{{Short description|Taxonomy of a dinosaur genus}} {{good article}} {{DISPLAYTITLE:Taxonomy of ''Allosaurus''}} {{Use mdy dates|date=April 2026}} [[File:Allosaurus fragilis USNM4734.jpg|thumb|The neotype specimen of ''Allosaurus fragilis'', USNM 4734]] The dinosaur genus ''Allosaurus'' has a complex taxonomic history, including multiple proposed species of which only a few are considered valid today. The genus was first described in 1877 by Othniel Charles Marsh from a very fragmentary specimen found in Colorado. For several decades, ''Allosaurus'' was known under the name ''Antrodemus'', until a 1976 publication re-established the name ''Allosaurus''. In 2023, another, much more complete specimen found in the same quarry was selected as the neotype (the specimen on which the taxon is based, replacing the inadequate original specimen).
Four species are considered potentially valid. Besides the well-known type species ''A. fragilis'', these are ''A. europaeus'', ''A. jimmadseni'', and ''A. anax''. ''A. europaeus'', based on a specimen from Portugal, is the only species found outside of North America. Its validity is contested, and some researchers found it to be a synonym of ''A. fragilis''. ''A. jimmadseni'' was described in 2020 and is known from multiple complete specimens. ''A. anax'' was described in 2024 based on a few bones that were previously assigned to the genus ''Saurophaganax''. Several other species are no longer accepted, and several other genera, such as ''Epanterias'', ''Creosaurus'', and ''Labrosaurus'', have been considered as synonyms of ''Allosaurus''. Fossils from Tanzania, Siberia, and Australia have been previously assigned but are no longer thought to belong to this genus.
==The genus ''Allosaurus''== [[File:Allosaurus holotype gilmore1920.png|thumb|upright=0.7|Casts of the holotype of ''Allosaurus fragilis'', YPM 1930, including a foot bone (1) and parts of two vertebrae (2 and 3)]] ''Allosaurus'' is among the best-known dinosaurs, both scientifically and in public perception.<ref name="paul2010" /> It is the most common theropod in the Morrison Formation, an important fossil deposit in the Western US, accounting for 70 to 75% of theropod specimens.<ref name="maidment2023" /> Additional specimens have been discovered in Portugal.<ref name="malafaia2025" /> Besides the type species, ''A. fragilis'', several additional species have been proposed, three of which are currently in use.<ref name="danison2024" /> Most specimens are not yet assigned to any particular species.<ref name="maidment2023" />
''Allosaurus'' was discovered during the Bone Wars, a feud between the two paleontologists Othniel Charles Marsh and Edward Drinker Cope that led to a surge of fossil discoveries in the Western US.<ref name="carpenter2002" /> Marsh coined the name ''Allosaurus fragilis'' in 1877 for a fragmentary specimen (the holotype specimen; the specimen on which the species was based).<ref name="marsh1877" /> According to Marsh, this specimen includes a lumbar vertebra, later identified as a middle {{Dinogloss|dorsal vertebra}} (back vertebra); a tail vertebra; and "feet bones", later identified as the proximal phalanx (finger bone) of digit III of the right foot.<ref name="marsh1877" /><ref name="madsen1976" /> James Henry Madsen, in his 1976 monograph "''Allosaurus Fragilis'': A Revised Osteology", noted that a tooth and a {{Dinogloss|humerus}} (upper arm bone) fragment are part of the same specimen.<ref name="madsen1976" />{{rp|11}} These fossils are part of the collection of the Peabody Museum of Natural History and have the specimen number YPM 1930. The genus name ''Allosaurus'' comes from the Greek words ''{{lang|grc-Latn|allos}}/{{lang|grc|αλλος}}'', meaning {{gloss|strange}} or {{gloss|different}}, and ''{{lang|grc-Latn|sauros}}/{{lang|grc|σαυρος}}'', meaning {{gloss|lizard}} or {{gloss|reptile}}.<ref name="liddell1980" /> Marsh chose the name {{gloss|different lizard}} because he believed that the vertebrae were different from those of other dinosaurs due to the deep concavities in their sides, giving them a lightweight construction. More complete specimens have later shown that these concavities were in fact internal chambers that were exposed due to damage of the bone.<ref name="creisler2003" /><ref name="marsh1877" /> The bones were collected in 1877 from the Morrison Formation by Marshall Felch in what is now known as Felch Quarry, in the Garden Park area, Colorado.<ref name="carpenter2002" />
{{multiple image |direction = horizontal |align = left |total_width = 300 |image1 = Osteology of the carnivorous Dinosauria in the United States National museum BHL40623194.jpg |alt1 = Page of a 1920 paper showing a skull in top and side views, with parts of the middle section of the skull reconstructed by plaster |image2 = Osteology of the carnivorous Dinosauria in the United States National museum BHL40623195.jpg |alt2 = The next page of the same paper, showing the same skull in the same views, but as labelled interpretive line drawings |footer = Skull of USNM 4734 as figured in Gilmore's 1920 monograph, where it was labelled ''Antrodemus valens''. The skull was reconstructed too short.}}
In a 1920 monograph, Charles W. Gilmore published the first comprehensive description of ''Allosaurus'', mostly based on the much more complete specimen USNM 4734. This specimen includes a skeleton with skull that Felch had discovered in the same quarry in which he already found the holotype. However, Gilmore used the name ''Antrodemus valens'' instead of ''Allosaurus fragilis'', an older name that was based on a single vertebra of uncertain origin, and which he thought had priority. ''Antrodemus'' then became the accepted name for the animal for more than 50 years, until Madsen published his 1976 monograph in which he returned to Marsh's name. Madsen's account was based on immense amounts of material from Cleveland-Lloyd Dinosaur Quarry as well as DINO 2560, a well-preserved skeleton from Dinosaur National Monument.<ref name="madsen1976" /><ref name="gilmore1920" />
