Thyreophoran stegosaurid dinosaur genus from Late Jurassic period
For the pachycephalosaurid of a similar name, see
Stegoceras
.
Stegosaurus
(
;
[2]
lit.
'
roof-lizard
'
) is a genus of
herbivorous
, four-legged,
armored dinosaur
from the
Late Jurassic
, characterized by the distinctive kite-shaped upright
plates
along their backs and
spikes
on their tails.
Fossils
of the genus have been found in the western
United States
and in
Portugal
, where they are found in
Kimmeridgian
- to
Tithonian
-aged
strata
, dating to between 155 and 145
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
,
Diplodocus
,
Camarasaurus
and
Allosaurus
, the latter of which may have preyed on it.
They were large, heavily built, herbivorous
quadrupeds
with rounded backs, short fore limbs, long hind limbs, and tails held high in the air. Due to their distinctive combination of broad, upright plates and tail tipped with spikes,
Stegosaurus
is one of the most recognizable kinds of dinosaurs. The function of this array of plates and spikes has been the subject of much speculation among scientists. Today, it is generally agreed that their spiked tails were most likely used for defense against predators, while their plates may have been used primarily for display, and secondarily for
thermoregulatory
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
, with the largest known specimens measuring about 7.5 metres (25 ft) long and weighing over 5 metric tons (5.5 short tons).
Stegosaurus
remains were first identified during the "
Bone Wars
" by
Othniel Charles Marsh
at
Dinosaur Ridge
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, on postal stamps, and in many other types of media.
History and naming
[
edit
]
Bone Wars and
Stegosaurus armatus
[
edit
]
Stegosaurus
, one of the many dinosaurs described in the
Bone Wars
, was first collected by
Arthur Lakes
and consisted of several caudal vertebrae, a dermal plate, and several additional postcranial elements that were collected north of
Morrison, Colorado
at Lakes’ YPM Quarry 5.
[3]
These first, fragmented bones (YPM 1850) became the
holotype
of
Stegosaurus armatus
when Yale paleontologist Othniel Charles Marsh described them in 1877.
[4]
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
(
tiles
) on a roof.
[4]
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
.
[3]
Because of this, the
International Code of Zoological Nomenclature
decided to replace the type species with the more well known species
Stegosaurus stenops
.
[5]
Marsh also incorrectly referred several fossils to
S. armatus
, including the dentary and teeth of the sauropod
Diplodocus
and putting sauropod limb bones and an
Allosaurus
tibia under YPM 1850.
[6]
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
in 1878.
[7]
Many later researchers have considered
Hypsirhophus
to be a
synonym
of
Stegosaurus
,
[8]
though Peter Galton (2010) suggested that it is distinct based on differences in the vertebrae.
[3]
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
.
[9]
Arthur Lakes made another discovery later in 1879 at
Como Bluff
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.
[6]
The majority of the fossils came from Quarry 13, including the type specimen of
Stegosaurus ungulatus (
YPM 1853), which was collected by Lakes and
William Harlow Reed
the same year and named by Marsh.
[6]
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.
[6]
The type specimen also preserved the pes, which was the namesake of the species, meaning "hoofed roofed lizard".
[10]
[3]
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.
[6]
[3]
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
.
[8]
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.
[8]
[3]
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.
[3]
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.
[10]
[8]
Marshall P. Felch collected the skeleton throughout 1885 and 1886 from Morrison Formation strata at his quarry in Garden Park, a town near
Canon City
,
Colorado
.
[11]
[8]
The skeleton was expertly unearthed by Felch, who first divided the skeleton into labeled blocks and prepared them separately.
[8]
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.
[10]
[12]
This first reconstruction, of
S. ungulatus
with missing parts filled in from
S. stenops
, was published by Marsh in 1891. (In 1893,
Richard Lydekker
mistakenly re-published Marsh's drawing under the label
Hypsirhophus
.)
[8]
Early skeletal mounts and plate interpretation
[
edit
]
The skeleton of
S. stenops
has since been deposited at the
National Museum of Natural History
in
Washington D. C.
, where it has been on display since 1915.
[13]
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.
[13]
The type specimen of
S. ungulatus
(YPM 1853) was incorporated into the first ever mounted skeleton of a stegosaur at the
Peabody Museum of Natural History
in 1910 by
Richard Swann Lull
. 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.
[14]
Additional specimens recovered from the same quarry by the United States
National Museum of Natural History
, including tail vertebrae and an additional large plate (USNM 7414), belong to the same individual as YPM 1853.
