HOME

TheInfoList



OR:

''Edmontosaurus annectens'' (meaning "connected lizard from
Edmonton Edmonton ( ) is the capital city of the Canadian province of Alberta. Edmonton is situated on the North Saskatchewan River and is the centre of the Edmonton Metropolitan Region, which is surrounded by Alberta's central region. The city anc ...
") is a
species In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...
of flat-headed and duck-billed (
hadrosaurid Hadrosaurids (), or duck-billed dinosaurs, are members of the ornithischian family Hadrosauridae. This group is known as the duck-billed dinosaurs for the flat duck-bill appearance of the bones in their snouts. The ornithopod family, which incl ...
)
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23  million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
from the very end of the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
Period, in what is now North America. Remains of ''E. annectens'' have been preserved in the Frenchman,
Hell Creek The Hell Creek Formation is an intensively studied division of mostly Upper Cretaceous and some lower Paleocene rocks in North America, named for exposures studied along Hell Creek, near Jordan, Montana. The formation stretches over portions of ...
, and
Lance Formation The Lance (Creek) Formation is a division of Late Cretaceous (dating to about 69 - 66 Ma) rocks in the western United States. Named after Lance Creek, Wyoming, the microvertebrate fossils and dinosaurs represent important components of the lat ...
s. All of these formations are dated to the late
Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interv ...
stage of the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', ...
Period, representing the last three million years before the extinction of the dinosaurs (between 68 and 66 
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago). ...
Holtz, Thomas R. Jr. (2012) ''Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages,'
Winter 2011 Appendix.
/ref>). ''E. annectens'' is also found in the
Laramie Formation The Laramie Formation is a geologic formation of Late Cretaceous (Maastrichtian) age, named by Clarence King in 1876 for exposures in northeastern Colorado, in the United States.King, C. 1876. Report of the Geological Exploration of the Fortieth ...
, and
magnetostratigraphy Magnetostratigraphy is a geophysical correlation technique used to date sedimentary and volcanic sequences. The method works by collecting oriented samples at measured intervals throughout the section. The samples are analyzed to determine their ' ...
suggests an age of 69-68 Ma for the Laramie Formation.*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. ''Edmontosaurus annectens'' is known from numerous specimens, including at least twenty partial to complete skulls, discovered in the U.S. states of
Montana Montana () is a U.S. state, state in the Mountain states, Mountain West List of regions of the United States#Census Bureau-designated regions and divisions, division of the Western United States. It is bordered by Idaho to the west, North ...
,
South Dakota South Dakota (; Sioux language, Sioux: , ) is a U.S. state in the West North Central states, North Central region of the United States. It is also part of the Great Plains. South Dakota is named after the Lakota people, Lakota and Dakota peo ...
,
North Dakota North Dakota () is a U.S. state in the Upper Midwest, named after the indigenous Dakota Sioux. North Dakota is bordered by the Canadian provinces of Saskatchewan and Manitoba to the north and by the U.S. states of Minnesota to the east, S ...
,
Wyoming Wyoming () is a state in the Mountain West subregion of the Western United States. It is bordered by Montana to the north and northwest, South Dakota and Nebraska to the east, Idaho to the west, Utah to the southwest, and Colorado to the sou ...
and
Colorado Colorado (, other variants) is a state in the Mountain states, Mountain West subregion of the Western United States. It encompasses most of the Southern Rocky Mountains, as well as the northeastern portion of the Colorado Plateau and the wes ...
and the
Canadian Canadians (french: Canadiens) are people identified with the country of Canada. This connection may be residential, legal, historical or cultural. For most Canadians, many (or all) of these connections exist and are collectively the source of ...
province of
Saskatchewan Saskatchewan ( ; ) is a province in western Canada, bordered on the west by Alberta, on the north by the Northwest Territories, on the east by Manitoba, to the northeast by Nunavut, and on the south by the U.S. states of Montana and North ...
. It had an extremely long and low
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, t ...
and was a large animal, up to approximately in length and in average asymptotic body mass, and may have been even larger. ''E. annectens'' exhibits one of the most striking examples of the "duckbill" snout common to hadrosaurs. It has a long taxonomic history, and specimens have at times been classified in the genera '' Diclonius'', '' Trachodon'', '' Hadrosaurus'', '' Claosaurus'', ''
Thespesius ''Thespesius'' (meaning "wondrous one") is a dubious genus of hadrosaurid dinosaur from the late Maastrichtian-age Upper Cretaceous Lance Formation of South Dakota. History In 1855 geologist Ferdinand Vandeveer Hayden sent a number of fossils ...
'', ''Anatosaurus'' and ''Anatotitan'', before being grouped together in ''
Edmontosaurus ''Edmontosaurus'' ( ) (meaning "lizard from Edmonton") 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 ro ...
''.


