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''Stegosaurus'' (; ) is a genus of
herbivorous A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpar ...
, four-legged, armored dinosaur from the
Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
, characterized by the distinctive kite-shaped upright plates along their backs and spikes on their tails.
Fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s of the genus have been found in the western
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and in
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, where they are found in Kimmeridgian- to early
Tithonian In the geological timescale, the Tithonian is the latest age of the Late Jurassic Epoch and the uppermost stage of the Upper Jurassic Series. It spans the time between 152.1 ± 4 Ma and 145.0 ± 4 Ma (million years ago). It is preceded by ...
-aged
strata In geology and related fields, a stratum ( : strata) is a layer of rock or sediment characterized by certain lithologic properties or attributes that distinguish it from adjacent layers from which it is separated by visible surfaces known as e ...
, dating to between 155 and 145 
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 ...
. Of the species that have been classified in the upper
Morrison Formation The Morrison Formation is a distinctive sequence of Upper Jurassic sedimentary rock found in the western United States which has been the most fertile source of dinosaur fossils in North America. It is composed of mudstone, sandstone, siltston ...
of the western US, only three are universally recognized: ''S. stenops'', ''S. ungulatus'' and ''S. sulcatus''. The remains of over 80 individual animals of this genus have been found. ''Stegosaurus'' would have lived alongside dinosaurs such as ''
Apatosaurus ''Apatosaurus'' (; meaning "deceptive lizard") is a genus of herbivorous sauropod dinosaur that lived in North America during the Late Jurassic period. Othniel Charles Marsh described and named the first-known species, ''A. ajax'', in 1 ...
'', ''
Diplodocus ''Diplodocus'' (, , or ) was a genus of diplodocid sauropod dinosaurs, whose fossils were first discovered in 1877 by S. W. Williston. The generic name, coined by Othniel Charles Marsh in 1878, is a neo-Latin term derived from Greek δ ...
'', ''
Brachiosaurus ''Brachiosaurus'' () is a genus of sauropod dinosaur that lived in North America during the Late Jurassic, about 154to 150million years ago. It was first Species description, described by American paleontologist Elmer S. Riggs in 1903 in pal ...
'', '' Ceratosaurus'', and ''
Allosaurus ''Allosaurus'' () is a genus of large carnosaurian theropod dinosaur that lived 155 to 145 million years ago during the Late Jurassic epoch ( Kimmeridgian to late Tithonian). The name "''Allosaurus''" means "different lizard" alludin ...
''; the latter two may have preyed on it. They were large, heavily built, herbivorous quadrupeds with rounded backs, short fore limbs, long hind limbs, and tails held high in the air. Due to their distinctive combination of broad, upright plates and tail tipped with spikes, ''Stegosaurus'' is one of the most recognizable kinds of dinosaurs. The function of this array of plates and spikes has been the subject of much speculation among scientists. Today, it is generally agreed that their spiked tails were most likely used for defense against predators, while their plates may have been used primarily for display, and secondarily for
thermoregulatory Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different. A thermoconforming organism, by contrast, simply adopts the surrounding temperatur ...
functions. ''Stegosaurus'' had a relatively low brain-to-body mass ratio. It had a short neck and a small head, meaning it most likely ate low-lying bushes and shrubs. One species, ''Stegosaurus ungulatus'', is one of the largest known of all the
stegosauria Stegosauria is a group of herbivorous ornithischian dinosaurs that lived during the Jurassic and early Cretaceous periods. Stegosaurian fossils have been found mostly in the Northern Hemisphere, predominantly in what is now North America, Europ ...
ns, reaching in length and in body mass, and some specimens indicate an even larger body size. ''Stegosaurus'' remains were first identified during the "
Bone Wars The Bone Wars, also known as the Great Dinosaur Rush, was a period of intense and ruthlessly competitive fossil hunting and discovery during the Gilded Age of American history, marked by a heated rivalry between Edward Drinker Cope (of the Acad ...
" 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 ...
at
Dinosaur Ridge Dinosaur Ridge is a segment of the Dakota Hogback in the Morrison Fossil Area National Natural Landmark located in Jefferson County, Colorado, near the town of Morrison and just west of Denver. The Dinosaur Ridge area is one of the world's mo ...
National Landmark. The first known skeletons were fragmentary and the bones were scattered, and it would be many years before the true appearance of these animals, including their posture and plate arrangement, became well understood. Despite its popularity in books and film, mounted skeletons of ''Stegosaurus'' did not become a staple of major natural history museums until the mid-20th century, and many museums have had to assemble composite displays from several different specimens due to a lack of complete skeletons. ''Stegosaurus'' is one of the better-known dinosaurs, and has been featured in film, postal stamps, and many other types of media.


History and naming


Bone Wars and ''Stegosaurus armatus''

''Stegosaurus'', one of the many dinosaurs described in the
Bone Wars The Bone Wars, also known as the Great Dinosaur Rush, was a period of intense and ruthlessly competitive fossil hunting and discovery during the Gilded Age of American history, marked by a heated rivalry between Edward Drinker Cope (of the Acad ...
, was first collected by Arthur Lakes and consisted of several caudal vertebrae, a dermal plate, and several additional postcranial elements that were collected north of
Morrison, Colorado The Town of Morrison is a home rule municipality in Jefferson County, Colorado, United States. The population was 428 at the 2010 census. Red Rocks Amphitheatre is located nearby. History This small foothills settlement is named after George ...
at Lakes’ YPM Quarry 5. These first, fragmented bones (YPM 1850) became 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 ''Stegosaurus armatus'' when Yale paleontologist Othniel Charles Marsh described them in 1877. Marsh initially believed the remains were from an aquatic turtle-like animal, and the basis for its
scientific name In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bo ...
, 'roof(ed) lizard' was due to his early belief that the plates lay flat over the animal's back, overlapping like the
shingles Shingles, also known as zoster or herpes zoster, is a viral disease characterized by a painful skin rash with blisters in a localized area. Typically the rash occurs in a single, wide mark either on the left or right side of the body or fac ...
(
tile Tiles are usually thin, square or rectangular coverings manufactured from hard-wearing material such as ceramic, stone, metal, baked clay, or even glass. They are generally fixed in place in an array to cover roofs, floors, walls, edges, or ...
s) on a roof. Though several more complete specimens have been attributed to ''Stegosaurus armatus'', preparation of the bones and analysis has discovered that this type specimen is actually dubious, which is not an ideal situation for the type species of a well-known genus like ''Stegosaurus''. Because of this, the
International Code of Zoological Nomenclature The International Code of Zoological Nomenclature (ICZN) is a widely accepted convention in zoology that rules the formal scientific naming of organisms treated as animals. It is also informally known as the ICZN Code, for its publisher, the I ...
decided to replace the type species with the more well known species ''Stegosaurus stenops''. Marsh also incorrectly referred several fossils to ''S. armatus'', including the dentary and teeth of the sauropod ''
Diplodocus ''Diplodocus'' (, , or ) was a genus of diplodocid sauropod dinosaurs, whose fossils were first discovered in 1877 by S. W. Williston. The generic name, coined by Othniel Charles Marsh in 1878, is a neo-Latin term derived from Greek δ ...
'' and putting sauropod limb bones and an ''
Allosaurus ''Allosaurus'' () is a genus of large carnosaurian theropod dinosaur that lived 155 to 145 million years ago during the Late Jurassic epoch ( Kimmeridgian to late Tithonian). The name "''Allosaurus''" means "different lizard" alludin ...
'' tibia under YPM 1850. On the other side of the Bone Wars,
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 interes ...
named '' Hypsirhophus discurus'' as another stegosaurian based on fragmentary fossils from Cope's Quarry 3 near the "Cope's Nipple" site in
Garden Park, Colorado Garden Park is a paleontological site in Fremont County, Colorado, known for its Jurassic dinosaurs and the role the specimens played in the infamous Bone Wars of the late 19th century. Located north of Cañon City, the name originates from the a ...
in 1878. Many later researchers have considered ''Hypsirhophus'' to be a
synonym A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are al ...
of ''Stegosaurus'', though Peter Galton (2010) suggested that it is distinct based on differences in the vertebrae. F. F. Hubbell, a collector for Cope, also found a partial ''Stegosaurus'' skeleton while digging at Como Bluff in 1877 or ‘78 that are now part of the ''Stegosaurus'' mount (AMNH 5752) at the
American Museum of Natural History 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 int ...
. Arthur Lakes made another discovery later in 1879 at
Como Bluff Como Bluff is a long ridge extending east–west, located between the towns of Rock River and Medicine Bow, Wyoming. The ridge is an anticline, formed as a result of compressional geological folding. Three geological formations, the Sundance, th ...
in
Albany County, Wyoming Albany County ( ) is a county in the U.S. state of Wyoming. As of the 2020 United States Census, the population was 37,066. Its county seat is Laramie, the site of the University of Wyoming. Its south border lies on the northern Colorado state ...
, the site also dating to the
Upper Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
of the
Morrison Formation The Morrison Formation is a distinctive sequence of Upper Jurassic sedimentary rock found in the western United States which has been the most fertile source of dinosaur fossils in North America. It is composed of mudstone, sandstone, siltston ...
, when he found several large ''Stegosaurus'' fossils in August of that year. The majority of the fossils came from Quarry 13, including the type specimen of ''Stegosaurus ungulatus (''YPM 1853), which was collected by Lakes and William Harlow Reed the same year and named by Marsh. The specimen was one of many found at the quarry, the specimen consisting of a partial skull, several vertebrae, an ischium, partial limbs, several plates, and four thagomizers, though eight thagomizers were referred based on a specimen preserved alongside the type. The type specimen also preserved the pes, which was the namesake of the species, meaning "hoofed roofed lizard". In 1881, he named a third species ''Stegosaurus'' "affinis", based only on a hip bone, though the fossil has since been lost and the species declared a nomen nudum. Later in 1887, Marsh described two more species of ''Stegosaurus'' from Como Bluff, ''Stegosaurus duplex,'' based on a partial vertebral column, partial pelvis, and partial left hindlimb (YPM 1858) from Reed's Quarry 11, though the species is now seen as synonymous with ''Stegosaurus ungulatus''. The other, ''Stegosaurus sulcatus'', was named based on a left forelimb, scapula, left femur, several vertebrae, and several plates and dermal armor elements (USNM V 4937) collected in 1883. ''Stegosaurus sulcatus'' most notably preserves a large spike that has been speculated to have been a shoulder spike that is used to diagnose the species. left, Type specimen of ''S. stenops'' on display at the National Museum of Natural History. The greatest ''Stegosaurus'' discovery came in 1885 with the discovery of a nearly complete, articulated skeleton of a subadult that included previously undiscovered elements like a complete skull, throat ossicles, and articulated plates. Marshall P. Felch collected the skeleton throughout 1885 and 1886 from Morrison Formation strata at his quarry in Garden Park, a town near Cañon City,
Colorado Colorado (, other variants) is a state in the 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 western edge of the ...
. The skeleton was expertly unearthed by Felch, who first divided the skeleton into labeled blocks and prepared them separately. The skeleton was shipped to Marsh in 1887, who named it ''Stegosaurus stenops'' ( "narrow-faced roof lizard") that year. Though it had not yet been completely prepared, the nearly complete and articulated type specimen of ''Stegosaurus stenops'' allowed Marsh to complete the first attempt at a reconstructed ''Stegosaurus'' skeleton. This first reconstruction, of ''S. ungulatus'' with missing parts filled in from ''S. stenops'', was published by Marsh in 1891. (In 1893,
Richard Lydekker Richard Lydekker (; 25 July 1849 – 16 April 1915) was an English naturalist, geologist and writer of numerous books on natural history. Biography Richard Lydekker was born at Tavistock Square in London. His father was Gerard Wolfe Lydekker ...
mistakenly re-published Marsh's drawing under the label ''Hypsirhophus''.)


