Allosaurus Robustus Holotype
   HOME

TheInfoList



OR:

''Allosaurus'' ( ) is an extinct
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
theropod Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
dinosaur that lived 155 to 145 million years ago during the
Late Jurassic The Late Jurassic is the third Epoch (geology), epoch of the Jurassic Period, and it spans the geologic time scale, geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic stratum, strata.Owen ...
period Period may refer to: Common uses * Period (punctuation) * Era, a length or span of time *Menstruation, commonly referred to as a "period" Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (o ...
(
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 154.8 ±0.8 Ma and 149.2 ±0.7 Ma (million years ago). The Kimmeridgian follows the Oxfordian ...
to late
Tithonian In the geological timescale, the Tithonian is the latest age (geology), age of the Late Jurassic Epoch and the uppermost stage (stratigraphy), stage of the Upper Jurassic Series. It spans the time between 149.2 ±0.7 annum, Ma and 143.1 ±0.6 (mi ...
ages). The first fossil remains that could definitively be ascribed to this
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
were described in
1877 Events January * January 1 – Queen Victoria is proclaimed Empress of India by the Royal Titles Act 1876, introduced by Benjamin Disraeli, the Prime Minister of the United Kingdom . * January 8 – Great Sioux War of 1876: Batt ...
by Othniel C. Marsh. The name "''Allosaurus''" means "different lizard", alluding to its lightweight , which Marsh believed were unique. The genus has a very complicated
taxonomy image:Hierarchical clustering diagram.png, 280px, Generalized scheme of taxonomy Taxonomy is a practice and science concerned with classification or categorization. Typically, there are two parts to it: the development of an underlying scheme o ...
and includes at least three valid
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
, the best known of which is ''A. fragilis''. The bulk of ''Allosaurus'' remains come from North America's
Morrison Formation The Morrison Formation is a distinctive sequence of Upper Jurassic sedimentary rock found in the western United States which has been the most fertile source of dinosaur fossils in North America. It is composed of mudstone, sandstone, siltston ...
, with material also known from the
Alcobaça Formation The Alcobaça Formation, previously known as the Guimarota Formation and also known as the Consolação Unit, is a geological Formation (geology), formation in Portugal. It dates back to the Kimmeridgian stage of the Late Jurassic. It is an impor ...
and
Lourinhã Formation The Lourinhã Formation () is a fossil-rich geological formation in western Portugal, named for the municipality of Lourinhã. The formation is mostly Late Jurassic in age (Kimmeridgian/Tithonian), with the top of the formation extending into the ...
in Portugal. It was known for over half of the 20th century as ''
Antrodemus ''Antrodemus'' ("chamber bodied") is a dubious genus of theropod dinosaur from the Upper Jurassic, probably the Morrison Formation, of Middle Park, Colorado. It contains one species, ''Antrodemus valens'', first described and named as a species ...
'', but a study of the abundant remains from the
Cleveland-Lloyd Dinosaur Quarry Jurassic National Monument, at the site of the Cleveland-Lloyd Dinosaur Quarry, well known for containing the densest concentration of Jurassic dinosaur fossils ever found, is a paleontological site located near Cleveland, Utah, in the San Rafa ...
returned the name "''Allosaurus''" to prominence. As one of the first well-known theropod dinosaurs, it has long attracted attention outside of paleontological circles. ''Allosaurus'' was a large
biped Bipedalism is a form of terrestrial locomotion where an animal moves by means of its two rear (or lower) limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' ' ...
al predator for its time. Its skull was light, robust, and equipped with dozens of sharp,
serrated Serration is a saw-like appearance or a row of sharp or tooth-like projections. A serrated cutting edge has many small points of contact with the material being cut. By having less contact area than a smooth blade or other edge, the applied pr ...
teeth. It averaged in length for ''A. fragilis'', with the largest specimens estimated as being long. Relative to the large and powerful legs, its three-fingered hands were small and the body was balanced by a long, muscular tail. It is classified in the family
Allosauridae Allosauridae is an extinct family of medium to large bipedal, carnivorous allosauroid theropod dinosaurs from the Late Jurassic. Allosauridae is a fairly old taxonomic group, having been first named by the American paleontologist Othniel Cha ...
. As the most abundant large predator of the Morrison Formation, ''Allosaurus'' was at the top of the food chain and probably preyed on large herbivorous dinosaurs such as
ornithopod Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (). They represent one of the most successful groups of herbivorous dinosaurs during the Cretaceous. The most primitive members of the group were bipedal and relatively sm ...
s,
stegosaurid Stegosauria is a group of herbivorous ornithischian dinosaurs that lived during the Jurassic and early Cretaceous periods. Stegosaurian fossils have been found mostly in the Northern Hemisphere (North America, Europe and Asia), Africa and South A ...
s, and
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their b ...
s. Scientists have debated whether ''Allosaurus'' had cooperative
social behavior Social behavior is behavior among two or more organisms within the same species, it encompasses any behavior in which one member affects another. Social behavior can be seen as similar to an exchange of goods, with the expectation that when you ...
and hunted in packs or was a solitary predator that forms congregations, with evidence supporting either side.


Discovery and history


Early discoveries and research

The discovery and early study of ''Allosaurus'' is complicated by the multiplicity of names coined 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 Aca ...
of the late 19th century. The first described
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 preserve ...
in this history was a bone obtained secondhand by Ferdinand V. Hayden in
1869 Events January * January 3 – Abdur Rahman Khan is defeated at Tinah Khan, and exiled from Afghanistan. * January 5 – Scotland's second oldest professional football team, Kilmarnock F.C., is founded. * January 20 – Elizabe ...
. It came from Middle Park, near
Granby, Colorado The Town of Granby is the List of municipalities in Colorado#Statutory town, Statutory Town that is the List of municipalities in Colorado, most populous municipality in Grand County, Colorado, Grand County, Colorado, United States. The town pop ...
, probably from
Morrison Formation The Morrison Formation is a distinctive sequence of Upper Jurassic sedimentary rock found in the western United States which has been the most fertile source of dinosaur fossils in North America. It is composed of mudstone, sandstone, siltston ...
rocks. The locals had identified such bones as "petrified horse hoofs". Hayden sent his specimen to
Joseph Leidy Joseph Mellick Leidy (September 9, 1823 – April 30, 1891) was an American paleontologist, parasitologist and anatomist. Leidy was professor of anatomy at the University of Pennsylvania, later becoming a professor of natural history at Swarth ...
, who identified it as half of a tail vertebra and tentatively assigned it to the European dinosaur genus '' Poekilopleuron'' as ''Poicilopleuron'' ''valens''. He later decided it deserved its own genus, ''Antrodemus''. ''Allosaurus'' itself is based on YPM 1930, a small collection of fragmentary bones including parts of three vertebrae, a rib fragment, a tooth, a toe bone, and the shaft of the right (upper arm bone).
Othniel Charles Marsh Othniel Charles Marsh (October 29, 1831 – March 18, 1899) was an American professor of paleontology. A prolific fossil collector, Marsh was one of the preeminent paleontologists of the nineteenth century. Among his legacies are the discovery or ...
gave these remains the name ''Allosaurus fragilis'' in 1877. ''Allosaurus'' comes from the
Greek Greek may refer to: Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group *Greek language, a branch of the Indo-European language family **Proto-Greek language, the assumed last common ancestor of all kno ...
words ''/'', meaning "strange" or "different", and ''/'', meaning "lizard" or "reptile". Marsh chose the name 'different lizard' because he believed that the vertebrae were different from those of other dinosaurs due to their lightweight construction. The species epithet ''fragilis'' is
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
for "fragile", again referring to the lightening features in the vertebrae. The bones were collected from the Morrison Formation of Garden Park, north of Cañon City. Marsh and
Edward Drinker Cope Edward Drinker Cope (July 28, 1840 – April 12, 1897) was an American zoologist, paleontology, paleontologist, comparative anatomy, comparative anatomist, herpetology, herpetologist, and ichthyology, ichthyologist. Born to a wealthy Quaker fam ...
, who were in scientific competition with each other, went on to coin several other genera based on similarly sparse material that would later figure in the taxonomy of ''Allosaurus''. These include Marsh's ''Creosaurus'' and ''Labrosaurus'', as well as Cope's ''Epanterias''. In their haste, Cope and Marsh did not always follow up on their discoveries (or, more commonly, those made by their subordinates). For example, after the discovery by Benjamin Mudge of the
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
specimen of ''Allosaurus'' in Colorado, Marsh elected to concentrate work in
Wyoming Wyoming ( ) is a landlocked U.S. state, state in the Mountain states, Mountain West subregion of the Western United States, Western United States. It borders Montana to the north and northwest, South Dakota and Nebraska to the east, Idaho t ...
. When work resumed at Garden Park in
1883 Events January * January 4 – ''Life'' magazine is founded in Los Angeles, California, United States. * January 10 – A fire at the Newhall Hotel in Milwaukee, Wisconsin, United States, kills 73 people. * January 16 – ...
, M. P. Felch found an almost complete ''Allosaurus'' and several partial skeletons. In addition, one of Cope's collectors, H. F. Hubbell, found a specimen in the
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, ...
area of Wyoming in
1879 Events January * January 1 ** The Specie Resumption Act takes effect. The United States Note is valued the same as gold, for the first time since the American Civil War. ** Brahms' Violin Concerto is premiered in Leipzig with Joseph Joachim ...
, but apparently did not mention its completeness and Cope never unpacked it. Upon unpacking it in
1903 Events January * January 1 – Edward VII is proclaimed Emperor of India. * January 10 – The Aceh Sultanate was fully annexed by the Dutch forces, deposing the last sultan, marking the end of the Aceh War that have lasted for al ...
(several years after Cope had died), it was found to be one of the most complete theropod specimens then known and the skeleton, now cataloged as
AMNH The American Museum of Natural History (AMNH) is a natural history museum on the Upper West Side of Manhattan in New York City. Located in Theodore Roosevelt Park, across the street from Central Park, the museum complex comprises 21 interco ...
5753, was put on public view in
1908 This is the longest year in either the Julian or Gregorian calendars, having a duration of 31622401.38 seconds of Terrestrial Time (or ephemeris time), measured according to the definition of mean solar time. Events January * January ...
. This is the well-known mount poised over a partial ''
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, a ...
'' skeleton as if
scavenging Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding be ...
it, illustrated as such in a painting by Charles R. Knight. Although notable as the first free-standing mount of a theropod dinosaur and often illustrated and photographed, it has never been scientifically described. The many names coined by Cope and Marsh complicated later research, with the situation further compounded by the terse descriptions they provided. Even at the time, authors such as Samuel W. Williston suggested that too many names had been coined. For example, Williston pointed out in
1901 December 13 of this year is the beginning of signed 32-bit Unix time, and is scheduled to end in January 19, 2038. Summary Political and military 1901 started with the unification of multiple British colonies in Australia on January ...
that Marsh had never been able to adequately distinguish ''Allosaurus'' from ''Creosaurus''. The most influential early attempt to sort out the situation was produced by Charles W. Gilmore in
1920 Events January * January 1 ** Polish–Soviet War: The Russian Red Army increases its troops along the Polish border from 4 divisions to 20. ** Kauniainen in Finland, completely surrounded by the city of Espoo, secedes from Espoo as its ow ...
. He came to the conclusion that the tail vertebra named ''Antrodemus'' by Leidy was indistinguishable from those of ''Allosaurus'' and that ''Antrodemus'' should be the preferred name because, as the older name, it had priority. ''Antrodemus'' became the accepted name for this familiar genus for over 50 years, until James H. Madsen published on the Cleveland-Lloyd specimens and concluded that ''Allosaurus'' should be used because ''Antrodemus'' was based on material with poor, if any, diagnostic features and locality information. For example, the
geological formation A geological formation, or simply formation, is a body of rock having a consistent set of physical characteristics (lithology) that distinguishes it from adjacent bodies of rock, and which occupies a particular position in the layers of rock expo ...
that the single bone of ''Antrodemus'' came from is unknown.


