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Inside the lacrimal bones were depressions that may have held glands , such as salt glands. The roof of the braincase was thin, perhaps to improve thermoregulation for the brain.
Allosaurus had nine vertebrae in the neck, 14 in the back, and five in the sacrum supporting the hips. Paul considered that to be too many and suggested 45 or less.
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. Of the three fingers, the innermost or thumb was the largest,  and diverged from the others.
The discovery and early study of Allosaurus is complicated by the 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 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 [ sic ] valens.
Allosaurus itself is based on YPM , a 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 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 resumed at Garden Park in , M.
Felch found an almost complete Allosaurus and several partial skeletons. Hubbell, found a specimen in the Como Bluff area of Wyoming in , but apparently did not mention its completeness, and Cope never unpacked it.
Upon unpacking in several years after Cope had died , it was found to be one of the most complete theropod specimens then known, and in the skeleton, now cataloged as AMNH , was put on public view.
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. 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.
Although sporadic work at what became known as the Cleveland-Lloyd Dinosaur Quarry in Emery County , Utah had taken place as early as , and the fossil site itself described by William L.
Stokes in ,  major operations did not begin there until 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 mortality around a waterhole, to getting trapped in a spring-fed pond or seep.
Skeletal remains from the quarry pertain to individuals of almost all ages and sizes, from less than 1 meter 3.
Such studies have covered topics including skeletal variation,  growth,   skull construction,  hunting methods,  the brain ,  and the possibility of gregarious living and parental care.
In the same team discovered a second Allosaurus , "Big Al Two", which is the best preserved skeleton of its kind to date. Pathologic bones included five ribs, five vertebrae, and four bones 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.
The infection was long lived, perhaps up to 6 months. There are currently four valid and one undescribed species of Allosaurus A.
This specimen was assigned to A. It differs from other species of Allosaurus in cranial details. 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.
To address this situation, Gregory S. This request is currently pending review. Creosaurus , Epanterias , and Labrosaurus are regarded as junior synonyms of Allosaurus.
One exception is Labrosaurus ferox , named in 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.
Several species initially classified within or referred to Allosaurus do not belong within the genus. Allosaurus valens is a new combination for Antrodemus valens used by Friedrich von Huene in ;  Antrodemus valens itself may also pertain to Allosaurus fragilis ,  as Gilmore suggested in However, they found that the specimen was undiagnostic, and thus A.
Allosaurus sibiricus was described in by A. Riabinin on the basis of a bone, later identified as a partial fourth metatarsal, from the Early Cretaceous of Buryatia , Russia.
Allosaurus meriani was a new combination by George Olshevsky for Megalosaurus meriani Greppin, , based on a tooth from the Late Jurassic of Switzerland.
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.
Paul for giant Morrison allosaur remains, and included in his conception Saurophagus maximus later Saurophaganax.
These remains had been known as Saurophagus , but that name was already in use, leading Chure to propose a substitute. There are also several species left over from the synonymizations of Creosaurus and Labrosaurus with Allosaurus.
Creosaurus potens was named by Lull in for a vertebra from the Early Cretaceous of Maryland. Glut listed it as a species of Allosaurus ,  it is now considered a dubious ceratosaurian related to Ceratosaurus.
Kurzanov and colleagues in designated six teeth from Siberia as Allosaurus sp. 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. Allosaurus was an allosaurid, a member of a family of large theropods within the larger group Carnosauria.
The family name Allosauridae was created for this genus in by Othniel Charles Marsh ,  but the term was largely unused until the s in favor of Megalosauridae , another family of large theropods that eventually became a wastebasket taxon.
Major publications using the name "Megalosauridae" instead of "Allosauridae" include Gilmore , ,  von Huene , ,  Romer , and ,   Steel, ,  and Walker , 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.
Below is a cladogram by Benson et al. Allosauridae is one of four families in Carnosauria; the other three are Neovenatoridae ,  Carcharodontosauridae and Sinraptoridae.
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.
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.
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.
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.
