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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 The SPIKES protocol is a method used in clinical medicine to break bad news to patients and families. As receiving bad news can cause distress and anxiety, clinicians need to deliver the news carefully. By using the SPIKES method for introducing a ...
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
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...
and in
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 ...
, where they are found in
Kimmeridgian In the geologic timescale, the Kimmeridgian is an age in the Late Jurassic Epoch and a stage in the Upper Jurassic Series. It spans the time between 157.3 ± 1.0 Ma and 152.1 ± 0.9 Ma (million years ago). The Kimmeridgian follows the Oxford ...
- to early Tithonian-aged strata, 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 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 1877, ...
'', ''
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 δι ...
'', ''
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 described by American paleontologist Elmer S. Riggs in 1903 from fossils found in th ...
'', ''
Ceratosaurus ''Ceratosaurus'' (from Greek κέρας/κέρατος, ' meaning "horn" and σαῦρος ' meaning "lizard") was a carnivorous theropod dinosaur in the Late Jurassic period ( Kimmeridgian to Tithonian). The genus was first described in 1 ...
'', 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
quadruped Quadrupedalism is a form of locomotion where four limbs are used to bear weight and move around. An animal or machine that usually maintains a four-legged posture and moves using all four limbs is said to be a quadruped (from Latin ''quattuor' ...
s 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 temperature ...
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 stegosaurians, 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 Ac ...
" 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, Colorado, Morrison and just west of Denver. The Dinosaur Ridge area is one ...
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 Ac ...
, was first collected by
Arthur Lakes Arthur Lakes (December 21, 1844—November 21, 1917) was an American geologist, artist, writer, teacher and Episcopalian minister. He captured much of his geological and palaeontological field work in sketches and watercolours. Lakes is credited ...
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 sever ...
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, 'roof(ed) lizard' was due to his early belief that the plates lay flat over the animal's back, overlapping like the shingles (
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 o ...
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 ...
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 δι ...
'' 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 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 are ...
in 1878. Many later researchers have considered ''Hypsirhophus'' to be a synonym 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. 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, the site also dating to the Upper Jurassic of the Morrison Formation, 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 William Harlow Reed (9 June 1848 – 24 April 1915) was an American fossil collector and pioneer. He served as a curator at the Museum of Geology at the University of Wyoming, Laramie. He collected for a while for Othniel Charles Marsh but left a ...
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 A canyon (from ; archaic British English spelling: ''cañon''), or gorge, is a deep cleft between escarpments or cliffs resulting from weathering and the erosive activity of a river over geologic time scales. Rivers have a natural tendency to cu ...
,
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 ...
. 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, ...
, 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 The Peabody Museum of Natural History at Yale University is among the oldest, largest, and most prolific university natural history museums in the world. It was founded by the philanthropist George Peabody in 1866 at the behest of his nephew Oth ...
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 unl ...
. 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 ...
'' 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 unl ...
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 The Peabody Museum of Natural History at Yale University is among the oldest, largest, and most prolific university natural history museums in the world. It was founded by the philanthropist George Peabody in 1866 at the behest of his nephew Oth ...
, 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 ''Iguana'' (, ) is a genus of herbivorous lizards that are native to tropical areas of Mexico, Central America, South America, and the Caribbean. The genus was first described in 1768 by Austrian naturalist Josephus Nicolaus Laurenti in his ...
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
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 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 of Pennsylvania, United States, and the county seat of Allegheny County. It is the most populous city in both Allegheny County and Western Pennsylvania, the second-most populous city in Pennsylva ...
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 18 ...
in 1902–03. The fossils included only a couple postcranial remains, though in the 1900s-1920s Carnegie crews at Dinosaur National Monument 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 The Denver Museum of Nature & Science is a municipal natural history and science museum in Denver, Colorado. It is a resource for informal science education in the Rocky Mountain region. A variety of exhibitions, programs, and activities help mus ...