===Neotype designation and diagnosis=== [[File:Allosaurus atrox (theropod dinosaur) (Morrison Formation, Upper Jurassic; Carnegie Quarry, Dinosaur National Monument, Utah, USA) 1 (48692087322).jpg|thumb|right|Skull of DINO 2560 at Dinosaur National Monument, which James Madsen suggested as the neotype (specimen that defines the species) of ''Allosaurus fragilis'' in 1976. This was not accepted, and the specimen USNM 4734 was instead designated as the neotype.]] The holotype specimen of ''Allosaurus fragilis'' is too fragmentary to distinguish it from related genera and therefore not diagnostic, and the same is true of the holotype of ''Antrodemus valens'', which consists of a part of a tail vertebra. Therefore, Madsen designated DINO 2560 as the neotype (replacement) specimen in his 1976 monograph, but this was not recognized because such a designation requires a formal ruling by the International Commission on Zoological Nomenclature (ICZN). In 2010, Gregory S. Paul and Kenneth Carpenter argued that the younger age and the proportionally longer skull of this specimen suggest that it might belong to a taxon other than ''A. fragilis''. Paul and Carpenter instead argued that USNM 4734 should become the neotype, and submitted a formal case to the ICZN to have the name ''A. fragilis'' officially transferred to this specimen in order to warrant stability of the genus. They argued that USNM 4734 is the ideal replacement specimen because it is from the same quarry as the holotype and therefore probably of the same species.<ref name="paul2010" /> The proposal has subsequently been supported by other comments and was ratified by the ICZN on December 29, 2023.<ref name="carrano2018" /><ref name="yun2019" /><ref name="iczn2023" />
The exact set of anatomical features used to distinguish ''Allosaurus'' from other genera varies between studies. Among the commonly cited features is the large horn that is formed by the {{Dinogloss|lacrimal bone}} and is situated above and in front of the eye. The lower jaw contains a bone known as the antarticular, which newly evolved in ''Allosaurus'' and is not found in other genera. The {{Dinogloss|paroccipital processes}} (a pair of bony struts that extend backwards and sidewards from the {{Dinogloss|braincase}} at the back of the skull) are extending ventrally (downwards) beyond the level of the {{Dinogloss|basal tubera}} (tubercle-like extensions on the underside of the braincase). Another distinguishing feature is found in the pelvis, where the lower end of the {{Dinogloss|ischium}} is suboval in side view.<ref name="malafaia2025" />
=== Inner systematics=== The relationships between individual ''Allosaurus'' specimens can be assessed using specimen-level phylogenetic analysis, in which a software calculates which of all possible cladograms (evolutionary trees) requires the least number of evolutionary steps to explain the differences between specimens. In 2024, Burigo and colleagues analyzed five skulls to conclude that ''A. europaeus'' is a separate species most closely related to ''A. jimmadseni''.<ref name="burigo2025" /> In 2025, Malafaia and colleagues used 17 skulls and instead concluded that the two ''A. europaeus'' skulls are more closely related to some specimens of ''A. fragilis'' than to each other, suggesting that ''A. europaeus'' is a synonym of ''A. fragilis''.<ref name="malafaia2025" />
{{clade |style=line-height:75%; |label1=''Allosaurus'' |1={{clade |label1=''A. fragilis'' |1={{clade |1=ML 415 (holotype of ''A. europaeus'') |2=USNM 8423 |3=AMNH 5753 |4=AMNH 666 |5=YPM VP 14554 |6=CM 11844 |7={{clade |1=USNM 4734 (neotype of ''A. fragilis'') |2={{clade |1=BYU 8901 |2={{clade |1=MNHN/UL.AND.# (assigned to ''A. europaeus'') |2=DINO 2560 }}}}}}}} |label2=''A. jimmadseni'' |2={{clade |1={{clade |1={{clade |1={{clade |1=DINO 11541 (holotype of ''A. jimmadseni'') |2=YPM VP 4944 }} |2={{clade |1=USNM 544100 |2={{clade |1=AMNH 600 |2=YPM VP 1890 }}}}}} |2={{clade |1=NMZ 1000005 ("Big Al II") |2=MOR 693 ("Big Al") }}}}}}}}}}
=== Outer systematics=== {{main|Allosauridae}} ''Allosaurus'' is the eponymous member several higher-level groups: Allosaurinae, Allosauridae, Allosauria, and Allosauroidea. The subfamily Allosaurinae was first used by Gregory S. Paul in a 1988 popular book and included ''Allosaurus'' and ''Chilantaisaurus'', but has not been widely used.<ref name="PaleobiologyDB" /><ref name="paul1988a" /> Allosauridae is a family that Marsh had named in 1878, one year after his description of ''A. fragilis''.<ref name="marsh1878" /> Originally, Allosauridae contained only ''Allosaurus'' itself. Although additional genera were later added, recent reviews have restricted the family to ''Allosaurus'' and ''Saurophaganax'',<ref name="holtz2004" /><ref name="carrano2012" /> with the latter no longer being recognized as a distinct genus.<ref name="danison2024" /> Allosauria is a higher-level group that typically includes Allosauridae and Carcharodontosauria. Allosauroidea is an even more inclusive group that also encompasses Metriacanthosauridae.<ref name="carrano2012" />
==Currently recognized species== thumb|right|''Allosaurus'' was a variable genus, one such area of variability being size.
