Allosaurus Totally Explained

Everything about Allosaurus totally explained

Allosaurus is a genus of large theropod dinosaur that lived 155 to 145 million years ago, in the late Jurassic period. The first remains that can definitely be ascribed to this genus were described in 1877 by Othniel Charles Marsh. As one of the first well-known theropod dinosaurs, it has long attracted attention outside of paleontological circles, and has been a lead dinosaur in several films and documentaries. Allosaurus was a large bipedal predator with a large skull, equipped with dozens of large, sharp teeth. It averaged 8.5 meters (30 ft) in length, though fragmentary remains suggest it could have reached over 12 meters (39 ft). Relative to the large and powerful hindlimbs, its three-fingered forelimbs were small, and the body was balanced by a long, heavy 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 is A. fragilis. The bulk of Allosaurus remains have come from North America's Morrison Formation, with material also from Portugal and possibly Tanzania. It was known for over half of the 20th century as Antrodemus, but 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 prominent large predator in the Morrison Formation, Allosaurus was at the top of the food chain, probably preying on contemporaneous large herbivorous dinosaurs. Potential prey included ornithopods, stegosaurids, and sauropods. While it's often thought of as preying on sauropod dinosaurs in groups, there's little evidence for cooperative social behavior in this genus, and individuals may have been aggressive toward each other instead. It may have attacked large prey by ambush, using its upper jaws like a hatchet.

Description

Allosaurus was a typical large theropod, having a massive skull on a short neck, a long tail and reduced forelimbs. Allosaurus fragilis, the best-known species, had an average length of 8.5 meters (28 ft), with the largest definitive Allosaurus specimen (AMNH 680) estimated at 9.7 meters long (32 ft), and an estimated weight of 2.3 metric tons (2.5 short tons). As with dinosaurs in general, weight estimates are debatable, and since 1980 have ranged between 1500 kilograms (3300 lb), 1000 to 4000 kilograms (2200 to 8800 lb), and 1010 kilograms (2230 lb) for modal adult weight (not maximum).
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 10.9 meters (36 ft) in length, Another potential specimen of Allosaurus, once assigned to the genus Epanterias (AMNH 5767), may have measured 12.1 meters in length (40 ft). 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 fourteen and seventeen teeth; the number of teeth doesn't exactly correspond to the size of the bone. Each dentary (the tooth-bearing bone of the lower jaw) had between fourteen and seventeen teeth, with an average count of sixteen. The teeth became shorter, more narrow, 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. 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 braincase was thin, perhaps to improve thermoregulation for the brain. The braincase and frontals may also have had a joint. The number of tail vertebrae is unknown and varied with individual size; James Madsen estimated about 50, 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, A furcula (wishbone) was also present, but has only been recognized since 1996; in some cases furculae were confused with gastralia. and had three fingers per hand, tipped with large, strongly curved and pointed claws. The wrist had a version of the semilunate carpal

Classification

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 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, 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 wasn't 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 Acrocanthosaurus and Yangchuanosaurus, are members of related families. 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 possible valid genera besides Allosaurus in the most recent review. He later decided it deserved its own genus, Antrodemus. Allosaurus itself is based on YPM 1930, a small collection of fragmentary bones including parts of three vertebrae, a rib fragment, a tooth, a toe bone, and, 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 saurus/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 were collected from the Morrison Formation of Garden Park, north of Cañon City. and Cope's Epanterias.
   In their haste, Cope and Marsh didn't 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 1883, M. P. Felch found an almost complete Allosaurus and several partial skeletons. This is the well-known mount poised over a partial Apatosaurus 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 the convoluted situation was produced by Charles W. Gilmore in 1920. He came to the conclusion that the tail vertebra dubbed 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.

Cleveland-Lloyd discoveries

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 J. Stokes in 1945, major operations didn't begin there until 1960. Under a cooperative effort involving nearly 40 institutions, thousands of bones were recovered between 1960 and 1965. 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 1 meter (3.3 ft) to 12 meters (39 ft) long, and the disarticulation is an advantage for describing bones usually found fused. 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.