, 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 Mesa County is a county located in the U.S. state of Colorado. As of the 2020 census, the population was 155,703. The county seat is Grand Junction. The county was named for the many large mesas in the area, including Grand Mesa. Mesa County ...
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 ...
, 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 An antorbital fenestra (plural: fenestrae) is an opening in the skull that is in front of the eye sockets. This skull character is largely associated with archosauriforms, first appearing during the Triassic Period. Among extant archosaurs, bird ...
, the hole between the nose and eye common to most archosaurs, 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 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 (5 
short tons The short ton (symbol tn) is a measurement unit equal to . It is commonly used in the United States, where it is known simply as a ton, although the term is ambiguous, the single word being variously used for short, long, and metric ton. The vari ...
) 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 to the sacrum, 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 ''Hesperosaurus'' (meaning "western lizard", from Ancient Greek language, Classical Greek (') "western" and (') "lizard") is an herbivorous stegosaurian dinosaur from the Kimmeridgian epoch of the Jurassic period, approximately 156 million year ...
'', 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 In anatomy, the atlas (C1) is the most superior (first) cervical vertebra of the spine and is located in the neck. It is named for Atlas of Greek mythology because, just as Atlas supported the globe, it supports the entire head. The atlas is t ...
. 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 (Latin for ''center'') may refer to: Places In Greenland * Nuuk Centrum, a district of Nuuk, Greenland * Centrum Lake, Greenland In the Netherlands * Amsterdam-Centrum, the inner-most borough of Amsterdam, Netherlands * Rotterdam Centrum, a borou ...
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 tangen ...
processes become more elevated
dorsa 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 ...
l. 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 ''Hesperosaurus'' (meaning "western lizard", from Ancient Greek language, Classical Greek (') "western" and (') "lizard") is an herbivorous stegosaurian dinosaur from the Kimmeridgian epoch of the Jurassic period, approximately 156 million year ...
'' there are two dorsosacrals, and only four fused sacrals, but in ''
Kentrosaurus ''Kentrosaurus'' ( ; ) is a genus of stegosaurid dinosaur from the Late Jurassic in Lindi Region of Tanzania. The type species is ''K. aethiopicus'', named and described by German palaeontologist Edwin Hennig in 1915. Often thought to be a " pri ...
'' 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 ...
become smaller, until the neural spines disappear at caudal 35. Around the middle of the tail, the neural spines become
bifurcated Bifurcation or bifurcated may refer to: Science and technology * Bifurcation theory, the study of sudden changes in dynamical systems ** Bifurcation, of an incompressible flow, modeled by squeeze mapping the fluid flow * River bifurcation, the for ...
, 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 ac ...
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 The triceps, or triceps brachii (Latin for "three-headed muscle of the arm"), is a large muscle on the back of the upper limb of many vertebrates. It consists of 3 parts: the medial, lateral, and long head. It is the muscle principally responsibl ...
'' muscle. Articulated with the scapula, the
coracoid A coracoid (from Greek κόραξ, ''koraks'', raven) is a paired bone which is part of the shoulder assembly in all vertebrates except therian mammals (marsupials and placentals). In therian mammals (including humans), a coracoid process is prese ...
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 Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct amphi ...
(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 Chirality is a property of asymmetry important in several branches of science. The word ''chirality'' is derived from the Greek (''kheir''), "hand", a familiar chiral object. An object or a system is ''chiral'' if it is distinguishable from i ...
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 Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. Alpha-keratin (α-keratin) is a type of keratin found in vertebrates. It is the key structural material making up scales, hair, nails, feathers, ho ...
sheaths.


Classification and species

Like the spikes and shields of ankylosaurs, 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, proto- ...
(2019) interpreted plates of an armored dinosaur from the
Lower Jurassic The Early Jurassic Epoch (in chronostratigraphy corresponding to the Lower Jurassic Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic-Jurassic extinction event, 201.3 Ma&nb ...
