Ceratosaurus

Ceratosaurus Temporal range: Late Jurassic, 153–148 Ma PreЄ Є O S D C P T J K Pg N ↓
Mounted cast of a juvenile skeleton, Dinosaur Discovery Museum
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Clade:	Dinosauria
Order:	Saurischia
Suborder:	Theropoda
Family:	†Ceratosauridae
Genus:	†Ceratosaurus Marsh, 1884
Species
†C. nasicornis Marsh, 1884 (type) †?C. dentisulcatus Madsen & Welles, 2000 †?C. magnicornis Madsen & Welles, 2000
Synonyms
Megalosaurus nasicornis (Marsh, 1884 [originally Ceratosaurus])

Ceratosaurus /ˌsɛrətoʊˈsɔːrəs/ meaning "horned lizard", in reference to the horn on its nose (Greek κερας/κερατος, keras/keratos meaning "horn" and σαυρος/sauros meaning "lizard"), was a large predatory theropod dinosaur from the Late Jurassic Period (Kimmeridgian to Tithonian), found in the Morrison Formation of North America, and the Lourinhã Formation of Portugal (and possibly the Tendaguru Formation in Tanzania).^[1] It was characterized by large jaws with blade-like teeth, a large, blade-like horn on the snout and a pair of hornlets over the eyes. The forelimbs were powerfully built but very short. The bones of the sacrum were fused (synsacrum) and the pelvic bones were fused together and to this structure^[2] (i.e. similar to modern birds). A row of small osteoderms was present down the middle of the back.

Description

Artist's impression of C. nasicornis

Ceratosaurus, at first glance, looked like a fairly typical theropod,^[3] however its skull was quite large in proportion to the rest of its body, and large nasal and brow horns^[4] and possessed a prominent nose horn formed from protuberances of the nasal bones.^[5] In addition to the large nasal horn, Ceratosaurus possessed smaller hornlike ridges in front of each eye, similar to those of Allosaurus, these ridges were formed by enlargement of the lacrimal bones.^[6] Uniquely among theropods, Ceratosaurus possessed dermal armor, in the form of small osteoderms running down the middle of its back.^[7] Its tail comprised about half of the body's total length and was thin and flexible with high vertebral spines.^[4]

The type specimen was an individual about 18 feet (5.5 m) long; it is not clear whether this animal was fully grown.^[7][8] David B. Norman (1985) estimated that the maximum length of Ceratosaurus was 20 ft (6.1 m),^[9] an assessment supported by a particularly large Ceratosaurus specimen from the Cleveland-Lloyd Quarry (UMNH 5728), discovered in the mid-1960s, which may have been 22 ft (6.7 m) long assuming similar proportions to the holotype.^[10]

Comparison of two specimens of C. nasicornis (red, pink) and other theropods from the Morrison Formation.

Marsh (1884) suggested that Ceratosaurus weighed about half as much as Allosaurus.^[4] In Predatory Dinosaurs of the World, published in 1988, Gregory S. Paul estimated that the C. nasicornis holotype skeleton came from an animal weighing about 524 kilograms (1,155 lb) while the large material from the Cleveland-Lloyd Quarry represents a much bigger and heavier individual, whose bulk he estimated at about 980 kilograms (2,160 lb).^[11]

The bigger specimen was later assigned by James H. Madsen and Samuel P. Welles to the new species C. dentisulcatus.^[12] A considerably lower figure was proposed by John Foster, a specialist on the Morrison Formation, in 2007. Foster used an equation provided by J.F. Anderson and colleagues to estimate mass from femur length, which yielded an approximate weight of 275 kilograms (606 lb) for C. magnicornis and 452 kilograms (996 lb) for C. dentisulcatus.^[6]

According to Rauhut (2000), Ceratosaurus can be distinguished based on the following features: a narrow rounded horn core centrally placed on the fused nasals, a median oval groove on nasals behind horn core, a premaxilla with three teeth, premaxillary teeth with reduced extent of mesial serrations, chevrons that are extremely long, a pubis with a large, rounded notch underneath the obturator foramen, small epaxial osteoderms.^[13]

Discovery and species

Ceratosaurus nasicornis skeleton restoration by O.C. Marsh from 1896, depicted in an erroneous upright position

Ceratosaurus is known from the Cleveland Lloyd Dinosaur Quarry in central Utah and the Dry Mesa Quarry in Colorado. The type species, described by O. C. Marsh in 1884 and redescribed by Gilmore in 1920, is Ceratosaurus nasicornis.