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 ''Thespesius'' (meaning "wondrous one") is a dubious genus of hadrosaurid dinosaur from the late Maastrichtian-age Upper Cretaceous Lance Formation of South Dakota. History In 1855 geologist Ferdinand Vandeveer Hayden sent a number of fossils ...
'', and '' Trachodon'', as well as ''
Edmontosaurus ''Edmontosaurus'' ( ) (meaning "lizard from Edmonton") 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 ro ...
''. References predating the 1980s typically use ''Anatosaurus'', ''Claosaurus'', ''Diclonius'', ''Thespesius'', or ''Trachodon'' for ''E. annectens'' fossils, depending on author and date.


Cope's ''Diclonius mirabilis''

The history of ''E. annectens'' predates both ''Edmontosaurus'' and ''annectens''. The first good specimen, the former
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of seve ...
(specimen on which a taxonomic name is based) 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 for American
paleontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of foss ...
Edward Drinker Cope Edward Drinker Cope (July 28, 1840 – April 12, 1897) was an American zoologist, paleontologist, comparative anatomist, herpetologist, and ichthyologist. Born to a wealthy Quaker family, Cope distinguished himself as a child prodigy intereste ...
. This specimen, found in
Hell Creek Formation The Hell Creek Formation is an intensively studied division of mostly Upper Cretaceous and some lower Paleocene rocks in North America, named for exposures studied along Hell Creek, near Jordan, Montana. The formation stretches over portions of ...
rocks, came from northeast of the
Black Hills The Black Hills ( lkt, Ȟe Sápa; chy, Moʼȯhta-voʼhonáaeva; hid, awaxaawi shiibisha) is an isolated mountain range rising from the Great Plains of North America in western South Dakota and extending into Wyoming, United States. Black Elk P ...
of
South Dakota South Dakota (; Sioux language, Sioux: , ) is a U.S. state in the West North Central states, North Central region of the United States. It is also part of the Great Plains. South Dakota is named after the Lakota people, Lakota and Dakota peo ...
and originally had extensive skin impressions. It was missing most of the pelvis and part of the torso due to a stream cutting through it. The bill had impressions of a horny sheath with a tooth-like series of interlocking points on the upper and lower jaws. When describing this specimen
AMNH The American Museum of Natural History (abbreviated as AMNH) is a natural history museum on the Upper West Side of Manhattan in New York City. In Theodore Roosevelt Park, across the street from Central Park, the museum complex comprises 26 inter ...
 5730, Cope assigned it to the species ''
Diclonius mirabilis ''Trachodon'' (meaning "rough tooth") is a dubious genus of hadrosaurid dinosaur based on teeth from the Campanian-age Upper Cretaceous Judith River Formation of Montana, U.S.Leidy, J. (1856). "Notice of remains of extinct reptiles and fishes ...
''. 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 and published by
Joseph Leidy Joseph Mellick Leidy (September 9, 1823 – April 30, 1891) was an American paleontologist, parasitologist and anatomist. Leidy was professor of anatomy at the University of Pennsylvania, later was a professor of natural history at Swarthmore ...
. 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. Leidy had come to recognize that his ''Trachodon'' was based on the remains of multiple kinds of dinosaurs, and though he had made some attempts to revise the genus, he had not yet made any formal declaration of his intentions. Cope's description promoted hadrosaurids as amphibious, contributing to this long-time image. His reasoning was that the teeth of the lower jaw were weakly connected to the bone and liable to break off if used to consume terrestrial food, and he described the beak as weak as well. However, aside from misidentifying several of the skull bones, by chance the lower jaws were missing the walls supporting the teeth from the inside; the teeth were actually well-supported.Lull and Wright, ''Hadrosaurian Dinosaurs of North America'', pp. 43. Cope intended to describe the skeleton as well as the skull, but his promised paper never appeared. It was purchased for the American Museum of Natural History in 1899, where it acquired its present designation AMNH 5730. Several years after Cope's description, his rival