(
Sinemurian In the geologic timescale, the Sinemurian is an age and stage in the Early or Lower Jurassic Epoch or Series. It spans the time between 199.3 ± 2 Ma and 190.8 ± 1.5 Ma (million years ago). The Sinemurian is preceded by the Hettangian and ...
-
Pliensbachian The Pliensbachian is an age (geology), age of the geologic timescale and stage (stratigraphy), stage in the stratigraphic column. It is part of the Early Jurassic, Early or Lower Jurassic epoch (geology), Epoch or series (stratigraphy), Series an ...
) Lower
Kota Formation The Kota Formation is a geological formation in India. The precise age of Kota Formation are uncertain, but it dates from the Early to Middle Jurassic, and is split into a Lower Member and Upper Member.Prasad GVR, and Manhas BK. 2007A new docodont ...
of
India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the so ...
as fossils of a member of
Ankylosauria 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. ...
; the author argued that this finding indicates a probable early Early Jurassic origin for both Ankylosauria and its sister group Stegosauria. The vast majority of stegosaurian dinosaurs thus far recovered belong to the
Stegosauridae Stegosauridae is a family of thyreophoran dinosaurs (armoured dinosaurs) within the suborder Stegosauria. The clade is defined as all species of dinosaurs more closely related to ''Stegosaurus'' than '' Huayangosaurus''.David B. Weishampel, Pete ...
, 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 sites ...
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 sout ...
. ''Stegosaurus'' frequently is discovered in its own clade in Stegosauridae called Stegosauridae, usually including the taxa ''
Wuerhosaurus ''Wuerhosaurus'' is a genus of stegosaurid dinosaur from the Early Cretaceous Period of western China and Inner Mongolia. As such, it was one of the last genera of stegosaurians known to have existed, since most others lived in the late Jurassic ...
'' and ''
Loricatosaurus ''Loricatosaurus'' (meaning "armored lizard") is a Stegosaurid genus from Callovian-age ( Middle Jurassic) rocks of England and France. Discovery and naming ''Loricatosaurus'' is known from remains previously assigned to '' Lexovisaurus'' tha ...
'', though ''
Hesperosaurus ''Hesperosaurus'' (meaning "western lizard", from Ancient Greek language, Classical Greek (') "western" and (') "lizard") is an herbivorous stegosaurian dinosaur from the Kimmeridgian epoch of the Jurassic period, approximately 156 million year ...
'' 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 In the geologic timescale, the Kimmeridgian is an age in the Late Jurassic Epoch and a stage in the Upper Jurassic Series. It spans the time between 157.3 ± 1.0 Ma and 152.1 ± 0.9 Ma (million years ago). The Kimmeridgian follows the Oxford ...
-lower Tithonian 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), 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 The Peabody Museum of Natural History at Yale University is among the oldest, largest, and most prolific university natural history museums in the world. It was founded by the philanthropist George Peabody in 1866 at the behest of his nephew Oth ...
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 unl ...
. 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 A canyon (from ; archaic British English spelling: ''cañon''), or gorge, is a deep cleft between escarpments or cliffs resulting from weathering and the erosive activity of a river over geologic time scales. Rivers have a natural tendency to cu ...
, 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 ''Hesperosaurus'' (meaning "western lizard", from Ancient Greek language, Classical Greek (') "western" and (') "lizard") is an herbivorous stegosaurian dinosaur from the Kimmeridgian epoch of the Jurassic period, approximately 156 million year ...
'' and ''
Wuerhosaurus ''Wuerhosaurus'' is a genus of stegosaurid dinosaur from the Early Cretaceous Period of western China and Inner Mongolia. As such, it was one of the last genera of stegosaurians known to have existed, since most others lived in the late Jurassic ...
'' 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 specime ...
becoming ''Stegosaurus mjosi'' and ''Stegosaurus homheni'', respectively. They regarded ''S. longispinus'' as
dubious Doubt is a mental state in which the mind remains suspended between two or more contradictory propositions, unable to be certain of any of them. Doubt on an emotional level is indecision between belief and disbelief. It may involve uncertainty ...
