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Allosaurus

Genus of theropod dinosaur

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Allosaurus (Big Al II) (specimen SMA 0005)
The_Wookies · CC BY 2.5

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Through Taxonomy (biology), Species and Social behavior

Allosaurus was a large bipedal predator for its time. Its skull was light, robust, and equipped with dozens of sharp, serrated teeth. It averaged 7.5–8 meters (25–26 ft) in length for A. fragilis, with the largest specimens of A. fragilis estimated as being 9 meters (30 ft) long. Relative to the large and powerful legs, its three-fingered hands were small and the body was balanced by a long, muscular tail. It is classified in the family Allosauridae. As the most abundant large predator of the Morrison Formation, Allosaurus was an apex predator of the food chain and probably preyed on large herbivorous dinosaurs such as ornithopods, stegosaurids, and sauropods. Scientists have debated whether Allosaurus had cooperative social behavior and hunted in packs or was a solitary predator that forms congregations, with evidence supporting both arguments.

History of discovery

Initial finds and naming of Allosaurus

Allosaurus was discovered during the Bone Wars, a feud between two American paleontologists, Othniel Charles Marsh and Edward Drinker Cope, that led to a surge of fossil discoveries in the Western US. The first described fossil in the taxonomic history of Allosaurus was a bone obtained secondhand by Ferdinand V. 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 [sic] valens. He later decided it deserved its own genus, Antrodemus.

Allosaurus itself is based on YPM 1930, a small collection of fragmentary bones including parts of three vertebrae, a rib fragment, a tooth, a toe bone, and the shaft of the right humerus (upper arm bone). Marsh gave these remains the name Allosaurus fragilis in 1877. Allosaurus comes from the Greek words allos/αλλος, meaning "strange" or "different", and sauros/σαυρος, meaning "lizard" or "reptile". Marsh chose the name 'different lizard' because he believed that the vertebrae were different from those of other dinosaurs due to their lightweight construction. The species epithet fragilis is Latin for "fragile", again referring to the lightening features in the vertebrae. The bones were uncovered by two of Marsh's collectors, Benjamin Mudge and Samuel W. Williston, in the autumn of 1877 at Felch Quarry, in the Garden Park area of Colorado. Marsh and his collectors were unsatisfied with the quality of the collected fossils, so he ordered to close the quarry that same autumn. Yet, Marsh named two new dinosaurs from these remains: Diplodocus and Allosaurus. In 1883, Marsh hired the original discoverer of the quarry, Marshall P. Felch, to continue excavations. Felch's subsequent discoveries made the quarry one of the prime sites of the Morrison, and included the holotype specimens of Ceratosaurus nasicornis, Stegosaurus stenops, and a mostly complete Allosaurus skeleton (USNM 4734) that would later be selected as the neotype specimen of Allosaurus fragilis (the single specimen the species is based on, replacing the inadequate holotype).

In 1879, one of Cope's collectors, H. F. Hubbell, found a specimen in the Como Bluff area, but apparently did not mention its completeness and Cope never unpacked it. Upon unpacking it in 1903 (several years after Cope had died), it was found to be one of the most complete theropod specimens then known and the skeleton, now cataloged as AMNH 5753, was put on public view in 1908. This is the well-known mount poised over a partial Apatosaurus skeleton as if scavenging it, illustrated as such in a painting by Charles R. Knight. Although notable as the first free-standing mount of a theropod dinosaur and often illustrated and photographed, it has never been scientifically described.

Renaming to Antrodemus and early discoveries at Dinosaur National Monument

The many names coined by Cope and Marsh complicated later research, with the situation further compounded by the terse descriptions they provided. Even at the time, authors such as Samuel W. Williston suggested that too many names had been coined. For example, Williston pointed out in 1901 that Marsh had never been able to adequately distinguish Allosaurus from Creosaurus. The most influential early attempt to sort out the situation was produced by Charles W. Gilmore in 1920. He came to the conclusion that the tail vertebra named Antrodemus by Leidy was indistinguishable from those of Allosaurus and that Antrodemus should be the preferred name because, as the older name, it had priority. Antrodemus became the accepted name for this familiar genus for over 50 years, until James H. Madsen published on the Cleveland-Lloyd specimens and concluded that Allosaurus should be used because Antrodemus was based on material with poor, if any, diagnostic features and locality information. For example, the geological formation that the single bone of Antrodemus came from is unknown.