===''Allosaurus fragilis''=== The type species of ''Allosaurus'', ''A. fragilis'', was named by Marsh in 1877, together with the genus ''Allosaurus''.<ref name="marsh1877" /> The name ''fragilis'' is Latin for {{gloss|fragile}} and refers to the lightening features in the vertebrae.<ref name="creisler2003" /> Because the specimen on which Marsh based the species on (the holotype, YPM 1930) is not diagnostic, the substantially more complete specimen USNM 4734 was designated as the neotype in 2023. USNM 4734 was discovered by Felch in the same quarry in which the holotype was discovered; it is therefore likely that both represent the same species.<ref name="paul2010" />
Malafaia and colleagues, in their 2025 study, proposed that ''A. fragilis'' can be diagnosed by four features found in the skull. There are two rows of {{Dinogloss|nutrient foramina}} (small openings for blood vessels, nerves, and similar tissues) on the outer surface of the {{Dinogloss|maxilla}} (upper jaw bone). The {{Dinogloss|jugal}} (bone of the cheek area) has a bulging on its lower edge, and in the braincase, the fossa (depression) on the {{Dinogloss|basioccipital}} below the {{Dinogloss|occipital condyle}} (the bony projection that connects with the vertebral column) has parallel margins and is less than 60% of the width of the occipital condyle. The lacrimal horn is triangular in shape.<ref name="malafaia2025" />
===''Allosaurus europaeus''=== {{multiple image |direction = horizontal |align = left |total_width = 300 |image1 = Allosaurus europaeus ML415 skull.png |alt1 = |image2 = A. europaeus material.png |alt2 = |footer = ''A. europaeus'' holotype skull with diagram showing preserved elements }} ''A. europaeus'' was named in 2006 by Octávio Mateus and colleagues based on a partial skull and three neck vertebrae (ML 415) from the Vale Frades beach in Lourinhã, Portugal. These authors also assigned a second specimen to this species, a partial skeleton that includes an {{Dinogloss|articulated}} hind limb and pelvis (MNHNUL/AND.001), which was found in 1988 near the village of Andrés in the District of Leiria. The specific name ''europaeus'' alludes to Europe. Mateus and colleagues defined their new species based on 17 features in the skull, such as a narrow lacrimal horn and a bifurcated rear end of the maxilla.<ref name="mateus2006" /> The status of ''A. europaeus'' is controversial, and some studies have argued that the species is a synonym of ''A. fragilis'',<ref name="malafaia2007" /> a ''nomen dubium'' (dubious name),<ref name="malafaia2010" /> or in need of re-evaluation.<ref name="evers2020" /> In a 2025 analysis, Malafaia and colleagues argued that all but one of the diagnostic features of ''A. europaeus'' fall within the variation range of ''A. fragilis''.<ref name="malafaia2025" />
In 2025, André Burigo and Mateus re-described the Vale Frades specimen and identified nine unique features in the skull and neck vertebrae that support the validity ''A. europaeus''. These authors further proposed that the species differs from ''A. fragilis'' in its longer lacrimal horns, a narrow crest on the nasal bone, the sizes of small pneumatic foramina (small openings) in the nasal bone, and the presence of additional laminae (sheets of bone) in the neck vertebrae.<ref name="burigo2025" /> These results were questioned by Malafaia and colleagues in 2025, although these authors cautioned that most of the diagnostic features listed by Burigo and Mateus were new and still have to be evaluated in a wider range of ''Allosaurus'' specimens.<ref name="malafaia2025" />
===''Allosaurus jimmadseni''=== thumb|Quarry map of DINO 11541, the holotype specimen of ''A. jimmadseni'' The name ''A. jimmadseni'' was first used in 2000, in the unpublished PhD thesis of Daniel Chure, and has since been used informally as a ''nomen nudum'' ({{gloss|naked name}}, a name that was invalidly published).<ref name="carrano2012" />{{rp|223}}<ref name="chure2020" /> ''A. jimmadseni'' was formally described by Chure and Mark Loewen in 2020. The name honors James Madsen for his work on the Cleveland-Lloyd Dinosaur Quarry and his influential 1976 monograph on the genus. A nearly complete specimen from Dinosaur National Monument, DINO 11541, was chosen as the holotype specimen, while several other specimens were assigned to the species, including MOR 693 ("Big Al") and SMA 0005 ("Big Al II"). Chure and Mark diagnosed the species by a unique combination of seven anatomical details, including a low and narrow crest that runs on either side of the skull on along the nasal bones, lacrimal horns that are higher than those of ''A. fragilis'', and the straight lower margin of the jugal.<ref name="chure2020" /> In 2024, Susannah Maidment found that ''A. fragilis'' and ''A. jimmadseni'' appear to have been contemporaneous but separated geographically, with ''A. fragilis'' concentrated in the south and east of the Morrison basin and ''A. jimmadseni'' in the north and west. Borth species occur together only at Dry Mesa quarry.<ref name="maidment2023" />
