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was found a short distance from a skeleton (specimen CM3018) identified as the new species "Apatosaurus louisae", named after Louise Carnegie, wife of Andrew Carnegie, who funded field research to find complete dinosaur skeletons in the American West. The skull was designated CM11162; it was very similar to the skull of "Diplodocus". Another smaller skeleton of "A.louisae" was found nearby CM11162 and CM3018. The skull was accepted as belonging to the "Apatosaurus" specimen by Douglass and Carnegie Museum director William H. Holland, although other scientistsmost notably Osbornrejected this identification. Holland defended his view in 1914 in an address to the
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Apatosaurus
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] |
6002
|
Paleontological Society of America, yet he left the Carnegie Museum mount headless. While some thought Holland was attempting to avoid conflict with Osborn, others suspected Holland was waiting until an articulated skull and neck were found to confirm the association of the skull and skeleton. After Holland's death in 1934, museum staff placed a cast of a "Camarasaurus" skull on the mount. While most other museums were using cast or sculpted "Camarasaurus" skulls on "Apatosaurus" mounts, the Yale Peabody Museum decided to sculpt a skull based on the lower jaw of a "Camarasaurus", with the cranium based on Marsh's 1891
|
Apatosaurus
|
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] |
6003
|
illustration of the skull. The skull also included forward-pointing nasalssomething different to any dinosaurand fenestrae differing from both the drawing and other skulls. No "Apatosaurus" skull was mentioned in literature until the 1970s when John Stanton McIntosh and David Berman redescribed the skulls of "Diplodocus" and "Apatosaurus". They found that though he never published his opinion, Holland was almost certainly correct, that "Apatosaurus" had a "Diplodocus"-like skull. According to them, many skulls long thought to pertain to "Diplodocus" might instead be those of "Apatosaurus". They reassigned multiple skulls to "Apatosaurus" based on associated and closely associated vertebrae. Even though they
|
Apatosaurus
|
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] |
6004
|
supported Holland, it was noted that "Apatosaurus" might have possessed a "Camarasaurus"-like skull, based on a disarticulated "Camarasaurus"-like tooth found at the precise site where an "Apatosaurus" specimen was found years before. On October20, 1979, after the publications by McIntosh and Berman, the first true skull of "Apatosaurus" was mounted on a skeleton in a museum, that of the Carnegie. In 1998 it was suggested that the Felch Quarry skull that Marsh had included in his 1896 skeletal restoration instead belonged to "Brachiosaurus". In 2011 the first specimen of "Apatosaurus" where a skull was found articulated with its cervical vertebrae
|
Apatosaurus
|
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] |
6005
|
was described. This specimen, CMCVP7180, was found to differ in both skull and neck features from "A.louisae", but shared many features of the cervical vertebrae with "A.ajax". Another well-preserved skull is Brigham Young University specimen 17096, a well-preserved skull and skeleton, with a preserved braincase. The specimen was found in Cactus Park Quarry in western Colorado. Almost all modern paleontologists agreed with Riggs that the two dinosaurs should be classified together in a single genus. According to the rules of the ICZN (which governs the scientific names of animals), the name "Apatosaurus", having been published first, has priority as the
|
Apatosaurus
|
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0.4306437373161316,
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] |
6006
|
official name; "Brontosaurus" was considered a junior synonym and was therefore long discarded from formal use. Despite this, at least one paleontologistRobert T. Bakkerargued in the 1990s that "A.ajax" and "A.excelsus" were in fact sufficiently distinct for the latter to merit a separate genus. In 2015 Emanuel Tschopp, Octávio Mateus, and Roger Benson released a paper on diplodocoid systematics, and proposed that genera could be diagnosed by thirteen differing characters, and species separated based on six. The minimum number for generic separation was chosen based on the fact that "A.ajax" and "A.louisae" differ in twelve characters, and "Diplodocus carnegiei" and
|
Apatosaurus
|
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] |
6007
|
"D.hallorum" differ in eleven characters. Thus, thirteen characters were chosen to validate the separation of genera. The six differing features for specific separation were chosen by counting the number of differing features in separate specimens generally agreed to represent one species, with only one differing character in "D.carnegiei" and "A.louisae", but five differing features in "B.excelsus". Therefore, Tschopp etal. argued that "Apatosaurus excelsus", originally classified as "Brontosaurus excelsus", had enough morphological differences from other species of "Apatosaurus" that it warranted being reclassified as a separate genus again. The conclusion was based on a comparison of 477 morphological characteristics across 81
|
Apatosaurus
|
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] |
6008
|
different dinosaur individuals. Among the many notable differences are the widerand presumably strongerneck of "Apatosaurus" species compared to "B.excelsus". Other species previously assigned to "Apatosaurus", such as "Elosaurus parvus" and "Eobrontosaurus yahnahpin" were also reclassified as "Brontosaurus". Some features proposed to separate "Brontosaurus" from "Apatosaurus" include: posterior dorsal vertebrae with the centrum longer than wide; the scapula rear to the acromial edge and the distal blade being excavated; the acromial edge of the distal scapular blade bearing a rounded expansion; and the ratio of the proximodistal length to transverse breadth of the astragalus 0.55 or greater. Sauropod expert Michael Daniel
|
Apatosaurus
|
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] |
6009
|
D'Emic pointed out that the criteria chosen were to an extent arbitrary and that they would require abandoning the name "Brontosaurus" again if newer analyses obtained different results. Mammal palaeontologist Donald Prothero criticized the mass media reaction to this study as superficial and premature, concluding that he would keep "Brontosaurus" in quotes and not treat the name as a valid genus. Many species of "Apatosaurus" have been designated from scant material. Marsh named as many species as he could, which resulted in many being based upon fragmentary and indistinguishable remains. In 2005 Paul Upchurch and colleagues published a study that