Cleveland-Lloyd discoveries and "Big Al"

Although sporadic work at what became known as Utah's
Cleveland-Lloyd Dinosaur Quarry Jurassic National Monument, at the site of the Cleveland-Lloyd Dinosaur Quarry, well known for containing the densest concentration of Jurassic dinosaur fossils ever found, is a paleontological site located near Cleveland, Utah, in the San Rafa ...
had taken place as early as
1927 Events January * January 1 – The British Broadcasting ''Company'' becomes the BBC, British Broadcasting ''Corporation'', when its Royal Charter of incorporation takes effect. John Reith, 1st Baron Reith, John Reith becomes the first ...
and the fossil site itself was described by William L. Stokes in
1945 1945 marked the end of World War II, the fall of Nazi Germany, and the Empire of Japan. It is also the year concentration camps were liberated and the only year in which atomic weapons have been used in combat. Events World War II will be ...
, major operations did not begin there until
1960 It is also known as the "Year of Africa" because of major events—particularly the independence of seventeen African nations—that focused global attention on the continent and intensified feelings of Pan-Africanism. Events January * Janu ...
. Madsen led a cooperative effort between 1960 and
1965 Events January–February * January 14 – The First Minister of Northern Ireland and the Taoiseach of the Republic of Ireland meet for the first time in 43 years. * January 20 ** Lyndon B. Johnson is Second inauguration of Lynd ...
involving nearly 40 institutions, during which thousands of bones were recovered from the site. The quarry is notable for the predominance of ''Allosaurus'' remains: the quarry preserves a minimum of 73 individual dinosaurs and at least 46 of those are ''A. fragilis.'' The bones found at the site were (no longer connected to each other) and well-mixed. The
taphonomy Taphonomy is the study of how organisms decay and become fossilized or preserved in the paleontological record. The term ''taphonomy'' (from Greek language, Greek , 'burial' and , 'law') was introduced to paleontology in 1940 by Soviet scientis ...
of the site has been intensely debated, with many mutually exclusive explanations proposed as to how it may have formed. Regardless of the actual cause, the great quantity of well-preserved ''Allosaurus'' remains has allowed this genus to be known in great detail, making it among the best-known of all theropods. Skeletal remains from the quarry pertain to individuals of almost all ages and sizes, from less than to long, and the disarticulation is an advantage for describing aspects of bones that otherwise would have been concealed by other bones. Because its fossils are common at both this quarry and others in the state, ''Allosaurus'' was designated as the
state fossil Most states in the US have designated a state fossil, many during the 1980s. It is common to designate a fossilized species, rather than a single specimen or a category of fossils. State fossils are distinct from other state emblems like state d ...
of Utah in
1988 1988 was a crucial year in the early history of the Internet—it was the year of the first well-known computer virus, the Morris worm, 1988 Internet worm. The first permanent intercontinental Internet link was made between the United State ...
. Significant finds would continue to be made after the Cleveland-Lloyd discoveries. In 1991, a Swiss team led by Kirby Siber discovered "Big Al" ( MOR 693): a 95% complete, partially articulated specimen that measured about 8 meters (about 26 ft) in length. The individual itself was below the average size for ''Allosaurus'', as it was a subadult estimated at only 87% grown. The excavation was taken over by a joint Museum of the Rockies and
University of Wyoming The University of Wyoming (UW) is a Public university, public land-grant university, land-grant research university in Laramie, Wyoming, United States. It was founded in March 1886, four years before the territory was admitted as the 44th state, ...
Geological Museum team. The Swiss team later excavated a second ''Allosaurus'', "Big Al Two". Both specimens are known to have multiple preserved injuries.


Portuguese discoveries

''Allosaurus'' fossils are known from at least three localities in Portugal. These specimens come from the
Lourinhã Lourinhã () is a municipality in the District of Lisbon and in the portuguese Oeste region. The population in 2011 was 25,735, in an area of 147.17 km2. The seat of the municipality is the town of Lourinhã, with a population of 8,800 inhab ...
,
Bombarral Bombarral () is a portuguese municipality in the Oeste region, historical province of Estremadura, and the Leiria district. The population in 2011 was 13,193, in an area of . It includes four civil parishes () that provide local services. Histo ...
, and Alcobaça formations, and, like the North American occurrences, date to the Kimmerdigian to Tithonian ages. The first specimen ( MNHNUL/AND.001), a partial skeleton that includes an articulated hindlimb and pelvis, was found in 1988 near the village of Andrés in the District of Leiria during the construction of a warehouse. Reported in 1999 and assigned to the species ''A. fragilis'', it was the first ''Allosaurus'' specimen to be found outside of North America. ''A. fragilis'' became the first dinosaur species known from both Europe and North America, suggesting faunal exchange between the two continents. This site has been worked on again between 2005 and 2010, resulting in the discovery of at least two more ''Allosaurus'' individuals, including many skull bones. In 2005, a single maxilla was reported from the Guimarota coal mine, a locality well-known for its fossils of Mesozoic mammals; this fossil was assigned to ''Allosaurus'' but not to any particular species. In 2006,
Octávio Mateus Octávio Mateus (born 1975) is a Portugal, Portuguese dinosaur paleontologist and biologist Professor of Paleontology at the Faculdade de Ciências e Tecnologia da NOVA University Lisbon, Universidade Nova de Lisboa. He graduated in University of ...
and colleagues reported a partial skull and three neck vertebrae ( ML 415) from the Vale Frades beach in
Lourinhã Lourinhã () is a municipality in the District of Lisbon and in the portuguese Oeste region. The population in 2011 was 25,735, in an area of 147.17 km2. The seat of the municipality is the town of Lourinhã, with a population of 8,800 inhab ...
. As this specimen differed from North American ''Allosaurus'' fossils, they assigned it to a new species, ''A. europaeus''. These authors also assigned the Andrés specimen to this species, though solely based on the fact that it was found in Portugal. The presence of a separate ''Allosaurus'' species in Europe would suggest that faunal interchange between the continents was interrupted, possibly due to the opening of the Atlantic Ocean. The status of ''A. europaeus'' was controversially discussed in the subsequent years, with different studies arguing that the species is a synonym of ''A. fragilis'', a ''
nomen dubium In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium,'' it may be impossible to determine whether a ...
'' (doubtful name), or in need of re-evaluation. In 2024, André Burigo and Mateus re-described the Vale Frades specimen and carried out further
fossil preparation Fossil preparation is a complex of tasks that can include excavating, revealing, conserving, and replicating the ancient remains and traces of organisms. It is an integral part of the science of paleontology, of museum exhibition, and the preser ...
that exposed additional bones. These authors identified nine unique features supporting the validity ''A. europaeus''. A 2025 study by Elisabete Malafaia and colleagues described the Andrés specimens in detail, including a newly discovered set of skull bones. Their analysis of the relationships between individual ''Allosaurus'' skulls instead suggested that ''A. europaeus'' is a synonym of ''A. fragilis'' because the Andrés specimens were more closely related to some North American specimens than to the Vale Frades specimen.


Taxonomy


Species

Seven species of ''Allosaurus'' have been named: ''A. anax'', ''A. amplus'', ''A. atrox'', ''A. europaeus'', the
type species In International_Code_of_Zoological_Nomenclature, 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 spe ...
''A. fragilis'', ''A. jimmadseni'' and ''A. lucasi''. Among these (excluding ''A. anax'', which was named in 2024), Chure and Loewen recovered only ''A. fragilis'', ''A. europaeus'', and the newly-named ''A. jimmadseni'' as being valid species. Some studies have suggested that ''A. europaeus'' does not show any unique characters compared to the North American species, though other authors have suggested that the species is valid and has a number of distinguishing characters. ''A. fragilis'' is the type species and was named by Marsh in 1877. It is known from the remains of at least 60 individuals, all found in the
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 154.8 ±0.8 Ma and 149.2 ±0.7 Ma (million years ago). The Kimmeridgian follows the Oxfordian ...
Tithonian In the geological timescale, the Tithonian is the latest age (geology), age of the Late Jurassic Epoch and the uppermost stage (stratigraphy), stage of the Upper Jurassic Series. It spans the time between 149.2 ±0.7 annum, Ma and 143.1 ±0.6 (mi ...
Upper Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
-age Morrison Formation of the United States, spread across
Colorado Colorado is a U.S. state, state in the Western United States. It is one of the Mountain states, sharing the Four Corners region with Arizona, New Mexico, and Utah. It is also bordered by Wyoming to the north, Nebraska to the northeast, Kansas ...
,
Montana Montana ( ) is a landlocked U.S. state, state in the Mountain states, Mountain West subregion of the Western United States. It is bordered by Idaho to the west, North Dakota to the east, South Dakota to the southeast, Wyoming to the south, an ...
,
New Mexico New Mexico is a state in the Southwestern United States, Southwestern region of the United States. It is one of the Mountain States of the southern Rocky Mountains, sharing the Four Corners region with Utah, Colorado, and Arizona. It also ...
,
Oklahoma Oklahoma ( ; Choctaw language, Choctaw: , ) is a landlocked U.S. state, state in the South Central United States, South Central region of the United States. It borders Texas to the south and west, Kansas to the north, Missouri to the northea ...
,
South Dakota South Dakota (; Sioux language, Sioux: , ) is a U.S. state, state in the West North Central states, North Central region of the United States. It is also part of the Great Plains. South Dakota is named after the Dakota people, Dakota Sioux ...
, Utah, and Wyoming. Details of the
humerus The humerus (; : humeri) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...
(upper arm) of ''A. fragilis'' have been used as diagnostic among Morrison theropods, but ''A. jimmadseni'' indicates that this is no longer the case at the species level. ''A. jimmadseni'' has been scientifically described based on two nearly complete skeletons. The first specimen to wear the identification was unearthed in Dinosaur National Monument in northeastern Utah, with the original "Big Al" individual subsequently recognized as belonging to the same species. This species differs from ''A. fragilis'' in several anatomical details, including a
jugal The jugal is a skull bone found in most reptiles, amphibians and birds. In mammals, the jugal is often called the malar or zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by species. Anatomy ...
(cheekbone) with a straight lower margin. Fossils are confined to the Salt Wash Member of the Morrison Formation, with ''A. fragilis'' only found in the higher Brushy Basin Member. However, stratigraphic work done by Suzannah Maidment found that both species were actually coeval and were instead segregated by geography, with ''A. fragilis'' mostly found in the southern parts of the Morrison Formation, while ''A. jimmadseni'' is largely found in the northern parts. The specific name ''jimmadseni'' is named in honor of Madsen, for his contributions to the taxonomy of the genus, notably for his 1976 work. ''A. fragilis'', ''A. jimmadseni'', ''A. anax'', ''A. amplus'', and ''A. lucasi'' are all known from remains discovered in the
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 154.8 ±0.8 Ma and 149.2 ±0.7 Ma (million years ago). The Kimmeridgian follows the Oxfordian ...
Tithonian In the geological timescale, the Tithonian is the latest age (geology), age of the Late Jurassic Epoch and the uppermost stage (stratigraphy), stage of the Upper Jurassic Series. It spans the time between 149.2 ±0.7 annum, Ma and 143.1 ±0.6 (mi ...
Upper Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
-age Morrison Formation of the United States, spread across
Colorado Colorado is a U.S. state, state in the Western United States. It is one of the Mountain states, sharing the Four Corners region with Arizona, New Mexico, and Utah. It is also bordered by Wyoming to the north, Nebraska to the northeast, Kansas ...
,
Montana Montana ( ) is a landlocked U.S. state, state in the Mountain states, Mountain West subregion of the Western United States. It is bordered by Idaho to the west, North Dakota to the east, South Dakota to the southeast, Wyoming to the south, an ...
,
New Mexico New Mexico is a state in the Southwestern United States, Southwestern region of the United States. It is one of the Mountain States of the southern Rocky Mountains, sharing the Four Corners region with Utah, Colorado, and Arizona. It also ...
,
Oklahoma Oklahoma ( ; Choctaw language, Choctaw: , ) is a landlocked U.S. state, state in the South Central United States, South Central region of the United States. It borders Texas to the south and west, Kansas to the north, Missouri to the northea ...
,
South Dakota South Dakota (; Sioux language, Sioux: , ) is a U.S. state, state in the West North Central states, North Central region of the United States. It is also part of the Great Plains. South Dakota is named after the Dakota people, Dakota Sioux ...
,
Utah Utah is a landlocked state in the Mountain states, Mountain West subregion of the Western United States. It is one of the Four Corners states, sharing a border with Arizona, Colorado, and New Mexico. It also borders Wyoming to the northea ...
and
Wyoming Wyoming ( ) is a landlocked U.S. state, state in the Mountain states, Mountain West subregion of the Western United States, Western United States. It borders Montana to the north and northwest, South Dakota and Nebraska to the east, Idaho t ...
. ''A. fragilis'' is regarded as the most common, known from the remains of at least 60 individuals. For a while in the late 1980s and early 1990s, it was common to recognize ''A. fragilis'' as the short-snouted species, with the long-snouted taxon being ''A. atrox.'' However, subsequent analysis of specimens from the Cleveland-Lloyd Dinosaur Quarry, Como Bluff, and Dry Mesa Quarry showed that the differences seen in the Morrison Formation material could be attributed to individual variation. A study of skull elements from the Cleveland-Lloyd site found wide variation between individuals, calling into question previous species-level distinctions based on such features as the shape of the lacrimal horns and the proposed differentiation of ''A. jimmadseni'' based on the shape of the
jugal The jugal is a skull bone found in most reptiles, amphibians and birds. In mammals, the jugal is often called the malar or zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by species. Anatomy ...
. ''A. anax'' was described and named in 2024 from several fossils representing various skeleton parts, the holotype being a
postorbital The ''postorbital'' is one of the bones in vertebrate skulls which forms a portion of the dermal skull roof and, sometimes, a ring about the orbit. Generally, it is located behind the postfrontal and posteriorly to the orbital fenestra. In some ve ...
numbered as OMNH 1771. This species is characterized by the lack of rugose ornamentation on the postorbital, the dorsal vertebrae with hourglass-shaped centra and
pneumatic Pneumatics (from Greek 'wind, breath') is the use of gas or pressurized air in mechanical systems. Pneumatic systems used in Industrial sector, industry are commonly powered by compressed air or compressed inert gases. A centrally located a ...
foramina, and other features of the postorbital, cervical vertebrae, and
fibula The fibula (: fibulae or fibulas) 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. ...
. The specific name comes from the Ancient Greek ἄναξ (''anax'', "king", "lord" or "tribal chief"), and is intended to be an updated reference to the now dubious saurischian genus ''
Saurophaganax ''Saurophaganax'' ("lord of lizard-eaters") is a dubious, chimeric genus of large saurischian dinosaur, possibly a sauropod, from the Late Jurassic ( Kimmeridgian) Morrison Formation of Oklahoma, United States. This taxon was historically co ...
'', to which the fossils were previously attributed. The ''Allosaurus'' material from Portugal has a controversial taxonomic research history. The Andrés ''Allosaurus'' specimens, consisting of very complete cranial and post-cranial remains, have been attributed to ''A. fragilis'', ''A.'' sp'', A. europaeus'' and ''A.'' cf. ''europaeus''. The Vale Frades ''Allosaurus'', consisting of a partial skull and cervical vertebrae and ribs, is the type specimen of ''A. europaeus'', although the validity of that species has been previously questioned. In 2024, a revised diagnosis of ''A. europaeus'' was published, confirming the validity of the species. The specific affinities of the Andrés specimens are still unclear. The issue of species and potential synonyms was historically complicated by the
type specimen In biology, a type is a particular wikt:en:specimen, specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally associated. In other words, a type is an example that serves to ancho ...
of ''Allosaurus fragilis'' ( YPM 1930) being extremely fragmentary, consisting of a few incomplete vertebrae, limb fragments, rib fragments, and a single tooth. Because of this, several scientists have interpreted the type specimen as potentially dubious, meaning the genus ''Allosaurus'' itself or at least the species ''A. fragilis'' would be a ''nomen dubium'' ("dubious name", based on a specimen too incomplete to compare to other specimens or to classify). To address this situation,
Gregory S. Paul Gregory Scott Paul (born December 24, 1954) is an American freelance researcher, author and illustrator who works in paleontology. He is best known for his work and research on theropoda, theropod dinosaurs and his detailed illustrations, both l ...
and
Kenneth Carpenter Kenneth Carpenter (born 21 September 1949) is an American paleontologist. He is the former director of the USU Eastern Prehistoric Museum and author or co-author of books on dinosaurs and Mesozoic life. His main research interests are armore ...
(
2010 The year saw a multitude of natural and environmental disasters such as the 2010 Haiti earthquake, the Deepwater Horizon oil spill, and the 2010 Chile earthquake. The 2009 swine flu pandemic, swine flu pandemic which began the previous year ...
) submitted a petition to the
ICZN The International Code of Zoological Nomenclature (ICZN) is a widely accepted convention in zoology that rules the formal scientific naming of organisms treated as animals. It is also informally known as the ICZN Code, for its formal author, t ...
to have the name ''A. fragilis'' officially transferred to the more complete specimen USNM 4734 (as a
neotype 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 associated. In other words, a type is an example that serves to anchor or centralizes ...
), a decision that was ratified by the ICZN on December 29, 2023. Teeth of indeterminate species of ''Allosaurus'' have been reported from Tönniesberg and Kahlberg in
Saxony Saxony, officially the Free State of Saxony, is a landlocked state of Germany, bordering the states of Brandenburg, Saxony-Anhalt, Thuringia, and Bavaria, as well as the countries of Poland and the Czech Republic. Its capital is Dresden, and ...
, Germany, dating to the upper Kimmeridigian.


