 ("Becky's Giant"),<ref name="census"/> suggest that ''E. annectens'' may have reached lengths of nearly {{convert|15|m|ft}} and weighed {{convert|15.87|MT|ST}},<ref name="PMA2014">{{cite journal |last1=Prieto-Márquez |first1=Albert |year=2014 |title=A juvenile Edmontosaurus from the late Maastrichtian (Cretaceous) of North America: Implications for ontogeny and phylogenetic inference in saurolophine dinosaurs |journal=Cretaceous Research |volume=50 |pages=282–303 |bibcode=2014CrRes..50..282P |doi=10.1016/j.cretres.2014.05.003}}</ref><ref>{{Cite journal |last1=Jiménez-Moreno |first1=Francisco Javier |last2=Ramírez-Velasco |first2=Ángel Alejandro |last3=Ocampo-Cornejo |first3=Patricio |last4=Velázquez-Castro |first4=Jorge |last5=Palomino-Merino |first5=Rodolfo |date=2025-07-02 |title=First Population Analysis in Hadrosauroid dinosaurs (Ornithopoda: Iguanodontia: Hadrosauroidea) |journal=Evolving Earth |at=100072 |issue=in press |doi=10.1016/j.eve.2025.100072 |issn=2950-1172 |doi-access=free}}</ref> 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.<ref name="census">{{cite journal |last1=Horner |first1=J.R. |author-link=Jack Horner (paleontologist) |last2=Goodwin |first2=M.B. |last3=Myhrvold |first3=N. |date=2011 |title=Dinosaur Census Reveals Abundant ''Tyrannosaurus'' and Rare Ontogenetic Stages in the Upper Cretaceous Hell Creek Formation (Maastrichtian), Montana, USA |journal=PLOS ONE |volume=6 |issue=2 |article-number=e16574 |bibcode=2011PLoSO...616574H |doi=10.1371/journal.pone.0016574 |pmc=3036655 |pmid=21347420 |doi-access=free}}</ref> 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 {{convert|36|-|39|ft}} in length and {{convert|5.6|MT|ST}} in average asymptotic body mass, while the largest individuals measured more than {{convert|6|MT|ST}} and even up to {{convert|6.6|-|7|MT|ST}}, based on the comparison between various specimens of different sizes from the Ruth Mason Dinosaur Quarry and other specimens from different localities.<ref name="Wosik2022">{{cite journal|last1=Wosik|first1=M.|last2=Evans|first2=D.C.|year=2022|title=Osteohistological and taphonomic life-history assessment of ''Edmontosaurus annectens'' (Ornithischia: Hadrosauridae) from the Late Cretaceous (Maastrichtian) Ruth Mason dinosaur quarry, South Dakota, United States, with implication for ontogenetic segregation between juvenile and adult hadrosaurids|journal=Journal of Anatomy|volume=241|issue=2|pages=272–296|doi=10.1111/joa.13679|pmid=35801524|pmc=9296034 |s2cid=250357069}}</ref> thumb|left|Tail vertebrae 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.<ref name=EDC83/> The skull was proportionally longer and lower than in any other known hadrosaurid. The toothless portion of the anterior mandible{{Ref|diastema|*}} was also relatively longer than in any hadrosaur.<ref name="RCMBS90">{{cite book |last=Chapman |first=Ralph E. |url=https://archive.org/details/dinosaursystemat00carp |title=Dinosaur Systematics: Perspectives and Approaches |author2=Brett-Surman, Michael K. |publisher=Cambridge University Press |year=1990 |isbn=0-521-43810-1 |editor-last=Carpenter |editor-first=Kenneth |editor-link=Kenneth Carpenter |location=Cambridge |pages=[https://archive.org/details/dinosaursystemat00carp/page/n187 163]–177 |chapter=Morphometric observations on hadrosaurid ornithopods |editor-last2=Currie |editor-first2=Philip J. |editor-link2=Philip J. Currie |url-access=limited}}</ref> The extreme length and breadth did not appear until an individual reached maturity, so many specimens lack the distinctive shape.<ref name=NCDE11/> The bones surrounding the large openings for the nostrils formed deep pockets around the openings. The eye sockets were rectangular and longer front to back than they were top to bottom, although this may have been exaggerated by postmortem crushing. The skull roof was flat and lacked a bony crest like that of ''E. regalis''. The quadrate bone that formed the articulation with the lower jaw was distinctly curved. The lower jaw was long, straight, and lacking the downward curve seen in other hadrosaurids, as well as possessing a heavy ridge running its length. The predentary was wide and shovel-like.<ref name=LW42b>Lull and Wright, ''Hadrosaurian Dinosaurs of North America'', pp. 157-159.</ref> The ridge on the lower jaw may have reinforced the long, slender structure.<ref name=LW42c>Lull and Wright, ''Hadrosaurian Dinosaurs of North America'', pp. 163-164.</ref>