|
Apatosaurus
|
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] |
6010
|
analyzed the species and specimen relationships of "Apatosaurus". They found that "A.louisae" was the most basal species, followed by FMNHP25112, and then a polytomy of "A.ajax", "A.parvus", and "A.excelsus". Their analysis was revised and expanded with many additional diplodocid specimens in 2015, which resolved the relationships of "Apatosaurus" slightly differently, and also supported separating "Brontosaurus" from "Apatosaurus". The cladogram below is the result of an analysis by Tschopp, Mateus, and Benson (2015). The authors analyzed most diplodocid type specimens separately to deduce which specimen belonged to which species and genus. "Apatosaurus" is a member of the family Diplodocidae, a clade
|
Apatosaurus
|
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] |
6011
|
of gigantic sauropod dinosaurs. The family includes some of the longest creatures ever to walk the earth, including "Diplodocus", "Supersaurus", and "Barosaurus". "Apatosaurus" is sometimes classified in the subfamily Apatosaurinae, which may also include "Suuwassea", "Supersaurus", and "Brontosaurus". Othniel Charles Marsh described "Apatosaurus" as allied to "Atlantosaurus" within the now-defunct group Atlantosauridae. In 1878 Marsh raised his family to the rank of suborder, including "Apatosaurus", "Atlantosaurus", "Morosaurus" (="Camarasaurus") and "Diplodocus". He classified this group within Sauropoda, a group he erected in the same study. In 1903 Elmer S. Riggs said the name Sauropoda would be a junior synonym of earlier
|
Apatosaurus
|
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] |
6012
|
names; he grouped "Apatosaurus" within Opisthocoelia. Sauropoda is still used as the group name. In 2011, John Whitlock published a study that placed "Apatosaurus" a more basal diplodocid, sometimes less basal than "Supersaurus". Cladogram of the Diplodocidae after Tschopp, Mateus, and Benson (2015). It was believed throughout the 19th and early 20th centuries that sauropods like "Apatosaurus" were too massive to support their own weight on dry land. It was theorized that they lived partly submerged in water, perhaps in swamps. More recent findings do not support this; sauropods are now thought to have been fully terrestrial animals. A study
|
Apatosaurus
|
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] |
6013
|
of diplodocid snouts showed that the square snout, large proportion of pits, and fine, subparallel scratches of the teeth of "Apatosaurus" suggests it was a ground-height, nonselective browser. It may have eaten ferns, cycadeoids, seed ferns, horsetails, and algae. Stevens and Parish (2005) speculate that these sauropods fed from riverbanks on submerged water plants. A 2015 study of the necks of "Apatosaurus" and "Brontosaurus" found many differences between them and other diplodocids, and that these variations may have shown that the necks of "Apatosaurus" and "Brontosaurus" were used for intraspecific combat. Various uses for the single claw on the forelimb
|
Apatosaurus
|
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] |
6014
|
of sauropods have been proposed. One suggestion is that they were used for defense, but their shape and size make this unlikely. It was also possible they were for feeding, but the most probable use for the claw was grasping objects such as tree trunks when rearing. Trackways of sauropods like "Apatosaurus" show that they may have had a range of around per day, and that they could potentially have reached a top speed of per hour. The slow locomotion of sauropods may be due to their minimal muscling, or to recoil after strides. A trackway of a juvenile has
|
Apatosaurus
|
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] |
6015
|
led some to believe that they were capable of bipedalism, though this is disputed. Diplodocids like "Apatosaurus" are often portrayed with their necks held high up in the air, allowing them to browse on tall trees. Some studies state diplodocid necks were less flexible than previously believed, because the structure of the neck vertebrae would not have allowed the neck to bend far upwards, and that sauropods like "Apatosaurus" were adapted to low browsing or ground feeding. Other studies by Taylor find that all tetrapods appear to hold their necks at the maximum possible vertical extension when in a normal,
|
Apatosaurus
|
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] |
6016
|
alert posture; they argue the same would hold true for sauropods barring any unknown, unique characteristics that set the soft tissue anatomy of their necks apart from that of other animals. "Apatosaurus", like "Diplodocus", would have held its neck angled upwards with the head pointing downwards in a resting posture. Kent Stevens and Michael Parrish (1999 and 2005) state "Apatosaurus" had a great feeding range; its neck could bend into a U-shape laterally. The neck's range of movement would have also allowed the head to feed at the level of the feet. Matthew Cobley et al. (2013) dispute this, finding
|
Apatosaurus
|
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] |
6017
|
that large muscles and cartilage would have limited movement of the neck. They state the feeding ranges for sauropods like "Diplodocus" were smaller than previously believed, and the animals may have had to move their whole bodies around to better access areas where they could browse vegetation. As such, they might have spent more time foraging to meet their minimum energy needs. The conclusions of Cobley etal. are disputed by Taylor, who analyzed the amount and positioning of intervertebral cartilage to determine the flexibility of the neck of "Apatosaurus" and "Diplodocus". He found that the neck of "Apatosaurus" was very
|
Apatosaurus
|
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] |
6018
|
flexible. Given the large body mass and long neck of sauropods like "Apatosaurus", physiologists have encountered problems determining how these animals breathed. Beginning with the assumption that, like crocodilians, "Apatosaurus" did not have a diaphragm, the dead-space volume (the amount of unused air remaining in the mouth, trachea, and air tubes after each breath) has been estimated at about for a specimen. Paladino calculates its tidal volume (the amount of air moved in or out during a single breath) at with an avian respiratory system, if mammalian, and if reptilian. On this basis, its respiratory system would likely have been