Synonyms

''Creosaurus'', ''Epanterias'', and ''Labrosaurus'' are provisionally regarded as junior synonyms of ''Allosaurus'', though the latter two require new analyses to clarify their specific status. Most of the species that are regarded as synonyms of ''A. fragilis'', or that were misassigned to the genus, are obscure and based on very scrappy remains. One exception is ''Labrosaurus ferox'', named in
1884 Events January * January 4 – The Fabian Society is founded in London to promote gradualist social progress. * January 5 – Gilbert and Sullivan's comic opera '' Princess Ida'', a satire on feminism, premières at the Savoy The ...
by Marsh for an oddly formed partial lower jaw, with a prominent gap in the tooth row at the tip of the jaw, and a rear section greatly expanded and turned down. Later researchers suggested that the bone was pathologic, showing an injury to the living animal, and that part of the unusual form of the rear of the bone was due to plaster reconstruction. It is now regarded as an example of ''A. fragilis''. In his
1988 1988 was a crucial year in the early history of the Internet—it was the year of the first well-known computer virus, the Morris worm, 1988 Internet worm. The first permanent intercontinental Internet link was made between the United State ...
book, ''Predatory Dinosaurs of the World'', the freelance artist & author Gregory S. Paul proposed that ''A. fragilis'' had tall pointed horns and a slender build compared to a postulated second species ''A. atrox'', as well as not being a different sex due to rarity. ''Allosaurus atrox'' was originally named by Marsh in
1878 Events January * January 5 – Russo-Turkish War: Battle of Shipka Pass IV – Russian and Bulgarian forces defeat the Ottoman Empire. * January 9 – Umberto I becomes King of Italy. * January 17 – Russo-Turkish War: ...
as the type species of its own genus, ''Creosaurus'', and is based on YPM 1890, an assortment of bones that includes a couple of pieces of the skull, portions of nine tail vertebrae, two hip vertebrae, an ilium, and ankle and foot bones. Although the idea of two common Morrison allosaur species was followed in some semi-technical and popular works, the
2000 2000 was designated as the International Year for the Culture of Peace and the World Mathematics, Mathematical Year. Popular culture holds the year 2000 as the first year of the 21st century and the 3rd millennium, because of a tende ...
thesis on Allosauridae noted that Charles Gilmore mistakenly reconstructed USNM 4734 as having a shorter skull than the specimens referred by Paul to ''atrox'', refuting supposed differences between USNM 4734 and putative ''A. atrox'' specimens like DINO 2560, AMNH 600, and AMNH 666. "Allosaurus agilis", seen in Zittel,
1887 Events January * January 11 – Louis Pasteur's anti-rabies treatment is defended in the Académie Nationale de Médecine, by Dr. Joseph Grancher. * January 20 ** The United States Senate allows the United States Navy to lease Pearl Har ...
, and Osborn,
1912 This year is notable for Sinking of the Titanic, the sinking of the ''Titanic'', which occurred on April 15. In Albania, this leap year runs with only 353 days as the country achieved switching from the Julian to Gregorian Calendar by skippin ...
, is a typographical error for ''A. fragilis.'' "Allosaurus ferox" is a typographical error by Marsh for ''A. fragilis'' in a figure caption for the partial skull YPM 1893 and YPM 1893 has been treated as a specimen of ''A fragilis''. Likewise, "Labrosaurus fragilis" is a typographical error by Marsh (
1896 Events January * January 2 – The Jameson Raid comes to an end as Jameson surrenders to the Boers. * January 4 – Utah is admitted as the 45th U.S. state. * January 5 – An Austrian newspaper reports Wilhelm Röntgen's dis ...
) for ''Labrosaurus ferox''. "A. whitei" is a ''
nomen nudum In Taxonomy (biology), taxonomy, a ''nomen nudum'' ('naked name'; plural ''nomina nuda'') is a designation which looks exactly like a scientific name of an organism, and may have originally been intended to be one, but it has not been published ...
'' coined by Pickering in 1996 for the complete ''Allosaurus'' specimens that Paul referred to ''A. atrox''. "Madsenius" was coined by David Lambert in
1990 Important events of 1990 include the Reunification of Germany and the unification of Yemen, the formal beginning of the Human Genome Project (finished in 2003), the launch of the Hubble Space Telescope, the separation of Namibia from South ...
, being based on remains from Dinosaur National Monument assigned to ''Allosaurus'' or ''Creosaurus'' (a synonym of ''Allosaurus''), and was to be described by paleontologist
Robert Bakker Robert Thomas Bakker (born March 24, 1945) is an American paleontologist who helped reshape modern theories about dinosaurs, particularly by adding support to the theory that some dinosaurs were endothermic (warm-blooded). Along with his mentor ...
as "Madsenius trux". However, "Madsenius" is now seen as yet another synonym of ''Allosaurus'' because Bakker's action was predicated upon the false assumption of USNM 4734 being distinct from long-snouted ''Allosaurus'' due to errors in Gilmore's
1920 Events January * January 1 ** Polish–Soviet War: The Russian Red Army increases its troops along the Polish border from 4 divisions to 20. ** Kauniainen in Finland, completely surrounded by the city of Espoo, secedes from Espoo as its ow ...
reconstruction of USNM 4734. "Wyomingraptor" was informally coined by Bakker for
allosaurid Allosauridae is an extinct family of medium to large bipedal, carnivorous allosauroid theropod dinosaurs from the Late Jurassic. Allosauridae is a fairly old taxonomic group, having been first named by the American paleontologist Othniel Charles ...
remains from the
Morrison Formation The Morrison Formation is a distinctive sequence of Upper Jurassic sedimentary rock found in the western United States which has been the most fertile source of dinosaur fossils in North America. It is composed of mudstone, sandstone, siltston ...
of the Late
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...
. The remains unearthed are labeled as ''Allosaurus'' and are housed in the Tate Geological Museum. However, there has been no official description of the remains and "Wyomingraptor" has been dismissed as a ''
nomen nudum In Taxonomy (biology), taxonomy, a ''nomen nudum'' ('naked name'; plural ''nomina nuda'') is a designation which looks exactly like a scientific name of an organism, and may have originally been intended to be one, but it has not been published ...
'', with the remains referable to ''Allosaurus''.