. 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 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 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''. * ''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 ...
'' in 1902. * ''Stegosaurus priscus'', described by Nopcsa in 1911, was reassigned to ''
Lexovisaurus ''Lexovisaurus'' is a genus of stegosaur from mid-to-Late Jurassic Europe, 165.7-164.7  mya. Fossils of limb bones and armor fragments have been found in middle to late Jurassic-aged strata of England. Discovery and taxonomy In the early ...
'', and is now the type species of ''
Loricatosaurus ''Loricatosaurus'' (meaning "armored lizard") is a Stegosaurid genus from Callovian-age ( Middle Jurassic) rocks of England and France. Discovery and naming ''Loricatosaurus'' is known from remains previously assigned to '' Lexovisaurus'' tha ...
''. * ''Stegosaurus longispinus'' was named by
Charles W. Gilmore Charles Whitney Gilmore (March 11, 1874 – September 27, 1945) was an American paleontologist who gained renown in the early 20th century for his work on vertebrate fossils during his career at the United States National Museum (now the N ...
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 Africa ...
is known solely from teeth and was described by Piveteau in 1926. The teeth were variously attributed to a stegosaur, the theropod ''
Majungasaurus ''Majungasaurus'' (; ) is a genus of abelisaurid theropod dinosaur that lived in Madagascar from 70 to 66 million years ago, at the end of the Cretaceous Period, making it one of the last known non-avian dinosaurs that went extinct during the ...
'', 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 incl ...
or even a
crocodylian Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period (Cenomanian stage) and are the closest living ...
, 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 Dong Zhiming (Chinese: 董枝明, Pinyin: ''Dǒng Zhimíng''; born January 1937) is a Chinese vertebrate paleontologist formerly employed at the Institute of Vertebrate Paleontology and Paleoanthropology (IVPP) in Beijing. He began working at th ...
. 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 Johnson County is a county in the north central part of the U.S. state of Wyoming. At the 2020 United States Census, the population was 8,447. The county seat is Buffalo. Kaycee is the only other incorporated town in the county. Johnson Count ...
. 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. 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 connects ...
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 i ...
) 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 wit ...
. 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 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 ''Hesperosaurus'' (meaning "western lizard", from Ancient Greek language, Classical Greek (') "western" and (') "lizard") is an herbivorous stegosaurian dinosaur from the Kimmeridgian epoch of the Jurassic period, approximately 156 million year ...
'' 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
pelycosaur Pelycosaur ( ) is an older term for basal or primitive Late Paleozoic synapsids, excluding the therapsids and their descendants. Previously, the term ''mammal-like reptile'' had been used, and pelycosaur was considered an order, but this is ...
s '' Dimetrodon'' and ''
Edaphosaurus ''Edaphosaurus'' (, meaning "pavement lizard" for dense clusters of teeth) is a genus of extinct edaphosaurid synapsids that lived in what is now North America and Europe around 303.4 to 272.5 million years ago, during the Late Carboniferous to ...
'' (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''). Additiona ...
'' 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 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 Histology, also known as microscopic anatomy or microanatomy, is the branch of biology which studies the microscopic anatomy of biological tissues. Histology is the microscopic counterpart to gross anatomy, which looks at larger structures vis ...
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 Display may refer to: Technology * Display device, output device for presenting information, including: ** Cathode ray tube, video display that provides a quality picture, but can be very heavy and deep ** Electronic visual display, output devi ...
. 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 glycogen body is an oval structure in the spinal cord of birds that is made of specialized cells that contain large amounts of glycogen. Housed within the synsacrum, the function of this structure is not known, but it does not seem to be rela ...
, a structure whose function is not definitely known, but which is postulated to facilitate the supply of glycogen 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 th ...