|
Apatosaurus
|
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] |
6019
|
parabronchi, with multiple pulmonary air sacs as in avian lungs, and a flow-through lung. An avian respiratory system would need a lung volume of about compared with a mammalian requirement of , which would exceed the space available. The overall thoracic volume of "Apatosaurus" has been estimated at , allowing for a , four-chambered heart and a lung capacity. That would allow about for the necessary tissue. Evidence for the avian system in "Apatosaurus" and other sauropods is also present in the pneumaticity of the vertebrae. Though this plays a role in reducing the weight of the animal, Wedel (2003)
|
Apatosaurus
|
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] |
6020
|
states they are also likely connected to air sacs, as in birds. James Spotila et al. (1991) concludes that the large body size of sauropods would have made them unable to maintain high metabolic rates because they would not have been able to release enough heat. They assumed sauropods had a reptilian respiratory system. Wedel says that an avian system would have allowed it to dump more heat. Some scientists state that the heart would have had trouble sustaining sufficient blood pressure to oxygenate the brain. Others suggest that the near-horizontal posture of the head and neck would have eliminated
|
Apatosaurus
|
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0.08751672506332397
] |
6021
|
the problem of supplying blood to the brain because it would not have been elevated. James Farlow (1987) calculates that an "Apatosaurus"-sized dinosaur about would have possessed of fermentation contents. Assuming "Apatosaurus" had an avian respiratory system and a reptilian resting-metabolism, Frank Paladino etal. (1997) estimate the animal would have needed to consume only about of water per day. A 1999 microscopic study of "Apatosaurus" and "Brontosaurus" bones concluded the animals grew rapidly when young and reached near-adult sizes in about 10years. In 2008, a study on the growth rates of sauropods was published by Thomas Lehman and Holly Woodward.
|
Apatosaurus
|
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] |
6022
|
They said that by using growth lines and length-to-mass ratios, "Apatosaurus" would have grown to 25t (25 long tons; 28 short tons) in 15years, with growth peaking at in a single year. An alternative method, using limb length and body mass, found "Apatosaurus" grew per year, and reached its full mass before it was about 70years old. These estimates have been called unreliable because the calculation methods are not sound; old growth lines would have been obliterated by bone remodelling. One of the first identified growth factors of "Apatosaurus" was the number of sacral vertebrae, which increased to five by
|
Apatosaurus
|
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] |
6023
|
the time of the creature's maturity. This was first noted in 1903 and again in 1936. Long-bone histology enables researchers to estimate the age that a specific individual reached. A study by Eva Griebeler etal. (2013) examined long-bone histological data and concluded the "Apatosaurus" sp.SMA0014 weighed , reached sexual maturity at 21years, and died aged 28. The same growth model indicated "Apatosaurus" sp.BYU 601–17328 weighed , reached sexual maturity at 19years, and died aged 31. Compared with most sauropods, a relatively large amount of juvenile material is known from "Apatosaurus". Multiple specimens in the OMNH are from juveniles of an
|
Apatosaurus
|
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] |
6024
|
undetermined species of "Apatosaurus"; this material includes partial shoulder and pelvic girdles, some vertebrae, and limb bones. OMNH juvenile material is from at least two different age groups and based on overlapping bones likely comes from more than three individuals. The specimens exhibit features that distinguish "Apatosaurus" from its relatives, and thus likely belong to the genus. Juvenile sauropods tend to have proportionally shorter necks and tails, and a more pronounced forelimb-hindlimb disparity than found in adult sauropods. An article published in 1997 reported research of the mechanics of "Apatosaurus" tails by Nathan Myhrvold and paleontologist Philip J. Currie. Myhrvold
|
Apatosaurus
|
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] |
6025
|
carried out a computer simulation of the tail, which in diplodocids like "Apatosaurus" was a very long, tapering structure resembling a bullwhip. This computer modeling suggested diplodocids were capable of producing a whiplike cracking sound of over 200 decibels, comparable to the volume of a cannon being fired. A pathology has been identified on the tail of "Apatosaurus", caused by a growth defect. Two caudal vertebrae are seamlessly fused along the entire articulating surface of the bone, including the arches of the neural spines. This defect might have been caused by the lack or inhibition of the substance that forms
|
Apatosaurus
|
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0.01996856927871704
] |
6026
|
intervertebral disks or joints. It has been proposed that the whips could have been used in combat and defense, but the tails of diplodocids were quite light and narrow compared to "Shunosaurus" and mamenchisaurids, and thus to injure another animal with the tail would severely injure the tail itself. The Morrison Formation is a sequence of shallow marine and alluvial sediments which, according to radiometric dating, dates from between 156.3mya at its base, and 146.8mya at the top, placing it in the late Oxfordian, Kimmeridgian, and early Tithonian stages of the Late Jurassic period. This formation is interpreted as originating
|
Apatosaurus
|
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] |
6027
|
in a locally semiarid environment with distinct wet and dry seasons. The Morrison Basin, where dinosaurs lived, stretched from New Mexico to Alberta and Saskatchewan; it was formed when the precursors to the Front Range of the Rocky Mountains started pushing up to the west. The deposits from their east-facing drainage basins were carried by streams and rivers and deposited in swampy lowlands, lakes, river channels, and floodplains. This formation is similar in age to the Lourinhã Formation in Portugal and the Tendaguru Formation in Tanzania. "Apatosaurus" was the second most common sauropod in the Morrison Formation ecosystem, after "Camarasaurus".