Formerly assigned species and fossils

Several species initially classified within or referred to ''Allosaurus'' do not belong within the genus. ''A. medius'' was named by Marsh in 1888 for various specimens from the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
Arundel Formation The Arundel Formation, also known as the Arundel Clay, is a clay-rich sedimentary geological formation, rock formation, within the Potomac Group, found in Maryland of the United States of America. It dates to the Early Cretaceous, and is of late ...
of
Maryland Maryland ( ) is a U.S. state, state in the Mid-Atlantic (United States), Mid-Atlantic region of the United States. It borders the states of Virginia to its south, West Virginia to its west, Pennsylvania to its north, and Delaware to its east ...
, although most of the remains were removed 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 for championing a non-Darwinian view of evolution, whereby mutation(s) could unlock p ...
to the new ornithopod species '' Dryosaurus grandis'', except for a tooth. It was transferred to ''Antrodemus'' by Oliver Hay in 1902, but Hay later clarified that this was an inexplicable error on his part. Gilmore considered the tooth nondiagnostic but transferred it to ''
Dryptosaurus ''Dryptosaurus'' ( ) is a genus of eutyrannosaurian theropod dinosaur that lived on the island continent of Appalachia approximately 67-66 million years ago during the end of the Maastrichtian age of the Late Cretaceous period. ''Dryptosaurus'' ...
'', as ''D. medius''. The referral was not accepted in the most recent review of basal tetanurans, and ''Allosaurus medius'' was simply listed as a dubious species of theropod. It may be closely related to ''
Acrocanthosaurus ''Acrocanthosaurus'' ( ; ) is a genus of carcharodontosaurid dinosaurs that existed in what is now North America during the Aptian and early Albian stages of the Early Cretaceous, from 113 to 110 million years ago. Like most dinosaur genera, ...
''. ''Allosaurus valens'' is a new combination for ''Antrodemus valens'' used by Friedrich von Huene in 1932; ''Antrodemus valens'' itself may also pertain to ''Allosaurus fragilis'', as Gilmore suggested in 1920. ''A. lucaris'', another Marsh name, was given to a partial skeleton in 1878. He later decided it warranted its own genus, ''Labrosaurus'', but this has not been accepted, and ''A. lucaris'' is also regarded as another specimen of ''A. fragilis''. ''Allosaurus lucaris'', is known mostly from vertebrae, sharing characters with ''Allosaurus''. Paul and Carpenter stated that the type specimen of this species, YPM 1931, was from a younger age than ''Allosaurus'', and might represent a different genus. However, they found that the specimen was undiagnostic, and thus ''A. lucaris'' was a ''nomen dubium''. ''Allosaurus sibiricus'' was described in 1914 by A. N. Riabinin on the basis of a bone, later identified as a partial fourth metatarsal, from the Early Cretaceous of
Buryatia Buryatia, officially the Republic of Buryatia, is a republic of Russia located in the Russian Far East. Formerly part of the Siberian Federal District, it has been administered as part of the Far Eastern Federal District since 2018. To its nort ...
, Russia. It was transferred to ''Chilantaisaurus'' in 1990, but is now considered a ''nomen dubium'' indeterminate beyond Theropoda. ''Allosaurus meriani'' was a new combination by George Olshevsky for ''
Megalosaurus ''Megalosaurus'' (meaning "great lizard", from Ancient Greek, Greek , ', meaning 'big', 'tall' or 'great' and , ', meaning 'lizard') is an extinct genus of large carnivorous theropod dinosaurs of the Middle Jurassic Epoch (Bathonian stage, 166 ...
meriani'' Greppin, 1870, based on a tooth from the Late Jurassic of Switzerland. However, a recent overview of ''Ceratosaurus'' included it in ''Ceratosaurus'' sp. '' Apatodon mirus'', based on a scrap of vertebra Marsh first thought to be a mammalian jaw, has been listed as a synonym of ''Allosaurus fragilis''. However, it was considered indeterminate beyond Dinosauria by Chure, and Mickey Mortimer believes that the synonymy of ''Apatodon'' with ''Allosaurus'' was due to correspondence to Ralph Molnar by John McIntosh, whereby the latter reportedly found a paper saying that Othniel Charles Marsh admitted that the ''Apatodon'' holotype was actually an allosaurid dorsal vertebra. ''A. amplexus'' was named by
Gregory S. Paul Gregory Scott Paul (born December 24, 1954) is an American freelance researcher, author and illustrator who works in paleontology. He is best known for his work and research on theropoda, theropod dinosaurs and his detailed illustrations, both l ...
for giant Morrison allosaur remains, and included in his conception ''Saurophagus maximus'' (later ''Saurophaganax''). ''A. amplexus'' was originally coined by Cope in 1878 as the type species of his new genus ''
Epanterias ''Epanterias'' is a dubious genus of theropod dinosaur from the Tithonian age Upper Jurassic upper Morrison Formation of Garden Park, Colorado. It was described by Edward Drinker Cope in 1878. The type species is ''Epanterias amplexus''. This ...
'', and is based on what is now AMNH 5767, parts of three vertebrae, a
coracoid A coracoid 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 present as part of the scapula, but this is n ...
, and a metatarsal. Following Paul's work, this species has been accepted as a synonym of ''A. fragilis''. A 2010 neotype designation by Greogry S. Paul and Kenneth Carpenter, however, suggested that ''Epanterias'' holotype is temporally younger than the ''A. fragilis'' type specimen, and that it is not the same taxon as the ''Allosaurus'' holotype. ''A. maximus'' was a new combination by David K. Smith for Chure's ''Saurophaganax maximus'', a taxon created by Chure in 1995 for giant allosaurid remains from the Morrison of Oklahoma. These remains had been known as ''Saurophagus'', but that name was already in use, leading Chure to propose a substitute. Smith, in his 1998 analysis of variation, concluded that ''S. maximus'' was not different enough from ''Allosaurus'' to be a separate genus, but did warrant its own species, ''A. maximus''. This reassignment was rejected in a review of basal tetanurans. A 2024 reassessment of fossil material assigned to ''Saurophaganax'' suggested that the holotype neural arch of this taxon could not confidently be assigned to a theropod, but that it exhibited some similarities to sauropods. Other ''Saurophaganax'' bones could be referred to diplodocid sauropods. As such, the researchers assigned the remaining theropod bones to a new species of ''Allosaurus'', ''A. anax''. There are also several species left over from the synonymizations of ''Creosaurus'' and ''Labrosaurus'' with ''Allosaurus''. '' Creosaurus potens'' was named by Lull in 1911 for a vertebra from the Early Cretaceous of Maryland. It is now regarded as a dubious theropod. ''Labrosaurus stechowi'', described in 1920 by Janensch based on isolated ''Ceratosaurus''-like teeth from the Tendaguru beds of Tanzania, was listed by Donald F. Glut as a species of ''Allosaurus'', is now considered a dubious ceratosaurian related to ''
Ceratosaurus ''Ceratosaurus'' (from Greek 'horn' and 'lizard') is a genus of carnivorous theropod dinosaur that lived in the Late Jurassic period (Kimmeridgian to Tithonian ages). The genus was first described in 1884 by American paleontologist Othni ...
''. ''L. sulcatus'', named by Marsh in 1896 for a Morrison theropod tooth, which like ''L. stechowi'' is now regarded as a dubious ''Ceratosaurus''-like ceratosaur. ''A. tendagurensis'' was named in 1925 by
Werner Janensch Werner Ernst Martin Janensch (11 November 1878 – 20 October 1969) was a German paleontologist and geologist. Biography Janensch was born at Herzberg (Elster). In addition to Friedrich von Huene, Janensch was probably Germany's most ...
for a partial shin (MB.R.3620) found in the Kimmeridgian-age
Tendaguru Formation The Tendaguru Formation, or Tendaguru Beds are a highly List of fossiliferous stratigraphic units in Tanzania, fossiliferous Lithostratigraphy, formation and Lagerstätte located in the Lindi Region of southeastern Tanzania. The formation represe ...
in
Mtwara Mtwara (Portuguese language, Portuguese: ''Montewara'') is the capital city of Mtwara Region in southeastern Tanzania. In the 1940s, it was planned and constructed as the export facility for the disastrous Tanganyika groundnut scheme, but was some ...
, Tanzania. Although tabulated as a tentatively valid species of ''Allosaurus'' in the second edition of the Dinosauria, subsequent studies place it as indeterminate beyond Tetanurae, either a carcharodontosaurian or megalosaurid. Although obscure, it was a large theropod, possibly around long and in weight. Kurzanov and colleagues in 2003 designated six teeth from Siberia as ''Allosaurus'' sp. (meaning the authors found the specimens to be most like those of ''Allosaurus'', but did not or could not assign a species to them). They were reclassified as an indeterminate theropod. Also, reports of ''Allosaurus'' in
Shanxi Shanxi; Chinese postal romanization, formerly romanised as Shansi is a Provinces of China, province in North China. Its capital and largest city of the province is Taiyuan, while its next most populated prefecture-level cities are Changzhi a ...
, China go back to at least 1982. These were interpreted as ''
Torvosaurus ''Torvosaurus'' () is a genus of large Megalosaurinae, megalosaurine Theropoda, theropod dinosaur that lived approximately 165 to 148 million years ago during the Callovian to Tithonian ages of the late Middle Jurassic, Middle and Late Jurassi ...
'' remains in 2012. An
astragalus Astragalus may refer to: * ''Astragalus'' (plant), a large genus of herbs and small shrubs *Astragalus (bone) The talus (; Latin for ankle or ankle bone; : tali), talus bone, astragalus (), or ankle bone is one of the group of foot bones known ...
(ankle bone) thought to belong to a species of ''Allosaurus'' was found at Cape Paterson, Victoria in Early Cretaceous beds in southeastern Australia. It was thought to provide evidence that Australia was a refugium for animals that had gone extinct elsewhere. This identification was challenged by Samuel Welles, who thought it more resembled that of an
ornithomimid Ornithomimidae (meaning "bird-mimics") is an extinct family of theropod dinosaurs which bore a superficial resemblance to modern ostriches. Ornithomimids were fast, omnivorous or herbivorous dinosaurs known mainly from the Late Cretaceous Period ...
, but the original authors defended their identification. With fifteen years of new specimens and research to look at, Daniel Chure reexamined the bone and found that it was not ''Allosaurus'', but could represent an allosauroid. Similarly, Yoichi Azuma and
Phil Currie Philip John Currie (born March 13, 1949) is a Canadian palaeontologist and museum curator who helped found the Royal Tyrrell Museum of Palaeontology in Drumheller, Alberta and is now a professor at the University of Alberta in Edmonton. In the ...
, in their description of ''
Fukuiraptor ''Fukuiraptor'' ("thief of Fukui") is a genus of medium-sized megaraptoran theropod dinosaur of the Early Cretaceous epoch (either Barremian or Aptian) that lived in what is now Japan. ''Fukuiraptor'' is known from the Kitadani Formation and ...
'', noted that the bone closely resembled that of their new genus. This specimen is sometimes referred to as " Allosaurus robustus", an informal museum name. It likely belonged to something similar to ''
Australovenator ''Australovenator'' (meaning "southern hunter") is a genus of megaraptoran theropod dinosaur from Cenomanian (Late Cretaceous)-age Winton Formation (dated to 95 million years ago) of Australia. Some specimens from the Albian-aged Eumeralla Forma ...
'', although one study considered it to belong to an
abelisaur Ceratosaurs are members of the clade Ceratosauria, a group of dinosaurs defined as all theropods sharing a more recent common ancestor with '' Ceratosaurus'' than with birds. The oldest known ceratosaur, '' Saltriovenator'', dates to the earlie ...
.


Description

''Allosaurus'' was a typical large
theropod Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
, having a massive skull on a short neck, a long, slightly sloping tail, and reduced forelimbs. ''Allosaurus fragilis'', the best-known species, had an average length of and mass of , with the largest definitive ''Allosaurus'' specimen (
AMNH The American Museum of Natural History (AMNH) is a natural history museum on the Upper West Side of Manhattan in New York City. Located in Theodore Roosevelt Park, across the street from Central Park, the museum complex comprises 21 interco ...
680) estimated at long, with an estimated weight of . In his 1976
monograph A monograph is generally a long-form work on one (usually scholarly) subject, or one aspect of a subject, typically created by a single author or artist (or, sometimes, by two or more authors). Traditionally it is in written form and published a ...
on ''Allosaurus'', James H. Madsen mentioned a range of bone sizes which he interpreted to show a maximum length of . As with dinosaurs in general, weight estimates are debatable, and since 1980 have ranged between , , and approximately for modal adult weight (not maximum). John Foster, a specialist on the Morrison Formation, suggests that is reasonable for large adults of ''A. fragilis'', but that is a closer estimate for individuals represented by the average-sized thigh bones he has measured. Using the subadult specimen nicknamed "Big Al", since assigned to the species ''Allosaurus jimmadseni'', researchers using computer modeling arrived at a best estimate of for the individual, but by varying parameters they found a range from approximately to approximately . A separate computational project estimated the adaptive optimum body mass in ''Allosaurus'' to be ~2,345 kg. ''A. europaeus'' has been measured up to in length and in body mass. Several gigantic specimens have been attributed to ''Allosaurus'', but may in fact belong to other genera. The dubious genus ''
Saurophaganax ''Saurophaganax'' ("lord of lizard-eaters") is a dubious, chimeric genus of large saurischian dinosaur, possibly a sauropod, from the Late Jurassic ( Kimmeridgian) Morrison Formation of Oklahoma, United States. This taxon was historically co ...
'' ( OMNH 1708) was estimated to reach around in length, and its single species was sometimes included in the genus ''Allosaurus'' as ''Allosaurus maximus''. However, a 2024 study concluded that some material assigned to ''Saurophaganax'' actually belonged to a
diplodocid Diplodocids, or members of the family Diplodocidae ("double beams"), are a group of sauropod dinosaurs. The family includes some of the longest creatures ever to walk the Earth, including '' Diplodocus'' and '' Supersaurus'', some of which may ha ...
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their b ...
with the material confidently assigned to
Allosauridae Allosauridae is an extinct family of medium to large bipedal, carnivorous allosauroid theropod dinosaurs from the Late Jurassic. Allosauridae is a fairly old taxonomic group, having been first named by the American paleontologist Othniel Cha ...
belonging to a new species of ''Allosaurus'', ''A. anax'', and the body mass of this species was tentatively estimated around based on fragmentary material. Another potential specimen of ''Allosaurus'', once assigned to the genus ''
Epanterias ''Epanterias'' is a dubious genus of theropod dinosaur from the Tithonian age Upper Jurassic upper Morrison Formation of Garden Park, Colorado. It was described by Edward Drinker Cope in 1878. The type species is ''Epanterias amplexus''. This ...
'' (AMNH 5767), may have measured in length. A more recent discovery is a partial skeleton from the Peterson Quarry in Morrison rocks of
New Mexico New Mexico is a state in the Southwestern United States, Southwestern region of the United States. It is one of the Mountain States of the southern Rocky Mountains, sharing the Four Corners region with Utah, Colorado, and Arizona. It also ...
; this large allosaurid was suggested to be a potential specimen of ''Saurophaganax'' prior to this taxon's 2024 reassessment. David K. Smith, examining ''Allosaurus'' fossils by quarry, found that the
Cleveland-Lloyd Dinosaur Quarry Jurassic National Monument, at the site of the Cleveland-Lloyd Dinosaur Quarry, well known for containing the densest concentration of Jurassic dinosaur fossils ever found, is a paleontological site located near Cleveland, Utah, in the San Rafa ...
(Utah) specimens are generally smaller than those from
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, ...
(Wyoming) or
Brigham Young University Brigham Young University (BYU) is a Private education, private research university in Provo, Utah, United States. It was founded in 1875 by religious leader Brigham Young and is the flagship university of the Church Educational System sponsore ...
's Dry Mesa Quarry (Colorado), but the shapes of the bones themselves did not vary between the sites. A later study by Smith incorporating Garden Park (Colorado) and
Dinosaur National Monument Dinosaur National Monument is an American national monument located on the southeast flank of the Uinta Mountains on the border between Colorado and Utah at the confluence of the Green River (Colorado River tributary), Green and Yampa River, Y ...
(Utah) specimens found no justification for multiple species based on skeletal variation; skull variation was most common and was gradational, suggesting individual variation was responsible. Further work on size-related variation again found no consistent differences, although the Dry Mesa material tended to clump together on the basis of the
astragalus Astragalus may refer to: * ''Astragalus'' (plant), a large genus of herbs and small shrubs *Astragalus (bone) The talus (; Latin for ankle or ankle bone; : tali), talus bone, astragalus (), or ankle bone is one of the group of foot bones known ...
, an ankle bone.
Kenneth Carpenter Kenneth Carpenter (born 21 September 1949) is an American paleontologist. He is the former director of the USU Eastern Prehistoric Museum and author or co-author of books on dinosaurs and Mesozoic life. His main research interests are armore ...
, using skull elements from the Cleveland-Lloyd site, found wide variation between individuals, calling into question previous species-level distinctions based on such features as the shape of the horns, and the proposed differentiation of ''A. jimmadseni'' based on the shape of the
jugal The jugal is a skull bone found in most reptiles, amphibians and birds. In mammals, the jugal is often called the malar or zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by species. Anatomy ...
. A study published by Motani et al., in 2020 suggests that ''Allosaurus'' was also sexually dimorphic in the width of the femur's head against its length.