[3]
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
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
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
wrote that the alternating pattern seen in
S. stenops
was probably due to shifting of the skeleton after death. He led the construction of the first ever
Stegosaurus
skeletal mount at the
Peabody Museum of Natural History
, which was depicted with paired plates. In 1914, Charles Gilmore argued against Lull's interpretation, noting that several specimens of
S. stenops
, including the now-completely prepared holotype, preserved the plates in alternating rows near the peak of the back, and that there was no evidence of the plates having shifted relative to the body during fossilization.
[8]
Gilmore and Lucas' interpretation became the generally accepted standard, and Lull's mount at the Peabody Museum was changed to reflect this in 1924.
[14]
Though considered one of the most distinctive types of dinosaur,
Stegosaurus
displays were missing from a majority of museums during the first half of the 20th century, due largely to the disarticulated nature of most fossil specimens. Until 1918, the only mounted skeleton of
Stegosaurus
in the world was O. C. Marsh's type specimen of
S. ungulatus
at the
Peabody Museum of Natural History
, which was put on display in 1910. However, this mount was dismantled in 1917 when the old Peabody Museum building was demolished.
[13]
This historically significant specimen was re-mounted ahead of the opening of the new Peabody Museum building in 1925.
[14]
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.
[13]
The aging mount was dismantled in 2003 and replaced with a cast in an updated pose in 2004.
[15]
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
described (and disputed) the popular misconception that the
Stegosaurus
had a "second brain" in its hips.
[16]
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
in 1940.
[17]
Plate arrangement
[
edit
]
One of the major subjects of books and articles about
Stegosaurus
is the plate arrangement.
[18]
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
,
[12]
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,
[19]
based on the arrangement of
iguana
dorsal spines.
- The plates were paired in a double row along the back, such as in Knight's 1901 reconstruction and the 1933 film
King Kong
.
- 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.
[20]
[21]
Second dinosaur rush
[
edit
]
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
, 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.
[22]
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.
[22]
[9]
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.
[9]
[23]
The AMNH mount is cast and on display at the Field Museum, which didn't collect any
Stegosaurus
skeletons during the Second Dinosaur Rush.
[23]
The Carnegie Museum in
Pittsburgh
on the other hand collected many
Stegosaurus
specimens, first at Freezout Hills in
Carbon County, Wyoming
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.
[24]
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.
[24]
Both the AMNH and CM material has been referred to
Stegosaurus ungulatus
.
[24]
[6]
Resurgent discoveries
[
edit
]
As part of the
Dinosaur Renaissance
and the resurgent interest in dinosaurs by museums and the public, fossils of
Stegosaurus
were once again being collected, though few have been fully described. An important discovery came in 1937 again at Garden Park by a high school teacher named Frank Kessler in while leading a nature hike. Kessler contacted the
Denver Museum of Nature and Science
, who sent paleontologist Robert Landberg.
[25]
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".
[25]
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.
[25]
The expedition was successful in finding a nearly complete
Stegosaurus
near the Kessler site by Bryan Small, who would become the eponym of the new site.
[25]
[26]
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.
[27]
[26]
[25]
The
Stegosaurus
skeletons have been mounted alongside an
Allosaurus
skeleton collected in Moffat County, Colorado originally in 1979.
[25]
1987 saw the discovery of a 40% complete
Stegosaurus
skeleton in
Rabbit Valley
in
Mesa County, Colorado
by Harold Bollan near the
Dinosaur Journey Museum
.
[28]
The skeleton was nicknamed the "Bollan
Stegosaurus"
and is in the collections of the Dinosaur Journey Museum.
[28]
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
.
[29]
2007 saw the description of a
Stegosaurus
specimen from the Upper Jurassic
Lourinha Formation
of
Portugal
, the specimen was placed as cf.
Stegosaurus ungulatus
by the describers.
[30]
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.
[30]
[27]
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 a quarry on the Red Canyon Ranch near
Shell, Wyoming
, and was excavated by crews from the Swiss
Sauriermuseum
in 2004 and later prepared by museum staff, who gave it the nickname Sarah after the landowner's daughter.
[27]
[31]
The skeleton had been excavated on private land and was available for purchase. The
Natural History Museum, London
worked with private donors, most notably Jeremy Herrmann, to find the funding and then arranged to purchase the specimen, which was given the new official museum collection specimen designation NHMUK PV R36730 and re-nicknamed Sophie after Jeremy Herrmann's daughter.