As mounted, the vertebral column of ''E. annectens'' includes twelve neck, twelve back, nine sacral, and at least thirty tail vertebrae.<ref name=LW42b/> The limb bones were longer and more lightly built than those of other hadrosaurids of comparable size. ''E. annectens'' had a distinctive pelvis, based on the proportions and form of the pubis bone.<ref name=RCMBS90/> ''E. annectens'', like other hadrosaurids, could move both on two legs and on four legs. It probably preferred to forage for food on four legs, but ran on two.<ref name="HWF04">{{cite book |last=Horner |first=John R. |author-link=Jack Horner (paleontologist) |url=https://archive.org/details/dinosauriandedit00weis |title=The Dinosauria |author2=Weishampel |first2=David B. |author-link2=David B. Weishampel |last3=Forster |first3=Catherine A. |author-link3=Catherine Forster |publisher=University of California Press |year=2004 |isbn=0-520-24209-2 |editor-last=Weishampel |editor-first=David B. |edition=2nd |location=Berkeley |pages=[https://archive.org/details/dinosauriandedit00weis/page/n456 438]–463 |chapter=Hadrosauridae |editor-last2=Osmólska |editor-first2=Halszka |editor-link2=Halszka Osmólska |editor-last3=Dodson |editor-first3=Peter |editor-link3=Peter Dodson |url-access=limited}}</ref> Henry Fairfield Osborn used the skeletons in the American Museum of Natural History to portray both quadrupedal and bipedal stances for ''E. annectens''.<ref name=HFO09a/>

In 2025, Sereno and colleagues described two well-preserved "mummy" specimens from Wyoming, revealing that ''Edmontosaurus annectens'' bore a fleshy crest, just like its older relative ''Edmontosaurus regalis'', over the neck and back, in addition to an interdigitating row of spikes over the hips and tail. The toes of the hindlimb are capped with large hooves.<ref name="sereno2026">{{cite journal |last1=Sereno |first1=Paul C. |last2=Saitta |first2=Evan T. |last3=Vidal |first3=Daniel |last4=Myhrvold |first4=Nathan |last5=Real |first5=María Ciudad |last6=Baumgart |first6=Stephanie L. |last7=Bop |first7=Lauren L. |last8=Keillor |first8=Tyler M. |last9=Eriksen |first9=Marcus |last10=Derstler |first10=Kraig |year= 2026|title=Duck-billed dinosaur fleshy midline and hooves reveal terrestrial clay-template 'mummification' |journal=Science |volume=391 | issue=6780 |article-number= eadw3536 |doi=10.1126/science.adw3536}}</ref><ref>{{Cite web |last=Dunham |first=Will |date=October 23, 2025 |title=Wyoming dinosaur 'mummies' reveal a surprise: hoofed feet |url=https://www.reuters.com/science/wyoming-dinosaur-mummies-reveal-surprise-hoofed-feet-2025-10-23/ |url-status=live |access-date=October 27, 2025 |website=Reuters}}</ref><ref>{{Cite news |last=Tamisiea |first=Jack |date=2025-10-23 |title=Two New Dinosaur Fossils Emerge From the 'Mummy Zone' |url=https://www.nytimes.com/2025/10/23/science/dinosaur-mummies-wyoming.html |access-date=2025-10-27 |work=The New York Times |language=en}}</ref><ref>{{Cite web |last=Niewijk |first=Grace |date=2025-10-24 |title=UChicago paleontologists unveil duck-billed dinosaur 'mummies' {{!}} University of Chicago News |url=https://news.uchicago.edu/story/uchicago-paleontologists-unveil-duck-billed-dinosaur-mummies |access-date=2025-10-27 |website=news.uchicago.edu |language=en}}</ref>