Skull

The skull and teeth of ''Allosaurus'' were modestly proportioned for a theropod of its size. Paleontologist
Gregory S. Paul Gregory Scott Paul (born December 24, 1954) is an American freelance researcher, author and illustrator who works in paleontology. He is best known for his work and research on theropoda, theropod dinosaurs and his detailed illustrations, both l ...
gives a length of for a skull belonging to an individual he estimates at long. Each
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammals h ...
(the bones that formed the tip of the snout) held five teeth with D-shaped cross-sections, and each
maxilla In vertebrates, the maxilla (: maxillae ) is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The two maxil ...
(the main tooth-bearing bones in the upper jaw) had between 14 and 17 teeth; the number of teeth does not exactly correspond to the size of the bone. Each
dentary In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone ...
(the tooth-bearing bone of the lower jaw) had between 14 and 17 teeth, with an average count of 16. The teeth became shorter, narrower, and more curved toward the back of the skull. All of the teeth had saw-like edges. They were shed easily, and were replaced continually, making them common fossils. Its skull was light, robust and equipped with dozens of sharp,
serrated Serration is a saw-like appearance or a row of sharp or tooth-like projections. A serrated cutting edge has many small points of contact with the material being cut. By having less contact area than a smooth blade or other edge, the applied pr ...
teeth. The skull had a pair of
horn Horn may refer to: Common uses * Horn (acoustic), a tapered sound guide ** Horn antenna ** Horn loudspeaker ** Vehicle horn ** Train horn *Horn (anatomy), a pointed, bony projection on the head of various animals * Horn (instrument), a family ...
s above and in front of the eyes. These horns were composed of extensions of the
lacrimal bone The lacrimal bones are two small and fragile bones of the facial skeleton; they are roughly the size of the little fingernail and situated at the front part of the medial wall of the orbit. They each have two surfaces and four borders. Several bon ...
s, and varied in shape and size. There were also lower paired ridges running along the top edges of the
nasal bone The nasal bones are two small oblong bones, varying in size and form in different individuals; they are placed side by side at the middle and upper part of the face and by their junction, form the bridge of the upper one third of the nose. Eac ...
s that led into the horns. The horns were probably covered in a
keratin Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. It is the key structural material making up Scale (anatomy), scales, hair, Nail (anatomy), nails, feathers, horn (anatomy), horns, claws, Hoof, hoove ...
sheath and may have had a variety of functions, including acting as sunshades for the eyes, being used for display, and being used in combat against other members of the same species (although they were fragile). There was a ridge along the back of the skull roof for muscle attachment, as is also seen in
tyrannosaurid Tyrannosauridae (or tyrannosaurids, meaning "tyrant lizards") is a family of coelurosaurian theropod dinosaurs that comprises two subfamilies containing up to fifteen genera, including the eponymous ''Tyrannosaurus''. The exact number of genera ...
s. Inside the lacrimal bones were depressions that may have held
gland A gland is a Cell (biology), cell or an Organ (biology), organ in an animal's body that produces and secretes different substances that the organism needs, either into the bloodstream or into a body cavity or outer surface. A gland may also funct ...
s, such as
salt gland The salt gland is an organ (anatomy), organ for excreting excess salt (chemistry), salts. It is found in the cartilaginous fishes subclass elasmobranchii (sharks, rays, and skates), seabirds, and some reptiles. Salt glands can be found in the r ...
s. Within the maxillae were sinuses that were better developed than those of more basal theropods such as ''
Ceratosaurus ''Ceratosaurus'' (from Greek 'horn' and 'lizard') is a genus of carnivorous theropod dinosaur that lived in the Late Jurassic period (Kimmeridgian to Tithonian ages). The genus was first described in 1884 by American paleontologist Othni ...
'' and ''
Marshosaurus ''Marshosaurus'' is a genus of medium-sized carnivorous Theropoda, theropod dinosaur, belonging to the family Piatnitzkysauridae, from the Late Jurassic Morrison Formation of Utah and possibly Colorado. Discovery and naming During the 1960s, o ...
''; they may have been related to the
sense of smell The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste. In humans, it ...
, perhaps holding something like
Jacobson's organ The vomeronasal organ (VNO), or Jacobson's organ, is the paired auxiliary olfactory (smell) sense organ located in the soft tissue of the nasal septum, in the nasal cavity just above the roof of the mouth (the hard palate) in various tetrapods. T ...
s. The roof of the braincase was thin, perhaps to improve
thermoregulation 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 ...
for the brain. The skull and lower jaws had joints that permitted motion within these units. In the lower jaws, the bones of the front and back halves loosely articulated, permitting the jaws to bow outward and increasing the animal's gape. The
braincase In human anatomy, the neurocranium, also known as the braincase, brainpan, brain-pan, or brainbox, is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the calv ...
and frontals may also have had a joint.


Postcranial skeleton

''Allosaurus'' had nine
vertebra Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spina ...
e in the neck, 14 in the back, and five in the
sacrum The sacrum (: sacra or sacrums), in human anatomy, is a triangular bone at the base of the spine that forms by the fusing of the sacral vertebrae (S1S5) between ages 18 and 30. The sacrum situates at the upper, back part of the pelvic cavity, ...
supporting the hips. The number of tail vertebrae is unknown and varied with individual size; James Madsen estimated about 50, while
Gregory S. Paul Gregory Scott Paul (born December 24, 1954) is an American freelance researcher, author and illustrator who works in paleontology. He is best known for his work and research on theropoda, theropod dinosaurs and his detailed illustrations, both l ...
considered that to be too many and suggested 45 or less. There were hollow spaces in the neck and
anterior Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...
back vertebrae. Such spaces, which are also found in modern theropods (that is, the birds), are interpreted as having held
air sacs Air sacs are spaces within an organism where there is the constant presence of air. Among modern animals, birds possess the most air sacs (9–11), with their extinct dinosaurian relatives showing a great increase in the pneumatization (presence ...
used in
respiration Respiration may refer to: Biology * Cellular respiration, the process in which nutrients are converted into useful energy in a cell ** Anaerobic respiration, cellular respiration without oxygen ** Maintenance respiration, the amount of cellul ...
. The rib cage was broad, giving it a barrel chest, especially in comparison to less derived theropods like ''Ceratosaurus''. ''Allosaurus'' had
gastralia Gastralia (: gastralium) are dermal bones found in the ventral body wall of modern crocodilians and tuatara, and many prehistoric tetrapods. They are found between the sternum and pelvis, and do not articulate with the vertebrae. In these reptil ...
(belly ribs), but these are not common findings, and they may have
ossified Ossification (also called osteogenesis or bone mineralization) in bone remodeling is the process of laying down new bone material by cells named osteoblasts. It is synonymous with bone tissue formation. There are two processes resulting in t ...
poorly. In one published case, the gastralia show evidence of injury during life. A
furcula The (Latin for "little fork"; : furculae) or wishbone is a forked bone found in most birds and some species of non-avian dinosaurs, and is either an interclavicle or formed by the fusion of the two clavicles. In birds, its primary function is ...
(wishbone) was also present, but has only been recognized since 1996; in some cases furculae were confused with gastralia. The ilium, the main hip bone, was massive, and the
pubic bone In vertebrates, the pubis or pubic bone () forms the lower and anterior part of each side of the hip bone. The pubis is the most forward-facing (ventral and anterior) of the three bones that make up the hip bone. The left and right pubic bones ar ...
had a prominent foot that may have been used for both muscle attachment and as a prop for resting the body on the ground. Madsen noted that in about half of the individuals from the
Cleveland-Lloyd Dinosaur Quarry Jurassic National Monument, at the site of the Cleveland-Lloyd Dinosaur Quarry, well known for containing the densest concentration of Jurassic dinosaur fossils ever found, is a paleontological site located near Cleveland, Utah, in the San Rafa ...
, independent of size, the pubes had not fused to each other at their foot ends. He suggested that this was a sexual characteristic, with females lacking fused bones to make egg-laying easier. This proposal has not attracted further attention, however. The forelimbs of ''Allosaurus'' were short in comparison to the hindlimbs (only about 35% the length of the hindlimbs in adults) and had three fingers per hand, tipped with large, strongly curved and pointed
claw A claw is a curved, pointed appendage found at the end of a toe or finger in most amniotes (mammals, reptiles, birds). Some invertebrates such as beetles and spiders have somewhat similar fine, hooked structures at the end of the leg or Arthro ...
s. The arms were powerful, and the forearm was somewhat shorter than the upper arm (1:1.2
ulna The ulna or ulnar bone (: ulnae or ulnas) is a long bone in the forearm stretching from the elbow to the wrist. It is on the same side of the forearm as the little finger, running parallel to the Radius (bone), radius, the forearm's other long ...
/
humerus The humerus (; : humeri) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...
ratio). The wrist had a version of the semilunate
carpal The carpal bones are the eight small bones that make up the wrist (carpus) that connects the hand to the forearm. The terms "carpus" and "carpal" are derived from the Latin carpus and the Greek καρπός (karpós), meaning "wrist". In huma ...
also found in more derived theropods like
maniraptora Maniraptora is a clade of coelurosaurian dinosaurs which includes the birds and the non-avian dinosaurs that were more closely related to them than to ''Ornithomimus velox''. It contains the major subgroups Avialae, Dromaeosauridae, Troodontidae, ...
ns. Of the three fingers, the innermost (or thumb) was the largest, and diverged from the others. The phalangeal formula is 2-3-4-0-0, meaning that the innermost finger (phalange) has two bones, the next has three, and the third finger has four. The legs were not as long or suited for speed as those of
tyrannosaurid Tyrannosauridae (or tyrannosaurids, meaning "tyrant lizards") is a family of coelurosaurian theropod dinosaurs that comprises two subfamilies containing up to fifteen genera, including the eponymous ''Tyrannosaurus''. The exact number of genera ...
s, and the claws of the toes were less developed and more
hoof The hoof (: hooves) is the tip of a toe of an ungulate mammal, which is covered and strengthened with a thick and horny keratin covering. Artiodactyls are even-toed ungulates, species whose feet have an even number of digits; the ruminants with ...
-like than those of earlier theropods. Each foot had three weight-bearing toes and an inner
dewclaw A dewclaw is a digit – vestigial in some animals – on the foot of many mammals, birds, and reptiles (including some extinct orders, like certain theropods). It commonly grows higher on the leg than the rest of the foot, such that in di ...
, which Madsen suggested could have been used for grasping in juveniles. There was also what is interpreted as the splint-like remnant of a fifth (outermost)
metatarsal The metatarsal bones or metatarsus (: metatarsi) are a group of five long bones in the midfoot, located between the tarsal bones (which form the heel and the ankle) and the phalanges ( toes). Lacking individual names, the metatarsal bones are ...
, perhaps used as a lever between the
Achilles tendon The Achilles tendon or heel cord, also known as the calcaneal tendon, is a tendon at the back of the lower leg, and is the thickest in the human body. It serves to attach the plantaris, gastrocnemius (calf) and soleus muscles to the calcane ...
and foot.


Skin

Skin impressions from ''Allosaurus'' have been described. One impression, from a juvenile specimen, measures 30 cm² and is associated with the anterior dorsal ribs/pectoral region. The impression shows small
scales Scale or scales may refer to: Mathematics * Scale (descriptive set theory), an object defined on a set of points * Scale (ratio), the ratio of a linear dimension of a model to the corresponding dimension of the original * Scale factor, a number ...
measuring 1–3 mm in diameter. A skin impression from the "Big Al Two" specimen, associated with the base of the tail, measures 20 cm × 20 cm and shows large scales measuring up to 2 cm in diameter. However, it has been noted that these scales are more similar to those of
sauropods Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their b ...
, and due to the presence of non-theropod remains associated with the tail of "Big Al Two" there is a possibility that this skin impression is not from ''Allosaurus''. Another ''Allosaurus'' fossil features a skin impression from the
mandible In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
, showing scales measuring 1–2 mm in diameter. The same fossil also preserves skin measuring 20 × 20 cm from the ventral side of the neck, showing scutate scales measuring 0.5 cm wide and 11 cm long. A small skin impression from an ''Allosaurus'' skull has been reported but never described.