===''Allosaurus anax''=== ''A. anax'' was described by Andy Danison and colleagues in 2024, based on a few bones that were previously included in the taxon ''Saurophaganax maximus'', which had been regarded as an allosaurid separate from ''Allosaurus''. Danison and colleagues found that ''Saurophaganax'' is a chimera, comprising the fossils of a diplodocid sauropod as well as ''Allosaurus'' fossils. One of these ''Allosaurus'' fossils, a {{Dinogloss|postorbital bone}} (OMNH 1771), became the holotype of ''A. anax'', while six additional specimens consisting of parts of cervical and dorsal vertebrae as well as {{Dinogloss|fibulae}} (calf bones) were assigned to the species. The name {{lang|grc|anax}} is Greek for {{gloss|king}}, and also alludes to the name ''Saurophaganax''. According to Danison and colleagues, the ''A. anax'' fossils are significantly larger than those of other ''Allosaurus'' species. Unique features of the species include the lack of a pronounced ornamentation of the postorbital and the hourglass-shaped dorsal centra (the vertebral bodies of the back vertebrae) that are penetrated by pneumatic foramina (small openings), among other features.<ref name="danison2024" />
==Previously assigned species and synonyms== ===''Poikilopleuron valens'' (''Antrodemus'')=== {{main|Antrodemus}} left|thumb|''Antrodemus valens'' holotype tail vertebra (above) compared to those of ''Allosaurus'' (below) In 1869, Ferdinand Vandeveer Hayden obtained a fossil secondhand from Middle Park, near Granby, Colorado, from Morrison Formation rocks. Hayden sent his specimen to Joseph Leidy, who identified it as half of a tail vertebra, and tentatively assigned it to the European theropod genus ''Poekilopleuron'', as the new species ''Poicilopleuron'' [sic] ''valens'', based on the shared presence of a large medullary cavity. He noted that if more differences from ''P. bucklandii'' were to be found, the new species might be assigned to its own genus, ''Antrodemus''.<ref name="leidy1870" /><ref name="gilmore1920" />{{rp|6}} In 1873, he amended his description and identified the species as ''Antrodemus valens''.<ref name="leidy1873" /> In his 1920 monograph, Gilmore concluded that ''Antrodemus valens'' was indistinguishable from ''Allosaurus'', and that ''Antrodemus valens'' should be the preferred name because, as the older name, it had priority.<ref name="gilmore1920" /> However, Gilmore was not fully confident about his choice as the ''Antrodemus'' specimen is not diagnostic. ''Antrodemus'' became the accepted name over ''Allosaurus'' for over 50 years, until Madsen, in his 1976 monograph, concluded that ''Allosaurus'' should be used because ''Antrodemus'' was based on material with poor, if any, diagnostic features and locality information. Madsen pointed out that Gilmore studied the ''Allosaurus'' holotype based on plaster casts sent to him by Richard Swann Lull, and might not have been aware of the existence of the humerus fragment, which is the most diagnostic of the ''Allosaurus'' holotype bones.<ref name="madsen1976" /> ''Antrodemus'' has since been treated as a ''nomen dubium''.<ref name="paul2010" /><ref name="rauhut2011" /> The specimen is catalogued as USNM 218.<ref name="paul2010" /> ''Allosaurus valens'' is a new combination for ''Antrodemus valens'' used by Friedrich von Huene in 1932.<ref name="huene1932" />{{rp|230}}
===''Laelaps trihedrodon''=== ''Laelaps trihedrodon'' was named by Cope in 1877 based on a right jaw with teeth that was discovered by a local, the school teacher Oramel W. Lucas, in Garden Park.<ref name="cope1877" /><ref name="chure2001" /><ref name="carpenter2002" /> Cope assigned the species to ''Laelaps'' (now ''Dryptosaurus''), a genus that he had established in 1866. Subsequently, Cope assigned some additional bones from the same locality to the species, which he also received from Lucas. As with many of Cope's species, the original description of ''L. trihedrodon'' is terse, and Cope never provided figures of any of the ''L. trihedrodon'' fossils. When Cope's collection was transferred to the American Museum of Natural History in 1903, the jaw, which is the holotype of the species, could not be located and is considered lost. Of the other specimens that Cope assigned to the species, only five incomplete tooth crowns (AMNH 5780) appear to survive. In a 1939 catalog, Oskar Kuhn listed the species as ''Antrodemus'' (?) ''trihedrodon'', while listing ''Allosaurus'' as a synonym of ''Antrodemus''.<ref name="kuhn1939" />{{rp|72}} In 2001, Chure argued that AMNH 5780 probably belongs to ''Allosaurus''.<ref name="chure2001" />
===''Labrosaurus''=== The genus ''Labrosaurus'' was named by Marsh in 1879, for a species he had described the year before as ''Allosaurus lucaris''.<ref name="marsh1879" /><ref name="marsh1878" /> The name ''Labrosaurus'' ({{gloss|greedy lizard}}) is derived from the Greek {{lang|grc|labros}} {{gloss|greedy}} or {{gloss|furious}} and {{lang|grc|sauros}} {{gloss|lizard}}.<ref name="creisler2003l" /> The taxonomic history of the genus is complex and includes several named species.<ref name="buffetaut2012" /><ref name="madsen2000" />{{rp|37–38}} In 1882, Marsh classified the genus as the only member of a new family, Labrosauridae;<ref name="marsh1882" /> today, this family is considered to be a synonym of Allosauridae.<ref name="carrano2012" />{{rp|292}} ''Labrosaurus'' is now generally thought to be a synonym of ''Allosaurus''.<ref name="carrano2012" /> Several additional species have been assigned to ''Labrosaurus''. ''L. ferox'' may be a synonym of ''Allosaurus fragilis'' or an indeterminate ''nomen dubium''.<ref name="holtz2004" /><ref name="madsen2000" /><ref name="paul2010" /> Other assigned species included ''L. sulcatus'', ''L. stechowi'', and ''L. meriani'', which may belong to ''Ceratosaurus'' or a related form.<ref name="buffetaut2012" /><ref name="madsen2000" /><ref name="rauhut2011" />