Early skeletal mounts and plate interpretation

The skeleton of ''S. stenops'' has since been deposited at the
National Museum of Natural History 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. ...
in
Washington D. C. ) , image_skyline = , image_caption = Clockwise from top left: the Washington Monument and Lincoln Memorial on the National Mall, United States Capitol, Logan Circle, Jefferson Memorial, White House, Adams Morgan, Na ...
, where it has been on display since 1915. Another mount was made for the NMNH in the form of a mounted composite skeleton consisting of several specimens referred to ''S. stenops'' that were collected at Quarry 13 at Como Bluff in 1887, the most complete being USNM 6531. The type specimen of ''S. ungulatus'' (YPM 1853) was incorporated into the first ever mounted skeleton of a stegosaur at the Peabody Museum of Natural History in 1910 by
Richard Swann Lull Richard Swann Lull (November 6, 1867 – April 22, 1957) was an American paleontologist and Sterling Professor at Yale University who is largely remembered now for championing a non-Darwinian view of evolution, whereby mutation(s) could ...
. It was initially mounted with paired plates set wide, above the base of the ribs, but was remounted in 1924 with two staggered rows of plates along the midline of the back.Revan, A. (2011)
Reconstructing an Icon: Historical Significance of the Peabody’s Mounted Skeleton of ''Stegosaurus'' and the Changes Necessary to Make It Correct Anatomically.
Doctoral dissertation, faculty of the Department of Geology and Geophysics, Yale University.
Additional specimens recovered from the same quarry by the United States
National Museum of Natural History 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. ...
, including tail vertebrae and an additional large plate (USNM 7414), belong to the same individual as YPM 1853. The next species of ''Stegosaurus'' to be named was ''S. marshi'' by Frederick Lucas in 1901. Lucas reclassified this species in the new genus ''
Hoplitosaurus ''Hoplitosaurus'' (meaning "Hoplite lizard") was a genus of armored dinosaur related to ''Polacanthus''. It was named from a partial skeleton found in the ?Barremian-age Lower Cretaceous Lakota Formation of Custer County, South Dakota. It is an ...
'' later that year. Lucas also re-examined the issue of the life appearance of ''Stegosaurus'', coming to the conclusion that the plates were arranged in pairs in two rows along the back, arranged above the bases of the ribs. Lucas commissioned
Charles R. Knight Charles Robert Knight (October 21, 1874 – April 15, 1953) was an American wildlife and paleoartist best known for his detailed paintings of dinosaurs and other prehistoric animals. His works have been reproduced in many books and are currently ...
to produce a life restoration of ''S. ungulatus'' based on his new interpretation. However, the following year, Lucas wrote that he now believed the plates were probably attached in staggered rows. In 1910,
Richard Swann Lull Richard Swann Lull (November 6, 1867 – April 22, 1957) was an American paleontologist and Sterling Professor at Yale University who is largely remembered now for championing a non-Darwinian view of evolution, whereby mutation(s) could ...
wrote that the alternating pattern seen in ''S. stenops'' was probably due to shifting of the skeleton after death. He led the construction of the first ever ''Stegosaurus'' skeletal mount at the Peabody Museum of Natural History, which was depicted with paired plates. In 1914, Charles Gilmore argued against Lull's interpretation, noting that several specimens of ''S. stenops'', including the now-completely prepared holotype, preserved the plates in alternating rows near the peak of the back, and that there was no evidence of the plates having shifted relative to the body during fossilization. Gilmore and Lucas' interpretation became the generally accepted standard, and Lull's mount at the Peabody Museum was changed to reflect this in 1924.


Plate arrangement

One of the major subjects of books and articles about ''Stegosaurus'' is the plate arrangement. The argument has been a major one in the history of dinosaur reconstruction. Four possible plate arrangements have been proposed over the years: * The plates lie flat along the back, as a shingle-like armor. This was Marsh's initial interpretation, which led to the name 'roof lizard'. As further and complete plates were found, their form showed they stood on edge, rather than lying flat. * By 1891, Marsh published a more familiar view of ''Stegosaurus'', with a single row of plates. This was dropped fairly early on (apparently because it was poorly understood how the plates were embedded in the skin and they were thought to overlap too much in this arrangement). It was revived, in somewhat modified form, in the 1980s, by Stephen Czerkas, based on the arrangement of iguana dorsal spines. * The plates were paired in a double row along the back, such as in Knight's 1901 reconstruction and the 1933 film ''
King Kong King Kong is a fictional giant monster resembling a gorilla, who has appeared in various media since 1933. He has been dubbed The Eighth Wonder of the World, a phrase commonly used within the franchise. His first appearance was in the novelizat ...
''. * Two rows of alternating plates. By the early 1960s, this had become (and remains) the prevalent idea, mainly because some ''S. stenops'' fossils in which the plates are still partially articulated show this arrangement. This arrangement is chiral and so demands that a specimen be distinguished from its distinct, hypothetical mirror-image form.