C. nasicornis holotype, Smithsonian Museum

Two further species were described in 2000: C. magnicornis (from the Fruita Paleontontological Area, outside Fruita, Colorado) and C. dentisulcatus.^[14] C. magnicornis has a slightly rounder horn but is otherwise highly similar to C. nasicornis; C. dentisulcatus is larger (6.7 meters), slightly more derived, and has an unknown horn shape (assuming it had them). The Portuguese remains have been ascribed to C. dentisulcatus (Mateus et al. 2000; 2006). However, the validity of C. magnicornis and C. dentisuclatus has been disputed by various palaeontologists,^[15][16][17] arguing that the differences from C. nasicornis are due to individual or ontogenetic (age related) variation; thus, they would be junior synonyms of C. nasicornis. C. meriani is a nomen dubium.^[18]

More additional species, including C. ingens and C. stechowi, have been described from less complete material. The former is now believed to be a dubious carcharodontosaurid instead, and the latter is a dubious ceratosaur and/or spinosaurid.^[19] Ceratosaurus is present in stratigraphic zones 2 and 4-6 of the Morrison Formation.^[20]

Classification

Partial fossil.

Cast of the hand of C. nasicornis (AMNH 27631).

Relatives of Ceratosaurus include Genyodectes, Elaphrosaurus, and the abelisaurs, such as Carnotaurus. The classification of Ceratosaurus and its immediate relatives has been under intense debate. Ceratosaurs are unique in their characters; they share some primitive traits with coelophysoids, but also share some derived traits with tetanuran theropods not found in coelophysians. Its closest relatives appear to be the abelisaurs.

In the past, Ceratosaurus, the abelisaurs, and the primitive coelophysoids were all grouped together and called Ceratosauria, defined as "theropods closer to Ceratosaurus than to Aves". Recent evidence, however, has shown large distinctions between the later, larger and more advanced ceratosaurs and earlier forms like Coelophysis. While considered distant from birds among the theropods, Ceratosaurus and its kin were still very bird-like, and even had a more avian tarsus (ankle joint) than Allosaurus.

The following is a cladogram based on the phylogenetic analysis conducted by Diego Pol and Oliver W. M. Rauhut in 2012,^[21] showing the relationships of Ceratosaurus:

Ceratosauria	Berberosaurus Deltadromeus Spinostropheus Limusaurus Elaphrosaurus  Neoceratosauria   Ceratosauridae  Ceratosaurus Genyodectes  Abelisauroidea  Noasauridae Abelisauridae

Paleobiology

Ceratosaurus and Dryosaurus.

Feeding

Ceratosaurus lived alongside dinosaurs such as Allosaurus, Torvosaurus, Apatosaurus, Diplodocus, Stegosaurus and Camarasaurus. Ceratosaurus reached lengths of 6.7 m (22 ft), and weighed up to 980 kilograms (2,160 lb). It was smaller than the other large carnivores of its time (allosaurs and Torvosaurus) and likely occupied a distinctly separate niche from them. Ceratosaurus fossils are noticeably less common than those of Allosaurus, but whether this implies Ceratosaurus being rarer is uncertain (animals with certain lifestyles are more biased toward fossilization than others). Ceratosaurus had a longer, more flexible body, with a deep tail shaped like that of a crocodilian.^[7] This suggests that it was a better swimmer than the stiffer Allosaurus. A recent study by Robert Bakker suggested that Ceratosaurus generally hunted aquatic prey, such as fish and crocodiles, although it had potential for feeding on large dinosaurs. The study also suggests that sometimes adults and juveniles ate together.^[22] This evidence is debatable, and Ceratosaurus tooth marks are very common on large, terrestrial dinosaur prey fossils. Scavenging from corpses, smaller predators, and after larger ones also likely accounted for some of its diet.

Ceratosaurus, Allosaurus and Torvosaurus appear to have had different ecological niches, based on anatomy and the location of fossils. Ceratosaurus and Torvosaurus may have preferred to be active around waterways, and had lower, thinner bodies that would have given them an advantage in forest and underbrush terrains, whereas Allosaurus were more compact, with longer legs, faster but less maneuverable, and seem to have preferred dry floodplains.^[23] Ceratosaurus, better known than Torvosaurus, differed noticeably from Allosaurus in functional anatomy by having a taller, narrower skull with large, broad teeth. Allosaurus was itself a potential food item to other carnivores, as illustrated by an Allosaurus pubic foot marked by the teeth of another theropod, probably Ceratosaurus or Torvosaurus. The location of the bone in the body (along the bottom margin of the torso and partially shielded by the legs), and the fact that it was among the most massive in the skeleton, indicates that the Allosaurus was being scavenged.^[24]

