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Species of Allosaurus

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Allosaurus was a variable genus, one such area of variability being size. How much variation was due to different species, and how much was due to other factors like age and sex, is debated.
Allosaurus size comparison image including Saurophaganax(A. maximus?)

There have been a number of potential species assigned to the carnosaurian dinosaur genus Allosaurus since its description in 1877 by Othniel Charles Marsh, but only a handful are still regarded as valid. Allosaurus was originally described from material from the Upper Jurassic Morrison Formation of the western United States of America; the type species A. fragilis became one of the best-known species of dinosaur.

The genus Allosaurus was part of the Marsh/Cope "Bone Wars" of the late 19th century, and its taxonomy became increasingly confused due to the competition, with several genera and species named by Cope and Marsh now regarded as synonyms of Allosaurus or A. fragilis. Since the description of Allosaurus, scientists have proposed additional species from such far-flung locales as Portugal, Siberia, and Tanzania, but only the one non-US species of Allosaurus is currently considered valid.

Query about type specimen[edit]

Mounted A. fragilis skeleton (USNM4734), which may become the new type specimen

The issue of synonyms is complicated by the type specimen of Allosaurus fragillis (catalogue number YPM 1930) being extremely fragmentary, consisting of a few incomplete vertebrae, limb bone fragments, rib fragments, and a tooth. Because of this, several scientists have noted that the type specimen, and thus the genus Allosaurus itself or at least the species A. fragillis, is technically a nomen dubium ("dubious name", based on a specimen too incomplete to compare to other specimens or to classify). In an attempt to fix this situation, Gregory S. Paul and Kenneth Carpenter (2010) submitted a petition to the ICZN to have the name A. fragillis officially transferred to the more complete specimen USNM4734 (as a neotype). This request is currently pending review.[1]

Potentially valid species[edit]

Saurophaganax, sometimes assigned to Allosaurus as A. maximus
Holotype material of Creosaurus atrox, more recently known as Allosaurus atrox

Eight species have been considered potentially valid since 1988 (Epanterias/A. amplexus,[2] A. atrox,[2] A. europaeus,[3] the type species A. fragilis,[4] the as-yet not formally described "A. jimmadseni",[5] A. lucasi,[6] Saurophaganax/A. maximus,[7] and A. amplus[8]).

A. fragilis is the type species and was named by Marsh in 1877.[9] It is known from the remains of at least sixty individuals, all found in the KimmeridgianTithonian Upper Jurassic-age Morrison Formation of the United States, spread across the states of Colorado, Montana, New Mexico, Oklahoma, South Dakota, Utah, and Wyoming.[4] Details of the humerus (upper arm) of A. fragilis have been used as diagnostic among Morrison theropods,[10] but the discovery of "A. jimmadseni" indicates that this will no longer be the case at the species level.[5]

A. amplexus was named by Gregory S. Paul for giant Morrison allosaur remains, and included in his conception Saurophagus maximus (later Saurophaganax).[2] A. amplexus was originally coined by Cope in 1878 as the type species of his new genus Epanterias,[11] and is based on what is now AMNH 5767, parts of three vertebrae, a coracoid, and a metatarsal.[12] Following Paul's work, this species has been accepted as a synonym of A. fragilis.[4] An 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. [13]

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. This specimen was assigned to A. fragilis,[14] 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.[3] However, more material may show it to be A. fragilis, as originally described.[15]

Skull of A. "jimmadseni" (DINO 11541) from Dinosaur National Monument, when it was still partially encased in matrix

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.[5][16] 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.[17]

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.[18] 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.[7] This reassignment was rejected in the most recent review of basal tetanurans.[4]

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.[2] 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 foot bones.[19] Although the idea of two common Morrison allosaur species was followed in some semi-technical and popular works,[20] 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.[5]

Biological variation[edit]

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.[21] A later study by Smith incorporating Garden Park (Colorado) and Dinosaur National Monument (Utah) specimens found no justification for multiple species based on skeletal variation; skull variation was most common and was gradational, suggesting individual variation was responsible.[7] 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.[22] Kenneth Carpenter, using skull elements from the Cleveland-Lloyd site, found wide variation between individuals, calling into question previous species-level distinctions based such features as the shape of the horns, and the proposed differentiation of "A. jimmadseni" based on the shape of the jugal.[23]

Invalid and synonymous species[edit]

The holotype dentary of Labrosaurus ferox, which may have been injured by the bite of another A. fragilis