Othniel Charles Marsh Othniel Charles Marsh (October 29, 1831 – March 18, 1899) was an American professor of Paleontology in Yale College and President of the National Academy of Sciences. He was one of the preeminent scientists in the field of paleontology. Among ...
published on a sizable lower jaw recovered by John Bell Hatcher in 1889 from
Lance Formation The Lance (Creek) Formation is a division of Late Cretaceous (dating to about 69 - 66 Ma) rocks in the western United States. Named after Lance Creek, Wyoming, the microvertebrate fossils and dinosaurs represent important components of the lat ...
rocks in
Niobrara County, Wyoming Niobrara County is a county in the U.S. state of Wyoming. As of the 2020 United States Census, the population was 2,467, making it the least populous county in Wyoming. Its county seat is Lusk. Its eastern boundary abuts the west lines of the ...
. Marsh named this partial jaw ''Trachodon longiceps''. 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 In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
, or tooth-bearing bone of the lower jaw, that is long, whereas Marsh's dentary is estimated at long. A second mostly complete skeleton (AMNH 5886) was found in 1904 in Hell Creek Formation rocks at Crooked Creek in central
Montana Montana () is a U.S. state, state in the Mountain states, Mountain West List of regions of the United States#Census Bureau-designated regions and divisions, division of the Western United States. It is bordered by Idaho to the west, North ...
by Oscar Hunter, a rancher. 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 A pistol is a handgun, more specifically one with the chamber integral to its gun barrel, though in common usage the two terms are often used interchangeably. The English word was introduced in , when early handguns were produced in Europe, ...
, and later sold it to Brown, who excavated it for the American Museum of Natural History in 1906. This specimen had a nearly complete
vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates, Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characterist ...
l 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''. 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 Henry Fairfield Osborn, Sr. (August 8, 1857 – November 6, 1935) was an American paleontologist, geologist and eugenics advocate. He was the president of the American Museum of Natural History for 25 years and a cofounder of the American E ...
described the tableau as representing the two animals feeding along a
marsh A marsh is a wetland that is dominated by herbaceous rather than woody plant species.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p Marshes can often be found a ...
, the standing individual having been startled by the approach of a ''
Tyrannosaurus ''Tyrannosaurus'' is a genus of large theropod dinosaur. The species ''Tyrannosaurus rex'' (''rex'' meaning "king" in Latin), often called ''T. rex'' or colloquially ''T-Rex'', is one of the best represented theropods. ''Tyrannosaurus'' live ...
''. Impressions of appropriate plant remains and shells based on associated fossils were included on the base of the group, including
ginkgo ''Ginkgo'' is a genus of non-flowering seed plants. The scientific name is also used as the English name. The order to which it belongs, Ginkgoales, first appeared in the Permian, 270 million years ago, and is now the only living genus withi ...
leaves, '' Sequoia'' cones, and
horsetail ''Equisetum'' (; horsetail, snake grass, puzzlegrass) is the only living genus in Equisetaceae, a family of ferns, which reproduce by spores rather than seeds. ''Equisetum'' is a " living fossil", the only living genus of the entire subclass ...
rushes.