In 1909, Earl Douglass from the Carnegie Museum discovered what should later become Dinosaur National Monument in Utah. Until 2022, Douglass and his team excavated over 700,000 lb (320,000 kg) of fossils of multiple dinosaur species from a single quarry, including several Allosaurus specimens. Among these finds is CM 11844, which was collected between 1913 and 1915 and comprises much of the skeleton and a fragmentary skull. Since 1938, this skeleton is on display at the Carnegie Museum. During the summer of 1924, the University of Utah uncovered DINO 2560, the best preserved Allosaurus specimen known at that time. The skull of this particularly large individual is on exhibit at the Dinosaur National Monument.

Cleveland-Lloyd discoveries and "Big Al"

Although sporadic work at what became known as Utah's Cleveland-Lloyd Dinosaur Quarry had taken place as early as 1927 and the fossil site itself was described by William L. Stokes in 1945, major operations did not begin there until 1960. Madsen led a cooperative effort between 1960 and 1965 involving nearly 40 institutions, during which thousands of bones were recovered from the site. The quarry is notable for the predominance of Allosaurus remains: the quarry preserves a minimum of 73 individual dinosaurs and at least 46 of those are A. fragilis. The great quantity of well-preserved Allosaurus remains has allowed this genus to be known in great detail, making it among the best-known of all theropods. Skeletal remains from the quarry pertain to individuals of almost all ages and sizes, from less than 1 metre (3.3 feet) to 12 metres (39 feet) long. Because its fossils are common at both this quarry and others in the state, Allosaurus was designated as the state fossil of Utah in 1988.

In the early 1990s, a Swiss team led by Kirby Siber set out for commercial fossil excavations at Howe Ranch Quarry, Wyoming. This quarry had originally been worked on in 1934 by Barnum Brown and his crew, who collected more than 30 tons of bones, mostly of sauropods. Because the Swiss team could not locate additional specimens in the quarry, they explored the surrounding area, where they discovered "Big Al" (MOR 693) in 1991: a 95% complete, partially articulated Allosaurus specimen. However, because the new site was located on public land, the excavation was taken over by a joint Museum of the Rockies and University of Wyoming Geological Museum team. The specimen, now on exhibit at the Museum of the Rockies, belonged to an individual of about 8 m (26 ft) in length. This was below the average size for Allosaurus, as it was a subadult estimated at only 87% grown. The Swiss team later excavated a second Allosaurus, "Big Al II" (SMA 0005), on private land on Howe Ranch, which is exhibited at the Aathal Dinosaur Museum in Switzerland.

In 1991, Brooks Britt argued that there were at least two species of Allosaurus: A robust species with a short and high skull and pointed lacrimal horns, and a more gracile species with a long and low skull and rounded lacrimal horns. The robust species is geologically younger from localities such as Dry Mesa Quarry and Garden Park, while the gracile species, found at the Cleveland-Lloyd and at Dinosaur National Monument, is older. Already in 1988, Gregory S. Paul made a similar distinction in a popular book, in which he referred to the gracile species as A. fragilis and to the robust species as A. atrox, using a species originally described by Marsh as Creosaurus atrox. However, a series of statistical analyses by David K. Smith between 1996 and 1999 suggested that the differences seen in the Morrison Formation material can be attributed to individual variation.

Portuguese and other European discoveries

Allosaurus is known from at least three localities in Portugal, from rocks of the Lourinhã, Bombarral, and Alcobaça formations. The first specimen (MNHNUL/AND.001), a partial skeleton that includes an articulated hind limb and pelvis, was found in 1988 near the village of Andrés in the District of Leiria during the construction of a warehouse. Reported in 1999 and assigned to the species A. fragilis, it was the first Allosaurus specimen to be found outside of North America. A. fragilis became the first dinosaur species known from both Europe and North America, suggesting faunal exchange between the two continents. This site has been worked on again between 2005 and 2010, resulting in the discovery of at least two more Allosaurus individuals, including many skull bones. In 2005, a single maxilla was reported from the Guimarota coal mine, a locality well-known for its fossils of Mesozoic mammals; this fossil was assigned to Allosaurus but not to any particular species.