|
Apatosaurus
|
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] |
6028
|
"Apatosaurus" may have been more solitary than other Morrison Formation dinosaurs. Fossils of the genus have only been found in the upper levels of the formation. Those of "Apatosaurus ajax" are known exclusively from the upper Brushy Basin Member, about 152–151 mya. "A.louisae" fossils are rare, known only from one site in the upper Brushy Basin Member; they date to the late Kimmeridgian stage, about 151mya. Additional "Apatosaurus" remains are known from similarly aged or slightly younger rocks, but they have not been identified as any particular species, and thus may instead belong to "Brontosaurus". The Morrison Formation records a
|
Apatosaurus
|
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] |
6029
|
time when the local environment was dominated by gigantic sauropod dinosaurs. Dinosaurs known from the Morrison Formation include the theropods "Allosaurus", "Ceratosaurus", "Ornitholestes", "Saurophaganax", and "Torvosaurus"; the sauropods "Brontosaurus", "Brachiosaurus", "Camarasaurus", and "Diplodocus"; and the ornithischians "Camptosaurus", "Dryosaurus", and "Stegosaurus". "Apatosaurus" is commonly found at the same sites as "Allosaurus", "Camarasaurus", "Diplodocus", and "Stegosaurus". "Allosaurus" accounted for 70–75% of theropod specimens and was at the top trophic level of the Morrison food web. Many of the dinosaurs of the Morrison Formation are of the same genera as those seen in Portuguese rocks of the Lourinhã Formationmainly "Allosaurus", "Ceratosaurus", and "Torvosaurus"or
|
Apatosaurus
|
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] |
6030
|
have a close counterpart"Brachiosaurus" and "Lusotitan", "Camptosaurus" and "Draconyx", and "Apatosaurus" and "Dinheirosaurus". Other vertebrates that are known to have shared this paleo-environment include ray-finned fishes, frogs, salamanders, turtles, sphenodonts, lizards, terrestrial and aquatic crocodylomorphans, and several species of pterosaur. Shells of bivalves and aquatic snails are also common. The flora of the period has been evidenced in fossils of green algae, fungi, mosses, horsetails, cycads, ginkgoes, and several families of conifers. Vegetation varied from river-lining forests of tree ferns with fern understory (gallery forests), to fern savannas with occasional trees such as the "Araucaria"-like conifer "Brachyphyllum". Apatosaurus Apatosaurus (;
|
Apatosaurus
|
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6031
|
Allosaurus Allosaurus () is a genus of carnivorous theropod dinosaur that lived 155 to 150 million years ago during the late Jurassic period (Kimmeridgian to early Tithonian). The name ""Allosaurus"" means "different lizard" alluding to its unique concave vertebrae (at the time of its discovery). It is derived from the Greek /"allos" ("different, other") and /"sauros" ("lizard / generic reptile"). The first fossil remains that could definitively be ascribed to this genus were described in 1877 by paleontologist Othniel Charles Marsh. As one of the first well-known theropod dinosaurs, it has long attracted attention outside of paleontological circles. Indeed, it
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Allosaurus
|
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6032
|
has been a top feature in several films and documentaries about prehistoric life. "Allosaurus" was a large bipedal predator. Its skull was large and equipped with dozens of sharp, serrated teeth. It averaged in length, though fragmentary remains suggest it could have reached over . Relative to the large and powerful hindlimbs, its three-fingered forelimbs were small, and the body was balanced by a long and heavily muscled tail. It is classified as an allosaurid, a type of carnosaurian theropod dinosaur. The genus has a complicated taxonomy, and includes an uncertain number of valid species, the best known of which
|
Allosaurus
|
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] |
6033
|
is "A. fragilis". The bulk of "Allosaurus" remains have come from North America's Morrison Formation, with material also known from Portugal and possibly Tanzania. It was known for over half of the 20th century as "Antrodemus", but a study of the copious remains from the Cleveland-Lloyd Dinosaur Quarry brought the name ""Allosaurus"" back to prominence and established it as one of the best-known dinosaurs. As the most abundant large predator in the Morrison Formation, "Allosaurus" was at the top of the food chain, probably preying on contemporaneous large herbivorous dinosaurs, and perhaps even other predators. Potential prey included ornithopods, stegosaurids,
|
Allosaurus
|
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] |
6034
|
and sauropods. Some paleontologists interpret "Allosaurus" as having had cooperative social behavior, and hunting in packs, while others believe individuals may have been aggressive toward each other, and that congregations of this genus are the result of lone individuals feeding on the same carcasses. Due to having a much weaker bite force and a much wider gape than most other large carnivorous dinosaurs, it may have attacked large prey by ambush, using its upper jaw like a hatchet. "Allosaurus" was a typical large theropod, having a massive skull on a short neck, a long tail and reduced forelimbs. "Allosaurus fragilis",
|
Allosaurus
|
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] |
6035
|
the best-known species, had an average length of , with the largest definitive "Allosaurus" specimen (AMNH 680) estimated at long, and an estimated weight of . In his 1976 monograph 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 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
|
Allosaurus
|
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] |
6036
|
individuals represented by the average-sized thigh bones he has measured. Using the subadult specimen nicknamed "Big Al", researchers using computer modelling arrived at a best estimate of for the individual, but by varying parameters they found a range from approximately to approximately . Several gigantic specimens have been attributed to "Allosaurus", but may in fact belong to other genera. The closely related genus "Saurophaganax" (OMNH 1708) reached perhaps in length, and its single species has sometimes been included in the genus "Allosaurus" as "Allosaurus maximus", though recent studies support it as a separate genus. Another potential specimen of "Allosaurus", once
|
Allosaurus
|
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] |
6037
|
assigned to the genus "Epanterias" (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; this large allosaurid may be another individual of "Saurophaganax". David K. Smith, examining "Allosaurus" fossils by quarry, found that the Cleveland-Lloyd Dinosaur Quarry (Utah) specimens are generally smaller than those from Como Bluff (Wyoming) or Brigham Young University'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 (Utah) specimens