====''Allosaurus lucaris''==== thumb|The holotype dentary of ''Labrosaurus ferox'', which may have been injured by the bite of another theropod The name ''Allosaurus lucaris'' was given by Marsh to a fragmentary skeleton in 1878.<ref name="marsh1878" /> This specimen (YPM 1931) stems from the Morrison Formation of Colorado<ref name="hay1908" /> or southern Wyoming.<ref name="paul2010" /> Originally, Marsh only mentioned one anterior dorsal vertebra. In 1879, he described additional material of the specimen including vertebrae and bones of the forelimb, and decided it warranted its own genus, ''Labrosaurus'', as ''L. lucaris''.<ref name="marsh1879" /><ref name="paul2010" /><ref name="hay1908" /> Marsh never published figures of these fossils.<ref name="hay1908" /> More recently, ''A. lucaris'' has often been regarded as a synonym of ''A. fragilis''.<ref name="holtz2004" /><ref name="malafaia2025" /><ref name="burigo2025" /> Paul and Carpenter argued in 2010 that the species is from a younger age than ''Allosaurus'' and might therefore represent a different genus. However, they found that the specimen was not diagnostic, and that ''A. lucaris'' is a ''nomen dubium''.<ref name="paul2010" />
====''Labrosaurus ferox''==== ''Labrosaurus ferox'' was named in 1884 by Marsh as a second species of ''Labrosaurus''.<ref name="marsh1884" /> It is based on a lower jaw (USNM 2315) discovered in Felch Quarry, the same quarry in which the ''Allosaurus'' holotype specimen was found.<ref name="paul2010" /> This jaw is oddly formed with a prominent gap in the tooth row at the tip of the jaw, and a rear section that is greatly expanded and down-turned.<ref name="marsh1884" /> Gilmore suggested in his 1920 monograph that the bone was pathological, showing an injury to the living animal.<ref name="gilmore1920" />{{rp|126}} In 2000, Madsen also suggested that the bone was pathological, and that the unusually expanded shape was partly due to inaccurate plaster reconstruction.<ref name="madsen2000" />{{rp|37}} It may be a specimen of ''A. fragilis'',<ref name="holtz2004" /><ref name="madsen2000" /> or an indeterminate specimen that may or may not belong to ''Allosaurus''.<ref name="paul2010" />
===''Epanterias amplexus''=== thumb|left|The type dorsal vertebra of ''Epanterias amplexus'' (A–C) and associated cervical vertebrae (D–I) and coracoid (J–L) ''Epanterias amplexus'' was named by Cope in 1878 based on a specimen from Quarry 2 in Garden Park, the same region in which Felch Quarry, the type locality of ''Allosaurus'', is located.<ref name="cope1878" /><ref name="chure2001" /> The name ''Epanterias'' translates to {{gloss|buttressed (vertebrae)}} and is derived from the Greek {{lang|grc|epi}} {{gloss|upon, on}} and {{lang|grc|anteris}} {{gloss|buttress}}, while the suffix ''ias'' means {{gloss|characterized by}}. The name ''amplexus'' is Latin for {{gloss|embracing}}.<ref name="creisler1992" /> The genus is based on AMNH 5767, which includes two anterior dorsal vertebrae, one of which preserves a mostly complete {{Dinogloss|neural arch}}; the fourth or fifth neck vertebra; an {{Dinogloss|axis}} (the second neck vertebra); a {{Dinogloss|coracoid}} (a bone of the shoulder girdle); and a possible metatarsal (foot bone).<ref name="osborn1921" />{{rp|283–284}}<ref name="paul2010" />
Cope thought that ''Epanterias'' was related to ''Camarasaurus'' and ''Streptospondylus'', and assigned it to the Camarasauridae,<ref name="cope1878" /><ref name="osborn1921" /> a group that was later assigned to Sauropoda. Subsequently, ''Epanterias'' appeared in lists of North American sauropods. In 1921, Henry Fairfield Osborn and Charles Craig Mook instead found ''Epanterias'' to be a large theropod, the largest known from the Morrison Formation at that time. These authors were the first to publish figures the fossils.<ref name="osborn1921" /> Robert T. Bakker considered ''Epanterias'' as a distinct genus of large allosaurid, which inspired Paul, who classified it as a species of ''Allosaurus'', ''Allosaurus amplexus''. Paul, in 1988, and Bakker, in 1990, suggested that large fossils from Oklahoma that would later become known as ''Saurophaganax maximus'' may be assigned to the same species.<ref name="paul1988a" /><ref name="bakker1990" /> In 1990, Bakker mentioned a third ''Epanterias amplexus'' locality, near Masonville, Colorado.<ref name="bakker1990" /> Other authors have considered ''E. amplexus'' as a synonym of ''Allosaurus fragilis''<ref name="holtz2004" /> or equated it with ''Allosaurus''.<ref name="carrano2012" /> In 2010, Paul and Carpenter noted that the ''E. amplexus'' specimen comes from higher in the Morrison Formation than the type specimen of ''Allosaurus fragilis'', and is therefore "probably a different taxon". They also stated that its holotype specimen is not diagnostic, and listed the taxon as a ''nomen dubium''.<ref name="paul2010" />