In 2006, Octávio Mateus and colleagues reported a partial skull and three neck vertebrae (ML 415) from the Vale Frades beach in Lourinhã. As this specimen differed from North American Allosaurus fossils, they assigned it to a new species, A. europaeus. These authors also assigned the Andrés specimen to this species, though solely based on the fact that it was found in Portugal. The presence of a separate Allosaurus species in Europe would suggest that faunal interchange between the continents was interrupted, possibly due to the opening of the Atlantic Ocean. The status of A. europaeus was controversially discussed in the subsequent years, with different studies arguing that the species is a synonym of A. fragilis, a nomen dubium (doubtful name), or in need of re-evaluation. In 2024, André Burigo and Mateus re-described the Vale Frades specimen and carried out further fossil preparation that exposed additional bones. These authors identified nine unique features supporting the validity A. europaeus. A 2025 study by Elisabete Malafaia and colleagues described the Andrés specimens in detail, including a newly discovered set of skull bones. Their analysis of the relationships between individual Allosaurus skulls instead suggested that A. europaeus is a synonym of A. fragilis because the Andrés specimens were more closely related to some North American specimens than to the Vale Frades specimen.

A. jimmadseni and A. anax

On July 15, 1990, George Engelmann discovered toe bones and some tail vertebrae weathering out of the rock while conducting an inventory of fossils at Dinosaur National Monument. Staff of the National Monument began excavating the new specimen later that year; the conditions were difficult because of its location in a steeply inclined rock face that, with ongoing excavation, became vertical. In 1994, much of the excavated skeleton was flown out via helicopter in single block weighing 2,700 kg (6,000 lb). The skull, which was still missing, was only located two years later, in 1996, with the help of novel radiological surveying techniques that detect gamma radiation from radioactive minerals which accumulate in bones during fossilization. The specimen, DINO 11541, is one of the most complete theropod skeletons recovered from the Late Jurassic. In 2000, Daniel Chure described the specimen in his PhD thesis, arguing that it represents a new species, A. jimmadseni. However, as the thesis did not meet the requirements of the International Commission on Zoological Nomenclature (ICZN), it remained an invalid nomen nudum ("naked name") until Chure and Mark Loewen formally described the species in 2020. These authors assigned several other specimens to A. jimmadseni, including the original "Big Al" individual (MOR 693). The name jimmadseni honors Madsen for his contributions to the taxonomy of the genus, notably his 1976 work.

The issue of species and potential synonyms was historically complicated by the type specimen of Allosaurus fragilis (YPM 1930) being extremely fragmentary. Because of this, several scientists have interpreted the type specimen as potentially dubious, meaning the genus Allosaurus itself or at least the species A. fragilis would be a nomen dubium ("dubious name", based on a specimen too incomplete to compare to other specimens or to classify). To address this situation, Gregory S. Paul and Kenneth Carpenter submitted a petition to the ICZN in 2010 to have the more complete specimen USNM 4734 selected as a neotype, a decision that was ratified by the ICZN on December 29, 2023. In 2014, Sebastian Dalman named the new species Allosaurus lucasi based on two specimens from the Tithonian of Colorado, but this species was not accepted by later authors.

In 1995, Chure created the taxon Saurophaganax maximus for giant allosaurid remains from western Oklahoma. These remains had been known as Saurophagus, but that name was already in use, leading Chure to propose a substitute. Smith, in his 1998 analysis of variation, concluded that S. maximus was not different enough from Allosaurus to be a separate genus, but did warrant its own species, A. maximus. This reassignment was rejected in a review of basal tetanurans. A 2024 reassessment by Andy Danison and colleagues suggested that the Saurophaganax specimen is a chimera that combines the bones of Allosaurus with those of a sauropod. The holotype of Saurophaganax itself, a neural arch, cannot be confidently be assigned to a theropod, making it a nomen dubium. The Allosaurus bones, however, were found to be distinct enough to warrant a new species of Allosaurus, A. anax. The name anax is Greek for 'king', and also alludes to the name change from Saurophagus to Saurophaganax.

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