Marsh's ''Claosaurus annectens''

The species now known as ''Edmontosaurus annectens'' was named in 1892 as ''Claosaurus annectens'' by
Othniel Charles Marsh Othniel Charles Marsh (October 29, 1831 – March 18, 1899) was an American professor of Paleontology in Yale College and President of the National Academy of Sciences. He was one of the preeminent scientists in the field of paleontology. Among ...
. This species is based on
USNM The National Museum of Natural History is a natural history museum administered by the Smithsonian Institution, located on the National Mall in Washington, D.C., United States. It has free admission and is open 364 days a year. In 2021, with 7. ...
 2414, a partial skull-roof and skeleton, with a second skull and skeleton, YPM 2182, designated the
paratype In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype nor a syntype). O ...
. Both were collected in 1891 by John Bell Hatcher from the late Maastrichtian-age Upper Cretaceous
Lance Formation The Lance (Creek) Formation is a division of Late Cretaceous (dating to about 69 - 66 Ma) rocks in the western United States. Named after Lance Creek, Wyoming, the microvertebrate fossils and dinosaurs represent important components of the lat ...
of
Niobrara County Niobrara County is a county in the U.S. state of Wyoming. As of the 2020 United States Census, the population was 2,467, making it the least populous county in Wyoming. Its county seat is Lusk. Its eastern boundary abuts the west lines of the ...
(then part of
Converse County Converse County is a county located in the U.S. state of Wyoming. As of the 2020 United States Census, the population was 13,751. Its county seat is Douglas. History Converse County was created in 1888 by the legislature of the Wyoming Territor ...
),
Wyoming Wyoming () is a state in the Mountain West subregion of the Western United States. It is bordered by Montana to the north and northwest, South Dakota and Nebraska to the east, Idaho to the west, Utah to the southwest, and Colorado to the sou ...
. This species has some historical footnotes attached: it is among the first dinosaurs to receive a skeletal restoration, and is the first hadrosaurid so restored; and YPM 2182 and UNSM 2414 are, respectively, the first and second essentially complete mounted dinosaur skeletons in the United States. YPM 2182 was put on display in 1901, and USNM 2414 in 1904. In the first decade of the twentieth century, two additional important specimens of ''C. annectens'' were recovered. The first, the "
Trachodon mummy The ''Edmontosaurus'' mummy AMNH 5060 is an exceptionally well-preserved fossil of a dinosaur in the collection of the American Museum of Natural History (AMNH). Discovered in 1908 in the United States near Lusk, Wyoming, it was the first dinos ...
" (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 The Natural History Museum in London is a museum that exhibits a vast range of specimens from various segments of natural history. It is one of three major museums on Exhibition Road in South Kensington, the others being the Science Museu ...
, but
Henry Fairfield Osborn Henry Fairfield Osborn, Sr. (August 8, 1857 – November 6, 1935) was an American paleontologist, geologist and eugenics advocate. He was the president of the American Museum of Natural History for 25 years and a cofounder of the American E ...
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, not as well-preserved but also found with skin impressions. They sold this specimen (SM 4036) to the
Senckenberg Museum The Naturmuseum Senckenberg is a museum of natural history, located in Frankfurt am Main. It is the second-largest of its type in Germany. The museum contains a large and diverse collection of birds with 90,000 bird skins, 5,050 egg sets, 17,0 ...
in Germany.


Canadian discoveries

''Edmontosaurus'' itself was coined in 1917 by
Lawrence Lambe Lawrence Morris Lambe (August 27, 1863 – March 12, 1919) was a Canadian geologist, palaeontologist, and ecologist from the Geological Survey of Canada (GSC). His published work, describing the diverse and plentiful dinosaur discoveries from ...
for two partial skeletons found in the
Horseshoe Canyon Formation The Horseshoe Canyon Formation is a stratigraphic unit of the Western Canada Sedimentary Basin in southwestern Alberta. It takes its name from Horseshoe Canyon, an area of badlands near Drumheller. The Horseshoe Canyon Formation is part of the ...
(formerly the lower Edmonton Formation) along the
Red Deer River The Red Deer River is a river in Alberta and a small portion of Saskatchewan, Canada. It is a major tributary of the South Saskatchewan River and is part of the larger Saskatchewan-Nelson system that empties into Hudson Bay. Red Deer Riv ...
of southern Alberta, Canada. The Horseshoe Canyon Formation is older than the rocks in which ''Claosaurus annectens'' was found. Lambe found that his new dinosaur compared best to Cope's ''Diclonius mirabilis''. In 1926, Charles Mortram Sternberg named ''Thespesius saskatchewanensis'' for NMC 8509, a skull and partial skeleton from the Wood Mountain plateau of southern
Saskatchewan Saskatchewan ( ; ) is a province in western Canada, bordered on the west by Alberta, on the north by the Northwest Territories, on the east by Manitoba, to the northeast by Nunavut, and on the south by the U.S. states of Montana and North ...
. He had collected this specimen in 1921, from rocks that were assigned to the Lance Formation, now the Frenchman Formation. NMC 8509 included an almost complete skull, numerous vertebrae, partial shoulder and hip girdles, and partial hind limbs, 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. At the time, ''T. saskatchewanensis'' was unusual because of its small size, estimated at in length.