===''Creosaurus atrox''=== right|thumb|Holotype material of ''Creosaurus atrox'', more recently known as ''Allosaurus atrox'' ''Creosaurus atrox'' was described by Marsh in 1878, in the same paper that introduced ''A. lucaris''.<ref name="marsh1878" /> The name ''Creosaurus'' ({{gloss|flesh lizard}}) derives from the Greek ''kreos'' {{gloss|flesh}} and ''sauros'' {{gloss|lizard}} and hints at the carnivorous diet of the animal.<ref name="creisler2003c" /> It is based on YPM 1890, which was collected by Samuel Wendell Williston at Como, Wyoming, and includes two skull bones (a jugal and a premaxilla with teeth), two {{Dinogloss|sacrals}} (hip vertebrae), and parts of the pelvis and hind limb, including an illium.<ref name="paul2010" /><ref name="marsh1878" /><ref name="gilmore1920" />{{rp|118}} Marsh found these fossils to be "in excellent preservation". He speculated that the animal preyed upon the Atlantosauridae (sauropods), and estimated its body length at about {{cvt|20|ft}}.<ref name="marsh1878" /> Marsh found ''Creosaurus'' to be closely related to ''Dryptosaurus'',<ref name="marsh1878" /> and in 1884 classified these and several other genera including ''Allosaurus'' within the Megalosauridae.<ref name="marsh1884" />
In 1901, Williston argued that Marsh has never been able to adequately distinguish ''Creosaurus'' from ''Allosaurus'', and that Marsh labelled the drawing of a dorsal vertebra as ''C. atrox'' in a 1884 paper<ref name="marsh1884" /> even though he had labelled the same drawing as ''A. fragilis'' in a 1879 paper.<ref name="marsh1879" /><ref name="williston1878" /> Osborn, in 1903, assigned two fragmentary skulls to ''Creosaurus'', but re-assigend them to ''Allosaurus'' in 1912.<ref name="osborn1903" /><ref name="osborn1912" /><ref name="gilmore1920" />{{rp|9}} Oliver Perry Hay, in 1908, also questioned the validity of ''Creosaurus'', as the ilium, which he assumed to be the only bone belonging to the holotype, is indistinguishable from that of ''Allosaurus''.<ref name="hay1908" /> In his 1920 monograph, Gilmore considered ''C. atrox'' as a synonym of ''Antrodemus'' (=''Allosaurus'').
thumb|left|Holotype vertebra of ''Creosaurus potens'' (left) compared to an equivalent vertebra of the ''Allosaurus'' specimen USNM 8367 (labelled as ''Antrodemus valens'') In 1911, Richard Swann Lull named a second species, ''Creosaurus potens'', based on a single vertebra (USNM 3049) that J. K. Murphy discovered in the Arundel Formation of Maryland.<ref name="lull1911" />{{rp|186}}<ref name="gilmore1920" />{{rp|116}} The word {{lang|la|potens}} is Latin for {{gloss|powerful}}.<ref name="lull1911" />{{rp|174}} Lull identified the vertebrae as a dorsal vertebra, although Gilmore, in his 1920 monograph, showed that it was from the front portion of the tail.<ref name="gilmore1920" />{{rp|117}} Lull stated that ''C. potens'' was much larger than ''Allosaurus medius'', which he recognized as a second species of large theropod from the Arundel Formation.<ref name="lull1911" />{{rp|174}} In 1921, Gilmore provisionally assigned ''C. potens'' to the genus ''Dryptosaurus'', as ''Dryptosaurus''? ''potens'' on the basis of the similarities of the vertebral centrum to the caudal vertebrae of ''Dryptosaurus''.<ref>{{Cite journal |last=Gilmore |first=Charles W. |date=1921 |title=The fauna of the Arundel formation of Maryland |url=https://doi.org/10.5479/si.00963801.59-2389.581 |journal=Proceedings of the United States National Museum |volume=59 |issue=2389 |pages=581–594 |doi=10.5479/si.00963801.59-2389.581 |issn=0096-3801|url-access=subscription }}</ref><ref name="gilmore1920" />{{rp|119}}
In his 1988 popular book, Paul assigned ''C. atrox'' to ''Allosaurus'', as ''Allosaurus atrox''. He suggested that ''A. fragilis'' had tall pointed horns and a slender build compared to ''A. atrox'', and that these differences were probably not due to sexual dimorphism because ''A. atrox'' is much rarer.<ref name="paul1988a" /> The idea of two common Morrison allosaur species was followed in some semi-technical and popular works.<ref name="lessem1993" /> However, Chure noted that 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.<ref name="burigo2025" /> Bakker, in a 1998 paper, referred to a "creosaur-type allosaurid" that is distinct from ''A. fragilis'', although this informal taxon did not include the ''Creosaurus'' holotype.<ref name="carrano2012" /> Matthew Carrano and colleagues, in a 2012 review, suggested that Bakker's taxon is mostly equivalent to the subsequently named species ''A. jimmadseni''.<ref name="carrano2012" /> In 2010, Paul and Carpenter considered ''C. atrox'' to be a ''nomen dubium''.<ref name="paul2010" />
===''Camptonotus amplus''=== In 1879, Marsh named a new genus of ornithischian dinosaur, ''Camptonotus'' (now ''Camptosaurus''), from the Morrison Formation of Como Bluff.<ref name="marsh1879b" /> Marsh's ''Camptonotus'' originally included two species: the type species ''C. dispar'', based on fragmentary skeletons from Quarry 13, as well as ''C. amplus'', based on a right foot (YPM VP.1879) from Quarry 1A ("Big Canyon Quarry"). Marsh initially stated that the ''C. amplus'' individual was three times as large as ''C. dispar'', but later declared that it was merely 50% larger, {{cvt|30|ft}} compared to {{cvt|20|ft}}. The ''C. amplus'' specimen included a foot claw that was deep and flattened side-to-side, rather than rounded as in ''C. dispar''.<ref name="galton2015" /><ref name="marsh1879b" />