Second Dinosaur Rush

After the end of the Bone Wars, many major institutions in the eastern United States were inspired by the depictions and finds by Marsh and Cope to assemble their own dinosaur fossil collections. The competition was foremost started by the
American Museum of Natural History 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 int ...
, the
Carnegie Museum of Natural History The Carnegie Museum of Natural History (abbreviated as CMNH) is a natural history museum in the Oakland neighborhood of Pittsburgh, Pennsylvania. It was founded by Pittsburgh-based industrialist Andrew Carnegie in 1896. Housing some 22 million ...
, and the
Field Museum of Natural History The Field Museum of Natural History (FMNH), also known as The Field Museum, is a natural history museum in Chicago, Illinois, and is one of the largest such museums in the world. The museum is popular for the size and quality of its educational ...
which all sent expeditions to the west to make their own dinosaur collections and mount skeletons in their fossil halls.Brinkman, P. D. (2010). The second Jurassic dinosaur rush. University of Chicago Press. The American Museum of Natural History was the first to launch an expedition in 1897, finding several assorted, but incomplete, ''Stegosaurus'' specimens at Bone Cabin Quarry in Como Bluff. These remains haven't been described and were mounted in 1932, the mount being a composite primarily of specimens AMNH 650 & 470 from Bone Cabin Quarry. The AMNH mount is cast and on display at the Field Museum, which didn't collect any ''Stegosaurus'' skeletons during the Second Dinosaur Rush. The Carnegie Museum in
Pittsburgh Pittsburgh ( ) is a city in the Commonwealth (U.S. state), Commonwealth of Pennsylvania, United States, and the county seat of Allegheny County, Pennsylvania, Allegheny County. It is the most populous city in both Allegheny County and Wester ...
on the other hand collected many ''Stegosaurus'' specimens, first at Freezout Hills in
Carbon County, Wyoming Carbon County is a county in the U.S. state of Wyoming. As of the 2020 United States Census, the population was 14,537. Its county seat is Rawlins. Its south border abuts the north line of Colorado. History Carbon County was organized in ...
in 1902–03. The fossils included only a couple postcranial remains, though in the 1900s-1920s Carnegie crews at
Dinosaur National Monument Dinosaur National Monument is an American national monument located on the southeast flank of the Uinta Mountains on the border between Colorado and Utah at the confluence of the Green and Yampa rivers. Although most of the monument area is i ...
discovered dozens of ''Stegosaurus'' specimens in one of the greatest single sites for the taxon.McIntosh, J. S. (1981). Annotated catalogue of the dinosaurs (Reptilia, Archosauria) in the collections of Carnegie Museum of Natural History. CM 11341, the most complete skeleton found at the quarry, was used for the basis of a composite ''Stegosaurus'' mount in 1940 along with several other specimens to finish the mount. A cranium (CM 12000) was also found by Carnegie crews, one of the few known. Both the AMNH and CM material has been referred to ''Stegosaurus ungulatus''.


Resurgent Discoveries

As part of the Dinosaur Renaissance and the resurgent interest in dinosaurs by museums and the public, fossils of ''Stegosaurus'' were once again being collected, though few have been fully described. An important discovery came in 1937 again at Garden Park by a high school teacher named Frank Kessler in while leading a nature hike. Kessler contacted the Denver Museum of Nature and Science, who sent paleontologist Robert Landberg. Landberg excavated the skeleton with the DMNS crews, recovering a 70% complete ''Stegosaurus'' skeleton along with turtles, crocodiles, and isolated dinosaur fossils at the quarry that would be nicknamed "The Kessler Site". Phillip Reinheimer, a steel worker, mounted the ''Stegosaurus'' skeleton at the DMNS in 1938. The skeleton remained mounted until 1989 when the museum curator of the DMNS began a revision of the museum's fossil hall and dispatched an expedition to find additional ''Stegosaurus'' remains. The expedition was successful in finding a nearly complete ''Stegosaurus'' near the Kessler site by Bryan Small, whose name would become the namesake of the new site. The "Small Quarry" ''Stegosaurus articulation and completeness clarified the position of plates and spikes on the back of ''Stegosaurus'' and the position and size of the throat ossicles found earlier first by Felch with the ''Stegosaurus stenops'' holotype, though like the ''S. stenops'' type, the fossils were flattened in a "roadkill" condition. The ''Stegosaurus'' skeletons have been mounted alongside an ''Allosaurus'' skeleton collected in Moffat County, Colorado originally in 1979. 1987 saw the discovery of a 40% complete ''Stegosaurus'' skeleton in Rabbit Valley in Mesa County, Colorado by Harold Bollan near the Dinosaur Journey Museum. The skeleton was nicknamed the "Bollan ''Stegosaurus"'' and is in the collections of the Dinosaur Journey Museum. At Jensen-Jensen Quarry, an articulated torso including several dorsal plates from a small individual were collected and briefly described in 2014, though the specimen was collected years before and is still in preparation at
Brigham Young University Brigham Young University (BYU, sometimes referred to colloquially as The Y) is a private research university in Provo, Utah. It was founded in 1875 by religious leader Brigham Young and is sponsored by the Church of Jesus Christ of Latter-d ...
. 2007 saw the description of a ''Stegosaurus'' specimen from the Upper Jurassic Lourinha Formation of
Portugal Portugal, officially the Portuguese Republic ( pt, República Portuguesa, links=yes ), is a country whose mainland is located on the Iberian Peninsula of Southwestern Europe, and whose territory also includes the Atlantic archipelagos of th ...
, the specimen was placed as cf. ''Stegosaurus ungulatus'' by the describers. The specimen is one of the few associated ''Stegosaurus'' skeletons known, though it only contains a tooth, 13 vertebrae, partial limbs, a cervical plate, and several assorted postcranial elements. Sophie the ''Stegosaurus'' is the best preserved ''Stegosaurus'' specimen, being 85% intact and containing 360 bones. Sophie was first discovered by Bob Simon in 2003 at the Red Canyon Quarry near
Shell, Wyoming Shell is a census-designated place (CDP) in Big Horn County, Wyoming, United States. The population was 83 at the 2010 Census. The community is named for the abundance of fossil shells located in the area. Nearby exposed formations such as the ...
and was excavated by crews from the Swiss Sauriermuseum in 2004.Siber, H. J., & Möckli, U. (2009)
The stegosaurs of the Sauriermuseum Aathal.
/ref> The skeleton was excavated on private land, so it was interned by US federal authorities who then gave Sophie to the
Natural History Museum, London 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 Museum an ...
where it was put on display in December of 2014 and later described in 2015. It is a young adult of undetermined sex, 5.8 m (19 ft) long and 2.9 m (9.5 ft) tall. The Sauriermuseum found several partial Stegosaurid skeletons throughout their excavations at Howe Quarry, Wyoming in the 1990s, though only Sophie has been described in detail. One skeleton collected at the site known as "Victoria" is very well preserved including many of the vertebrae preserved in semi-articulation and next to an ''
Allosaurus ''Allosaurus'' () is a genus of large carnosaurian theropod dinosaur that lived 155 to 145 million years ago during the Late Jurassic epoch ( Kimmeridgian to late Tithonian). The name "''Allosaurus''" means "different lizard" alludin ...
'' skeleton found nicknamed "Big Al II".


Description

The quadrupedal ''Stegosaurus'' is one of the most easily identifiable dinosaur genera, due to the distinctive double row of kite-shaped plates rising vertically along the rounded back and the two pairs of long spikes extending horizontally near the end of the tail. ''S. stenops'' reached in length and in body mass, while ''S. ungulatus'' reached in length and in body mass. Some large individuals may have reached in length and in body mass. Most of the information known about ''Stegosaurus'' comes from the remains of mature animals; more recently, though, juvenile remains of ''Stegosaurus'' have been found. One subadult specimen, discovered in 1994 in Wyoming, is long and high, and is estimated to have weighed 1.5-2.2 metric tons (1.6-2.4 short tons) while alive. It is on display in the University of Wyoming Geological Museum.


Skull

The long and narrow
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, th ...
was small in proportion to the body. It had a small antorbital fenestra, the hole between the nose and eye common to most
archosaur Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avia ...
s, including modern birds, though lost in extant crocodylians. The skull's low position suggests that ''Stegosaurus'' may have been a browser of low-growing vegetation. This interpretation is supported by the absence of front teeth and their likely replacement by a horny beak or
rhamphotheca The beak, bill, or rostrum is an external anatomical structure found mostly in birds, but also in turtles, non-avian dinosaurs and a few mammals. A beak is used for eating, preening, manipulating objects, killing prey, fighting, probing for food, ...
. The lower jaw had flat downward and upward extensions that would have completely hidden the teeth when viewed from the side, and these probably supported a turtle-like beak in life. The presence of a beak extended along much of the jaws may have precluded the presence of cheeks in these species. Such an extensive beak was probably unique to ''Stegosaurus'' and some other advanced stegosaurids among ornithischians, which usually had beaks restricted to the jaw tips.Barrett, P.M. (2001). Tooth wear and possible jaw action of ''Scelidosaurus harrisonii'' Owen and a review of feeding mechanisms in other thyreophoran dinosaurs. Pp. 25-52 in Carpenter, K. (ed.): ''The Armored Dinosaurs''. Bloomington: Indiana University Press. Other researchers have interpreted these ridges as modified versions of similar structures in other ornithischians which might have supported fleshy cheeks, rather than beaks. Stegosaurian teeth were small, triangular, and flat; wear facets show that they did grind their food. Despite the animal's overall size, the braincase of ''Stegosaurus'' was small, being no larger than that of a dog. A well-preserved ''Stegosaurus'' braincase allowed
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 ...
to obtain, in the 1880s, a cast of the brain cavity or endocast of the animal, which gave an indication of the brain size. The endocast showed the brain was indeed very small, the smallest proportionally of all dinosaur endocasts then known. The fact that an animal weighing over 4.5 
metric tons 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 ...
(5  short tons) could have a brain of no more than contributed to the popular old idea that all dinosaurs were unintelligent, an idea now largely rejected. Actual brain anatomy in ''Stegosaurus'' is poorly known, but the brain itself was small even for a dinosaur.