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; textbooks and encyclopedias drew a distinction between the "''
Iguanodon ''Iguanodon'' ( ; meaning 'iguana-tooth'), named in 1825, is a genus of iguanodontian dinosaur. While many species have been classified in the genus ''Iguanodon'', dating from the late Jurassic Period to the early Cretaceous Period of Asia, ...
''-like" ''Claosaurus annectens'' and the "duck-billed" ''Hadrosaurus'' (based on Cope's ''Diclonius mirabilis''), while Hatcher explicitly identified ''C. annectens'' as synonymous with the hadrosaurid represented by those same duck-billed skulls, the two differentiated only by individual variation or distortion from pressure. Hatcher's revision, published in 1902, was sweeping: he considered almost all hadrosaurid genera then known as synonyms of ''Trachodon''. This included '' Cionodon'', ''Diclonius'', ''Hadrosaurus'', '' Ornithotarsus'', ''
Pteropelyx ''Pteropelyx'' (meaning "winged pelvis") is a Nomen dubium, dubious genus of Late Cretaceous hadrosaurid dinosaur from the Judith River Formation of Montana, named by Edward Drinker Cope in 1889. Historically, several species were assigned to it, ...
'', and ''Thespesius'', as well as '' Claorhynchus'' and ''
Polyonax ''Polyonax'' (meaning "master over many") was a genus of ceratopsid dinosaur from the late Maastrichtian-age Upper Cretaceous Denver Formation of Colorado, United States. Founded upon poor remains, it is today regarded as a dubious name. Hi ...
'', fragmentary genera now thought to be horned dinosaurs. 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. Charles W. Gilmore in 1915 reassessed hadrosaurids and recommended that ''Thespesius'' 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 The Judith River Formation is a fossil-bearing geologic formation in Montana, and is part of the Judith River Group. It dates to the Late Cretaceous, between 79 and 75.3 million years ago, corresponding to the "Judithian" land vertebrate age. I ...
and its equivalents. In regards to ''Claosaurus annectens'', he recommended that it be considered the same as ''Thespesius occidentalis''. A multiplicity of names resumed, with the American Museum duckbills being known as ''Diclonius mirabilis'', ''Trachodon mirabilis'', ''Trachodon annectens'', ''Claosaurus'', or ''Thespesius''.