A number of species assigned to Allosaurus are no longer recognized as valid, for one reason or another. A. lucaris, another Marsh name, was given to a partial skeleton in 1878.[19] He later decided it warranted its own genus, Labrosaurus,[24] but this has not been accepted, and A. lucaris is also regarded as another specimen of A. fragilis.[4] Allosaurus lucaris, is known mostly from vertebrae, sharing characters with Allosaurus.[25] 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.[1]

Antrodemus valens holotype tail vertebra (above) compared to same of Allosaurus (below)

Several species coined in genera other than Allosaurus are also now thought to be synonymous with A. fragilis. Labrosaurus ferox was 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.[26] Later researchers suggested that the bone was pathologic, showing an injury to the living animal,[27] and that part of the unusual form of the rear of the bone was due to plaster reconstruction.[28] It is recognized as most likely a specimen of A. fragilis.[4][28]

Allosaurus valens is a new combination for Antrodemus valens used by Friedrich von Huene in 1932;[5] Antrodemus valens itself may also pertain to Allosaurus fragilis,[4] as Gilmore suggested in 1920.[27]

Misassigned species[edit]

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,[29] although most of the remains were removed by Richard Swann Lull to the new ornithopod species Dryosaurus grandis, except for a tooth.[30] Gilmore considered the tooth nondiagnostic but transferred it to Dryptosaurus, as D. medius.[27] The referral was not accepted in the most recent review, and Allosaurus medius was simply listed as a dubious species of theropod.[4] It may be closely related to Acrocanthosaurus.[31]

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.[32] It was transferred to Chilantaisaurus in 1990,[33] but is now considered a nomen dubium indeterminate beyond Theropoda.[34]

Allosaurus meriani was a new combination by George Olshevsky for Megalosaurus meriani Greppin, 1870, based on a tooth from the Late Jurassic of Switzerland.[35][36] However, a recent overview of Ceratosaurus included it in Ceratosaurus sp.[28]

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.[37][38] However, it was considered indeterminate beyond Dinosauria by Dan Chure,[5] and Mickey Mortimer believes that the synonymy of Apatodon with Allosaurus was due to 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.[39]

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.[30] It is now regarded as a dubious theropod.[4] Labrosaurus stechowi, described in 1920 by Janensch based on isolated Ceratosaurus-like teeth from the Tendaguru beds of Tanzania,[40] Although Donald F. Glut listed it as a species of Allosaurus,[38] it is now considered a dubious ceratosaurian related to Ceratosaurus.[28][41] L. sulcatus, named by Marsh in 1896 for a Morrison theropod tooth,[42] which like L. stechowi is now regarded as a dubious Ceratosaurus-like ceratosaur.[28][41]

A. tendagurensis tibia, Naturkunde Museum Berlin

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.[43] Although tabulated as a tentatively valid species of Allosaurus in the second edition of the Dinosauria,[4] subsequent studies place it as indeterminate beyond Tetanurae, either a carcharodontosaurian or megalosaurid.[44][45] Although obscure, it was a large theropod, possibly around 10 meters long (33 ft) and 2.5 metric tons (2.8 short tons) in weight.[46]

Typographical errors and nomina nuda[edit]

"Allosaurus agilis", seen in Zittel, 1887, and Osborn, 1912, is a typographical error for A. fragilis.[5] "Allosaurus ferox" is a typographical error by Marsh for A. fragilis in a figure caption for the partial skull YPM 1893,[42] and YPM 1893 has been treated as a specimen of A fragilis.[4] Likewise, "Labrosaurus fragilis" is a typographical error by Marsh (1896) for Labrosaurus ferox.[28] "A. whitei" is a nomen nudum coined by Pickering in 1996 for the complete Allosaurus specimens that Paul referred to A. atrox.[5]

Specimens misassigned to Allosaurus though not described as new species[edit]

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).[47] Also, reports of Allosaurus in Shanxi, China go back to at least 1982.[48]

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.[49] This identification was challenged by Samuel Welles, who thought it more resembled that of an ornithomimid,[50] but the original authors defended their identification.[51] 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.[52] Similarly, Yoichi Azuma and Phil Currie, in their description of Fukuiraptor, noted that the bone closely resembled that of their new genus.[53] This specimen is sometimes referred to as "Allosaurus robustus", an informal museum name.[16] It may have belonged to something similar to, or the same as, Australovenator,[54] or it may represent an abelisaur.[55] A speculative "polar" or "dwarf allosaur" was used for the "Spirits of the Ice Forest" episode of Walking with Dinosaurs.


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