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Ceratosaurus

Genus of theropod dinosaur from the Late Jurassic period

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Cast of Ceratosaurus from the Cleveland Lloyd Quarry, Utah, on display at the Natural History Museum of Utah, Salt Lake City.
Jens Lallensack · CC BY-SA 4.0

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The Garden Park specimen remains the most complete skeleton known from the genus and only a handful of additional specimens have been described since. Two additional species, Ceratosaurus dentisulcatus and Ceratosaurus magnicornis, were described in 2000 from two fragmentary skeletons from the Cleveland-Lloyd Quarry of Utah and from the vicinity of Fruita, Colorado. These species are no longer accepted by most paleontologists, and all three skeletons possibly represent different growth stages of the same species. In 1999, the discovery of the first juvenile specimen was reported. In 2000, a partial specimen was excavated and described from the Lourinhã Formation of Portugal, providing evidence for the presence of the genus outside of North America. Fragmentary remains have also been reported from Tanzania, Uruguay, and Switzerland, although their assignment to Ceratosaurus is currently not accepted by most paleontologists.

Ceratosaurus was a medium-sized theropod. The original specimen is estimated to be 5.3 m (17 ft) or 5.69 m (18.7 ft) long, while the specimen described as C. dentisulcatus was larger, at around 7 m (23 ft) long. Ceratosaurus was characterized by deep jaws that supported proportionally very long, blade-like teeth, a prominent, ridge-like horn on the midline of the snout, and a pair of horns over the eyes. The forelimbs were very short, but remained fully functional. The hand had four fingers with claws on the first three. The tail was deep from top to bottom. A row of small osteoderms (skin bones) was present down the middle of the neck, back, and tail. Additional osteoderms were present at unknown positions on the animal's body.

Ceratosaurus gives its name to Ceratosauria, a clade of theropod dinosaurs that diverged early on from the evolutionary lineage leading to modern birds. Within Ceratosauria, some paleontologists proposed it to be most closely related to Genyodectes from Argentina, which shares the strongly elongated teeth. The geologically older genus Proceratosaurus from England, although originally described as a presumed antecedent of Ceratosaurus, was later found to be an early tyrannosauroid. Ceratosaurus shared its habitat with other large theropod genera, including Torvosaurus and Allosaurus, and it has been suggested that these theropods occupied different ecological niches to reduce competition. Ceratosaurus may have preyed upon plant-eating dinosaurs; while some paleontologists suggested that it hunted aquatic prey such as fish, this has been rejected by more recent studies. The nasal horn was probably not used as a weapon as was originally suggested by Marsh, but more likely was used solely for display.

History of discovery

Holotype specimen of C. nasicornis

The first specimen, holotype USNM 4735, was discovered and excavated by farmer Marshall Parker Felch in 1883 and 1884. Found in articulation, with the bones still connected to each other, it was nearly complete, including the skull. Significant missing parts include an unknown number of vertebrae, all but the last ribs of the trunk, the humeri (upper arm bones), the distal finger bones of both hands, most of the right arm, most of the left leg, and most of the feet. The specimen was found encased in hard sandstone, leading to the skull and spine being heavily distorted during fossilization. The site of discovery, located in the Garden Park area north of Cañon City, Colorado, and known as the Felch Quarry, is regarded as one of the richest fossil sites of the Morrison Formation. Numerous dinosaur fossils had been recovered from this quarry even before the discovery of Ceratosaurus, most notably a nearly complete specimen of Allosaurus (USNM 4734) in 1883 and 1884.

After excavation, the specimen was shipped to the Peabody Museum of Natural History in New Haven, where it was studied by Marsh, who described it as the new genus and species Ceratosaurus nasicornis in 1884. The name Ceratosaurus may be translated as "horn lizard" (from the Greek words κερας / κερατος, keras / keratos—"horn" and σαυρος/sauros—"lizard") and nasicornis with "nose horn" (from the Latin words nasus—"nose" and cornu—"horn"). Given the completeness of the specimen, the newly described genus was, at the time, the best-known theropod discovered in America. In 1898 and 1899, the specimen was transferred to the National Museum of Natural History in Washington, DC, along with many other fossils originally described by Marsh. Only part of this material was fully prepared when it arrived in Washington. Subsequent preparation lasted from 1911 to the end of 1918. Packaging and shipment from New Haven to Washington caused some damage to the Ceratosaurus specimen. In 1920, Charles Gilmore published an extensive redescription of this and the other theropod specimens received from New Haven, including the nearly complete Allosaurus specimen recovered from the same quarry.