"Big Al"

One of the more significant Allosaurus finds was the 1991 discovery of "Big Al" (MOR 693), a 95% complete, partially articulated specimen that 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 Swiss team, led by Kirby Siber. The same team later excavated a second Allosaurus, "Big Al Two", which is the best preserved skeleton of its kind to date. The specimen was described by Breithaupt in 1996. recent work has followed a "one species" interpretation, A. europaeus was found in the Kimmeridgian-age Porto Novo Member of the Lourinhã Formation, but may be the same as A. fragilis. A. tendagurensis was found in Kimmeridgian-age rocks of Tendaguru, in Mtwara, Tanzania. Although the most recent review tentatively accepted it as a valid species of Allosaurus, it may be a more basal tetanuran, or simply a dubious theropod. Later researchers suggested that the bone was pathologic, showing an injury to the living animal, It is now regarded as an example of A. fragilis. Siberia, and Switzerland, The Morrison Formation is interpreted as a semiarid environment with distinct wet and dry seasons, and flat floodplains. Vegetation varied from river-lining forests of conifers, tree ferns, and ferns, to fern savannas with rare trees.
The Morrison Formation has been a rich fossil hunting ground, holding fossils of green alage, fungi, mosses, horsetails, ferns, cycads, ginkgoes, and several families of conifers. Other fossils discovered include bivalves, snails, ray-finned fishes, frogs, salamanders, turtles, sphenodonts, lizards, terrestrial and aquatic crocodylomorphans, several species of pterosaur, numerous dinosaur species, and early mammals such as docodonts, multituberculates, symmetrodonts, and triconodonts. Such dinosaurs as the theropods Ceratosaurus, Ornitholestes, and Torvosaurus, the sauropods Apatosaurus, Brachiosaurus, Camarasaurus, and Diplodocus, and the ornithischians Camptosaurus, Dryosaurus, and Stegosaurus are known from the Morrison. The Late Jurassic formations of Portugal where Allosaurus is present are interpreted as having been similar to the Morrison but with a stronger marine influence. Many of the dinosaurs of the Morrison Formation are the same genera as those seen in Portuguese rocks (mainly Allosaurus, Ceratosaurus, Torvosaurus, and Apatosaurus), or have a close counterpart (Brachiosaurus and Lusotitan, Camptosaurus and Draconyx). Allosaurus coexisted with fellow large theropods Ceratosaurus and Torvosaurus in both the United States and Portugal. Ceratosaurus, better known than Torvosaurus, differed noticeably from Allosaurus in functional terms by having a taller, narrower skull with large, broad teeth.

Paleobiology

Life history

The wealth of Allosaurus fossils, from nearly all ages of individuals, allows scientists to study how the animal grew and how long its lifespan may have been. Remains may reach as far back in the lifespan as eggs—crushed eggs from Colorado have been suggested as those of Allosaurus.
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.

Feeding

Paleontologists accept Allosaurus as an active predator of large animals. Sauropods seem to be likely candidates as both live prey and as objects of scavenging, based on the presence of scrapings on sauropod bones fitting allosaur teeth well and the presence of shed allosaur teeth with sauropod bones. 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. 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. The original authors noted that Allosaurus itself has no modern equivalent, that the tooth row is well-suited to such an attack, and that articulations in the skull cited by their detractors as problematic actually helped protect the palate 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 wouldn't 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. 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.

Social behavior

Allosaurus has long been regarded in the semitechnical and popular literature as an animal that preyed on sauropods and other large dinosaurs by hunting in groups.
Recent research suggests that Allosaurus and other theropods were like other diapsids and had largely aggressive instead of cooperative interactions with other members of their own species. One study noted 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 diapsids (including lizards, crocodiles, and birds) very rarely cooperate to hunt in such a way. Many modern diapsid predators are 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 them when aggregated at feeding sites. This suggests that, for example, the Cleveland-Lloyd quarry shows Allosaurus individuals were drawn together to feed on other disabled or dead allosaurs, and sometimes were killed in the process, thus accumulating. 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. 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. Allosaurus has been depicted in popular culture since the early years of the 20th century. It is top predator in both Arthur Conan Doyle's 1912 novel, The Lost World, and the 1925 film adaptation, the first full-length motion picture to feature dinosaurs (it isn't to be confused with Tyrannosaurus, which also appears in the film). It later became the starring dinosaur of the 1956 film The Beast of Hollow Mountain, and the 1969 film The Valley of Gwangi, two genre combinations of living dinosaurs with Westerns. In The Valley of Gwangi, Gwangi is billed as an Allosaurus, although Ray Harryhausen based his model for the creature on Charles R. Knight's depiction of a Tyrannosaurus. Harryhausen sometimes confuses the two, stating in a DVD interview "They're both meat eaters, they're both tyrants... one was just a bit larger than the other." In nonfictional presentations, Allosaurus appears in the second and fifth episodes of the BBC television series Walking with Dinosaurs, and the Walking with Dinosaurs special The Ballad of Big Al chronicles the life of the Allosaurus specimen nicknamed "Big Al".

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