[32]
The mounted skeleton went on display in December 2014 and was scientifically described in 2015.
[27]
It is a young adult of undetermined sex, 5.8 m (19 ft) long and 2.9 m (9.5 ft) tall.
[27]
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
skeleton found nicknamed "Big Al II".
[31]
Description
[
edit
]
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 6.5 m (21.3 ft) in length and 3.5 metric tons (3.9 short tons) in body mass, while
S. ungulatus
reached 7 m (23.0 ft) in length and 3.8 metric tons (4.2 short tons) in body mass.
[33]
Some large individuals may have reached 7.5 m (25 ft) in length and 5.0?5.3 metric tons (5.5?5.8 short tons) in body mass.
[34]
[35]
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 4.6 m (15.1 ft) long and 2 m (6.6 ft) high, and is estimated to have weighed 1.5-2.2 metric tons (1.6-2.4 short tons)
[36]
while alive. It is on display in the University of Wyoming Geological Museum.
[37]
Skull
[
edit
]
The long and narrow
skull
was small in proportion to the body. It had a small
antorbital fenestra
, 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.
[38]
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.
[39]
[40]
Other researchers have interpreted these ridges as modified versions of similar structures in other ornithischians which might have supported fleshy cheeks, rather than beaks.
[41]
Stegosaurian teeth were small, triangular, and flat; wear facets show that they did grind their food.
[42]
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
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
) could have a brain of no more than 80 g (2.8 oz) contributed to the popular old idea that all dinosaurs were unintelligent, an idea now largely rejected.
[43]
Actual brain anatomy in
Stegosaurus
is poorly known, but the brain itself was small even for a dinosaur.
[44]
Skeleton
[
edit
]
In
Stegosaurus stenops
there are 27 bones in the
vertebral column
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
, two greater than
Huayangosaurus
, although
Miragaia
preserves 17 cervicals and an unknown number of dorsals. The first cervical vertebra is the
axis bone
, which is connected and often fused to the
atlas bone
. Farther posteriorly, the proportionately larger the cervicals become, although they do not change greatly in anything other than size. Past the first few dorsals, the
centrum
of the bones become more elongate front-to-back, and the
transverse
processes become more elevated
dorsal
. The sacrum of
S. stenops
includes four sacral vertebrae, but one of the dorsals is also incorporated into the structure. In some specimens of
S. stenops
, a caudal is also incorporated, as a caudosacral. In
Hesperosaurus
there are two dorsosacrals, and only four fused sacrals, but in
Kentrosaurus
there may be as many as seven vertebrae in the sacrum, with both dorsosacrals and caudosacrals.
S. stenops
preserves 46 caudal vertebrae, and up to 49, and along the series both the centrums and the
neural spines
become smaller, until the neural spines disappear at caudal 35. Around the middle of the tail, the neural spines become
bifurcated
, meaning they are divided near the top.
[27]
With multiple well-preserved skeletons,
S. stenops
preserves all regions of the body, including the limbs. The
scapula
(shoulder blade) is sub-rectangular, with a robust blade. Though it is not always perfectly preserved, the
acromion
ridge is slightly larger than in
Kentrosaurus
. The blade is relatively straight, although it curves towards the back. There is a small bump on the back of the blade, that would have served as the base of the
triceps
muscle. Articulated with the scapula, the
coracoid
is sub-circular.
[27]
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.
[45]
All four limbs were supported by pads behind the toes.
[46]
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 1 m (3.3 ft) above the ground.
[26]
Plates
[
edit
]
The most recognizable features of
Stegosaurus
are its dermal plates, which consisted of between 17 and 22 separate plates and flat spines.
[14]
These were highly modified
osteoderms
(bony-cored scales), similar to those seen in crocodiles and many lizards today. They were not directly attached to the animal's skeleton, instead arising from the skin. The largest plates were found over the hips and could measure over 60 cm (24 in) wide and 60 cm (24 in) tall.
[14]
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.
[3]
Many of the plates are manifestly
chiral
[20]
[21]
and no two plates of the same size and shape have been found for an individual; however plates have been correlated between individuals. Well preserved integumentary impressions of the plates of
Hesperosaurus
show a smooth surface with long and parallel, shallow grooves. This indicates that the plates were covered in
keratinous
sheaths.