==Classification== [[File:Edmontosaurus annectens USNM 2414.jpg|thumb|''E. annectens'' holotype, Smithsonian National Museum of Natural History]] thumb|upright|Most known complete ''Edmontosaurus'' skulls (''E. annectens'' from lower middle to right). ''E. annectens'' was a saurolophine, or "flat-headed", hadrosaurid. This group was historically known as Hadrosaurinae.<ref name=PM2010>{{cite journal |last=Prieto-Márquez |first=Alberto |year=2010 |title=Global phylogeny of Hadrosauridae (Dinosauria: Ornithopoda) using parsimony and Bayesian methods |journal=Zoological Journal of the Linnean Society |volume=159 |issue=2 |pages=435–502 |doi=10.1111/j.1096-3642.2009.00617.x|doi-access=free }}</ref> Species now considered to be synonymous with ''Edmontosaurus annectens'' were long recognized as closely related to both the genus<ref name=LML20>{{cite journal |last=Lambe |first=Lawrence M. |author-link=Lawrence Lambe |year=1920 |title=The hadrosaur ''Edmontosaurus'' from the Upper Cretaceous of Alberta |journal=Department of Mines, Geological Survey Memoirs |volume=120 |pages=1–79}}</ref> and the species.<ref name=JBH02>{{cite journal |last=Hatcher |first=John B. |author-link=John Bell Hatcher |title=The genus and species of the Trachodontidae (Hadrosauridae, Claosauridae) Marsh |journal=Annals of the Carnegie Museum |year=1902 |volume=1 |issue=3 |pages=377–386 |doi=10.5962/p.331063 |s2cid=251485258 |doi-access=free }}</ref> 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,<ref name=JBH02/> to Jack Horner, David B. Weishampel, and Catherine Forster in 2004,<ref name=HWF04/> and most recently Nicolás Campione and David Evans,<ref name=NCDE11/> have proposed that the large, flat-headed specimens most recently classified as ''Anatotitan copei'' belong to ''E. annectens''.

''E. annectens'' was also historically classified in an independent genus, ''Anatosaurus'', following the influential 1942 revision of Hadrosauridae by Richard Swann Lull and Nelda Wright, until it was reclassified as a species of ''Edmontosaurus'' by Michael K. Brett-Surman.<ref name=RCMBS90/> With the discovery that ''A. copei'' and ''E. annectens'' most likely represent the same species, some paleontologists have proposed using ''Anatosaurus'' as a valid genus name for ''E. annectens''.<ref name="Holtz2008"/>

The cladogram below follows Godefroit ''et al.'' (2012) analysis.<ref>{{cite journal | last1 = Godefroit | first1 = P. | last2 = Bolotsky | first2 = Y. L. | last3 = Lauters | first3 = P. | year = 2012 | title = A New Saurolophine Dinosaur from the Latest Cretaceous of Far Eastern Russia | journal = PLOS ONE | volume = 7 | issue = 5| article-number = e36849 | doi = 10.1371/journal.pone.0036849 | pmid = 22666331 | pmc=3364265| bibcode = 2012PLoSO...736849G | doi-access = free }}</ref>