Skeleton

In ''Stegosaurus stenops'' there are 27 bones in the
vertebral column The vertebral column, also known as the backbone or spine, is part of the axial skeleton. The vertebral column is the defining characteristic of a vertebrate in which the notochord (a flexible rod of uniform composition) found in all chordate ...
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 language, Greek roots, describe something in its standard anatomical position. Th ...
to the
sacrum The sacrum (plural: ''sacra'' or ''sacrums''), in human anatomy, is a large, triangular bone at the base of the spine that forms by the fusing of the sacral vertebrae (S1S5) between ages 18 and 30. The sacrum situates at the upper, back part o ...
, a varying number of vertebrae in the sacrum, with four in most subadults, and around 46 caudal (tail) vertebrae. The presacrals are divided into cervical (neck) and dorsal (back) vertebrae, with around 10 cervicals and 17 dorsals, the total number being one greater than in '' Hesperosaurus'', two greater than ''
Huayangosaurus ''Huayangosaurus'' is a genus of stegosaurian dinosaur from the Middle Jurassic of China. The name derives from "Huayang" (華陽), an alternate name for Sichuan (the province where it was discovered), and "saurus", meaning "lizard". It lived ...
'', although '' Miragaia'' preserves 17 cervicals and an unknown number of dorsals. The first cervical vertebra is the axis bone, which is connected and often fused to the atlas bone. Farther posteriorly, the proportionately larger the cervicals become, although they do not change greatly in anything other than size. Past the first few dorsals, the centrum of the bones become more elongate front-to-back, and the
transverse Transverse may refer to: *Transverse engine, an engine in which the crankshaft is oriented side-to-side relative to the wheels of the vehicle * Transverse flute, a flute that is held horizontally * Transverse force (or ''Euler force''), the tange ...
processes become more elevated dorsal. The sacrum of ''S. stenops'' includes four sacral vertebrae, but one of the dorsals is also incorporated into the structure. In some specimens of ''S. stenops'', a caudal is also incorporated, as a caudosacral. In '' Hesperosaurus'' there are two dorsosacrals, and only four fused sacrals, but in '' Kentrosaurus'' there may be as many as seven vertebrae in the sacrum, with both dorsosacrals and caudosacrals. ''S. stenops'' preserves 46 caudal vertebrae, and up to 49, and along the series both the centrums and the
neural spines 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 characteristic i ...
become smaller, until the neural spines disappear at caudal 35. Around the middle of the tail, the neural spines become bifurcated, meaning they are divided near the top. With multiple well-preserved skeletons, ''S. stenops'' preserves all regions of the body, including the limbs. The
scapula The scapula (plural scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on eith ...
(shoulder blade) is sub-rectangular, with a robust blade. Though it is not always perfectly preserved, the
acromion In human anatomy, the acromion (from Greek: ''akros'', "highest", ''ōmos'', "shoulder", plural: acromia) is a bony process on the scapula (shoulder blade). Together with the coracoid process it extends laterally over the shoulder joint. The ...
ridge is slightly larger than in ''Kentrosaurus''. The blade is relatively straight, although it curves towards the back. There is a small bump on the back of the blade, that would have served as the base of the '' triceps'' muscle. Articulated with the scapula, the coracoid is sub-circular. The hind feet each had three short toes, while each fore foot had five toes; only the inner two toes had a blunt hoof. The phalangeal formula is 2-2-2-2-1, meaning the innermost finger of the fore limb has two bones, the next has two, etc. All four limbs were supported by pads behind the toes. The fore limbs were much shorter than the stocky hind limbs, which resulted in an unusual posture. The tail appears to have been held well clear of the ground, while the head of ''Stegosaurus'' was positioned relatively low down, probably no higher than above the ground.


Plates

The most recognizable features of ''Stegosaurus'' are its dermal plates, which consisted of between 17 and 22 separate plates and flat spines. These were highly modified osteoderms (bony-cored scales), similar to those seen in crocodiles and many lizards today. They were not directly attached to the animal's skeleton, instead arising from the skin. The largest plates were found over the hips and could measure over wide and tall. In a 2010 review of ''Stegosaurus'' species, Peter Galton suggested that the arrangement of the plates on the back may have varied between species, and that the pattern of plates as viewed in profile may have been important for species recognition. Galton noted that the plates in ''S. stenops'' have been found articulated in two staggered rows, rather than paired. Fewer ''S. ungulatus'' plates have been found, and none articulated, making the arrangement in this species more difficult to determine. However, the type specimen of ''S. ungulatus'' preserves two flattened spine-like plates from the tail that are nearly identical in shape and size, but are mirror images of each other, suggesting that at least these were arranged in pairs. Many of the plates are manifestly chiral and no two plates of the same size and shape have been found for an individual; however plates have been correlated between individuals. Well preserved integumentary impressions of the plates of ''Hesperosaurus'' show a smooth surface with long and parallel, shallow grooves. This indicates that the plates were covered in keratinous sheaths.


Classification and species

Like the spikes and shields of
ankylosaur Ankylosauria is a group of herbivorous dinosaurs of the order Ornithischia. It includes the great majority of dinosaurs with armor in the form of bony osteoderms, similar to turtles. Ankylosaurs were bulky quadrupeds, with short, powerful limbs. ...
s, the bony plates and spines of stegosaurians evolved from the low-keeled osteoderms characteristic of basal thyreophorans.Norman, David (2001). "''Scelidosaurus'', the earliest complete dinosaur" in ''The Armored Dinosaurs'', pp 3-24. Bloomington: Indiana University Press. .
Galton Sir Francis Galton, FRS FRAI (; 16 February 1822 – 17 January 1911), was an English Victorian era polymath: a statistician, sociologist, psychologist, anthropologist, tropical explorer, geographer, inventor, meteorologist, prot ...
(2019) interpreted plates of an armored dinosaur from the Lower Jurassic ( Sinemurian- Pliensbachian) Lower Kota Formation of
India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area, the List of countries and dependencies by population, second-most populous ...
as fossils of a member of Ankylosauria; the author argued that this finding indicates a probable early Early Jurassic origin for both Ankylosauria and its
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
Stegosauria. The vast majority of stegosaurian dinosaurs thus far recovered belong to the Stegosauridae, which lived in the later part of the Jurassic and early Cretaceous, and which were defined by
Paul Sereno Paul Callistus Sereno (born October 11, 1957) is a professor of paleontology at the University of Chicago and a National Geographic "explorer-in-residence" who has discovered several new dinosaur species on several continents, including at si ...
as all stegosaurians more closely related to ''Stegosaurus'' than to ''Huayangosaurus''. This group is widespread, with members across the Northern Hemisphere,
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
and possibly
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the sou ...
. ''Stegosaurus'' frequently is discovered in its own clade in Stegosauridae called Stegosauridae, usually including the taxa '' Wuerhosaurus'' and '' Loricatosaurus'', though '' Hesperosaurus'' is sometimes found in the group. in 2017, Raven and Maidment published a new phylogenetic analysis, including almost every known stegosaurian genus:


Species

Many of the species initially described have since been considered to be invalid or synonymous with earlier named species, leaving two well-known and one poorly known species. Confirmed ''Stegosaurus'' remains have been found in the Morrison Formation's stratigraphic zones 2–6, with additional remains possibly referrable to ''Stegosaurus'' recovered from stratigraphic zone 1. * ''Stegosaurus ungulatus'', meaning "hoofed roof lizard", was named by Marsh in 1879, from remains recovered at
Como Bluff Como Bluff is a long ridge extending east–west, located between the towns of Rock River and Medicine Bow, Wyoming. The ridge is an anticline, formed as a result of compressional geological folding. Three geological formations, the Sundance, th ...
, Wyoming (Quarry 12, near Robber's Roost). It might be synonymous with ''S. stenops''. At , it was the longest species within the genus ''Stegosaurus''. A fragmentary ''Stegosaurus'' specimen discovered in Portugal and dating from the upper Kimmeridgian-lower
Tithonian In the geological timescale, the Tithonian is the latest age of the Late Jurassic Epoch and the uppermost stage of the Upper Jurassic Series. It spans the time between 152.1 ± 4 Ma and 145.0 ± 4 Ma (million years ago). It is preceded by ...
stage has been tentatively assigned to this species. ''Stegosaurus ungulatus'' can be distinguished from ''S. stenops'' by the presence of longer hind limbs, proportionately smaller, more pointed plates with wide bases and narrow tips, and by several small, flat, spine-like plates just before the spikes on the tail. These spine-like plates appear to have been paired, due to the presence of at least one pair that are identical but mirrored. ''S. ungulatus'' also appears to have had longer legs (femora) and hip bones than other species. The type specimen of ''S. ungulatus'' was discovered with eight spikes, though they were scattered away from their original positions. These have often been interpreted as indicating that the animal had four pairs of tail spikes. No specimens have been found with complete or articulated sets of tail spikes, but no additional specimens have been found that preserve eight spikes together. It is possible the extra pair of spikes came from a different individual, and though no other extra bones were found with the specimen, these may be found if more digging were done at the original site. Specimens from other quarries (such as a tail from Quarry 13, now forming part of the composite skeleton AMNH 650 at the
American Museum of Natural History 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 int ...
), referred to ''S. ungulatus'' on the basis of their notched tail vertebrae, are preserved with only four tail spikes. The type specimen of ''S. ungulatus'' (YPM 1853) was incorporated into the first ever mounted skeleton of a stegosaur at the Peabody Museum of Natural History in 1910 by
Richard Swann Lull Richard Swann Lull (November 6, 1867 – April 22, 1957) was an American paleontologist and Sterling Professor at Yale University who is largely remembered now for championing a non-Darwinian view of evolution, whereby mutation(s) could ...
. It was initially mounted with paired plates set wide, above the base of the ribs, but was remounted in 1924 with two staggered rows of plates along the midline of the back. Additional specimens recovered from the same quarry by the United States
National Museum of Natural History 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. ...
, including tail vertebrae and an additional large plate (USNM 7414), belong to the same individual as YPM 1853. * ''Stegosaurus stenops'', meaning "narrow-faced roof lizard", was named by Marsh in 1887, with the holotype having been collected by Marshall Felch at Garden Park, north of Cañon City, Colorado, in 1886. This is the best-known species of ''Stegosaurus'', mainly because its remains include at least one complete articulated skeleton. It had proportionately large, broad plates and rounded tail plates. Articulated specimens show that the plates were arranged alternating in a staggered double row. ''S. stenops'' is known from at least 50 partial skeletons of adults and juveniles, one complete skull, and four partial skulls. It was shorter than other species, at . Found in the Morrison Formation, Colorado, Wyoming, and Utah. * ''Stegosaurus sulcatus'', meaning "furrowed roof lizard", was described by Marsh in 1887 based on a partial skeleton. It has traditionally been considered a synonym of ''S. armatus'', though more recent studies suggest it is not. ''S. sulcatus'' is distinguished mainly by its unusually large, furrowed spikes with very large bases. A spike associated with the type specimen, originally thought to be a tail spike, may in fact come from the shoulder or hip, since its base is much larger than the corresponding tail vertebrae. A review published by Maidment and colleagues in 2008 regarded it as an indeterminate species possibly not even belonging to ''Stegosaurus'' at all, but to a different genus. Peter Galton suggested it should be considered a valid species due to its unique spikes. Susannah Maidment and colleagues in 2008 proposed extensive alterations to the taxonomy of ''Stegosaurus''. They advocated synonymizing ''S. stenops'' and ''S. ungulatus'' with ''S. armatus'', and sinking '' Hesperosaurus'' and '' Wuerhosaurus'' into ''Stegosaurus'', with their
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 specim ...
becoming ''Stegosaurus mjosi'' and ''Stegosaurus homheni'', respectively. They regarded ''S. longispinus'' as dubious. Thus, their conception of ''Stegosaurus'' would include three valid species (''S. armatus'', ''S. homheni'', and ''S. mjosi'') and would range from the
Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
of North America and Europe to the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145& ...
of Asia. However, this classification scheme was not followed by other researchers, and a 2017 cladistic analysis co-authored by Maidment with Thomas Raven rejects the synonymy of ''Hesperosaurus'' with ''Stegosaurus''. In 2015, Maidment ''et al.'' revised their suggestion due to the recognition by Galton of ''S. armatus'' as a ''nomen dubium'' and its replacement by ''S. stenops'' as type species.


Doubtful species and junior synonyms

* ''Stegosaurus armatus'', meaning "armored roof lizard", was the first species to be found and the original type species named by O.C. Marsh in 1877. It is known from a partial skeleton, and more than 30 fragmentary specimens have been referred to it. However, the
type specimen In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to anchor or centralizes th ...
was very fragmentary, consisting only of a partial tail, hips, and leg, parts of some back vertebrae, and a single fragmentary plate (the presence of which was used to give the animal its name). No other plates or spikes were found, and the entire front half of the animal appears not to have been preserved. Because the type specimen is very fragmentary, it is extremely difficult to compare it with other species based on better specimens, and it is now generally considered to be a ''nomen dubium''. Because of this, it was replaced by ''S. stenops'' as the type species of ''Stegosaurus'' in a ruling of the
ICZN The International Code of Zoological Nomenclature (ICZN) is a widely accepted convention in zoology that rules the formal scientific naming of organisms treated as animals. It is also informally known as the ICZN Code, for its publisher, the I ...
in 2013. * ''Stegosaurus'' "affinis", named by Marsh in 1881, is only known from a pubis which has since been lost. Because Marsh did not provide an adequate description of the bone with which to distinguish a new species, this name is considered a ''
nomen nudum In taxonomy, a ''nomen nudum'' ('naked name'; plural ''nomina nuda'') is a designation which looks exactly like a scientific name of an organism, and may have originally been intended to be one, but it has not been published with an adequate desc ...
''. * ''Diracodon laticeps'' was described by Marsh in 1881, from some jawbone fragments.
Bakker Bakker is a common Dutch surname equivalent to English Baker. In 2007 it was the seventh most common name in the Netherlands (55,273 people). although others note that the material is not diagnostic and is only referable to ''Stegosaurus'' sp., making it a ''nomen dubium''. * ''Stegosaurus duplex'', meaning "two plexus roof lizard" (in allusion to the greatly enlarged neural canal of the sacrum which Marsh characterized as a "posterior brain case"), was named by Marsh in 1887 (including the holotype specimen). The disarticulated bones were actually collected in 1879 by Edward Ashley at Como Bluff. Marsh initially distinguished it from ''S. ungulatus'' based on the fact that each sacral (hip) vertebra bore its own rib, which he claimed was unlike the anatomy of ''S. ungulatus''; however, the sacrum of ''S. ungulatus'' had not actually been discovered. Marsh also suggested that ''S. duplex'' may have lacked armor, since no plates or spikes were found with the specimen, though a single spike may actually have been present nearby, and re-examination of the site maps has shown that the entire specimen was found highly disarticulated and scattered. It is generally considered a synonym of ''S. ungulatus'' today, and parts of the specimen were actually incorporated into the Peabody Museum ''S. ungulatus'' skeletal mount in 1910.


Reassigned species

* ''Stegosaurus marshi'', which was described by Lucas in 1901, was renamed ''
Hoplitosaurus ''Hoplitosaurus'' (meaning "Hoplite lizard") was a genus of armored dinosaur related to ''Polacanthus''. It was named from a partial skeleton found in the ?Barremian-age Lower Cretaceous Lakota Formation of Custer County, South Dakota. It is an ...
'' in 1902. * ''Stegosaurus priscus'', described by Nopcsa in 1911, was reassigned to '' Lexovisaurus'', and is now the type species of '' Loricatosaurus''. * ''Stegosaurus longispinus'' was named by Charles W. Gilmore in 1914 based on a fragmentary postcranial skeleton that has largely been lost. It is now the type species of the genus ''Alcovasaurus,'' though it has been referred to ''Miragaia''. * ''Stegosaurus madagascariensis'' from
Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Afric ...
is known solely from teeth and was described by Piveteau in 1926. The teeth were variously attributed to a stegosaur, the theropod '' Majungasaurus'', a
hadrosaur 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 includ ...
or even a crocodylian, but is now considered a possible ankylosaur. * ''Stegosaurus homheni'' is an alternative combination for the Chinese Cretaceous stegosaur '' Wuerhosaurus homheni'', which was described based on a partial postcranial skeleton in 1973 by Dong Zhiming. It was referred to ''Stegosaurus'' in 2008 by Maidment ''et al'', but some still consider the species to be in its own genus. * ''Stegosaurus mjosi'' was described as '' Hesperosaurus mjosi'' by Carpenter ''et al'' in 2001 based on a partial skull and incomplete postcranial skeleton from the Morrison Formation of Johnson County, Wyoming. The species was referred to ''Stegosaurus'' mostly by Maidment ''et al'' starting in 2008, but ''Hesperosaurus'' has been the more popular combination since the discovery of more remains.