[47]
Classification and species
[
edit
]
Like the spikes and shields of
ankylosaurs
, the
bony plates
and spines of stegosaurians evolved from the low-keeled osteoderms characteristic of basal thyreophorans.
[48]
Galton
(2019) interpreted plates of an armored dinosaur from the
Lower Jurassic
(
Sinemurian
-
Pliensbachian
) Lower
Kota Formation
of
India
as fossils of a member of
Ankylosauria
; the author argued that this finding indicates a probable early Early Jurassic origin for both Ankylosauria and its
sister group
Stegosauria.
[49]
The vast majority of stegosaurian dinosaurs thus far recovered belong to the
Stegosauridae
, which lived in the later part of the Jurassic and early Cretaceous, and which were defined by
Paul Sereno
as all stegosaurians more closely related to
Stegosaurus
than to
Huayangosaurus
.
[50]
This group is widespread, with members across the Northern Hemisphere,
Africa
and possibly
South America
.
[51]
Stegosaurus
frequently is discovered in a clade within the Stegosauridae called Stegosaurinae, usually including the taxa
Wuerhosaurus
and
Hesperosaurus
.
[52]
The
cladogram
below displays the results of the "preferred tree"
phylogenetic analysis
of Raven
et al
. (2023), showing the position of the Stegosaurinae within Stegosauria and
Eurypoda
.
[53]
In 2017, Raven and Maidment published a phylogenetic analysis including almost every known stegosaurian genus.
[54]
Their dataset was expanded upon in the following years with additional taxa. In their 2024 description of stegosaur fossil material from China's
Hekou Group
, Li
et al
. used a modified version of the dataset of Raven and Maidment to analyze the phylogenetic relations of the Stegosauria:
[1]
Species
[
edit
]
Many of the species initially described have since been considered to be invalid or synonymous with earlier named species,
[6]
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.
[55]
- Stegosaurus stenops
, meaning "narrow-faced roof lizard", was named by Marsh in 1887,
[11]
with the holotype having been collected by Marshall Felch at Garden Park, north of
Canon City
, Colorado, in 1886. This is the best-known species of
Stegosaurus
, mainly because its remains include at least one complete articulated skeleton. It had proportionately large, broad plates and rounded tail plates. Articulated specimens show that the plates were arranged alternating in a staggered double row.
S. stenops
is known from at least 50 partial skeletons of adults and juveniles, one complete skull, and four partial skulls. It was shorter than other species, at 6.5 m (21 ft).
[33]
Found in the Morrison Formation, Colorado, Wyoming, and Utah.
[41]
- Stegosaurus ungulatus
, meaning "hoofed roof lizard", was named by Marsh in 1879,
[10]
from remains recovered at
Como Bluff
, Wyoming (Quarry 12, near Robber's Roost).
[6]
It might be synonymous with
S. stenops
.
[27]
At 7 m (23.0 ft),
[33]
it was the longest species within the genus
Stegosaurus
. A fragmentary
Stegosaurus
specimen discovered in Portugal and dating from the upper
Kimmeridgian
-lower
Tithonian
stage has been tentatively assigned to this species.
[30]
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.
[3]
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.
[6]
The type specimen of
S. ungulatus
(YPM 1853) was incorporated into the first ever mounted skeleton of a stegosaur at the
Peabody Museum of Natural History
in 1910 by
Richard Swann Lull
. 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.
[14]
Additional specimens recovered from the same quarry by the United States
National Museum of Natural History
, including tail vertebrae and an additional large plate (USNM 7414), belong to the same individual as YPM 1853.
[3]
- Stegosaurus sulcatus
, meaning "furrowed roof lizard", was described by Marsh in 1887 based on a partial skeleton.
[11]
It has traditionally been considered a synonym of
S. armatus
,
[41]
though more recent studies suggest it is not.
[3]
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.
[23]
[56]
Peter Galton suggested it should be considered a valid species due to its unique spikes.
[3]
Susannah Maidment and colleagues in 2008 proposed extensive alterations to the taxonomy of
Stegosaurus
. They advocated synonymizing
S. stenops
and
S. ungulatus
with
S. armatus
, and sinking
Hesperosaurus
and
Wuerhosaurus
into
Stegosaurus
, with their
type species
becoming
Stegosaurus mjosi
and
Stegosaurus homheni
, respectively. They regarded
S. longispinus
as
dubious
. Thus, their conception of
Stegosaurus
would include three valid species (
S. armatus
,
S. homheni
, and
S. mjosi
) and would range from the
Late Jurassic
of North America and Europe to the
Early Cretaceous
of Asia.