|
Allosaurus
|
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] |
6038
|
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, an ankle bone. Kenneth Carpenter, 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 skull and teeth of
|
Allosaurus
|
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] |
6039
|
"Allosaurus" were modestly proportioned for a theropod of its size. Paleontologist Gregory S. Paul gives a length of for a skull belonging to an individual he estimates at long. Each premaxilla (the bones that formed the tip of the snout), held five teeth with D-shaped cross-sections, and each maxilla (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 (the tooth-bearing bone of the lower jaw) had between 14 and 17 teeth, with an average count of 16. The teeth
|
Allosaurus
|
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] |
6040
|
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. The skull had a pair of horns above and in front of the eyes. These horns were composed of extensions of the lacrimal bones, and varied in shape and size. There were also lower paired ridges running along the top edges of the nasal bones that led into the horns. The horns were probably covered in a keratin sheath and may have had a variety of functions, including acting
|
Allosaurus
|
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] |
6041
|
as sunshades for the eye, 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 tyrannosaurids. Inside the lacrimal bones were depressions that may have held glands, such as salt glands. Within the maxillae were sinuses that were better developed than those of more basal theropods such as "Ceratosaurus" and "Marshosaurus"; they may have been related to the sense of smell, perhaps holding something like Jacobson's organ. The roof of the
|
Allosaurus
|
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] |
6042
|
braincase was thin, perhaps to improve thermoregulation 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 and frontals may also have had a joint. "Allosaurus" had nine vertebrae in the neck, 14 in the back, and five in the sacrum supporting the hips. The number of tail vertebrae is unknown and varied with individual size; James Madsen estimated about 50, while Gregory S. Paul considered that
|
Allosaurus
|
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] |
6043
|
to be too many and suggested 45 or less. There were hollow spaces in the neck and anterior back vertebrae. Such spaces, which are also found in modern theropods (that is, the birds), are interpreted as having held air sacs used in respiration. The rib cage was broad, giving it a barrel chest, especially in comparison to less derived theropods like "Ceratosaurus". "Allosaurus" had gastralia (belly ribs), but these are not common findings, and they may have ossified poorly. In one published case, the gastralia show evidence of injury during life. A furcula (wishbone) was also present, but has only
|
Allosaurus
|
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] |
6044
|
been recognized since 1996; in some cases furculae were confused with gastralia. The ilium, the main hip bone, was massive, and the pubic bone 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, 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
|
Allosaurus
|
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] |
6045
|
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 claws. The arms were powerful, and the forearm was somewhat shorter than the upper arm (1:1.2 ulna/humerus ratio). The wrist had a version of the semilunate carpal also found in more derived theropods like maniraptorans. 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
|
Allosaurus
|
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] |
6046
|
bones, the next has three, and the third finger has four. The legs were not as long or suited for speed as those of tyrannosaurids, and the claws of the toes were less developed and more hoof-like than those of earlier theropods. Each foot had three weight-bearing toes and an inner dewclaw, 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, perhaps used as a lever between the Achilles tendon and foot. The discovery and early study of "Allosaurus" is complicated by the
|
Allosaurus
|
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] |
6047
|
multiplicity of names coined during the Bone Wars of the late 19th century. The first described fossil in this history was a bone obtained secondhand by Ferdinand Vandiveer Hayden in 1869. It came from Middle Park, near Granby, Colorado, probably from Morrison Formation rocks. The locals had identified such bones as "petrified horse hoofs". Hayden sent his specimen to Joseph Leidy, 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
|
Allosaurus
|
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] |
6048
|
small collection of fragmentary bones including parts of three vertebrae, a rib fragment, a tooth, a toe bone, and, most useful for later discussions, the shaft of the right humerus (upper arm). Othniel Charles Marsh gave these remains the formal name "Allosaurus fragilis" in 1877. "Allosaurus" comes from the Greek "allos/αλλος", meaning "strange" or "different" and "sauros/σαυρος", meaning "lizard" or "reptile". It was named 'different lizard' because its vertebrae were different from those of other dinosaurs known at the time of its discovery. The species epithet "fragilis" is Latin for "fragile", referring to lightening features in the vertebrae. The bones
|
Allosaurus
|
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] |
6049
|
were collected from the Morrison Formation of Garden Park, north of Cañon City. Marsh and Edward Drinker Cope, who were in scientific competition, 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", and 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 type specimen of "Allosaurus" in Colorado, Marsh elected to concentrate work in Wyoming; when work
|
Allosaurus
|
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] |
6050
|
resumed at Garden Park in 1883, 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 area of Wyoming in 1879, but apparently did not mention its completeness, and Cope never unpacked it. Upon unpacking in 1903 (several years after Cope had died), it was found to be one of the most complete theropod specimens then known, and in 1908 the skeleton, now cataloged as AMNH 5753, was put on public view. This is the well-known mount poised over a partial "Apatosaurus"
|
Allosaurus
|
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] |
6051
|
skeleton as if scavenging it, illustrated as such 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 multiplicity of early names complicated later research, with the situation compounded by the terse descriptions provided by Marsh and Cope. Even at the time, authors such as Samuel Wendell Williston suggested that too many names had been coined. For example, Williston pointed out in 1901 that Marsh had never been able to adequately distinguish "Allosaurus" from "Creosaurus". The most influential early attempt to sort out