Classification

''Allosaurus'' was an allosaurid, a member of a
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
of large theropods within the larger group
Carnosauria Carnosauria is an extinct group of carnivorous theropod dinosaurs that lived during the Jurassic and Cretaceous periods. While Carnosauria was historically considered largely synonymous with Allosauroidea, some recent studies have revived Ca ...
. The family name
Allosauridae Allosauridae is an extinct family of medium to large bipedal, carnivorous allosauroid theropod dinosaurs from the Late Jurassic. Allosauridae is a fairly old taxonomic group, having been first named by the American paleontologist Othniel Cha ...
was created for this genus in 1878 by
Othniel Charles Marsh Othniel Charles Marsh (October 29, 1831 – March 18, 1899) was an American professor of paleontology. A prolific fossil collector, Marsh was one of the preeminent paleontologists of the nineteenth century. Among his legacies are the discovery or ...
, but the term was largely unused until the 1970s in favor of
Megalosauridae Megalosauridae is a monophyletic Family (taxonomy), family of Carnivore, carnivorous theropod dinosaurs within the group Megalosauroidea. Appearing in the Middle Jurassic, megalosaurids were among the first major radiation of large theropod dino ...
, another family of large theropods that eventually became a
wastebasket taxon Wastebasket taxon (also called a wastebin taxon, dustbin taxon or catch-all taxon) is a term used by some taxonomists to refer to a taxon that has the purpose of classifying organisms that do not fit anywhere else. They are typically defined by e ...
. This, along with the use of ''Antrodemus'' for ''Allosaurus'' during the same period, is a point that needs to be remembered when searching for information on ''Allosaurus'' in publications that predate James Madsen's 1976 monograph. Major publications using the name "Megalosauridae" instead of "Allosauridae" include Gilmore, 1920, von Huene, 1926,
Romer Romer, Römer, Roemer, or similar may refer to: * Romer (surname), includes a list of people with the name * Romer (tool), a cartographic device also known as a reference card * Rømer scale, a disused temperature scale named after Ole Rømer * ...
, 1956 and 1966, Steel, 1970, and
Walker Walker or The Walker may refer to: People *Walker (given name) *Walker (surname) *Walker (Brazilian footballer) (born 1982), Brazilian footballer Places In the United States *Walker, Arizona, in Yavapai County *Walker, Mono County, California * ...
, 1964. Following the publication of Madsen's influential monograph, Allosauridae became the preferred family assignment, but it too was not strongly defined. Semi-technical works used Allosauridae for a variety of large theropods, usually those that were larger and better-known than megalosaurids. Typical theropods that were thought to be related to ''Allosaurus'' included '' Indosaurus'', '' Piatnitzkysaurus'', '' Piveteausaurus'', '' Yangchuanosaurus'', ''
Acrocanthosaurus ''Acrocanthosaurus'' ( ; ) is a genus of carcharodontosaurid dinosaurs that existed in what is now North America during the Aptian and early Albian stages of the Early Cretaceous, from 113 to 110 million years ago. Like most dinosaur genera, ...
'', ''
Chilantaisaurus ''Chilantaisaurus'' (" lizard") is a genus of large theropod dinosaur, possibly a neovenatorid or a primitive coelurosaur, from the Late Cretaceous Ulansuhai Formation of China (Turonian age, about 92 million years ago). The type species, ''C. ta ...
'', '' Compsosuchus'', ''
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 ...
'', and '' Szechuanosaurus''. Given modern knowledge of theropod diversity and the advent of cladistic study of
evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
ary relationships, none of these theropods is now recognized as an allosaurid, although several, like ''Acrocanthosaurus'' and ''Yangchuanosaurus'', are members of closely related families. Below is a cladogram based on the analysis of Benson et al. in 2010. Allosauridae is one of four families in Allosauroidea; the other three are
Neovenatoridae Carnosauria is an extinct group of carnivorous theropod dinosaurs that lived during the Jurassic and Cretaceous periods. While Carnosauria was historically considered largely synonymous with Allosauroidea, some recent studies have revived Carn ...
,
Carcharodontosauridae Carcharodontosauridae (carcharodontosaurids; from the Greek καρχαροδοντόσαυρος, ''carcharodontósauros'': "shark-toothed lizards") is a group of carnivorous theropod dinosaurs. In 1931, Ernst Stromer named Carcharodontosaurida ...
and Sinraptoridae. Allosauridae has at times been proposed as ancestral to the
Tyrannosauridae Tyrannosauridae (or tyrannosaurids, meaning "tyrant lizards") is a family of coelurosaurian theropod dinosaurs that comprises two subfamilies containing up to fifteen genera, including the eponymous ''Tyrannosaurus''. The exact number of genera ...
(which would make it
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
), one example being Gregory S. Paul's ''Predatory Dinosaurs of the World'', but this has been rejected, with tyrannosaurids identified as members of a separate branch of theropods, the
Coelurosauria Coelurosauria (; from Greek, meaning "hollow-tailed lizards") is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs. Coelurosauria is a subgroup of theropod dinosaurs that includes compsognathids, tyra ...
. Allosauridae is the smallest of the carnosaur families, with only ''Saurophaganax'' and a currently unnamed French
allosauroid Carnosauria is an extinct group of carnivorous theropod dinosaurs that lived during the Jurassic and Cretaceous periods. While Carnosauria was historically considered largely synonymous with Allosauroidea, some recent studies have revived Carn ...
accepted as possible valid
genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
besides ''Allosaurus'' in the most recent review. Another genus, ''Epanterias'', is a potential valid member, but it and ''Saurophaganax'' may turn out to be large examples of ''Allosaurus''. Some reviews have kept the genus ''
Saurophaganax ''Saurophaganax'' ("lord of lizard-eaters") is a dubious, chimeric genus of large saurischian dinosaur, possibly a sauropod, from the Late Jurassic ( Kimmeridgian) Morrison Formation of Oklahoma, United States. This taxon was historically co ...
'' and included ''Epanterias'' with ''Allosaurus''. The controversial ''Saurophaganax'', initially recognized as a large ''Allosaurus''-like theropod, has had a controversial taxonomic history. In 2019, Rauhut and Pol noted that its taxonomic placement within
Allosauroidea Carnosauria is an extinct group of carnivorous theropod dinosaurs that lived during the Jurassic and Cretaceous periods. While Carnosauria was historically considered largely synonymous with Allosauroidea, some recent studies have revived Carn ...
is unstable, being recovered as a sister taxon of
Metriacanthosauridae Metriacanthosauridae (Greek for "moderately-spined lizards") is an extinct family of allosauroid theropod dinosaurs that lived in Europe and Asia from the Middle Jurassic to the Early Cretaceous. The family is split into two subgroups: Metriacan ...
or Allosauria, or even as the basalmost carcharodontosaurian. In 2024, ''Saurophaganax'' was reassessed as a dubious, chimeric taxon with the holotype being so fragmentary that it could only be confidently referred to the
Saurischia Saurischia ( , meaning "reptile-hipped" from the Greek ' () meaning 'lizard' and ' () meaning 'hip joint') is one of the two basic divisions of dinosaurs (the other being Ornithischia), classified by their hip structure. Saurischia and Ornithi ...
, and some specimens more likely belonging to a
diplodocid Diplodocids, or members of the family Diplodocidae ("double beams"), are a group of sauropod dinosaurs. The family includes some of the longest creatures ever to walk the Earth, including '' Diplodocus'' and '' Supersaurus'', some of which may ha ...
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their b ...
.


Paleobiology


Life history

The wealth of ''Allosaurus'' fossils, from nearly all ages of individuals, allows scientists to study how the animal grew and how long its lifespan may have been. Remains may reach as far back in the lifespan as
egg An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the ...
s—crushed eggs from Colorado have been suggested as those of ''Allosaurus''. Based on
histological Histology, also known as microscopic anatomy or microanatomy, is the branch of biology that studies the microscopic anatomy of biological tissue (biology), tissues. Histology is the microscopic counterpart to gross anatomy, which looks at large ...
analysis of limb bones, bone deposition appears to stop at around 22 to 28 years, which is comparable to that of other large theropods like ''
Tyrannosaurus ''Tyrannosaurus'' () is a genus of large theropod dinosaur. The type species ''Tyrannosaurus rex'' ( meaning 'king' in Latin), often shortened to ''T. rex'' or colloquially t-rex, is one of the best represented theropods. It lived througho ...
''. From the same analysis, its maximum growth appears to have been at age 15, with an estimated growth rate of about 150 kilograms (330 lb) per year. Medullary bone tissue (endosteally derived, ephemeral, mineralization located inside the medulla of the long bones in gravid female birds) has been reported in at least one ''Allosaurus'' specimen, a shin bone from the Cleveland-Lloyd Quarry. Today, this bone tissue is only formed in female birds that are laying eggs, as it is used to supply
calcium Calcium is a chemical element; it has symbol Ca and atomic number 20. As an alkaline earth metal, calcium is a reactive metal that forms a dark oxide-nitride layer when exposed to air. Its physical and chemical properties are most similar to it ...
to shells. Its presence in the ''Allosaurus'' individual has been used to establish sex and show it had reached reproductive age. However, other studies have called into question some cases of medullary bone in dinosaurs, including this ''Allosaurus'' individual. Data from extant birds suggested that the medullary bone in this ''Allosaurus'' individual may have been the result of a bone pathology instead. However, with the confirmation of medullary tissue indicating sex in a specimen of ''Tyrannosaurus'', it may be possible to ascertain whether or not the ''Allosaurus'' in question was indeed female. The discovery of a juvenile specimen with a nearly complete hindlimb shows that the legs were relatively longer in juveniles, and the lower segments of the leg (shin and foot) were relatively longer than the thigh. These differences suggest that younger ''Allosaurus'' were faster and had different hunting strategies than adults, perhaps chasing small prey as juveniles, then becoming ambush hunters of large prey upon adulthood. The
thigh bone The femur (; : femurs or femora ), or thigh bone is the only bone in the thigh — the region of the lower limb between the hip and the knee. In many four-legged animals the femur is the upper bone of the hindleg. The top of the femur fits in ...
became thicker and wider during growth, and the cross-section less circular, as muscle attachments shifted, muscles became shorter, and the growth of the leg slowed. These changes imply that juvenile legs has less predictable stresses compared with adults, which would have moved with more regular forward progression. Conversely, the skull bones appear to have generally grown isometrically, increasing in size without changing in proportion.