Paleobiology


Posture and movement

Soon after its discovery, Marsh considered ''Stegosaurus'' to have been bipedal, due to its short forelimbs. He had changed his mind, however, by 1891, after considering the heavy build of the animal. Although ''Stegosaurus'' is undoubtedly now considered to have been quadrupedal, some discussion has occurred over whether it could have reared up on its hind legs, using its tail to form a tripod with its hind limbs, to browse for higher foliage. This has been proposed by
Bakker Bakker is a common Dutch surname equivalent to English Baker. In 2007 it was the seventh most common name in the Netherlands (55,273 people).Carpenter Carpentry is a skilled trade and a craft in which the primary work performed is the cutting, shaping and installation of building materials during the construction of buildings, ships, timber bridges, concrete formwork, etc. Carpenters t ...
. A study by Mallison (2010) found support for a rearing up posture in ''Kentrosaurus'', though not for ability for the tail to act as a tripod. ''Stegosaurus'' had short fore limbs in relation to its hind limbs. Furthermore, within the hind limbs, the lower section (comprising the
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it conn ...
and
fibula The fibula or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. Its upper extremity ...
) was short compared with the
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates ...
. This suggests it could not walk very fast, as the stride of the back legs at speed would have overtaken the front legs, giving a maximum speed of . Tracks discovered by Matthew Mossbrucker (Morrison Natural History Museum, Colorado) suggest that ''Stegosaurus'' lived and traveled in multiple-age herds. One group of tracks is interpreted as showing four or five baby stegosaurs moving in the same direction, while another has a juvenile stegosaur track with an adult track overprinting it. As the plates would have been obstacles during copulation, it is possible the female stegosaur laid on her side as the male entered her from above and behind. Another suggestion is that the female would stand on all fours but squat down the fore limbs and raise the tail up and out of the male's way as he supports his fore limbs on her hips. However, their reproductive organs still could not touch as there is no evidence of muscle attachments for a mobile penis nor a
baculum The baculum (also penis bone, penile bone, or ''os penis'', ''os genitale'' or ''os priapi'') is a bone found in the penis of many placental mammals. It is absent from the human penis, but present in the penises of some primates, such as the ...
in male dinosaurs.


Plate function

The function of ''Stegosaurus plates has been much debated. Marsh suggested that they functioned as some form of armor, though Davitashvili (1961) disputed this, claiming that they were too fragile and ill-placed for defensive purposes, leaving the animal's sides unprotected. Nevertheless, others have continued to support a defensive function. Bakker suggested in 1986 that the plates were covered in horn comparing the surface of the fossilized plates to the bony cores of horns in other animals known or thought to bear horns. Christiansen and Tschopp (2010), having studied a well-preserved specimen of '' Hesperosaurus'' with skin impressions, concluded that the plates were covered in a keratin sheath which would have strengthened the plate as a whole and provided it with sharp cutting edges. Bakker stated that ''Stegosaurus'' could flip its osteoderms from one side to another to present a predator with an array of spikes and blades that would impede it from closing sufficiently to attack the ''Stegosaurus'' effectively. He contends that they had insufficient width for them to stand erect easily in such a manner as to be useful in display without continuous muscular effort. Mobility of the plates, however, has been disputed by other paleontologists. Another possible function of the plates is they may have helped to control the body temperature of the animal, in a similar way to the sails of the pelycosaurs ''
Dimetrodon ''Dimetrodon'' ( or ,) meaning "two measures of teeth,” is an extinct genus of non-mammalian synapsid that lived during the Cisuralian (Early Permian), around 295–272 million years ago (Mya). It is a member of the family Sphenacodont ...
'' and '' Edaphosaurus'' (and modern elephant and rabbit ears). The plates had blood vessels running through grooves and air flowing around the plates would have cooled the blood. Buffrénil, et al. (1986) found "extreme vascularization of the outer layer of bone", which was seen as evidence that the plates "acted as thermoregulatory devices". Likewise, 2010 structural comparisons of ''Stegosaurus'' plates to ''
Alligator An alligator is a large reptile in the Crocodilia order in the genus ''Alligator'' of the family Alligatoridae. The two extant species are the American alligator (''A. mississippiensis'') and the Chinese alligator (''A. sinensis''). Additional ...
'' osteoderms seem to support the conclusion that the potential for a thermoregulatory role in the plates of ''Stegosaurus'' definitely exists. The thermoregulation hypothesis has been seriously questioned, since other stegosaurs such as ''Kentrosaurus'', had more low
surface area The surface area of a solid object is a measure of the total area that the surface of the object occupies. The mathematical definition of surface area in the presence of curved surfaces is considerably more involved than the definition of ...
spikes than plates, implying that cooling was not important enough to require specialized structural formations such as plates. However, it has also been suggested that the plates could have helped the animal increase heat absorption from the sun. Since a cooling trend occurred towards the end of the Jurassic, a large ectothermic reptile might have used the increased surface area afforded by the plates to absorb radiation from the sun. Christiansen and Tschopp (2010) state that the presence of a smooth, insulating keratin covering would have hampered thermoregulation, but such a function cannot be entirely ruled out as extant cattle and ducks use horns and beaks to dump excess heat despite the keratin covering. Histological surveys of plate microstructure attributed the vascularization to the need to transport nutrients for rapid plate growth. The vascular system of the plates have been theorized to have played a role in threat displaying as ''Stegosaurus'' could have pumped blood into them, causing them to " blush" and give a colorful, red warning. However, Christiansen and Tschopp (2010) consider this unlikely, as stegosaur plates were covered in horn rather than skin. The plates' large size suggests that they may have served to increase the apparent height of the animal, either to intimidate enemies or to impress other members of the same species in some form of sexual display. A 2015 study of the shapes and sizes of ''Hesperosaurus'' plates suggested that they were
sexually dimorphic Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...
, with wide plates belonging to males and taller plates belonging to females. Christiansen and Tschopp (2010) proposed that the display function would have been reinforced by the horny sheath which would have increased the visible surface and such horn structures are often brightly colored. Some have suggested that plates in stegosaurs were used to allow individuals to identify members of their species. The use of exaggerated structures in dinosaurs as species identification has been questioned, as no such function exists in modern species.


Thagomizer (tail spikes)

Debate has been going on about whether the tail spikes were used for display only, as posited by Gilmore in 1914 or used as a weapon. Robert Bakker noted the tail was likely to have been much more flexible than that of other dinosaurs, as it lacked ossified tendons, thus lending credence to the idea of the tail as a weapon. However, as Carpenter has noted, the plates overlap so many tail vertebrae, movement would be limited. Bakker also observed that ''Stegosaurus'' could have maneuvered its rear easily, by keeping its large hind limbs stationary and pushing off with its very powerfully muscled but short forelimbs, allowing it to swivel deftly to deal with attack. More recently, a study of the tail spikes by McWhinney ''et al.'', which showed a high incidence of trauma-related damage, lends more weight to the position that the spikes were indeed used in combat. This study showed that 9.8% of ''Stegosaurus'' specimens examined had injuries to their tail spikes. Additional support for this idea was a punctured tail vertebra of an ''Allosaurus'' into which a tail spike fits perfectly. ''S. stenops'' had four dermal spikes, each about long. Discoveries of articulated stegosaur armor show, at least in some species, these spikes protruded horizontally from the tail, not vertically as is often depicted. Initially, Marsh described ''S. ungulatus'' as having eight spikes in its tail, unlike ''S. stenops''. However, recent research re-examined this and concluded this species also had four.


"Second brain"

At one time, stegosaurs were described as having a "second brain" in their hips. Soon after describing ''Stegosaurus'', Marsh noted a large canal in the hip region of the spinal cord, which could have accommodated a structure up to 20 times larger than the famously small brain. This has led to the influential idea that dinosaurs like ''Stegosaurus'' had a "second brain" in the tail, which may have been responsible for controlling reflexes in the rear portion of the body. This "brain" was proposed to have given a ''Stegosaurus'' a temporary boost when it was under threat from predators. This space, however, is more likely to have served other purposes. The sacro-lumbar expansion is not unique to stegosaurs, nor even ornithischians. It is also present in birds. In their case, it contains what is called the glycogen body, a structure whose function is not definitely known, but which is postulated to facilitate the supply of
glycogen Glycogen is a multibranched polysaccharide of glucose that serves as a form of energy storage in animals, fungi, and bacteria. The polysaccharide structure represents the main storage form of glucose in the body. Glycogen functions as one of ...
to the animal's
nervous system In biology, the nervous system is the highly complex part of an animal that coordinates its actions and sensory information by transmitting signals to and from different parts of its body. The nervous system detects environmental changes ...
. It also may function as a balance organ, or reservoir of compounds to support the nervous system.