{{clade| style=font-size:85%;line-height:85% |1={{clade |1=''Bactrosaurus'' |label2=Hadrosauridae |2={{clade |label1=Hadrosaurinae |1={{clade |1=''Hadrosaurus'' |2=''Lophorhothon'' }} |label2=Saurolophidae |2={{clade |1=Lambeosaurinae |label2=Saurolophinae |2={{clade |1=''Wulagasaurus'' |2={{clade |label1=Brachylophosaurini |1={{clade |1=''Acristavus'' |2={{clade |1=''Maiasaura'' |2=''Brachylophosaurus'' }} }} |2={{clade |1=''Kritosaurus'' |2={{clade |1={{clade |1=''Gryposaurus latidens'' |2={{clade |1=''Gryposaurus notabilis'' |2=''Gryposaurus monumentensis'' }} }} |2={{clade |label1=Saurolophini |1={{clade |1=''Prosaurolophus'' |2={{clade |1=''Saurolophus angustirostris'' |2=''Saurolophus osborni'' }} }} |label2=Edmontosaurini |2={{clade |1={{clade |1=''Kerberosaurus'' |2=''Kundurosaurus'' }} |2={{clade |1='''''Edmontosaurus annectens''''' |2=''Edmontosaurus regalis'' }} }} }} }} }} }} }} }} }} }} }}

==Paleobiology== thumb|Close-up of teeth 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 {{Convert|13|ft|0}} above the ground. Like other hadrosaurs, they could have moved both bipedally and quadrupedally.<ref name=HWF04/>

The extensive depressions surrounding its nasal openings may have hosted nasal diverticula. These postulated diverticula would have taken the form of inflatable soft-tissue sacs. Such sacs could be used for both visual and auditory signals.<ref name=JAH75>{{cite journal |last=Hopson |first=James A. |year=1975 |title=The evolution of cranial display structures in hadrosaurian dinosaurs |journal=Paleobiology |volume=1 |issue=1 |pages=21–43 |doi=10.1017/S0094837300002165 |bibcode=1975Pbio....1...21H |s2cid=88689241 }}</ref>

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.<ref>{{Cite web |last=Naish |first=Darren |author-link=Darren Naish |date=April 17, 2018 |title=Enough with the "Duck-Billed Dinosaurs" |url=https://www.scientificamerican.com/blog/tetrapod-zoology/enough-with-the-duck-billed-dinosaurs |access-date=August 24, 2024 |website=scientificamerican.com}}</ref><ref>{{Cite web |last=Black |first=Riley |date=June 14, 2012 |title=Shovel-Beaked, Not Duck-Billed |url=https://www.smithsonianmag.com/science-nature/shovel-beaked-not-duck-billed-122016600/ |access-date=August 24, 2024 |website=smithsonianmag.com}}</ref>

===Growth=== [[File:Edmontosaurus annectens HMNS.jpg|alt=|left|thumb|Mounted skeletons of a juvenile and adult ''E. annectens'' at the Houston Museum of Natural Science, 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.<ref name=NCDE11/> 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.<ref name="Wosik2022"/>

==Paleoecology== [[File:DMNS Edmontosaurus.png|thumb|The damage to the tail vertebrae of this ''E. annectens'' skeleton (on display at the Denver Museum of Nature and Science) indicates that it may have been bitten by a ''Tyrannosaurus''.]] 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.<ref name=NCDE11/>