|
Allosaurus
|
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] |
6052
|
the convoluted situation was produced by Charles W. Gilmore in 1920. He came to the conclusion that the tail vertebra named "Antrodemus" by Leidy was indistinguishable from those of "Allosaurus", and "Antrodemus" thus should be the preferred name because as the older name it had priority. "Antrodemus" became the accepted name for this familiar genus for over fifty years, until James 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 that the single bone of "Antrodemus"
|
Allosaurus
|
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] |
6053
|
came from is unknown). ""Antrodemus"" has been used informally for convenience when distinguishing between the skull Gilmore restored and the composite skull restored by Madsen. Although sporadic work at what became known as the Cleveland-Lloyd Dinosaur Quarry in Emery County, Utah had taken place as early as 1927, and the fossil site itself described by William L. Stokes in 1945, major operations did not begin there until 1960. Under a cooperative effort involving nearly 40 institutions, thousands of bones were recovered between 1960 and 1965. The quarry is notable for the predominance of "Allosaurus" remains, the condition of the specimens,
|
Allosaurus
|
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] |
6054
|
and the lack of scientific resolution on how it came to be. The majority of bones belong to the large theropod "Allosaurus fragilis" (it is estimated that the remains of at least 46 "A. fragilis" have been found there, out of at minimum 73 dinosaurs), and the fossils found there are disarticulated and well-mixed. Nearly a dozen scientific papers have been written on the taphonomy of the site, suggesting numerous mutually exclusive explanations for how it may have formed. Suggestions have ranged from animals getting stuck in a bog, to becoming trapped in deep mud, to falling victim to drought-induced
|
Allosaurus
|
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] |
6055
|
mortality around a waterhole, to getting trapped in a spring-fed pond or seep. Regardless of the actual cause, the great quantity of well-preserved "Allosaurus" remains has allowed this genus to be known in detail, making it among the best-known 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 bones usually found fused. The period since Madsen's monograph has been marked by a great expansion in studies dealing with topics concerning "Allosaurus" in life (paleobiological and paleoecological topics). Such studies have covered
|
Allosaurus
|
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] |
6056
|
topics including skeletal variation, growth, skull construction, hunting methods, the brain, and the possibility of gregarious living and parental care. Reanalysis of old material (particularly of large 'allosaur' specimens), new discoveries in Portugal, and several very complete new specimens have also contributed to the growing knowledge base. In 1991 "Big Al" (MOR 693), a 95% complete, partially articulated specimen of "Allosaurus" was discovered. It measured about 8 meters (about 26 ft) in length. MOR 693 was excavated near Shell, Wyoming, by a joint Museum of the Rockies and University of Wyoming Geological Museum team. This skeleton was discovered by a
|
Allosaurus
|
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] |
6057
|
Swiss team, led by Kirby Siber. In 1996 the same team discovered a second "Allosaurus", "Big Al Two", which is the best preserved skeleton of its kind to date. The completeness, preservation, and scientific importance of this skeleton gave "Big Al" its name; the individual itself was below the average size for "Allosaurus fragilis", and was a subadult estimated at only 87% grown. The specimen was described by Breithaupt in 1996. Nineteen of its bones were broken or showed signs of infection, which may have contributed to "Big Al's" death. Pathologic bones included five ribs, five vertebrae, and four bones
|
Allosaurus
|
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] |
6058
|
of the feet; several damaged bones showed osteomyelitis, a bone infection. 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. Al had an infection on the first phalanx on the third toe that was afflicted by an involucrum. The infection was long lived, perhaps up to 6 months. Big Al Two is also known to have multiple injuries. There are currently four valid and one undescribed species of "Allosaurus" ("A. amplus", "A. europaeus", the
|
Allosaurus
|
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] |
6059
|
type species "A. fragilis", the as-yet not formally described "A. jimmadseni", and "A. lucasi"). "A. fragilis" is the type species and was named by Marsh in 1877. It is known from the remains of at least sixty individuals, all found in the Kimmeridgian–Tithonian Upper Jurassic-age Morrison Formation of the United States, spread across the states of Colorado, Montana, New Mexico, Oklahoma, South Dakota, Utah, and Wyoming. Details of the humerus (upper arm) of "A. fragilis" have been used as diagnostic among Morrison theropods, but the discovery of "A. jimmadseni" indicates that this will no longer be the case at the
|
Allosaurus
|
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] |
6060
|
species level. Daniel Chure's work on Morrison allosaurid remains has been responsible, directly or indirectly, for "A. jimmadseni" and "A. maximus". "A. jimmadseni" is the proposed name for a new species of Morrison allosaur, based on a nearly complete skeleton and skull. It differs from "A. fragilis" in several anatomical details including a jugal or cheekbone with a straight lower margin, and is confined to the Salt Wash Member of the Morrison Formation, while "A. fragilis" is only present in the higher Brushy Basin Member. "A. fragilis", "A. jimmadseni", "A. amplus", and "A. lucasi" are all known from remains discovered
|
Allosaurus
|
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0.053760360926389694,
0.009680540300905704,
0.3169574439525604,
-0.252501517534256
] |
6061
|
in the Kimmeridgian–Tithonian Upper Jurassic-age Morrison Formation of the United States, spread across the states of Colorado, Montana, New Mexico, Oklahoma, South Dakota, Utah and Wyoming. "A. fragilis" is regarded as the most common, known from the remains of at least sixty 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
|
Allosaurus
|
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] |
6062
|
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. "A. europaeus" was found in the Kimmeridgian-age Porto Novo Member of the Lourinhã Formation, but may be the same as "A. fragilis". "Allosaurus" material from Portugal was first reported in 1999 on the basis of MHNUL/AND.001, a partial skeleton including a quadrate, vertebrae, ribs, gastralia, chevrons, part of the hips, and hindlimbs.