In an 1892 paper, Marsh published the first skeletal reconstruction of Ceratosaurus, which depicts the animal at 22 ft (6.7 m) in length and 12 ft (3.7 m) in height. As noted by Gilmore in 1920, the trunk was depicted much too long in this reconstruction, incorporating at least six dorsal vertebrae too many. This error was repeated in several subsequent publications, including the first life reconstruction, which was drawn in 1899 by Frank Bond under the guidance of Charles R. Knight, but not published until 1920. A more accurate life reconstruction, published in 1901, was produced by Joseph M. Gleeson, again under Knight's supervision. The holotype was mounted by Gilmore in 1910 and 1911. Since then, it was exhibited at the National Museum of Natural History. Most early reconstructions show Ceratosaurus in an upright posture, with the tail dragging on the ground. Gilmore's mount of the holotype, in contrast, was ahead of its time. Inspired by the thigh bones, which were found angled against the lower leg, he depicted the mount as a running animal with a horizontal posture and a tail that did not make contact with the ground. Because of the strong flattening of the fossils, Gilmore did not mount the specimen as a free-standing skeleton but as a bas-relief within an artificial wall. With the bones being partly embedded in a plaque, scientific access was limited. In the course of the renovation of the museum's dinosaur exhibition between 2014 and 2019, the specimen was dismantled and freed from the encasing plaque. In the new exhibition, the original skeleton was replaced by a free-standing cast that is lying on its back fighting a Stegosaurus. The original bones are stored in the museum collection to allow full access for scientists.

Additional finds in North America

After the discovery of the holotype of C. nasicornis, a significant Ceratosaurus find was not made until the early 1960s, when paleontologist James Madsen and his team unearthed a fragmentary, disarticulated skeleton including the skull (UMNH VP 5278) in the Cleveland-Lloyd Dinosaur Quarry in Utah. This find represents one of the largest-known Ceratosaurus specimens. A second, articulated specimen including the skull (MWC 1) was discovered in 1976 by Thor Erikson, the son of paleontologist Lance Erikson, near Fruita, Colorado. A fairly complete specimen, it lacks lower jaws, forearms, and gastralia. The skull, although reasonably complete, was found disarticulated and is strongly flattened sideways. Although it was a large individual, it had not yet reached adult size, as indicated by unfused sutures between the skull bones. Scientifically accurate three-dimensional reconstructions of the skull for use in museum exhibits were produced using a complicated process including molding and casting of the individual original bones, correction of deformities, reconstruction of missing parts, assembly of the bone casts into their proper position, and painting to match the original color of the bones.

Both the Fruita and Cleveland-Lloyd specimens were described by Madsen and Samuel Paul Welles in a 2000 monograph, with the Utah specimen being assigned to the new species C. dentisulcatus and the Colorado specimen being assigned to the new species C. magnicornis. The name dentisulcatus refers to the parallel grooves present on the inner sides of the premaxillary teeth and the first three teeth of the lower jaw in that specimen. Magnicornis points to the larger nasal horn. As of 2025, both species are generally considered as synonyms of C. nasicornis. Brooks Britt and colleagues, in 2000, claimed that the C. nasicornis holotype was in fact a juvenile individual, with the two larger species representing the adult state of a single species. Oliver Rauhut, in 2003, and Matthew Carrano and Scott Sampson, in 2008, considered the anatomical differences between these species to represent ontogenetic (age-related) or individual variation.

A further specimen (BYU 881-12893) was discovered in 1992 in the Agate Basin Quarry southeast of Moore, Utah. The specimen includes the front half of a skull, seven fragmentary pelvic dorsal vertebrae, and an articulated pelvis and sacrum. In 1996, a Ceratosaurus skeleton belonging to a juvenile individual was discovered near Bone Cabin Quarry in Wyoming. This specimen is 34% smaller than the C. nasicornis holotype and consists of a complete skull as well as 30% of the remainder of the skeleton including a complete pelvis. The specimen was bought by the North American Museum of Ancient Life, but because this museum is private rather than a public repository, the specimen has not been scientifically described. The museum sold the specimen to a private owner in 2024. During an auction at Sotheby's on July 16, 2025, the specimen was resold at a price of $30,510,000.

Besides these five skeletal finds, fragmentary Ceratosaurus remains have been reported from various localities from stratigraphic zones 2 and 4-6 of the Morrison Formation, including some of the major fossil sites of the formation. Dinosaur National Monument, Utah, yielded an isolated right premaxilla (DNM 972). A large shoulder blade (scapulocoracoid) was reported from Como Bluff in Wyoming. Another specimen stems from the Dry Mesa Quarry of Colorado and includes a left scapulocoracoid, as well as fragments of vertebrae and limb bones. In Mygatt Moore Quarry, Colorado, the genus is known from teeth.