. 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 A coracoid (from Greek κόραξ, ''koraks'', raven) is a paired bone which is part of the shoulder assembly in all vertebrates except therian mammals (marsupials and placentals). In therian mammals (including humans), a coracoid process is prese ...
, 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 Histology, also known as microscopic anatomy or microanatomy, is the branch of biology which studies the microscopic anatomy of biological tissues. Histology is the microscopic counterpart to gross anatomy, which looks at larger structures vis ...
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 Gigantothermy (sometimes called ectothermic homeothermy or inertial homeothermy) is a phenomenon with significance in biology and paleontology, whereby large, bulky ectothermic animals are more easily able to maintain a constant, relatively high bod ...
metabolism, on par with that of modern reptiles. This was uncovered using the spectroscopy of lipoxidation signals, which are byproducts of oxidative phosphorylation 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 Ornithischia () is an extinct order of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the Greek st ...
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
gastrolith A gastrolith, also called a stomach stone or gizzard stone, is a rock held inside a gastrointestinal tract. Gastroliths in some species are retained in the muscular gizzard and used to grind food in animals lacking suitable grinding teeth. In oth ...
s—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 (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" 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, da ...
'' and ''
Plateosaurus ''Plateosaurus'' (probably meaning "broad lizard", often mistranslated as "flat lizard") is a genus of plateosaurid dinosaur that lived during the Late Triassic period, around 214 to 204 million years ago, in what is now Central and Northern Eur ...
'' 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-like conifer ''
Brachyphyllum ''Brachyphyllum'' (meaning "short leaf") is a form genus of fossil coniferous plant foliage. Plants of the genus have been variously assigned to several different conifer groups including Araucariaceae and Cheirolepidiaceae. They are known from ...
''. 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 ''Harpactognathus'' (meaning "seizing/grasping jaw") is a genus of pterosaur found in the Late Jurassic-age Morrison Formation of Albany County, Wyoming, United States. It is based on NAMAL 101, a partial skull consisting of the snout, recovere ...
'' and ''
Mesadactylus ''Mesadactylus'' ('mesa finger') is an extinct genus of pterosaur from the Kimmeridgian-Tithonian-age Upper Jurassic Morrison Formation of Colorado, United States. The genus was named in 1989 by James Jensen and Kevin Padian. The type species ...
'', numerous dinosaur species, and early mammals such as
docodonts Docodonta is an order of extinct mammaliaforms that lived during the Mesozoic, from the Middle Jurassic to Early Cretaceous. They are distinguished from other early mammaliaforms by their relatively complex molar teeth, from which the order ge ...
(like ''
Docodon ''Docodon'' (meaning 'beam tooth') was a mammaliaform from the Late Jurassic of western North America. It was the first docodontan cynodont to be named. Description ''Docodon'' was the first docodontan cynodont found and named, and later gave ...
''),
multituberculates Multituberculata (commonly known as multituberculates, named for the multiple tubercles of their teeth) is an extinct order of rodent-like mammals with a fossil record spanning over 130 million years. They first appeared in the Middle Jurassic, a ...
, 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 ''Saurophaganax'' ("lord of lizard-eaters") is a genus of large allosaurid dinosaur from the Morrison Formation of Late Jurassic (latest Kimmeridgian age, about 151 million years ago) Oklahoma, United States.Turner, C.E. and Peterson, F., (1999) ...
'', ''
Torvosaurus ''Torvosaurus'' () is a genus of carnivorous megalosaurid theropod dinosaur that lived approximately 165 to 148 million years ago during the late Middle and Late Jurassic period (Callovian to Tithonian stages) in what is now Colorado, Portuga ...
'', ''
Ceratosaurus ''Ceratosaurus'' (from Greek κέρας/κέρατος, ' meaning "horn" and σαῦρος ' meaning "lizard") was a carnivorous theropod dinosaur in the Late Jurassic period ( Kimmeridgian to Tithonian). The genus was first described in 1 ...