|
Allosaurus
|
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] |
6063
|
This specimen was assigned to "A. fragilis", but the subsequent discovery of a partial skull and neck (ML 415) near Lourinhã, in the Kimmeridgian-age Porto Novo Member of the Lourinhã Formation, spurred the naming of the new species "A. europaeus". It differs from other species of "Allosaurus" in cranial details. However, more material may show it to be "A. fragilis", as originally described. The issue of species and potential synonyms is complicated by the type specimen of "Allosaurus fragilis" (catalog number YPM 1930) being extremely fragmentary, consisting of a few incomplete vertebrae, limb bone fragments, rib fragments, and a tooth.
|
Allosaurus
|
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] |
6064
|
Because of this, several scientists have interpreted the type specimen as potentially dubious, and thus 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 and Kenneth Carpenter (2010) submitted a petition to the ICZN to have the name ""A. fragilis"" officially transferred to the more complete specimen USNM4734 (as a neotype). This request is currently pending review. "Creosaurus", "Epanterias", and "Labrosaurus" are regarded as junior synonyms of "Allosaurus". Most of
|
Allosaurus
|
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] |
6065
|
the species that are regarded as synonyms of "A. fragilis", or that were misassigned to the genus, are obscure and were based on scrappy remains. One exception is "Labrosaurus ferox", named in 1884 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
|
Allosaurus
|
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0.14326675236225128,
0.3794768154621124,
0.01747852936387062
] |
6066
|
now regarded as an example of "A. fragilis." In his 1988 book "Predatory Dinosaurs of the World", the freelance dinosaurologist Gregory Paul proposed that "A. fragilis" had tall pointed horns and a slender build compared to a postulated second species "A. atrox", and was not a different gender due to rarity. "Allosaurus atrox" was originally named by Marsh in 1878 as the type species of its own genus, "Creosaurus", and is based on YPM 1890, an assortment of bones including a couple of pieces of the skull, portions of nine tail vertebrae, two hip vertebrae, an illium, and ankle and
|
Allosaurus
|
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] |
6067
|
foot bones. Although the idea of two common Morrison allosaur species was followed in some semi-technical and popular works, the 2000 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, and Osborn, 1912, 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
|
Allosaurus
|
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] |
6068
|
1893, and YPM 1893 has been treated as a specimen of "A fragilis." Likewise, "Labrosaurus fragilis" is a typographical error by Marsh (1896) for "Labrosaurus ferox". "A. whitei" is a nomen nudum coined by Pickering in 1996 for the complete "Allosaurus" specimens that Paul referred to "A. atrox". 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 Arundel Formation of Maryland, although most of the remains were removed by Richard Swann Lull to the new ornithopod species "Dryosaurus grandis",
|
Allosaurus
|
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] |
6069
|
except for a tooth. Gilmore considered the tooth nondiagnostic but transferred it to "Dryptosaurus", as "D. medius". The referral was not accepted in the most recent review, and "Allosaurus medius" was simply listed as a dubious species of theropod. It may be closely related to "Acrocanthosaurus". "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
|
Allosaurus
|
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] |
6070
|
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, Russia. It
|
Allosaurus
|
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] |
6071
|
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 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 Dan Chure, and Mickey Mortimer believes that the synonymy of "Apatodon" with "Allosaurus" was due to
|
Allosaurus
|
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] |
6072
|
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 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", and is based on what is now AMNH 5767, parts of three vertebrae, a coracoid, and a metatarsal. Following Paul's work, this species has been accepted as a synonym of
|
Allosaurus
|
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] |
6073
|
"A. fragilis". A 2010 study by Paul and Kenneth Carpenter, however, indicates that "Epanterias" is temporally younger than the "A. fragilis" type specimen, so is a separate species at minimum. "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
|
Allosaurus
|
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] |
6074
|
separate genus, but did warrant its own species, "A. maximus". This reassignment was rejected in the most recent review of basal tetanurans. 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, Although Donald F. Glut listed it as a species of "Allosaurus", it is now considered a dubious ceratosaurian related
|
Allosaurus
|
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] |
6075
|
to "Ceratosaurus". "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 for a partial shin (MB.R.3620) found in the Kimmeridgian-age Tendaguru Formation in Mtwara, 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 10 meters long (33 ft) and 2.5 metric tons (2.8 short tons) in
|
Allosaurus
|
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] |
6076
|
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). Also, reports of "Allosaurus" in Shanxi, China go back to at least 1982. An astragalus (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
|
Allosaurus
|
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] |
6077
|
thought it more resembled that of an ornithomimid, 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, in their description of "Fukuiraptor", 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 may have belonged to something similar to, or the same as, "Australovenator", or it may represent an abelisaur. "Allosaurus" was an
|
Allosaurus
|
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] |
6078
|
allosaurid, a member of a family of large theropods within the larger group Carnosauria. The family name Allosauridae was created for this genus in 1878 by Othniel Charles Marsh, but the term was largely unused until the 1970s in favor of Megalosauridae, another family of large theropods that eventually became a wastebasket taxon. 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,
|
Allosaurus
|
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] |
6079
|