Feeding

Most paleontologists accept ''Allosaurus'' as an active predator of large animals. There is dramatic evidence for allosaur attacks on ''Stegosaurus'', including an ''Allosaurus'' tail vertebra with a partially healed puncture wound that fits a ''Stegosaurus''
tail spike A thagomizer () is the distinctive arrangement of spike-shaped Osteoderm, osteoderms on the tails of some Stegosauria, stegosaurian dinosaurs. These spikes are believed to have been a defensive measure against predators. The arrangement of spike ...
, and a ''Stegosaurus'' neck plate with a U-shaped wound that correlates well with an ''Allosaurus'' snout.
Sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their b ...
s seem to be likely candidates as both live prey and as objects of
scavenging Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding be ...
, based on the presence of scrapings on sauropod bones fitting allosaur teeth well and the presence of shed allosaur teeth with sauropod bones. However, as Gregory Paul noted in 1988, ''Allosaurus'' was probably not a predator of fully grown sauropods, unless it hunted in packs, as it had a modestly sized skull and relatively small teeth, and was greatly outweighed by contemporaneous sauropods. Another possibility is that it preferred to hunt juveniles instead of fully grown adults. Research in the 1990s and the first decade of the 21st century may have found other solutions to this question.
Robert T. Bakker Robert Thomas Bakker (born March 24, 1945) is an American paleontologist who helped reshape modern theories about dinosaurs, particularly by adding support to the theory that some dinosaurs were endothermic (warm-blooded). Along with his mentor ...
, comparing ''Allosaurus'' to
Cenozoic The Cenozoic Era ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterized by the dominance of mammals, insects, birds and angiosperms (flowering plants). It is the latest of three g ...
saber-toothed carnivorous mammals, found similar adaptations, such as a reduction of jaw muscles and increase in neck muscles, and the ability to open the jaws extremely wide. Although ''Allosaurus'' did not have saber teeth, Bakker suggested another mode of attack that would have used such neck and jaw adaptations: the short teeth in effect became small serrations on a
saw A saw is a tool consisting of a tough blade, Wire saw, wire, or Chainsaw, chain with a hard toothed edge used to cut through material. Various terms are used to describe toothed and abrasive saws. Saws began as serrated materials, and when man ...
-like cutting edge running the length of the upper jaw, which would have been driven into prey. This type of jaw would permit slashing attacks against much larger prey, with the goal of weakening the victim. Similar conclusions were drawn by another study using
finite element analysis Finite element method (FEM) is a popular method for numerically solving differential equations arising in engineering and mathematical models, mathematical modeling. Typical problem areas of interest include the traditional fields of structural ...
on an ''Allosaurus'' skull. According to their biomechanical analysis, the skull was very strong but had a relatively small bite force. By using jaw muscles only, it could produce a bite force of 805 to 8,724 N, but the skull could withstand nearly 55,500 N of vertical force against the tooth row. The authors suggested that ''Allosaurus'' used its skull like a machete against prey, attacking open-mouthed, slashing flesh with its teeth, and tearing it away without splintering bones, unlike ''Tyrannosaurus'', which is thought to have been capable of damaging bones. They also suggested that the architecture of the skull could have permitted the use of different strategies against different prey; the skull was light enough to allow attacks on smaller and more agile ornithopods, but strong enough for high-impact ambush attacks against larger prey like stegosaurids and sauropods. Their interpretations were challenged by other researchers, who found no modern analogs to a hatchet attack and considered it more likely that the skull was strong to compensate for its open construction when absorbing the stresses from struggling prey. The original authors noted that ''Allosaurus'' itself has no modern equivalent, that the tooth row is well-suited to such an attack, and that articulations in the skull cited by their detractors as problematic actually helped protect the
palate The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly sep ...
and lessen stress. Another possibility for handling large prey is that theropods like ''Allosaurus'' were "flesh grazers" which could take bites of flesh out of living sauropods that were sufficient to sustain the predator so it would not have needed to expend the effort to kill the prey outright. This strategy would also potentially have allowed the prey to recover and be fed upon in a similar way later. An additional suggestion notes that ornithopods were the most common available dinosaurian prey, and that ''Allosaurus'' may have subdued them by using an attack similar to that of modern big cats: grasping the prey with their forelimbs, and then making multiple bites on the throat to crush the trachea. This is compatible with other evidence that the forelimbs were strong and capable of restraining prey. Studies done by Stephen Lautenschager et al. from the University of Bristol also indicate ''Allosaurus'' could open its jaws quite wide and sustain considerable muscle force. When compared with ''Tyrannosaurus'' and the therizinosaurid '' Erlikosaurus'' in the same study, it was found that ''Allosaurus'' had a wider gape than either; the animal was capable of opening its jaws to a 92-degree angle at maximum. The findings also indicate that large carnivorous dinosaurs, like modern carnivores, had wider jaw gapes than herbivores. A biomechanical study published in 2013 by Eric Snively and colleagues found that ''Allosaurus'' had an unusually low attachment point on the skull for the longissimus capitis superficialis neck muscle compared to other theropods such as ''
Tyrannosaurus ''Tyrannosaurus'' () is a genus of large theropod dinosaur. The type species ''Tyrannosaurus rex'' ( meaning 'king' in Latin), often shortened to ''T. rex'' or colloquially t-rex, is one of the best represented theropods. It lived througho ...
''. This would have allowed the animal to make rapid and forceful vertical movements with the skull. The authors found that vertical strikes as proposed by Bakker and Rayfield are consistent with the animal's capabilities. They also found that the animal probably processed carcasses by vertical movements in a similar manner to
falcon Falcons () are birds of prey in the genus ''Falco'', which includes about 40 species. Some small species of falcons with long, narrow wings are called hobbies, and some that hover while hunting are called kestrels. Falcons are widely distrib ...
s, such as
kestrel The term kestrel (from , derivative from , i.e. ratchet) is the common name given to several species of predatory birds from the falcon genus ''Falco''. Kestrels are most easily distinguished by their typical hunting behaviour which is to hover ...
s: The animal could have gripped prey with the skull and feet, then pulled back and up to remove flesh. This differs from the prey-handling envisioned for tyrannosaurids, which probably tore flesh with lateral shakes of the skull, similar to crocodilians. In addition, ''Allosaurus'' was able to "move its head and neck around relatively rapidly and with considerable control", at the cost of power. Other aspects of feeding include the eyes, arms, and legs. The shape of the skull of ''Allosaurus'' limited potential
binocular vision Binocular vision is seeing with two eyes. The Field_of_view, field of view that can be surveyed with two eyes is greater than with one eye. To the extent that the visual fields of the two eyes overlap, #Depth, binocular depth can be perceived. Th ...
to 20° of width, slightly less than that of modern
crocodilia Crocodilia () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchia ...
ns. As with crocodilians, this may have been enough to judge prey distance and time attacks. The arms, compared with those of other theropods, were suited for both grasping prey at a distance or clutching it close, and the articulation of the claws suggests that they could have been used to hook things. Finally, the top speed of ''Allosaurus'' has been estimated at per hour. A paper on the cranio-dental morphology of ''Allosaurus'' and how it worked has deemed the hatchet jaw attack unlikely, reinterpreting the unusually wide gape as an adaptation to allow ''Allosaurus'' to deliver a muscle-driven bite to large prey, with the weaker jaw muscles being a trade-off to allow for the widened gape. Sauropod carrion may also have been important to large theropods in the Morrison Formation. Forensic techniques indicate that sauropod carcasses were targeted by ''Allosaurus'' at all stages of decomposition, indicating that late-stage decay pathogens were not a significant deterrent. A survey of sauropod bones from the Morrison Formation also reported widespread bite marks on sauropod bones in low-economy regions, which suggests that large theropods scavenged large sauropods when available, with the scarcity of such bite marks on the remains of smaller bones being potentially attributable to much more complete consumption of smaller or adolescent sauropods and on ornithischians, which would have been more commonly taken as live prey. A single dead adult ''Barosaurus'' or ''Brachiosaurus'' would have had enough calories to sustain multiple large theropods for weeks or months, though the vast majority of the Morrison's sauropod fossil record consisted of much smaller-bodied taxa such as ''Camarasaurus lentus'' or ''Diplodocus''. It has also been argued that disabled individuals such as Big Al and Big Al II were physically incapable of hunting due to their numerous injuries but were able to survive nonetheless as scavengers of giant sauropod-falls, Interestingly, a recent review of paleopathologies in theropods may support this conclusion. The researchers found a positive association between allosaurids and fractures to the appendicular skeleton, while tyrannosaurs had a statistically negative association with these types of injuries. The fact that allosaurs were more likely to survive and heal even when severe fractures limited their locomotion abilities can be explained, in part, by different resource accessibility paradigms for the two groups, as allosauroids generally lived in sauropod-inhabited ecosystems, some of which, including the Morrison, have been interpreted as arid and highly water-stressed environments; however, the water-stressed nature of the Morrison has been heavily criticized in several more recent works on the basis of fossil evidence for the presence of extensive forest cover and aquatic ecosystems.


Social behavior

It has been speculated since the 1970s that ''Allosaurus'' preyed on sauropods and other large dinosaurs by hunting in groups. Such a depiction is common in semitechnical and popular dinosaur literature.
Robert T. Bakker Robert Thomas Bakker (born March 24, 1945) is an American paleontologist who helped reshape modern theories about dinosaurs, particularly by adding support to the theory that some dinosaurs were endothermic (warm-blooded). Along with his mentor ...
has extended social behavior to parental care, and has interpreted shed allosaur teeth and chewed bones of large prey animals as evidence that adult allosaurs brought food to lairs for their young to eat until they were grown, and prevented other carnivores from scavenging on the food. However, there is actually little evidence of gregarious behavior in theropods, and social interactions with members of the same species would have included antagonistic encounters, as shown by injuries to gastralia and bite wounds to skulls (the pathologic lower jaw named ''Labrosaurus ferox'' is one such possible example). Such head-biting may have been a way to establish dominance in a pack or to settle territorial disputes. Although ''Allosaurus'' may have hunted in packs, it has been argued that ''Allosaurus'' and other theropods had largely aggressive interactions instead of cooperative interactions with other members of their own species. The study in question noted that cooperative hunting of prey much larger than an individual predator, as is commonly inferred for theropod dinosaurs, is rare among vertebrates in general, and modern
diapsid Diapsids ("two arches") are a clade of sauropsids, distinguished from more primitive eureptiles by the presence of two holes, known as temporal fenestrae, in each side of their skulls. The earliest traditionally identified diapsids, the araeosc ...
carnivores (including lizards, crocodiles, and birds) rarely cooperate to hunt in such a way. Instead, they are typically territorial and will kill and cannibalize intruders of the same species, and will also do the same to smaller individuals that attempt to eat before they do when aggregated at feeding sites. According to this interpretation, the accumulation of remains of multiple ''Allosaurus'' individuals at the same site; e.g., in the Cleveland–Lloyd Quarry, are not due to pack hunting, but to the fact that ''Allosaurus'' individuals were drawn together to feed on other disabled or dead allosaurs, and were sometimes killed in the process. This could explain the high proportion of juvenile and subadult allosaurs present, as juveniles and subadults are disproportionally killed at modern group feeding sites of animals like crocodiles and
Komodo dragon The Komodo dragon (''Varanus komodoensis''), also known as the Komodo monitor, is a large reptile of the monitor lizard family Varanidae that is endemic to the Indonesian islands of Komodo (island), Komodo, Rinca, Flores, Gili Dasami, and Gili ...
s. The same interpretation applies to Bakker's lair sites. There is some evidence for cannibalism in ''Allosaurus'', including ''Allosaurus'' shed teeth found among rib fragments, possible tooth marks on a shoulder blade, and cannibalized allosaur skeletons among the bones at Bakker's lair sites. On the other hand, pathological analysis done by Foth ''et al.'' argued evidence of surviving serious injuries may support gregariousness in ''Allosaurus''.


Brain and senses

The brain of ''Allosaurus'', as interpreted from spiral
CT scan A computed tomography scan (CT scan), formerly called computed axial tomography scan (CAT scan), is a medical imaging technique used to obtain detailed internal images of the body. The personnel that perform CT scans are called radiographers or ...
ning of an
endocast An endocast is the internal cast of a hollow object, often referring to the cranial vault in the study of brain development in humans and other organisms. Endocasts can be artificially made for examining the properties of a hollow, inaccessible ...
, was more consistent with
crocodilia Crocodilia () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchia ...
n brains than those of the other living
archosaur Archosauria () or archosaurs () is a clade of diapsid sauropsid tetrapods, with birds and crocodilians being the only extant taxon, extant representatives. Although broadly classified as reptiles, which traditionally exclude birds, the cladistics ...
s, birds. The structure of the
vestibular apparatus The vestibular system, in vertebrates, is a sensory system that creates the sense of balance and spatial orientation for the purpose of coordinating motor coordination, movement with balance. Together with the cochlea, a part of the auditory sys ...
indicates that the skull was held nearly horizontal, as opposed to strongly tipped up or down. The structure of the
inner ear The inner ear (internal ear, auris interna) is the innermost part of the vertebrate ear. In vertebrates, the inner ear is mainly responsible for sound detection and balance. In mammals, it consists of the bony labyrinth, a hollow cavity in the ...
was like that of a crocodilian, indicating that ''Allosaurus'' was more adapted to hear lower frequencies and would have had difficulty hearing subtle sounds. The
olfactory bulb The olfactory bulb (Latin: ''bulbus olfactorius'') is a neural structure of the vertebrate forebrain involved in olfaction, the sense of smell. It sends olfactory information to be further processed in the amygdala, the orbitofrontal cortex (OF ...
s were large and well suited for detecting odors, but were typical for an animal of its size.


Paleopathology

The specimen "Big Al" is well-known for its numerous injuries. Nineteen of its bones were broken or showed signs of serious infection, which may have contributed to "Big Al's" death. Pathologic bones included five ribs, five vertebrae, and four bones of the feet. A particular problem for the living animal was infection and trauma to the right foot that probably affected movement and may have also predisposed the other foot to injury because of a change in gait. "Big Al" had an infection on the first phalanx on the third toe that was afflicted by an
involucrum An involucrum (plural involucra) is a layer of new bone growth outside existing bone. There are two main contexts: * In pyogenic osteomyelitis where it is a layer of living bone that has formed around dead bone. It can be identified by radiogr ...
. The infection was long-lived, perhaps up to six months. ''Allosaurus'' was one of only two theropods examined in a 2001 study to exhibit a tendon avulsion, and in both cases the avulsion occurred on the forelimb. When the researchers looked for stress fractures, they found that ''Allosaurus'' had a significantly greater number of stress fractures than ''
Albertosaurus ''Albertosaurus'' (; meaning "Alberta lizard") is a genus of large tyrannosaurid theropod dinosaur that lived in northwestern North America during the early to middle Maastrichtian age of the Late Cretaceous period, about 71 million yea ...
'', ''
Ornithomimus ''Ornithomimus'' (; "bird mimic") is a genus of ornithomimid theropod dinosaurs from the Campanian and Maastrichtian ages of Late Cretaceous Western North America. ''Ornithomimus'' was a swift, bipedal dinosaur which fossil evidence indicates wa ...
'' or '' Archaeornithomimus''. Of the 47 hand bones the researchers studied, three were found to contain stress fractures. Of the feet, 281 bones were studied and 17 were found to have stress fractures. The stress fractures in the foot bones "were distributed to the
proximal Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
phalanges The phalanges (: phalanx ) are digit (anatomy), digital bones in the hands and foot, feet of most vertebrates. In primates, the Thumb, thumbs and Hallux, big toes have two phalanges while the other Digit (anatomy), digits have three phalanges. ...
" and occurred across all three weight-bearing toes in "statistically indistinguishable" numbers. Since the lower end of the third metatarsal would have contacted the ground first while an allosaur was running, it would have borne the most stress. If the allosaurs' stress fractures were caused by damage accumulating while walking or running this bone should have experience more stress fractures than the others. The lack of such a bias in the examined ''Allosaurus'' fossils indicates an origin for the stress fractures from a source other than running. The authors conclude that these fractures occurred during interaction with prey, like an allosaur trying to hold struggling prey with its feet. The abundance of stress fractures and avulsion injuries in ''Allosaurus'' provide evidence for "very active" predation-based rather than scavenging diets. The left
scapula The scapula (: 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 either side ...
and
fibula The fibula (: fibulae or fibulas) 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. ...
of an ''Allosaurus fragilis'' specimen cataloged as USNM 4734 are both pathological, both probably due to healed fractures. The specimen USNM 8367 preserved several pathological gastralia which preserve evidence of healed fractures near their middle. Some of the fractures were poorly healed and "formed pseudoarthroses". A specimen with a fractured rib was recovered from the Cleveland-Lloyd Quarry. Another specimen had fractured ribs and fused vertebrae near the end of the tail. An apparent subadult male ''Allosaurus fragilis'' was reported to have extensive pathologies, with a total of fourteen separate injuries. Additionally, a subadult ''Allosaurus'' individual that suffered from
spondyloarthropathy Spondyloarthritis (SpA), also known as spondyloarthropathy, is a collection of syndromes connected by genetic predisposition and clinical symptoms. The best-known subtypes are enteropathic arthritis (EA), psoriatic arthritis (PsA), ankylosing ...
has been discovered in Dana Quarry in Wyoming. This finding represents the first known fossil evidence of spondyloarthropathy occurring in a theropod. Other pathologies reported in ''Allosaurus'' include: * Willow breaks in two ribs * Healed fractures in the
humerus The humerus (; : humeri) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...
and
radius In classical geometry, a radius (: radii or radiuses) of a circle or sphere is any of the line segments from its Centre (geometry), center to its perimeter, and in more modern usage, it is also their length. The radius of a regular polygon is th ...
* Distortion of
joint A joint or articulation (or articular surface) is the connection made between bones, ossicles, or other hard structures in the body which link an animal's skeletal system into a functional whole.Saladin, Ken. Anatomy & Physiology. 7th ed. McGraw- ...
surfaces in the foot, possibly due to
osteoarthritis Osteoarthritis is a type of degenerative joint disease that results from breakdown of articular cartilage, joint cartilage and underlying bone. A form of arthritis, it is believed to be the fourth leading cause of disability in the world, affect ...
or developmental issues *
Osteopetrosis Osteopetrosis, literally , also known as marble bone disease or Albers-Schönberg disease, is an extremely rare inherited disorder whereby the bones harden, becoming denser, in contrast to more prevalent conditions like osteoporosis, in which ...
along the endosteal surface of a
tibia The tibia (; : tibiae or tibias), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two Leg bones, bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outsi ...
. * Distortions of the joint surfaces of the tail vertebrae, possibly due to
osteoarthritis Osteoarthritis is a type of degenerative joint disease that results from breakdown of articular cartilage, joint cartilage and underlying bone. A form of arthritis, it is believed to be the fourth leading cause of disability in the world, affect ...
or developmental issues * " tensive '
neoplastic A neoplasm () is a type of abnormal and excessive growth of tissue. The process that occurs to form or produce a neoplasm is called neoplasia. The growth of a neoplasm is uncoordinated with that of the normal surrounding tissue, and persists ...
'
ankylosis Ankylosis () is a stiffness of a joint due to abnormal adhesion and rigidity of the bones of the joint, which may be the result of injury or disease. The rigidity may be complete or partial and may be due to inflammation of the Tendon, tendinous ...
of caudals", possibly due to physical trauma, as well as the fusion of chevrons to centra * Coossification of vertebral centra near the end of the tail *
Amputation Amputation is the removal of a Limb (anatomy), limb or other body part by Physical trauma, trauma, medical illness, or surgery. As a surgical measure, it is used to control pain or a disease process in the affected limb, such as cancer, malign ...
of a chevron and foot bone, both possibly a result of bites * " tensive exostoses" in the first phalanx of the third toe * Lesions similar to those caused by
osteomyelitis Osteomyelitis (OM) is the infectious inflammation of bone marrow. Symptoms may include pain in a specific bone with overlying redness, fever, and weakness. The feet, spine, and hips are the most commonly involved bones in adults. The cause is ...
in two
scapulae The scapula (: 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 either side ...
*
Bone spurs An exostosis, also known as a osteochondroma, is a benign chondrogenic lesions derived from aberrant cartilage from the perichondral ring. Exostoses can cause chronic pain ranging from mild to moderate, but rarely severe, depending on the shape, s ...
in a
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammals h ...
,
ungual An ungual (from Latin ''unguis'', i.e. ''nail'') is a highly modified distal toe bone which ends in a hoof, claw, or nail. Elephants and ungulates have ungual phalanges, as did the sauropod Sauropoda (), whose members are known as sauropods (; ...
, and two
metacarpals In human anatomy, the metacarpal bones or metacarpus, also known as the "palm bones", are the appendicular skeleton, appendicular bones that form the intermediate part of the hand between the phalanges (fingers) and the carpal bones (wrist, wris ...
* Exostosis in a pedal phalanx possibly attributable to an infectious disease * A metacarpal with a round depressed fracture