Growth and metabolism

Juveniles of ''Stegosaurus'' have been preserved, probably showing the growth of the genus. The two juveniles are both relatively small, with the smaller individual being long, and the larger having a length of . The specimens can be identified as not mature because they lack the fusion of the
scapula The scapula (plural scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on eith ...
and coracoid, and the lower hind limbs. Also, the pelvic region of the specimens are similar to ''Kentrosaurus'' juveniles. One 2009 study of ''Stegosaurus'' specimens of various sizes found that the plates and spikes had delayed histological growth in comparison to the skeleton and when the dinosaur reached maturity, growth in the osteoderms may have increased. A 2013 study concluded, based on the rapid deposition of highly vascularised fibrolamellar bone, that ''Kentrosaurus'' had a quicker growth rate than ''Stegosaurus'', contradicting the general rule that larger dinosaurs grew faster than smaller ones. A 2022 study by Wiemann and colleagues of various dinosaur genera including ''Stegosaurus'' suggests that it had an
ectothermic An ectotherm (from the Greek () "outside" and () "heat") is an organism in which internal physiological sources of heat are of relatively small or of quite negligible importance in controlling body temperature.Davenport, John. Animal Life ...
(cold blooded) or gigantothermic metabolism, on par with that of modern reptiles. This was uncovered using the
spectroscopy Spectroscopy is the field of study that measures and interprets the electromagnetic spectra that result from the interaction between electromagnetic radiation and matter as a function of the wavelength or frequency of the radiation. Matter ...
of lipoxidation signals, which are byproducts of
oxidative phosphorylation Oxidative phosphorylation (UK , US ) or electron transport-linked phosphorylation or terminal oxidation is the metabolic pathway in which cells use enzymes to oxidize nutrients, thereby releasing chemical energy in order to produce adenosine t ...
and correlate with metabolic rates. They suggested that such metabolisms may have been common for ornithischian dinosaurs in general, with the group evolving towards ectothermy from an ancestor with an
endothermic In thermochemistry, an endothermic process () is any thermodynamic process with an increase in the enthalpy (or internal energy ) of the system.Oxtoby, D. W; Gillis, H.P., Butler, L. J. (2015).''Principle of Modern Chemistry'', Brooks Cole. ...
(warm blooded) metabolism.


Diet

''Stegosaurus'' and related genera were herbivores. However, their teeth and jaws are very different from those of other herbivorous ornithischian dinosaurs, suggesting a different feeding strategy that is not yet well understood. The other ornithischians possessed teeth capable of grinding plant material and a jaw structure capable of movements in planes other than simply orthal (i.e. not only the fused up-down motion to which stegosaur jaws were likely limited). Unlike the sturdy jaws and grinding teeth common to its fellow ornithischians, ''Stegosaurus'' (and all stegosaurians) had small, peg-shaped teeth that have been observed with horizontal wear facets associated with tooth-food contact and their unusual jaws were probably capable of only orthal (up-down) movements. Their teeth were "not tightly pressed together in a block for efficient grinding", and no evidence in the fossil record of stegosaurians indicates use of gastroliths—the stone(s) some dinosaurs (and some present-day bird species) ingested—to aid the grinding process, so how exactly ''Stegosaurus'' obtained and processed the amount of plant material required to sustain its size remains "poorly understood". The stegosaurians were widely distributed geographically in the late Jurassic. Palaeontologists believe it would have eaten plants such as mosses, ferns, horsetails, cycads, and conifers or fruits. One hypothesized feeding behavior strategy considers them to be low-level browsers, eating low-growing fruit of various nonflowering plants, as well as foliage. This scenario has ''Stegosaurus'' foraging at most 1 m above the ground. Conversely, if ''Stegosaurus'' could have raised itself on two legs, as suggested by Bakker, then it could have browsed on vegetation and fruits quite high up, with adults being able to forage up to above the ground. A detailed computer analysis of the biomechanics of ''Stegosauruss feeding behavior was performed in 2010, using two different three-dimensional models of ''Stegosaurus'' teeth given realistic physics and properties. Bite force was also calculated using these models and the known skull proportions of the animal, as well as simulated tree branches of different size and hardness. The resultant bite forces calculated for ''Stegosaurus'' were 140.1
newtons The newton (symbol: N) is the unit of force in the International System of Units (SI). It is defined as 1 kg⋅m/s, the force which gives a mass of 1 kilogram an acceleration of 1 metre per second per second. It is named after Isaac Newton in r ...
(N), 183.7 N, and 275 N (for anterior, middle and posterior teeth, respectively), which means its bite force was less than half that of a Labrador retriever. ''Stegosaurus'' could have easily bitten through smaller green branches, but would have had difficulty with anything over 12 mm in diameter. ''Stegosaurus'', therefore, probably browsed primarily among smaller twigs and foliage, and would have been unable to handle larger plant parts unless the animal was capable of biting much more efficiently than predicted in this study. However, a 2016 study indicates that ''Stegosaurus'' bite strength was stronger than previously believed. Comparisons were made between it (represented by a specimen known as "
Sophie Sophie is a version of the female given name Sophia, meaning "wise". People with the name Born in the Middle Ages * Sophie, Countess of Bar (c. 1004 or 1018–1093), sovereign Countess of Bar and lady of Mousson * Sophie of Thuringia, Duchess o ...
" from the United Kingdom's Natural History Museum) and two other herbivorous dinosaurs; ''
Erlikosaurus ''Erlikosaurus'' (meaning " Erlik's lizard") is a genus of therizinosaurid that lived in Asia during the Late Cretaceous period. The fossils, a skull and some post-cranial fragments, were found in the Bayan Shireh Formation of Mongolia in 1972, ...
'' and '' Plateosaurus'' to determine if all three had similar bite forces and similar niches. Based on the results of the study, it was revealed that the subadult ''Stegosaurus'' specimen had a bite similar in strength to that of modern herbivorous mammals, in particular, cattle and sheep. Based on this data, it is likely ''Stegosaurus'' also ate woodier, tougher plants such as cycads, perhaps even acting as a means of spreading cycad seeds.


Paleoecology

The Morrison Formation is interpreted as a semiarid environment with distinct wet and dry seasons, and flat floodplains. Vegetation varied from river-lining forests of conifers, tree ferns, and ferns (gallery forests), to fern savannas with occasional trees such as the
Araucaria ''Araucaria'' (; original pronunciation: .ɾawˈka. ɾja is a genus of evergreen coniferous trees in the family Araucariaceae. There are 20 extant species in New Caledonia (where 14 species are endemic, see New Caledonian ''Araucaria ...
-like conifer '' Brachyphyllum''. The flora of the period has been revealed by fossils of green algae, fungi, mosses, horsetails, ferns, cycads, ginkoes, and several families of conifers. Animal fossils discovered include bivalves, snails, ray-finned fishes, frogs, salamanders, turtles like ''
Glyptops ''Glyptops'' (Greek for "grooved face") is an extinct genus of pleurosternid freshwater turtle known from the Late Jurassic of North America. Taxonomy The type species, ''Glyptops plicatulus'', was first described as ''Compsemys plicatulus'' by ...
'', sphenodonts, lizards, terrestrial and aquatic crocodylomorphans like '' Hoplosuchus'', several species of
pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 ...
s such as '' Harpactognathus'' and '' Mesadactylus'', numerous dinosaur species, and early mammals such as docodonts (like '' Docodon''), multituberculates, symmetrodonts, and triconodonts. Dinosaurs that lived alongside ''Stegosaurus'' included theropods ''
Allosaurus ''Allosaurus'' () is a genus of large carnosaurian theropod dinosaur that lived 155 to 145 million years ago during the Late Jurassic epoch ( Kimmeridgian to late Tithonian). The name "''Allosaurus''" means "different lizard" alludin ...
'', '' Saurophaganax'', '' Torvosaurus'', '' Ceratosaurus'', '' Marshosaurus'', '' Stokesosaurus'', '' Ornitholestes'', '' Coelurus'' and '' Tanycolagreus''. Sauropods dominated the region, and included''
Brachiosaurus ''Brachiosaurus'' () is a genus of sauropod dinosaur that lived in North America during the Late Jurassic, about 154to 150million years ago. It was first Species description, described by American paleontologist Elmer S. Riggs in 1903 in pal ...
'', ''
Apatosaurus ''Apatosaurus'' (; meaning "deceptive lizard") is a genus of herbivorous sauropod dinosaur that lived in North America during the Late Jurassic period. Othniel Charles Marsh described and named the first-known species, ''A. ajax'', in 1 ...
'', ''
Diplodocus ''Diplodocus'' (, , or ) was a genus of diplodocid sauropod dinosaurs, whose fossils were first discovered in 1877 by S. W. Williston. The generic name, coined by Othniel Charles Marsh in 1878, is a neo-Latin term derived from Greek δ ...
'', ''
Camarasaurus ''Camarasaurus'' ( ) was a genus of quadrupedal, herbivorous dinosaurs and is the most common North American sauropod fossil. Its fossil remains have been found in the Morrison Formation, dating to the Late Jurassic epoch ( Kimmeridgian to ...
'', and ''
Barosaurus ''Barosaurus'' ( ) was a giant, long-tailed, long-necked, plant-eating sauropod dinosaur closely related to the more familiar ''Diplodocus''. Remains have been found in the Morrison Formation from the Upper Jurassic Period of Utah and South Da ...
''. Other ornithischians included '' Camptosaurus'', '' Gargoyleosaurus'', ''
Dryosaurus ''Dryosaurus'' ( , meaning 'tree lizard', Greek ' () meaning 'tree, oak' and () meaning 'lizard'; the name reflects the forested habitat, not a vague oak-leaf shape of its cheek teeth as is sometimes assumed) is a genus of an ornithopod dinosaur ...
'', '' Othnielosaurus'' and '' Drinker''.Foster, J. (2007). "Appendix." Jurassic West: The Dinosaurs of the Morrison Formation and Their World. Indiana University Press. pp. 327-329. ''Stegosaurus'' is commonly found at the same sites as ''Allosaurus'', ''Apatosaurus'', ''Camarasaurus'', and'' Diplodocus''. ''Stegosaurus'' may have preferred drier settings than these other dinosaurs.