[23]
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
.
[3]
[57]
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.
[27]
In 2024, Li and colleagues described specimen
GSAU
201201, a partial skeleton of a stegosaur from the upper
Hekou Group
of
Gansu Province
, China (discovered in
c.
2000-04
), which dates to the
Aptian
?
Albian
ages of the
Early Cretaceous
. The specimen consists of three articulated cervical vertebrae with associated ribs, three dorsal vertebrae, thirteen ribs, a right forelimb including a partial humerus, ulna, and radius, and one dermal plate. Although certain features of the fossil material are different when compared to
Wuerhosaurus
and
Stegosaurus stenops
, Li
et al
. considered the new specimen as
Stegosaurus
sp. Fossils of the ankylosaur
Taohelong
were also found in the same layers of the Hekou Group.
[1]
Doubtful species and junior synonyms
[
edit
]
- 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.
[4]
It is known from a partial skeleton, and more than 30 fragmentary specimens have been referred to it.
[41]
However, the
type specimen
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.
[6]
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.
[5]
- 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
.
[8]
- Diracodon laticeps
was described by Marsh in 1881, from some jawbone fragments.
[58]
Bakker
resurrected
D. laticeps
in 1986 as a senior synonym of
S. stenops
,
[59]
although others note that the material is not diagnostic and is only referable to
Stegosaurus
sp., making it a
nomen dubium
.
[6]
- 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.
[6]
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.
[3]
Reassigned species
[
edit
]
- Stegosaurus marshi
, which was described by Lucas in 1901, was renamed
Hoplitosaurus
in 1902.
[60]
- Stegosaurus priscus
, described by Nopcsa in 1911, was reassigned to
Lexovisaurus
,
[41]
and is now the type species of
Loricatosaurus
.
[23]
- Stegosaurus longispinus
was named by
Charles W. Gilmore
in 1914 based on a fragmentary postcranial skeleton that has largely been lost.
[61]
[8]
It is now the type species of the genus
Alcovasaurus,
though it has been referred to
Miragaia
.
[62]
[61]
- Stegosaurus madagascariensis
from
Madagascar
is known solely from teeth and was described by Piveteau in 1926. The teeth were variously attributed to a stegosaur, the theropod
Majungasaurus
,
[63]
a
hadrosaur
or even a
crocodylian
, but is now considered a possible ankylosaur.
[23]
- 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
.
[64]
It was referred to
Stegosaurus
in 2008 by Maidment
et al
,
[23]
but some still consider the species to be in its own genus.
[65]
[66]
- Stegosaurus mjosi
was described as
Hesperosaurus mjosi
by Carpenter
et al
in 2001 based on a partial skull and incomplete postcranial skeleton from the Morrison Formation of
Johnson County, Wyoming
. The species was referred to
Stegosaurus
mostly by Maidment
et al
starting in 2008,
[67]
[23]
but
Hesperosaurus
has been the more popular combination since the discovery of more remains.
[68]
Paleobiology
[
edit
]
Posture and movement
[
edit
]
Soon after its discovery, Marsh considered
Stegosaurus
to have been bipedal, due to its short forelimbs.
[69]
He had changed his mind, however, by 1891, after considering the heavy build of the animal.
[12]
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.
[41]
This has been proposed by
Bakker
[59]
[70]
and opposed by
Carpenter
.
[26]
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.
[71]
Stegosaurus
had short fore limbs in relation to its hind limbs. Furthermore, within the hind limbs, the lower section (comprising the
tibia
and
fibula
) was short compared with the
femur
. 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 15.3?17.9 km/h (9.5?11.1 mph).
[72]
[42]
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.
[73]
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.
[74]
Plate function
[
edit
]
The function of
Stegosaurus'
plates has been much debated. Marsh suggested that they functioned as some form of armor,
[69]
though Davitashvili (1961) disputed this, claiming that they were too fragile and ill-placed for defensive purposes, leaving the animal's sides unprotected.
[75]
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.
[76]
Christiansen and Tschopp (2010), having studied a well-preserved specimen of
Hesperosaurus
with skin impressions, concluded that the plates were covered in a keratin sheath which would have strengthened the plate as a whole and provided it with sharp cutting edges.
[47]
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.
[76]
Mobility of the plates, however, has been disputed by other paleontologists.
