Allosaurus
Allosaurus is a genus of large 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”. It is derived from the Greek ἄλλος/allos(“different, other”) and σαῦρος/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. These remains became known as Antrodemus. As one of the first well-known theropod dinosaurs, it has long attracted attention outside of paleontological circles. Indeed, it 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 8.5 m (28 ft) in length, though fragmentary remains suggest it could have reached over 12 m (39 ft). 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 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 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, 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. It may have attacked large prey by ambush, using its upper jaw 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 m (28 ft), with the largest definitive Allosaurus specimen (AMNH 680) estimated at 9.7 meters (32 feet) long, and an estimated weight of 2.3 metric tons (2.5 short tons). In his 1976 monograph on Allosaurus, James Madsen mentioned a range of bone sizes which he interpreted to show a maximum length of 12 to 13 m (39 to 43 ft). As with dinosaurs in general, weight estimates are debatable, and since 1980 have ranged between 1,500 kilograms (3,300 pounds), 1,000 to 4,000 kg (2,200 to 8,800 lb), and 1,010 kilograms (2,230 pounds) for modal adult weight (not maximum). John Foster, a specialist on the Morrison Formation, suggests that 1,000 kg (2,200 lb) is reasonable for large adults of A. fragilis, but that 700 kg (1,500 lb) is a closer estimate for 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 1,500 kilograms (3,300 lb) for the individual, but by varying parameters they found a range from approximately 1,400 kilograms (3,100 lb) to approximately 2,000 kilograms (4,400 lb).
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 m (36 ft) 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 assigned to the genus Epanterias (AMNH 5767), may have measured 12.1 meters (40 feet) 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.
Skull
The skull and teeth of Allosaurus were modestly proportioned for a theropod of its size. Paleontologist Gregory S. Paul gives a length of 845 mm (33.3 in) for a skull belonging to an individual he estimates at 7.9 m (26 ft) 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 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 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 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.
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, 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 Acrocanthosaurus and Yangchuanosaurus, are members of closely related families.
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 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.
Species and taxonomy
There are currently four valid and one undescribed species of Allosaurus (A. amplus, A. europaeus, the type species A. fragilis, the as-yet not formally described “A. jimmadseni”, and A. lucasi).
A. fragilis, “A. jimmadseni”, A. amplus, and A. lucasi are all known from remains discovered 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 Quarry, Como Bluff, and Dry Mesa Quarry showed that the differences seen in the Morrison Formation material could be attributed to individual variation. A study of skull elements from the Cleveland-Lloyd site found wide variation between individuals, calling into question previous species-level distinctions based on such features as the shape of the lacrimal horns, and the proposed differentiation of “A. jimmadseni” based on the shape of the jugal. A. europaeus was found in the Kimmeridgian-age Porto Novo Member of the Lourinhã Formation, but may be the same as A. fragilis.
Allosaurus tendagurensis was found in Kimmeridgian-age rocks of Tendaguru, in Mtwara, Tanzania. Subsequent studies classified it as a non-coelurosaurian tetanuran, either a megalosaurid or carcharodontosaur. Although obscure, it was a large theropod, possibly around 10 meters (33 feet) long and 2.5 metric tons (2.8 short tons) in weight.