Paleoecology

''Allosaurus'' was the most common large theropod in the vast tract of Western American fossil-bearing rock known as the
Morrison Formation The Morrison Formation is a distinctive sequence of Upper Jurassic sedimentary rock found in the western United States which has been the most fertile source of dinosaur fossils in North America. It is composed of mudstone, sandstone, siltston ...
, accounting for 70 to 75% of theropod specimens, and as such was at the top
trophic level The trophic level of an organism is the position it occupies in a food web. Within a food web, a food chain is a succession of organisms that eat other organisms and may, in turn, be eaten themselves. The trophic level of an organism is the ...
of the Morrison food chain. The Morrison Formation is interpreted as a
semiarid A semi-arid climate, semi-desert climate, or steppe climate is a dry climate sub-type. It is located on regions that receive precipitation below potential evapotranspiration, but not as low as a desert climate. There are different kinds of sem ...
environment with distinct wet and
dry season The dry season is a yearly period of low rainfall, especially in the tropics. The weather in the tropics is dominated by the tropical rain belt, which moves from the northern to the southern tropics and back over the course of the year. The t ...
s, and flat
floodplain A floodplain or flood plain or bottomlands is an area of land adjacent to a river. Floodplains stretch from the banks of a river channel to the base of the enclosing valley, and experience flooding during periods of high Discharge (hydrolog ...
s. Vegetation varied from river-lining forests of
conifer Conifers () are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a sin ...
s,
tree fern Tree ferns are arborescent (tree-like) ferns that grow with a trunk (botany), trunk elevating the fronds above ground level, making them trees. Many extant tree ferns are members of the order Cyatheales, to which belong the families Cyatheaceae ( ...
s, and
fern The ferns (Polypodiopsida or Polypodiophyta) are a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. They differ from mosses by being vascular, i.e., having specialized tissue ...
s (
gallery forest A gallery forest is one formed as a corridor along rivers or wetlands, projecting into landscapes that are otherwise only sparsely treed such as savannas, grasslands, or deserts. The gallery forest maintains a more temperate microclimate above th ...
s), to fern
savanna A savanna or savannah is a mixed woodland-grassland (i.e. grassy woodland) biome and ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach th ...
s with occasional trees such as the ''
Araucaria ''Araucaria'' (; original pronunciation: .ɾawˈka. ɾja is a genus of evergreen coniferous trees in the family Araucariaceae. While today they are largely confined to the Southern Hemisphere, during the Jurassic and Cretaceous they were glo ...
''-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 Morrison Formation has been a rich fossil hunting ground. The flora of the period has been revealed by fossils of
green algae The green algae (: green alga) are a group of chlorophyll-containing autotrophic eukaryotes consisting of the phylum Prasinodermophyta and its unnamed sister group that contains the Chlorophyta and Charophyta/ Streptophyta. The land plants ...
, fungi,
moss Mosses are small, non-vascular plant, non-vascular flowerless plants in the taxonomic phylum, division Bryophyta (, ) ''sensu stricto''. Bryophyta (''sensu lato'', Wilhelm Philippe Schimper, Schimp. 1879) may also refer to the parent group bryo ...
es,
horsetails ''Equisetum'' (; horsetail) is the only living genus in Equisetaceae, a family of vascular plants that reproduce by spores rather than seeds. ''Equisetum'' is a "living fossil", the only living genus of the entire subclass Equisetidae, which f ...
, ferns,
cycad Cycads are seed plants that typically have a stout and woody (ligneous) trunk (botany), trunk with a crown (botany), crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants o ...
s,
ginkgo ''Ginkgo'' is a genus of non-flowering seed plants, assigned to the gymnosperms. The scientific name is also used as the English common name. The order to which the genus belongs, Ginkgoales, first appeared in the Permian, , and ''Ginkgo'' is n ...
es, and several families of
conifer Conifers () are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a sin ...
s. Animal fossils discovered include
bivalve Bivalvia () or bivalves, in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class (biology), class of aquatic animal, aquatic molluscs (marine and freshwater) that have laterally compressed soft bodies enclosed b ...
s,
snail A snail is a shelled gastropod. The name is most often applied to land snails, terrestrial molluscs, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class Gas ...
s,
ray-finned fishes Actinopterygii (; ), members of which are known as ray-finned fish or actinopterygians, is a class (biology), class of Osteichthyes, bony fish that comprise over 50% of living vertebrate species. They are so called because of their lightly built ...
, frogs,
salamander Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All t ...
s, turtles, sphenodonts, lizards, terrestrial and aquatic
crocodylomorphs Crocodylomorpha is a group of pseudosuchian archosaurs that includes the crocodilians and their extinct relatives. They were the only members of Pseudosuchia to survive the end-Triassic extinction. Extinct crocodylomorphs were considerably more ...
, several species of
pterosaur Pterosaurs are 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 million to 66 million years ago). Pterosaurs are the earli ...
, numerous dinosaur species, and early
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s such as Docodonta, docodonts, Multituberculata, multituberculates, Symmetrodonta, symmetrodonts, and Triconodonta, triconodonts. Dinosaurs known from the Morrison include the theropods ''
Ceratosaurus ''Ceratosaurus'' (from Greek 'horn' and 'lizard') is a genus of carnivorous theropod dinosaur that lived in the Late Jurassic period (Kimmeridgian to Tithonian ages). The genus was first described in 1884 by American paleontologist Othni ...
'', ''Ornitholestes'', ''Tanycolagreus'', and ''
Torvosaurus ''Torvosaurus'' () is a genus of large Megalosaurinae, megalosaurine Theropoda, theropod dinosaur that lived approximately 165 to 148 million years ago during the Callovian to Tithonian ages of the late Middle Jurassic, Middle and Late Jurassi ...
'', the
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their b ...
s ''Haplocanthosaurus'', ''Camarasaurus'', ''Cathetosaurus'', ''Brachiosaurus'', ''Suuwassea'', ''
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, a ...
'', ''Brontosaurus'', ''Barosaurus'', ''Diplodocus'', ''Supersaurus'', ''Amphicoelias'', and ''Maraapunisaurus'', and the ornithischians ''Camptosaurus'', ''Dryosaurus'', and ''Stegosaurus''. ''Allosaurus'' is commonly found at the same sites as ''Apatosaurus'', ''Camarasaurus'', ''Diplodocus'', and ''Stegosaurus''. The Late Jurassic formations of Portugal where ''Allosaurus'' is present are interpreted as having been similar to the Morrison, but with a stronger Ocean, marine influence. Many of the dinosaurs of the Morrison Formation are the same genera as those seen in Portuguese rocks (mainly ''Allosaurus'', ''Ceratosaurus'', ''Torvosaurus'', and ''Stegosaurus''), or have a close counterpart (''Brachiosaurus'' and ''Lusotitan'', ''Camptosaurus'' and ''Draconyx''). ''Allosaurus'' coexisted with fellow large theropods ''
Ceratosaurus ''Ceratosaurus'' (from Greek 'horn' and 'lizard') is a genus of carnivorous theropod dinosaur that lived in the Late Jurassic period (Kimmeridgian to Tithonian ages). The genus was first described in 1884 by American paleontologist Othni ...
'' and ''
Torvosaurus ''Torvosaurus'' () is a genus of large Megalosaurinae, megalosaurine Theropoda, theropod dinosaur that lived approximately 165 to 148 million years ago during the Callovian to Tithonian ages of the late Middle Jurassic, Middle and Late Jurassi ...
'' in both the United States and Portugal. The three appear to have had different ecological niches, based on anatomy and the location of fossils. ''Ceratosaurus'' and ''Torvosaurus'' may have preferred to be active around waterways, and had lower, thinner bodies that would have given them an advantage in forest and underbrush terrains, whereas ''Allosaurus'' was more compact, with longer legs, faster but less maneuverable, and seems to have preferred dry floodplains. ''Ceratosaurus'', better known than ''Torvosaurus'', differed noticeably from ''Allosaurus'' in functional anatomy by having a taller, narrower skull with large, broad teeth. ''Allosaurus'' was itself a potential food item to other carnivores, as illustrated by an ''Allosaurus'' Pubis (bone), pubic foot marked by the teeth of another theropod, probably ''
Ceratosaurus ''Ceratosaurus'' (from Greek 'horn' and 'lizard') is a genus of carnivorous theropod dinosaur that lived in the Late Jurassic period (Kimmeridgian to Tithonian ages). The genus was first described in 1884 by American paleontologist Othni ...
'' or ''
Torvosaurus ''Torvosaurus'' () is a genus of large Megalosaurinae, megalosaurine Theropoda, theropod dinosaur that lived approximately 165 to 148 million years ago during the Callovian to Tithonian ages of the late Middle Jurassic, Middle and Late Jurassi ...
''. The location of the bone in the body (along the bottom margin of the torso and partially shielded by the legs), and the fact that it was among the most massive in the skeleton, indicates that the ''Allosaurus'' was being scavenged. A bone assemblage in the Upper Jurassic Mygatt-Moore Quarry preserves an unusually high occurrence of theropod bite marks, most of which can be attributed to ''Allosaurus'' and ''Ceratosaurus'', while others could have been made by ''Torvosaurus'' given the size of the striations. While the position of the bite marks on the herbivorous dinosaurs is consistent with predation or early access to remains, bite marks found on ''Allosaurus'' material suggest scavenging, either from the other theropods or from another ''Allosaurus''. The unusually high concentration of theropod bite marks compared to other assemblages could be explained either by a more complete utilization of resources during a dry season by theropods, or by a collecting bias in other localities.


Notes


References


External links


Specimens, discussion, and references pertaining to ''Allosaurus fragilis''
at The Theropod Database

(), from Pioneer: Utah's Online Library
Restoration of MOR 693 ("Big Al")
an

at Scott Hartman's Skeletal Drawing website

() {{Authority control Allosauridae Dinosaur genera Kimmeridgian dinosaurs Tithonian dinosaurs Lourinhã Formation Morrison Formation Dinosaurs of the United States Fossil taxa described in 1877 Taxa named by Othniel Charles Marsh