These changes imply that juvenile legs has less predictable stresses compared with adults, which would have moved with more regular forward progression.
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.
Bakker , comparing Allosaurus to Cenozoic sabre-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 sabre teeth, Bakker suggested another mode of attack that would have used such neck and jaw adaptations: 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 on an Allosaurus skull. According to their biomechanical analysis, the skull was very strong but had a relatively small bite force.
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.
This strategy would also potentially have allowed the prey to recover and be fed upon in a similar way later.
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 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. They also found that the animal probably processed carcasses by vertical movements in a similar manner to falcons , such as kestrels: This differs from the prey-handling envisioned for tyrannosaurids, which probably tore flesh with lateral shakes of the skull, similar to crocodilians.
Other aspects of feeding include the eyes, arms, and legs. As with crocodilians, this may have been enough to judge prey distance and time attacks.
A new 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.
It has been speculated since the s that Allosaurus preyed on sauropods and other large dinosaurs by hunting in groups. Bakker 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.
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 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. 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 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 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 , 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 the forelimb have special behavioral significance since while injuries to the feet could be caused by running or migration , resistant prey items are the most probable source of injuries to the hand.
Allosaurus was one of only two theropods examined in the 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 , Ornithomimus or Archaeornithomimus.
Of the 47 hand bones the researchers studied, 3 were found to contain stress fractures. Of the feet, bones were studied and 17 found to have stress fractures.
The stress fractures in the foot bones "were distributed to the proximal 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.
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 and fibula of an Allosaurus fragilis specimen catalogued as USNM are both pathological, both probably due to healed fractures.
The specimen USNM 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. The specimen MOR had pathologies on five ribs, the sixth neck vertebra the third eighth and thirteenth back vertebrae, the second tail vertebra and its chevron, the gastralia right scapula, manual phalanx I left ilium metatarsals III and V, the first phalanx of the third toe and the third phalanx of the second.
The ilium had "a large hole The near end of the first phalanx of the third toe was afflicted by an involucrum. Other pathologies reported in Allosaurus include: Willow breaks in two ribs.
Healed fractures in the humerus and radius. Distortion of joint surfaces in the foot possibly due to osteoarthritis or developmental issues.
Osteopetrosis along the endosteal surface of a tibia. Coossification of vertebral centra near the end of the tail. The island is in the shape of a reverse tear-drop, with high mountains occupying the northern end and flat plain to the south.
The variations in altitude create an array of micro-climates, with tropical rainforest dominating the lower elevations and temperate rainforest occupying the high elevations to the north.
At the center of the island is a huge artificial lagoon spanning the Tyrannosaur and Maiasaur paddocks which feeds into a river that flows to the north, through the Dilophosaur paddock and ends in an artificial waterfall which masks the entrance to a utility tunnel.
In the novel InGen has built an extensive infrastructure to house visitors and contain 15 different breeds of dinosaurs, centered in the northeast sector of Isla Nublar.
The Visitor Center is located at the base of the mountains in its own compound. There is a separate hotel, or lodge, for visitor accommodation with a landscaped pool area.
The visitor center contains exhibit space for educational purposes, the laboratories, the operations room, a garage where the electric touring cars are housed, and a hatchery where visitors can observe newborn dinosaurs.
John Hammond also possesses a private bungalow in a secluded area near the Visitors Center compound. Two docks - at the east and north ends of the island - manage supply shipments from the mainland.
A theme-park river ride is under construction, as well as an aviary to house the pterosaurs. There is also a mountain-top luxury restaurant still under construction when the events in the novel take place.
Criss-crossing the island are miles of underground concrete maintenance tunnels which connect the paddocks. The novel ends with the Costa Rican airforce dropping a napalm bomb on the island, decimating all the dinosaurs.
Isla Nublar as depicted in Jurassic Park is a large, mountainous island of lush tropical rainforest, miles km west of Costa Rica.
Maps of the island used in the movie, and promotional material for the film, depict it in the same reverse tear drop shape described in the novel.