''Anatosaurus'' to the present

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 American Museum duckbills, Marsh's ''Claosaurus annectens'', and several other species by creating a new generic name. They created the new genus '' Anatosaurus'' ("duck lizard", because of its wide, duck-like beak;
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power ...
''anas'' = duck +
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
''sauros'' = lizard) and made Marsh's species the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen( ...
, calling it ''Anatosaurus annectens''. To this genus, they also assigned Marsh's ''Trachodon longiceps'', 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''.Lull and Wright, ''Hadrosaurian Dinosaurs of North America'', pp. 154–164. Lull and Wright decided to remove the American Museum 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 A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of seve ...
of the species, with Brown's (AMNH 5886) as the plesiotype. ''Anatosaurus'' would come to be called the "classic duck-billed dinosaur." 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 The International Commission on Zoological Nomenclature (ICZN) is an organization dedicated to "achieving stability and sense in the scientific naming of animals". Founded in 1895, it currently comprises 26 commissioners from 20 countries. Orga ...
, 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'' (
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power ...
''anas'' ("duck") and the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
''
Titan Titan most often refers to: * Titan (moon), the largest moon of Saturn * Titans, a race of deities in Greek mythology Titan or Titans may also refer to: Arts and entertainment Fictional entities Fictional locations * Titan in fiction, fictiona ...
'', 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). Because the type species of ''Anatosaurus'' (''A. annectens'') was sunk into ''Edmontosaurus'', the name ''Anatosaurus'' is abandoned as a junior synonym of ''Edmontosaurus''. Of the remaining species of ''Anatosaurus'', ''A. saskatchewanensis'' and ''A. edmontoni'' were assigned to ''Edmontosaurus'' as well, and ''A. longiceps'' went to ''Anatotitan'', as either a second species or as a synonym of ''A. copei''. ''A. longiceps'' may be a synonym of ''E. annectens'', though it has also been treated as a ''nomen dubium'' by some.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. The conception of ''Edmontosaurus'' that emerged included three valid species: the type ''E. regalis'', ''E. annectens'' (including ''Anatosaurus edmontoni'', emended to ''edmontonensis''), and ''E. saskatchewanensis''. The debate about the proper taxonomy of the ''A. copei'' specimens continues to the present: 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. In 2007 another "mummy" was announced; nicknamed "
Dakota Dakota may refer to: * Dakota people, a sub-tribe of the Sioux ** Dakota language, their language Dakota may also refer to: Places United States * Dakota, Georgia, an unincorporated community * Dakota, Illinois, a town * Dakota, Minnesota ...
", it was discovered in 1999 by
Tyler Lyson Tyler R. Lyson (born 1982 or 1983) is an American paleontologist. He is the discoverer of the dinosaur fossil Dakota, a fossilized mummified hadrosaur. He has done significant research on the evolution of turtles and on the rise of mammals after th ...
, and came from the
Hell Creek Formation The Hell Creek Formation is an intensively studied division of mostly Upper Cretaceous and some lower Paleocene rocks in North America, named for exposures studied along Hell Creek, near Jordan, Montana. The formation stretches over portions of ...
of
North Dakota North Dakota () is a U.S. state in the Upper Midwest, named after the indigenous Dakota Sioux. North Dakota is bordered by the Canadian provinces of Saskatchewan and Manitoba to the north and by the U.S. states of Minnesota to the east, S ...
. In a 2011 study by Nicolás Campione and David Evans, the authors conducted the first-ever
morphometric Morphometrics (from Greek μορϕή ''morphe'', "shape, form", and -μετρία ''metria'', "measurement") or morphometry refers to the quantitative analysis of ''form'', a concept that encompasses size and shape. Morphometric analyses are co ...
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 can be distinguished from skulls of ''E. regalis'' by the elongate snout and other details of skull anatomy.