[77]
Another possible function of the plates is they may have helped to control the body temperature of the animal,
[77]
in a similar way to the sails of the
pelycosaurs
Dimetrodon
and
Edaphosaurus
(and modern elephant and rabbit ears). The plates had blood vessels running through grooves and air flowing around the plates would have cooled the blood.
[78]
Buffrenil, et al. (1986) found "extreme vascularization of the outer layer of bone",
[79]
[77]
which was seen as evidence that the plates "acted as thermoregulatory devices".
[79]
Likewise, 2010 structural comparisons of
Stegosaurus
plates to
Alligator
osteoderms seem to support the conclusion that the potential for a thermoregulatory role in the plates of
Stegosaurus
definitely exists.
[80]
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.
[81]
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.
[47]
Histological
surveys of plate microstructure attributed the vascularization to the need to transport nutrients for rapid plate growth.
[79]
[82]
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.
[83]
However, the stegosaur plates were covered in horn rather than skin.
[47]
The plates' large size suggests that they may have served to increase the apparent height of the animal, either to intimidate enemies
[8]
or to impress other members of the same species in some form of sexual
display
.
[75]
A 2015 study of the shapes and sizes of
Hesperosaurus
plates suggested that they were
sexually dimorphic
, with wide plates belonging to males and taller plates belonging to females.
[29]
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.
[47]
Some have suggested that plates in stegosaurs were used to allow individuals to identify members of their species.
[82]
The use of exaggerated structures in dinosaurs as species identification has been questioned, as no such function exists in modern species.
[84]
Thagomizer (tail spikes)
[
edit
]
There has been debate about whether the tail spikes were only used for display, as posited by Gilmore in 1914,
[8]
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.
[26]
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.
[59]
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.
[85]
Additional support for this idea was a punctured tail vertebra of an
Allosaurus
into which a tail spike fits perfectly. The damage shows that the spike entered at an angle from below and displaced a piece of the
process
upward, remodeled bone on the underside of the process shows that an infection developed.
[86]
S. stenops
had four dermal spikes, each about 60?90 cm (2.0?3.0 ft) 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.
[26]
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.
[6]
Growth and metabolism
[
edit
]
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 1.5 m (4.9 ft) long, and the larger having a length of 2.6 m (8.5 ft). The specimens can be identified as not mature because they lack the fusion of the
scapula
and
coracoid
, and the lower hind limbs. Also, the
pelvic region
of the specimens are similar to
Kentrosaurus
juveniles.
[87]
One 2009 study of
Stegosaurus
specimens of various sizes found that the plates and spikes had slower
histological
growth than the skeleton at least until the dinosaur reached its mature size.
[88]
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.
[89]
A 2022 study by Wiemann and colleagues of various dinosaur genera including
Stegosaurus
suggests that it had an
ectothermic
(cold blooded) or
gigantothermic
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
(warm blooded) metabolism.
[90]
Diet
[
edit
]
Stegosaurus
and related genera were herbivores. However, their teeth and jaws are very different from those of other herbivorous
ornithischian
dinosaurs, suggesting a different feeding strategy that is not yet well understood. The other ornithischians possessed teeth capable of grinding plant material and a jaw structure capable of movements in planes other than simply orthal (i.e. not only the fused up-down motion to which stegosaur jaws were likely limited). Unlike the sturdy jaws and grinding teeth common to its fellow ornithischians,
Stegosaurus
(and all stegosaurians) had small, peg-shaped teeth that have been observed with horizontal wear facets associated with tooth-food contact
[91]
and their unusual jaws were probably capable of only orthal (up-down) movements.
[41]
Their teeth were "not tightly pressed together in a block for efficient grinding",
[92]
and no evidence in the fossil record of stegosaurians indicates use of
gastroliths
?the stone(s) some dinosaurs (and some present-day bird species) ingested?to aid the grinding process, so how exactly
Stegosaurus
obtained and processed the amount of plant material required to sustain its size remains "poorly understood".
[92]
The stegosaurians were widely distributed geographically in the late Jurassic.
[41]
Palaeontologists believe it would have eaten plants such as mosses, ferns, horsetails, cycads, and conifers.
[93]
One hypothesized feeding behavior strategy considers them to be low-level browsers, eating low-growing foliage of various nonflowering plants. This scenario has
Stegosaurus
foraging at most 1 m above the ground.