The Lancian time interval was the last interval before the Cretaceous–Paleogene extinction event that killed off the non-avian dinosaurs. ''Edmontosaurus'' was one of the most common dinosaurs of the interval. Robert Bakker reports that it made up one-seventh of the large dinosaur sample, with most of the remaining five-sixths made up of ''Triceratops''.<ref name="RTB86b">Bakker, Robert T. (1986). ''The Dinosaur Heresies''. p. 438.</ref> The coastal plain ''Triceratops''–''Edmontosaurus'' association, dominated by ''Triceratops'', extended from Colorado to Saskatchewan.<ref name="DJC01">{{cite book |last=Lehman |first=Thomas M. |title=Mesozoic Vertebrate Life |publisher=Indiana University Press |year=2001 |isbn=0-253-33907-3 |editor=Tanke, Darren |location=Bloomington and Indianapolis |pages=[https://archive.org/details/mesozoicvertebra0000unse/page/310 310–328] |chapter=Late Cretaceous dinosaur provinciality |editor-last2=Carpenter |editor-first2=Kenneth |editor-link2=Kenneth Carpenter |chapter-url=https://archive.org/details/mesozoicvertebra0000unse/page/310}}</ref> 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''.<ref name="DBWetal04">{{cite book |last1=Weishampel |first1=David B. |author-link=David B. Weishampel |url=https://archive.org/details/dinosauriandedit00weis |title=The Dinosauria |last2=Barrett |first2=Paul M. |last3=Coria |first3=Rodolfo A. |last4=Le Loueff |first4=Jean |author5=Xu Xing |author6=Zhao Xijin |last7=Sahni |first7=Ashok |last8=Gomani |first8=Elizabeth M.P. |last9=Noto |first9=Christopher N. |publisher=University of California Press |year=2004 |isbn=0-520-24209-2 |editor-last1=Weishampel |editor-first1=David B. |edition=2nd |location=Berkeley |pages=[https://archive.org/details/dinosauriandedit00weis/page/n535 517]–606 |chapter=Dinosaur distribution |editor-last2=Dodson |editor-first2=Peter |editor-link2=Peter Dodson |editor-last3=Osmólska |editor-first3=Halszka |editor-link3=Halszka Osmólska |url-access=limited}}</ref><ref name="HCFF">{{cite web |last=Bigelow |first=Phillip |date=July 21, 2010 |title=Cretaceous "Hell Creek Faunal Facies"; Late Maastrichtian |url=http://www.scn.org/~bh162/hellcreek2.html |archive-url=https://web.archive.org/web/20070124225117/http://www.scn.org/~bh162/hellcreek2.html |archive-date=January 24, 2007 |access-date=November 8, 2011 |work=Seattle Community Network}}</ref> thumb|left|The Hell Creek Formation is well exposed in the badlands in the vicinity of Fort Peck Reservoir. 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.<ref name=DAR89b>Russell, Dale A. (1989). ''An Odyssey in Time: Dinosaurs of North America''. pp. 175–180.</ref>

The Lance Formation, as typified by exposures approximately {{convert|62|mi}} 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.<ref name=DAR89c>Russell, Dale A. (1989). ''An Odyssey in Time: Dinosaurs of North America''. pp. 180–181.</ref> The climate was humid and subtropical, with conifers, palmettos, and ferns in the swamps, and conifers, ash, live oak, and shrubs in the forests.<ref name="KD94">{{cite book |last=Derstler |first=Kraig |title=The Dinosaurs of Wyoming |publisher=Wyoming Geological Association |year=1994 |editor-last=Nelson |editor-first=Gerald E. |series=Wyoming Geological Association Guidebook, 44th Annual Field Conference |pages=127–146 |chapter=Dinosaurs of the Lance Formation in eastern Wyoming}}</ref> 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".<ref name=DAR89c/>

==See also== {{Portal|Dinosaurs}} * Timeline of hadrosaur research

==Notes== {{Ref|taxonomy|*}} Many of the original references deal with specimens or species that were not assigned to ''E. annectens'' until later. This is particularly true with the specimens long known, chronologically, as ''Diclonius mirabilis'', ''Anatosaurus copei'', and ''Anatotitan copei''. {{note|diastema|*}} This toothless section is also known as a ''diastema''.

==References== {{Reflist|2}}

Category:Saurolophinae Category:Dinosaur species Category:Maastrichtian dinosaurs Category:Hell Creek fauna Category:Lance Formation Category:Frenchman Formation Category:Scollard Formation Category:Taxa named by Othniel Charles Marsh Category:Fossil taxa described in 1890 Category:Dinosaurs of Canada Category:Dinosaurs of the United States