The species, under the name ''Camptosaurus amplus'', was accepted as valid in several studies in the late 1990s and early 2000s.<ref name="galton2015" /> In the late 1990s, Bakker first suggested that the foot instead belonged to an allosaurid. In 2015, Galton comprehensively reviewed the species, confirming that the foot is probably that of an allosaurid. He also found that a fragmentary skull (YPM VP.1892) from the same quarry probably belonged to the same individual, while the flattened claw instead belonged to a juvenile sauropod dinosaur. He tentatively assigned the species to ''Allosaurus'', as ?''Allosaurus amplus''.<ref name="galton2015" /> Subsequent authors noted that Galton did not suggest any difference between ''A. amplus'' and ''A. fragilis'', and that the former therefore cannot be assigned to any particular species of ''Allosaurus''.<ref name="yun2019" /><ref name="burigo2025" /> Burigo and colleagues, in their 2025 study, considered the species a ''nomen dubium''.<ref name="galton2015" />
===''Allosaurus medius''=== thumb|upright=0.45|Holotype tooth of ''A. medius'' ''A. medius'' was named by Marsh in 1888 for various fossils from the Early Cretaceous Arundel Formation of Maryland.<ref name="marsh1888" /><ref name="weishampel2004" />{{rp|556}} These include teeth, limb, and foot bones, and, according to Marsh, indicate an animal {{cvt|10|to|12|ft}} in length. In 1911, Lull moved most of the remains to the new ornithopod species ''Dryosaurus grandis'', except for a tooth which he considered to be the holotype of ''A. medius''. He assigned several other fossils from the Arundel to ''A. medius'', including teeth, vertebrae, and phalanges. Lull therefore recognized two species of large theropod from the Arundel, ''A. medius'' and ''Creosaurus potens'', the latter of which he named in the same publication.<ref name="lull1911" /><ref name="gilmore1920" />{{rp|119–121}} Gilmore, in his 1920 monograph, considered the holotype tooth to be not diagnostic but provisionally transferred it to ''Dryptosaurus'' based on its geographical context, noting that this assignment "is seriously in doubt". He also expressed doubt that two large theropods were present in the formation, albeit retaining both names. Gilmore in 1921 also moved the other parts of the original ''A. medius'' material, which Lull had assigned to ''Dryosaurus'', to a new species of ''Ornithomimus'', which he named ''Ornithomimus affinis''.<ref name="gilmore1920" />{{rp|119–121}} ''A. medius'' has been listed under several genera by different authors, creating the combinations ''Antrodemus medius'', ''Dryptosaurus medius'', and ''Labrosaurus medius''.<ref name="kuhn1939" />{{rp|74–75}} The 2004 edition of the encyclopedia The Dinosauria listed ''A. medius'' as a dubious species of theropod.<ref name="holtz2004" />
===''Allosaurus sibiricus''=== ''Allosaurus sibiricus'' was described in 1914 by Anatoly Riabinin on the basis of a bone, later identified as a partial fourth metatarsal, from the Early Cretaceous of Buryatia, Russia.<ref name="riabinin1914" /><ref name="carrano2012" /> In 1990, Ralph Molnar and colleagues transferred it to ''Chilantaisaurus'', as ''Chilantaisaurus''? ''sibiricus'', based on its age and locality.<ref name="molnar1990" /> It is now considered a ''nomen dubium'', indeterminate beyond Theropoda.<ref name="carrano2012" />
===''Allosaurus tendagurensis''=== thumb|left|''A. tendagurensis'' tibia, Naturkunde Museum Berlin ''A. tendagurensis'' was named in 1925 by Werner Janensch for a tibia (MB.R.3620) found in the Kimmeridgian-Tithonian Tendaguru Formation in Mtwara, Tanzania.<ref name="janensch1925" /> Missing portions of the bone had been restored, which possibly made the bone more elongate than it originally was.<ref name="carrano2012" />{{rp|252}} Janensch also assigned tail vertebrae to this species, which was not accepted by later studies.<ref name="carrano2012" /> Although tabulated as a tentatively valid species of ''Allosaurus'' in the second edition of ''The Dinosauria'',<ref name="holtz2004" /> subsequent studies placed it as indeterminate beyond Tetanurae, either a carcharodontosaurian or megalosaurid.<ref name="rauhut2005" /><ref name="rauhut2011" /><ref name="carrano2012" />{{rp|252}} Carrano and colleagues, in 2012, noted that the tibia is not very similar to that of ''Allosaurus''.<ref name="carrano2012" />{{rp|252}}
===''Saurophaganax maximus''=== {{main|Saurophaganax}} The fossils that would later become known as ''Saurophaganax'' were discovered by John Willis Stovall in 1931–1931 in the Morrison Formation of Oklahoma. In a popular magazine, Stovall named these fossils ''Saurophagus maximus''. In 1953, Charles Lewis Camp and colleagues pointed out that the name ''Saurophagus'' had already been given to a modern bird, the great kiskadee, and that Stovall's description was too terse to comply with the nomenclatural rules of the ICZN. Consequently, ''Saurophagus maximus'' was considered a ''nomen nudum''.<ref name="danison2024" /> In 1995, Chure properly described the taxon and renamed it ''Saurophaganax''.<ref name="chure1995" />