[94]
Conversely, if
Stegosaurus
could have raised itself on two legs, as suggested by Bakker, then it could have browsed on vegetation quite high up, with adults being able to forage up to 6 m (20 ft) above the ground.
[42]
A detailed computer analysis of the biomechanics of
Stegosaurus'
s 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.
[95]
However, a 2016 study indicates that
Stegosaurus
's
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
and
Plateosaurus
to determine if all three had similar bite forces and similar niches. Based on the results of the study, it was revealed that the subadult
Stegosaurus
specimen had a bite similar in strength to that of modern herbivorous mammals, in particular, cattle and sheep. Based on this data, it is likely
Stegosaurus
also ate woodier, tougher plants such as cycads, perhaps even acting as a means of spreading cycad seeds.
[96]
"Second brain"
[
edit
]
Casts of the and brain cavity of
S. ungulatus
(left), and brain cavity marked with red (right)
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.
[42]
This space, however, is more likely to have served other purposes. The sacro-lumbar expansion is not unique to stegosaurs, nor even ornithischians. It is also present in birds. In their case, it contains what is called the
glycogen body
, a structure whose function is not definitely known, but which is postulated to facilitate the supply of
glycogen
to the animal's
nervous system
.
[97]
It also may function as a balance organ, or reservoir of compounds to support the nervous system.
[98]
Paleoecology
[
edit
]
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
. 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
, sphenodonts, lizards, terrestrial and aquatic crocodylomorphs like
Hoplosuchus
, several species of
pterosaurs
such as
Harpactognathus
and
Mesadactylus
, numerous dinosaur species, and early mammals such as
docodonts
(like
Docodon
),
multituberculates
,
symmetrodonts
, and
triconodonts
.
[99]
Dinosaurs that lived alongside
Stegosaurus
included theropods
Allosaurus
,
Saurophaganax
,
Torvosaurus
,
Ceratosaurus
,
Marshosaurus
,
Stokesosaurus
,
Ornitholestes
,
Coelurus
and
Tanycolagreus
. Sauropods dominated the region, and included
Brontosaurus
,
Brachiosaurus
,
Apatosaurus
,
Diplodocus
,
Camarasaurus
, and
Barosaurus
. Other ornithischians included
Camptosaurus
,
Gargoyleosaurus
,
Dryosaurus
,
Othnielosaurus
and
Drinker
.
[100]
Stegosaurus
is commonly found at the same sites as
Allosaurus
,
Apatosaurus
,
Camarasaurus
, and
Diplodocus
.
Stegosaurus
may have preferred drier settings than these other dinosaurs.
[101]
Cultural significance
[
edit
]
One of the most recognizable of all dinosaurs,
[42]
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 the French science illustrator Auguste-Michel Jobin,
[102]
[103]
which appeared in the November 1884 issue of
Scientific American
and elsewhere, and which depicted the dinosaur amid a speculative Morrison age Jurassic landscape. Jobin 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.
[104]
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.
[104]
Artist
Charles R. Knight
published his first illustration of
Stegosaurus ungulatus
based on Marsh's skeletal reconstruction in a November 1897 issue of
The Century Magazine
. This illustration would later go on to form the basis of the stop-motion puppet used in the 1933 film
King Kong
. 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
, who painted
Stegosaurus
this way in his "Age of Reptiles" mural at the Peabody Museum in 1947.
[105]
Stegosaurus
made its major public debut as a paper mache model commissioned by the U.S.
National Museum of Natural History
for the 1904
Louisiana Purchase Exposition
. The model was based on Knight's latest miniature with the double row of staggered plates,
[13]
and was exhibited in the United States Government Building at the exposition in
St. Louis
before being relocated to
Portland, Oregon
for the
Lewis and Clark Centennial Exposition
in 1905. The model was moved to the
Smithsonian
National Museum of Natural History (now the
Arts and Industries Building
) in
Washington, D.C.
along with other prehistory displays, and to the current National Museum of Natural History building in 1911. Following renovations to the museum in the 2010s, the model was moved once again for display at the
Museum of the Earth
in
Ithaca, New York
.
[106]
See also
[
edit
]
References
[
edit
]
- ^
a
b
c
Li, Ning; Li, Daqing; Peng, Guangzhao; You, Hailu (2024).
"The first stegosaurian dinosaur from Gansu Province, China"
.
Cretaceous Research
.
158
(in press). 105852.
Bibcode
:
2024CrRes.15805852L
.
doi
:
10.1016/j.cretres.2024.105852
.
S2CID
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