In popular culture

One of the most recognizable of all dinosaurs, ''Stegosaurus'' has been depicted on film, in cartoons and comics and as children's toys. Due to the fragmentary nature of most early ''Stegosaurus'' fossil finds, it took many years before reasonably accurate restorations of this dinosaur could be produced. The earliest popular image of ''Stegosaurus'' was an engraving produced by A. Tobin for the November 1884 issue of ''
Scientific American ''Scientific American'', informally abbreviated ''SciAm'' or sometimes ''SA'', is an American popular science magazine. Many famous scientists, including Albert Einstein and Nikola Tesla, have contributed articles to it. In print since 1845, it ...
'', which included the dinosaur amid a speculative Morrison age landscape. Tobin restored the ''Stegosaurus'' as bipedal and long-necked, with the plates arranged along the tail and the back covered in spikes. This covering of spikes might have been based on a misinterpretation of the teeth, which Marsh had noted were oddly shaped, cylindrical, and found scattered, such that he thought they might turn out to be small dermal spines.Debus, A. A. (2009). ''Prehistoric Monsters: The Real and Imagined Creatures of the Past That We Love to Fear''. McFarland. Marsh published his more accurate skeletal reconstruction of ''Stegosaurus'' in 1891, and within a decade ''Stegosaurus'' had become among the most-illustrated types of dinosaur. Artist
Charles R. Knight Charles Robert Knight (October 21, 1874 – April 15, 1953) was an American wildlife and paleoartist best known for his detailed paintings of dinosaurs and other prehistoric animals. His works have been reproduced in many books and are currently ...
published his first illustration of ''Stegosaurus ungulatus'' based on Marsh's skeletal reconstruction in a November 1897 issue of '' The Century Magazine''. This illustration would later go on to form the basis of the stop-motion puppet used in the 1933 film ''
King Kong King Kong is a fictional giant monster resembling a gorilla, who has appeared in various media since 1933. He has been dubbed The Eighth Wonder of the World, a phrase commonly used within the franchise. His first appearance was in the novelizat ...
''. Like Marsh's reconstruction, Knight's first restoration had a single row of large plates, though he next used a double row for his more well-known 1901 painting, produced under the direction of Frederic Lucas. Again under Lucas, Knight revised his version of ''Stegosaurus'' again two years later, producing a model with a staggered double row of plates. Knight would go on to paint a stegosaur with a staggered double plate row in 1927 for the
Field Museum of Natural History The Field Museum of Natural History (FMNH), also known as The Field Museum, is a natural history museum in Chicago, Illinois, and is one of the largest such museums in the world. The museum is popular for the size and quality of its educational ...
, and was followed by Rudolph F. Zallinger, who painted ''Stegosaurus'' this way in his "Age of Reptiles" mural at the Peabody Museum in 1947.Moore, R. (2014). ''Dinosaurs by the Decades: A Chronology of the Dinosaur in Science and Popular Culture''. ABC-CLIO. ''Stegosaurus'' made its major public debut as a paper mache model commissioned by the U.S.
National Museum of Natural History 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. ...
for the 1904
Louisiana Purchase Exposition The Louisiana Purchase Exposition, informally known as the St. Louis World's Fair, was an World's fair, international exposition held in St. Louis, Missouri, United States, from April 30 to December 1, 1904. Local, state, and federal funds tota ...
. The model was based on Knight's latest miniature with the double row of staggered plates, and was exhibited in the United States Government Building at the exposition in St. Louis before being relocated to
Portland, Oregon Portland (, ) is a port city in the Pacific Northwest and the largest city in the U.S. state of Oregon. Situated at the confluence of the Willamette and Columbia rivers, Portland is the county seat of Multnomah County, the most populous ...
for the Lewis and Clark Centennial Exposition in 1905. The model was moved to the Smithsonian National Museum of Natural History (now the
Arts and Industries Building The Arts and Industries Building is the second oldest (after The Castle) of the Smithsonian museums on the National Mall in Washington, D.C. Initially named the National Museum, it was built to provide the Smithsonian with its first proper faci ...
) in Washington, D.C. along with other prehistory displays, and to the current National Museum of Natural History building in 1911. Following renovations to the museum in the 2010s, the model was moved once again for display at the Museum of the Earth in
Ithaca, New York Ithaca is a city in the Finger Lakes region of New York, United States. Situated on the southern shore of Cayuga Lake, Ithaca is the seat of Tompkins County and the largest community in the Ithaca metropolitan statistical area. It is named ...
. The popularity of ''Stegosaurus'' is owed partly to its prominent display in natural history museums. Though considered one of the most distinctive types of dinosaur, ''Stegosaurus'' displays were missing from a majority of museums during the first half of the 20th century, due largely to the disarticulated nature of most fossil specimens. Until 1918, the only mounted skeleton of ''Stegosaurus'' in the world was O. C. Marsh's type specimen of ''S. ungulatus'' at the Peabody Museum of Natural History, which was put on display in 1910. However, this mount was dismantled in 1917 when the old Peabody Museum building was demolished. This historically significant specimen was re-mounted ahead of the opening of the new Peabody Museum building in 1925. 1918 saw the completion of the second ''Stegosaurus'' mount, and the first depicting ''S. stenops''. This mount was created under the direction of Charles Gilmore at the U.S. National Museum of Natural History. It was a composite of several skeletons, primarily USNM 6531, with proportions designed to closely follow the ''S. stenops'' type specimen, which had been on display in relief nearby since 1918. The aging mount was dismantled in 2003 and replaced with a cast in an updated pose in 2004. A third mounted skeleton of ''Stegosaurus'', referred to ''S. stenops'', was put on display at the
American Museum of Natural History 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 int ...
in 1932. Mounted under the direction of Charles J. Long, the American Museum mount was a composite consisting of partial remains filled in with replicas based on other specimens. In his article about the new mount for the museum's journal,
Barnum Brown Barnum Brown (February 12, 1873 – February 5, 1963), commonly referred to as Mr. Bones, was an American paleontologist. Named after the circus showman P. T. Barnum, he discovered the first documented remains of '' Tyrannosaurus'' during a career ...
described (and disputed) the popular misconception that the ''Stegosaurus'' had a "second brain" in its hips. Another composite mount, using specimens referred to ''S. ungulatus'' collected from
Dinosaur National Monument Dinosaur National Monument is an American national monument located on the southeast flank of the Uinta Mountains on the border between Colorado and Utah at the confluence of the Green and Yampa rivers. Although most of the monument area is i ...
between 1920 and 1922, was put on display at the
Carnegie Museum of Natural History The Carnegie Museum of Natural History (abbreviated as CMNH) is a natural history museum in the Oakland neighborhood of Pittsburgh, Pennsylvania. It was founded by Pittsburgh-based industrialist Andrew Carnegie in 1896. Housing some 22 million ...
in 1940.McGinnis, H. J. (1984). ''Carnegie's Dinosaurs''. Pittsburgh: Carnegie Institute.


See also

*
Timeline of stegosaur research This timeline of stegosaur research is a chronological listing of events in the history of paleontology focused on the stegosaurs, the iconic plate-backed, spike-tailed herbivorous eurypod dinosaurs that predominated during the Jurassic period. ...


References


External links


Europe's First Stegosaurus Boosts Pangaea Theory
{{Featured article Stegosaurs Dinosaurs of Europe Dinosaurs of the Morrison Formation Late Jurassic dinosaurs of North America Symbols of Colorado Fossil taxa described in 1877 Taxa named by Othniel Charles Marsh Paleontology in Colorado Ornithischian genera