According to this model, high mountains occupy the northern and western ends of the island, while the Park itself is built in the north-eastern sector surrounded by a mile long perimeter fence.
The Visitor Center and its support buildings are located within a self-contained sector in the center of the island, nestled at the base of the northern mountains and surrounded by an electrified perimeter fence.
The Visitor Center, in the style of a rustic African lodge, features a two-storey rotunda at its center with incomplete exhibit space across both levels.
The fossilized remains of a Tyrannosaurus rex and an Alamosaurus hang from the ceiling as the centerpiece of the room.
There is also a large restaurant with outdoor terrace and gift-shop. The restaurant is served by an industrial kitchen overseen by a gourmet chef.
The Velociraptors are held in a specially built containment unit near to the Visitor Center with a viewing platform and guard tower.
There is also a shelter for emergencies and the maintenance shed where the generator that controls the power grid on the island is located.
In the film the visitors reach the island by helicopter and fly through a verdant mountain valley before landing on the helipad at the base of a waterfall.
As in the novel InGen has constructed an infrastructure of electrified fences, concrete moats and electrified roadway by which the visitor vehicles are guided on their tours of the various dinosaur paddocks.
There are 12 species of dinosaur with their own paddocks, as compared to the 15 in the novel. During the luncheon scene in the Visitor Center, slides in the background show future attractions under construction in the park.
These include a "Jungle River Ride", an aviary, and a separate restaurant facility. Grant and the children must scale, was built in the Waimea Valley on Oahu.
Isla Nublar is not shown in The Lost World: No details are provided as to the fate of Isla Nublar after the events in Jurassic Park , although it is implied that Isla Nublar is abandoned and the dinosaurs on the island have been exterminated.
This is why InGen seeks to recoup some of its losses from Jurassic Park by capturing the surviving dinosaurs on Isla Sorna and bringing them to San Diego.
John Hammond reveals that Isla Nublar was the "showroom" for the tourists, while Isla Sorna, 87 miles further west, was the secret breeding-ground where the vast majority of the dinosaurs were cloned and nurtured on an industrial scale before being moved to Nublar.
The facility in San Diego, an enormous amphitheater, was built before John Hammond decided on relocating the park to Isla Nublar.
Jurassic World contradicts the assertion that Jurassic Park was demolished and the dinosaurs destroyed, due to the T.
Paul and Amanda Kirby had been instructed by the mercenaries they hired to find someone familiar with the island and they erroneously believed Dr.
Grant had been on Isla Sorna before. When informed about this after they are stranded on Isla Sorna, Grant angrily reveals to them he had never been on the island before and Billy Brennan corrects Paul that Grant had been on Isla Nublar and not Isla Sorna.
Jurassic World revives Isla Nublar as the main setting for a Jurassic movie for the first time since the original Jurassic Park.
This time it is the site of a fully operational and wildly successful version of the original park: It is also discovered that the new park was running for ten years, starting in Simon Masrani mentions that he purchased InGen in order to fulfill the dying wish of John Hammond that the park should be operational.
In Jurassic World , the new Visitor Center complex is in the middle of the island, the focal point of a network of chain shops and restaurants with a broad Main Street in the middle.
The Visitor Center itself is a much enlarged version of the original in Jurassic Park , with a multi-story rotunda that features dinosaur holograms , interactive displays and a life-size statue of John Hammond.
The laboratory on view to visitors is called the Hammond Creation Lab. An enormous lagoon where the Mosasaurus lives forms the centerpiece of the visitor complex, around which the hotels, train terminal and shopping village are centered.
There is also an amphitheater overlooking the lagoon for live viewings of Mosasaurus feedings similar to SeaWorld ; the seating rows can be lowered beneath the surface to view the creature behind glass.
Visitors are transported to Isla Nublar on large ferries and shuttled to the Visitors Center via a high-speed light rail that ends in a massive terminal near the Visitor Center.
The operations center sits just beyond the tourist area, within the foothills of the mountains.