'', ''
Marshosaurus ''Marshosaurus'' is a genus of medium-sized carnivorous theropod dinosaur, belonging to the Megalosauroidea, from the Late Jurassic Morrison Formation of Utah and possibly Colorado. Description ''Marshosaurus'' was medium-sized for a thero ...
'', ''
Stokesosaurus ''Stokesosaurus'' (meaning "Stokes' lizard") is a genus of small (around in length), carnivorous early tyrannosauroid theropod dinosaurs from the late Jurassic period of Utah, United States. History From 1960 onwards Utah geologist William ...
'', ''
Ornitholestes ''Ornitholestes'' (meaning "bird robber") is a small theropod dinosaur of the late Jurassic (Brushy Basin Member of the Morrison Formation, middle Kimmeridgian age, about 154 million years agoTurner, C.E. and Peterson, F., (1999). "Biostratigrap ...
'', ''
Coelurus ''Coelurus'' ( ) is a genus of coelurosaurian dinosaur from the Late Jurassic period (mid-late Kimmeridgian faunal stage, 155–152 million years ago). The name means "hollow tail", referring to its hollow tail vertebrae ( Greek κο ...
'' and ''
Tanycolagreus ''Tanycolagreus'' is a genus of coelurosaurian theropod from the Late Jurassic of North America. Discovery and naming In 1995 Western Paleontological Laboratories, Inc. uncovered the partial skeleton of a small theropod at the Bone Cabin Quarr ...
''. 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 described by American paleontologist Elmer S. Riggs in 1903 from fossils found in th ...
'', ''
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 1877, ...
'', ''
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 δι ...
'', '' Camarasaurus'', 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 ''Camptosaurus'' ( ) is a genus of plant-eating, beaked ornithischian dinosaurs of the Late Jurassic period of western North America and possibly also Europe. The name means 'flexible lizard' ( Greek (') meaning 'bent' and (') meaning 'li ...
'', ''
Gargoyleosaurus ''Gargoyleosaurus'' (meaning "gargoyle lizard") is one of the earliest ankylosaurs known from reasonably complete fossil remains. The holotype was discovered in 1995 at the Bone Cabin Quarry West locality, in Albany County, Wyoming in exposures ...
'', ''
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 ''The Century Magazine'' was an illustrated monthly magazine first published in the United States in 1881 by The Century Company of New York City, which had been bought in that year by Roswell Smith and renamed by him after the Century Associatio ...
''. 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, and was followed by
Rudolph F. Zallinger Rudolph Franz Zallinger (; November 12, 1919 – August 1, 1995) was an American-based Austrians, Austrian-Russians, Russian artist. His most notable works include his mural ''The Age of Reptiles'' (1947) at Yale University's Peabody Museum of Na ...
, 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 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 St. Louis () is the second-largest city in Missouri, United States. It sits near the confluence of the Mississippi and the Missouri Rivers. In 2020, the city proper had a population of 301,578, while the bi-state metropolitan area, which e ...
before being relocated to
Portland, Oregon Portland (, ) is a port city in the Pacific Northwest and the list of cities in Oregon, largest city in the U.S. state of Oregon. Situated at the confluence of the Willamette River, Willamette and Columbia River, Columbia rivers, Portland is ...
for the
Lewis and Clark Centennial Exposition The Lewis and Clark Centennial Exposition, commonly also known as the Lewis and Clark Exposition, and officially known as the Lewis and Clark Centennial and American Pacific Exposition and Oriental Fair, was a worldwide exposition held in Portlan ...
in 1905. The model was moved to the Smithsonian National Museum of Natural History (now the Arts and Industries Building) 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, ...
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 a ...
. 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 The Peabody Museum of Natural History at Yale University is among the oldest, largest, and most prolific university natural history museums in the world. It was founded by the philanthropist George Peabody in 1866 at the behest of his nephew Oth ...
, 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 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 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