In 1896, Marsh used the name Labrosaurus sulcatus in a caption for a figure depicting a single tooth (YPM 1936). Because Marsh did not provide any description, the name might be a nomen nudum. Later authors noted that the tooth came from Quarry 9 in Como Bluff, and that its assignment to Labrosaurus was baseless as it did not resemble other Labrosaurus teeth recognized at that time. The tooth has characteristic longitudinal grooves, which resulted in the assignment of several other, similar teeth from Tanzania and Switzerland to Labrosaurus. Subsequent studies identified the tooth as belonging to Ceratosaurus, either from the front portion of the lower jaw or the premaxilla of the upper jaw.

Finds outside North America

From 1909 to 1913, German expeditions of the Berlin Museum für Naturkunde uncovered a diverse dinosaur fauna from the Tendaguru Formation in German East Africa, in what is now Tanzania. Although commonly considered the most important African dinosaur locality, large theropod dinosaurs are only known through few and very fragmentary remains. In 1920, German paleontologist Werner Janensch assigned three dorsal vertebrae of a juvenile individual from the quarry "TL" to Ceratosaurus, as Ceratosaurus sp. (of uncertain species). In 1925, Janensch named a new species of Ceratosaurus, C. roechlingi, based on fragmentary remains from the quarry "Mw" encompassing a quadrate bone, a fibula, fragmentary caudal vertebrae, and other fragments. This specimen stems from an individual substantially larger than the C. nasicornis holotype.

In their 2000 monograph, Madsen and Welles confirmed the assignment of these finds to Ceratosaurus. In addition, they ascribed several teeth to the genus, which had originally been described by Janensch as a possible species of Labrosaurus, Labrosaurus (?) stechowi. Other authors questioned the assignment of any of the Tendaguru finds to Ceratosaurus, noting that none of these specimens displays features diagnostic for that genus. In 2011, Rauhut found both C. roechlingi and Labrosaurus (?) stechowi to be possible ceratosaurids, but found them to be undiagnostic at genus level and designated them as nomina dubia (doubtful names). In 1990, Timothy Rowe and Jacques Gauthier mentioned yet another Ceratosaurus species from Tendaguru, Ceratosaurus ingens, which purportedly was erected by Janensch in 1920 and was based on 25 isolated, very large teeth up to 15 cm (5.9 in) in length. However, Janensch assigned this species to Megalosaurus, not to Ceratosaurus. Therefore, this name might be a simple copying error. Rauhut, in 2011, showed that Megalosaurus ingens was not closely related to either Megalosaurus or Ceratosaurus, but possibly represents a carcharodontosaurid instead.

In 2000 and 2006, paleontologists led by Octávio Mateus described a find from the Lourinhã Formation of central-west Portugal (ML 352) as a new specimen of Ceratosaurus, consisting of a right femur (thigh bone), a left tibia (shin bone), and several isolated teeth recovered from the cliffs of Valmitão beach, between the municipalities of Lourinhã and Torres Vedras. The bones were found embedded in yellow to brown, fine-grained sandstones, which were deposited by rivers as floodplain deposits and belong to the lower levels of the Porto Novo Member, which is thought to be late Kimmeridgian in age. Additional bones of this individual (SHN (JJS)-65), including a left femur, a right tibia, and a partial left fibula (calf bone), were since exposed due to progressing cliff erosion. Although initially part of a private collection, these additional elements became officially curated after the private collection was donated to the Sociedade de História Natural in Torres Vedras and were described in detail in 2015. The specimen was ascribed to the species Ceratosaurus dentisulcatus by Mateus and colleagues in 2006. A 2008 review by Carrano and Sampson confirmed the assignment to Ceratosaurus, but concluded that the assignment to any specific species is not possible at present. In 2015, Elisabete Malafaia and colleagues, who questioned the validity of C. dentisulcatus, assigned the specimen to Ceratosaurus aff. Ceratosaurus nasicornis.

Other reports include a single tooth found in Moutier, Switzerland. The tooth was originally included in Megalosaurus meriani by Jean-Baptiste Greppin in 1870, but transferred to the genus Labrosaurus, as Labrosaurus (?) stechowi, by Janensch in 1920. The tooth was assigned to Ceratosaurus sp. (of unknown species) by Madsen and Welles in their 2000 study, and to an indeterminate member of Ceratosauria by Carrano and colleagues in 2012. In 2008, Matías Soto and Daniel Perea described teeth from the Tacuarembó Formation in Uruguay, including a presumed premaxillary tooth crown. This shows vertical striations on its inner side and lacks denticles on its front edge. These features are, in this combination, only known from Ceratosaurus. The authors, however, stressed that an assignment to Ceratosaurus is infeasible because the remains are scant and note that the assignment of the European and African material to Ceratosaurus has to be viewed with caution. In 2020, Soto and colleagues described additional Ceratosaurus teeth from the same formation that further support their earlier interpretation.

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