Description

The skull and skeleton of ''E. annectens'' are well-known.
Edward Drinker Cope Edward Drinker Cope (July 28, 1840 – April 12, 1897) was an American zoologist, paleontologist, comparative anatomist, herpetologist, and ichthyologist. Born to a wealthy Quaker family, Cope distinguished himself as a child prodigy intereste ...
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 , although to be fair to Cope a dozen
vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates, Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characterist ...
e, the
hips In vertebrate anatomy, hip (or "coxa"Latin ''coxa'' was used by Celsus in the sense "hip", but by Pliny the Elder in the sense "hip bone" (Diab, p 77) in medical terminology) refers to either an anatomical region or a joint. The hip region is ...
, 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. The hip height of this specimen is estimated as approximately . 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 The National Museum of Natural History is a natural history museum administered by the Smithsonian Institution, located on the National Mall in Washington, D.C., United States. It has free admission and is open 364 days a year. In 2021, with 7. ...
 2414 and YPM 2182, measure long and long, respectively. ''E. annectens'' may have weighed about 6.6 
metric ton The tonne ( or ; symbol: t) is a unit of mass equal to 1000  kilograms. It is a non-SI unit accepted for use with SI. It is also referred to as a metric ton to distinguish it from the non-metric units of the short ton ( United State ...
s (7.3 
tons Tons can refer to: * Tons River, a major river in India * Tamsa River, locally called Tons in its lower parts (Allahabad district, Uttar pradesh, India). * the plural of ton, a unit of mass, force, volume, energy or power :* short ton, 2,000 poun ...
) when fully grown. Recently-found specimens that are still under study at the
Museum of the Rockies Museum of the Rockies is a museum in Bozeman, Montana. Originally affiliated with Montana State University in Bozeman, and now also, the Smithsonian Institution, the museum is largely known for its paleontological collections. The Museum houses ...
, namely MOR 1142 ("X-rex") and MOR 1609 ("Becky's Giant"), suggest that ''E. annectens'' may have reached lengths of nearly and weighed , potentially making it one of the largest hadrosaurids; however, 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 , 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 A goose (plural, : geese) is a bird of any of several waterfowl species in the family (biology), family Anatidae. This group comprises the genera ''Anser (bird), Anser'' (the grey geese and white geese) and ''Branta'' (the black geese). Some o ...
in side view, and to a short-billed spoonbill in top view. The skull was longer and lower proportionally than in any other known hadrosaurid. The toothless portion of the
anterior Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bon ...
was relatively longer than in any hadrosaur. The extreme length and breadth did not appear until an individual reached maturity, so many specimens lack the distinctive shape. The bones surrounding the large openings for the
nostril A nostril (or naris , plural ''nares'' ) is either of the two orifices of the nose. They enable the entry and exit of air and other gasses through the nasal cavities. In birds and mammals, they contain branched bones or cartilages called t ...
s formed deep pockets around the openings. The eye sockets were rectangular and longer front to back than top to bottom, although this may have been exaggerated by postmortem crushing. The skull roof was flat and lacked a bony crest, and the
quadrate bone The quadrate bone is a skull bone in most tetrapods, including amphibians, sauropsids (reptiles, birds), and early synapsids. In most tetrapods, the quadrate bone connects to the quadratojugal and squamosal bones in the skull, and forms uppe ...
that formed the articulation with the lower jaw was distinctly curved. The lower jaw was long and straight, lacking the downward curve seen in other hadrosaurids, and possessing a heavy ridge running its length. The predentary was wide and shovel-like.Lull and Wright, ''Hadrosaurian Dinosaurs of North America'', pp. 157-159. The ridge on the lower jaw may have reinforced the long, slender structure.Lull and Wright, ''Hadrosaurian Dinosaurs of North America'', pp. 163-164. As mounted, the vertebral column of ''E. annectens'' includes twelve neck, twelve back, nine sacral, and at least thirty tail vertebrae. 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 In vertebrates, the pubic region ( la, pubis) is the most forward-facing (ventral and anterior) of the three main regions making up the coxal bone. The left and right pubic regions are each made up of three sections, a superior ramus, inferior ra ...
. ''E. annectens'', like other hadrosaurids, could move both on
two legs In sports (particularly association football), a two-legged tie is a contest between two teams which comprises two matches or "legs", with each team as the home team in one leg. The winning team is usually determined by aggregate score, the sum ...
and on four legs. It probably preferred to forage for food on four legs, but ran on two.
Henry Fairfield Osborn Henry Fairfield Osborn, Sr. (August 8, 1857 – November 6, 1935) was an American paleontologist, geologist and eugenics advocate. He was the president of the American Museum of Natural History for 25 years and a cofounder of the American E ...
used the skeletons in the American Museum of Natural History to portray both quadrupedal and bipedal stances for ''E. annectens''.


Classification

''E. annectens'' was a saurolophine or "flat-headed"
hadrosaurid Hadrosaurids (), or duck-billed dinosaurs, are members of the ornithischian family Hadrosauridae. This group is known as the duck-billed dinosaurs for the flat duck-bill appearance of the bones in their snouts. The ornithopod family, which incl ...
; this group was historically known as Hadrosaurinae. Species now considered to be synonyms of ''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 genera. On the other side of the issue, other authors, from John Bell Hatcher in 1902, to
Jack Horner Jack Horner may refer to: *''Little Jack Horner'', a nursery rhyme People * Jack Horner (baseball) (1863–1910), American professional baseball player *Jack Horner (journalist) (1912–2005), Gordon John Horner, Minnesota sportscaster * Jack B. H ...
, David B. Weishampel, and Catherine Forster in 2004, and most recently Nicolás Campione and David Evans, 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 {{Short pages monitor