David K. Smith, in a 1998 analysis of variation within ''Allosaurus'', concluded that ''S. maximus'' was not different enough from ''Allosaurus'' to be a separate genus, but did warrant its own species, ''A. maximus''.<ref name="smith1998" /> This reassignment was rejected in the 2004 edition of ''The Dinosauria''.<ref name="holtz2004" /> A 2024 reassessment by Andy Danison and colleagues suggested that the holotype neural arch of ''Saurophaganax,'' along with some other material might have been from a sauropod. Other ''Saurophaganax'' bones were assigned to diplodocid sauropods.<ref>{{Cite journal |last=Danison |first=Andrew |last2=Wedel |first2=Mathew |last3=Barta |first3=Daniel |last4=Woodward |first4=Holly |last5=Flora |first5=Holley |last6=Lee |first6=Andrew |last7=Snively |first7=Eric |date=December 21, 2024 |title=Chimerism in specimens referred to Saurophaganax maximus reveals a new species of Allosaurus (Dinosauria, Theropoda) |url=https://doi.org/10.18435/vamp29404 |journal=Vertebrate Anatomy Morphology Palaeontology |volume=12 |doi=10.18435/vamp29404 |issn=2292-1389|doi-access=free }}</ref> The remaining bones were confirmed as those of a theropod, and assigned to a new species of ''Allosaurus'', ''A. anax''.<ref name="danison2024" />
===''Allosaurus lucasi''=== ''Allosaurus lucasi'' was named in 2014 by Sebastian Dalman for two specimens from the top of the Morrison Formation (Tithonian) in Colorado. The name ''lucasi'' honors Spencer G. Lucas for his contributions to vertebrate paleontology. The specimens were excavated in 1953 by a field crew of the Yale Peabody Museum led by Joseph T. Gregory at McElmo Canyon in what is now Canyons of the Ancients National Monument, Colorado. The holotype specimen (YPM VP 57589) is a fragmentary skeleton of an adult individual that includes disarticulated bones of the skull and postcranium, including an almost complete left foot. The second specimen (YPM VP 57726) consists of two skull bones (dentary and {{Dinogloss|splenial}}) of a juvenile individual. According to Dalman, ''A. lucasi'' differs from ''A. fragilis'' in several features, such as its shorter premaxilla and the shorter branches of the {{Dinogloss|quadratojugal}} that contacted the jugal and the {{Dinogloss|quadrate}}. Dalman suggested that ''A. lucasi'' was comparatively large, and possibly the geologically youngest known species of ''Allosaurus''.<ref name="dalman2014" /> Later authors did not accept this proposed species. Chure and Loewen declared the species as "invalid" in 2020,<ref name="chure2020" />{{rp|5}} while Burigo and colleagues considered it a ''nomen dubium'' in 2025.<ref name="burigo2025" /><ref name="malafaia2025" />
==Typographical errors and ''nomina nuda''== "Allosaurus agilis", seen in Osborn, 1912, is a typographical error for ''A. fragilis.''<ref name="olshevsky1978"/> "Allosaurus ferox" is a typographical error by Marsh for ''A. fragilis'' made in a footnote in 1896.<ref name="hay1908" /> Likewise, "Labrosaurus fragilis" is a typographical error by Marsh (1896) for ''Labrosaurus ferox''.<ref name="hay1908" /><ref name="madsen2000" />{{rp|37}} "Labrosaurus huene" is a ''nomen nudum'' used by von Huene in 1956 and 1958 for a tooth from the Upper Jurassic of Sichuan, China.<ref name="madsen2000" />{{rp|37}} "Allosaurus whitei" was coined by Pickering in 1995, and has been listed as a synonym of ''A. fragilis''.<ref name="malafaia2025" />
==Other misassigned specimens== Several fragmentary specimens from Europe and Asia have been assigned to ''Allosaurus'', but Burigo and colleagues, in a 2025 review, found that only the Portuguese material as well as some teeth from Germany described in 2016 are indeed assignable to the genus. Other teeth from Germany and France, as well as tracks from England, cannot be assigned to any particular genus.<ref name="burigo2025" /> A specimen from the Late Cretaceous Jobu Formation of Kumamoto, Japan, consisting of vertebrae, limb bones, and teeth, was assigned to ''Allosaurus'' by Minoru Tamura and colleagues in 1991.<ref name="tamura1991" /> Burigo and colleagues instead assigned this specimen to ''Segnosaurus''.<ref name="burigo2025" /> In 2003, Kurzanov and colleagues assigned six teeth (PIN 4874/2) from Siberia to ''Allosaurus'' sp.<ref name="kurzanov2003" /> Carrano and colleagues, in 2012, noted that these teeth are indeterminate beyond Theropoda.<ref name="carrano2012" />{{rp|261}}
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. In their 1981 description of the specimen, Ralph Molnar and colleagues suggested that it might indicate that Australia was a refugium for animals that had gone extinct elsewhere.<ref name="molnar1981" /> This identification was challenged by Samuel Welles in 1983, who thought it more resembled that of an ornithomimid,<ref name="welles1983" /> but the original authors defended their identification in 1985.<ref name="molnar1985" /> In 1998, 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.<ref name="chure1998" /> Similarly, Yoichi Azuma and Phil Currie, in their 2000 description of ''Fukuiraptor'', noted that the bone closely resembled that of their new genus.<ref name="azuma2000" /> It may have belonged to ''Australovenator'' or a similar taxon,<ref name="hocknull2009" /> or it may represent an abelisaur.<ref name="agnolin2010" />
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Category:Allosauridae Category:Reptile taxonomy