von Huene, 1926, Romer, 1956 and 1966, Steel, 1970, and Walker, 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", "Chilantaisaurus", "Compsosuchus", "Stokesosaurus", and "Szechuanosaurus". Given modern knowledge of theropod diversity and the advent of cladistic study of evolutionary relationships, none of these theropods is now recognized as an allosaurid, although several, like
|
Allosaurus
|
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] |
6080
|
"Acrocanthosaurus" and "Yangchuanosaurus", are members of closely related families. Below is a cladogram by Benson "et al." in 2010. Allosauridae is one of four families in Carnosauria; the other three are Neovenatoridae, Carcharodontosauridae and Sinraptoridae. Allosauridae has at times been proposed as ancestral to the Tyrannosauridae (which would make it paraphyletic), one recent 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. Allosauridae is the smallest of the carnosaur families, with only "Saurophaganax" and a currently unnamed French allosauroid accepted as
|
Allosaurus
|
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] |
6081
|
possible valid genera 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". Recent reviews have kept the genus "Saurophaganax" and included "Epanterias" with "Allosaurus". 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 eggs—crushed eggs from Colorado have been suggested as those of "Allosaurus". Based on histological analysis of limb bones, bone deposition
|
Allosaurus
|
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] |
6082
|
appears to stop at around 22 to 28 years, which is comparable to that of other large theropods like "Tyrannosaurus". 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
|
Allosaurus
|
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] |
6083
|
to supply calcium 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 gender 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
|
Allosaurus
|
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] |
6084
|
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 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
|
Allosaurus
|
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] |
6085
|
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. 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, and a "Stegosaurus" neck plate with a U-shaped wound that correlates well with an "Allosaurus" snout. Sauropods seem to be likely candidates as both live prey and as objects of scavenging, based on the presence of
|
Allosaurus
|
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] |
6086
|
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 first decade of the 21st century may have found other solutions to this question. Robert T. Bakker, comparing "Allosaurus" to Cenozoic sabre-toothed carnivorous
|
Allosaurus
|
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] |
6087
|
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 sabre 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-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
|
Allosaurus
|
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] |
6088
|
using finite element analysis 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
|
Allosaurus
|
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] |
6089
|
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 analogues 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
|
Allosaurus
|
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] |
6090
|
such an attack, and that articulations in the skull cited by their detractors as problematic actually helped protect the palate 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
|
Allosaurus
|
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] |
6091
|
dinosaurian prey, and that allosaurs 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
|
Allosaurus
|
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] |
6092
|
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". 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
|
Allosaurus
|
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] |
6093
|
are consistent with the animal's capabilities. They also found that the animal probably processed carcasses by vertical movements in a similar manner to falcons, such as kestrels: 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.
|
Allosaurus
|
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] |
6094
|
The shape of the skull of "Allosaurus" limited potential binocular vision to 20° of width, slightly less than that of modern crocodilians. 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 30 to 55 kilometers per hour (19 to 34 miles per hour). A new paper on
|
Allosaurus
|
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] |
6095
|
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. 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 has extended social behavior to parental care, and has interpreted shed allosaur teeth and chewed bones of
|
Allosaurus
|
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0.042861297726631165
] |
6096
|
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
|
Allosaurus
|
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] |
6097
|
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 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
|
Allosaurus
|
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] |
6098
|
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 dragons. The same interpretation applies to Bakker's lair sites.
|
Allosaurus
|
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] |
6099
|
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. The brain of "Allosaurus", as interpreted from spiral CT scanning of an endocast, was more consistent with crocodilian brains than those of the other living archosaurs, birds. The structure of the vestibular apparatus indicates that the skull was held nearly horizontal, as opposed to strongly tipped up or down. The structure of the inner ear was like that of a crocodilian, and so "Allosaurus" probably could
|
Allosaurus
|
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] |
6100
|
have heard lower frequencies best, and would have had trouble with subtle sounds. The olfactory bulbs were large and seem to have been well suited for detecting odors, although the area for evaluating smells was relatively small. In 2001, Bruce Rothschild and others published a study examining evidence for stress fractures and tendon avulsions in theropod dinosaurs and the implications for their behavior. Since stress fractures are caused by repeated trauma rather than singular events they are more likely to be caused by the behavior of the animal than other kinds of injury. Stress fractures and tendon avulsions occurring in
|
Allosaurus
|
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0.22110721468925476,
-0.03977452591061592,
-0.48162841796875,
-0.1142064779996872,
-0.03292124718427658,
0.09931060671806335,
0.24924050271511078,
0.03370467573404312
] |
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