Nasal horn

Skull cast showing large nasal horn

Marsh (1884) considered the nasal horn of Ceratosaurus to be a "most powerful weapon" for both offensive and defensive purposes, and Gilmore (1920) concurred with this analysis.^[4][7] However, this interpretation is now generally considered unlikely.^[25] Norman (1985) believed that the horn was "probably not for protection against other predators," but might instead have been used for intraspecific combat among male ceratosaurs contending for breeding rights.^[9] Paul (1988) suggested a similar function, and illustrated two Ceratosaurus engaged in a non-lethal butting contest.^[11] Rowe and Gauthier (1990) went further, suggesting that the nasal horn of Ceratosaurus was "probably used for display purposes alone" and played no role in physical confrontations.^[5] If used for display, it is likely that the horn would have been brightly colored.^[6]

Paleopathology

In 2001, Bruce Rothschild and others published a study examining evidence for stress fractures in theropod dinosaurs. They examined a single foot bone referred to Ceratosaurus and found that it had a stress fracture.^[26]

The holotype specimen of Ceratosaurus nasicornis, USMN 4735 was found with its left metatarsals II to IV fused together. Whether or not this fusion was pathological or natural to the species became controversial when Baur in 1890 speculated that the fusion was the result of a healed fracture. An analysis by Tanke and Rothschild suggests that the fusion was indeed pathological.^[27]

An unidentified species of Ceratosaurus preserved a broken tooth that showed signs of further wear received after the break.^[27]

In popular culture

Restoration by J. M. Gleeson from 1901, made under supervision of Charles R. Knight.

Ceratosaurus has appeared in several films, including the first live action film to feature dinosaurs, D. W. Griffith's Brute Force (1914).^[28] In the Rite of Spring segment of Fantasia (1940), Ceratosaurus are shown as opportunistic predators attacking Stegosaurus and sauropods trapped in mud. In The Animal World (1956) a Ceratosaurus kills a Stegosaurus in battle, but is soon attacked by another Ceratosaurus trying to steal a meal. This scene ends with both Ceratosaurus falling to their deaths off the edge of a high cliff.

A Ceratosaurus battles a Triceratops in the 1966 remake of One Million Years B.C.. Ceratosaurus is also featured in The Land That Time Forgot (1975) where it battles a Triceratops, and its sequel The People That Time Forgot (1977) in which Patrick Wayne's character rescues a cavegirl from two pursuing Ceratosaurus by driving the dinosaurs off with smoke bombs (after having failed to frighten them off by firing shots in the air once the Ceratosaurus' attention had been shifted to Patrick Wayne's party of explorers). A Ceratosaurus made a brief appearance in the film Jurassic Park III in which it is repelled from attacking the main characters by a large mound of Spinosaurus dung. This dinosaur also appears in the television documentary When Dinosaurs Roamed America, a Ceratosaurus makes a few appearances as a predator, killing Dryosaurus and eating it, a different one is shown chasing the same Dryosaurus but is later killed and eaten by an Allosaurus. Ceratosaurus is also featured in episodes of Jurassic Fight Club where it is seen as a rival to Allosaurus and preying on Stegosaurus.

See also

• Timeline of ceratosaur research

References

1. Mateus, O.; M.T. Antunes (2015). "Ceratosaurus sp. (Dinosauria: Theropoda) in the Late Jurassic of Portugal". =31st International Geological Congress. Rio de Janeiro, Brazil.
2. Sereno 1997
3. Marsh, O.C. (1892). "Restorations of Claosaurus and Ceratosaurus". American Journal of Science 44 (262): 343–349. doi:10.2475/ajs.s3-44.262.343.
4. Marsh, O.C. (1884). "Principal characters of American Jurassic dinosaurs, part VIII: The order Theropoda" (PDF). American Journal of Science 27 (160): 329–340. doi:10.2475/ajs.s3-27.160.329.
5. Rowe, T.; Gauthier, J. (1990). "Ceratosauria". In Weishampel, D.B.; Dodson, P.; Osmólska, H. The Dinosauria. University of California Press. pp. 151–168. ISBN 0-520-06726-6.
6. Foster, John (2007). "Gargantuan to Minuscule: The Morrison Menagerie, Part II". Jurassic West: The Dinosaurs of the Morrison Formation and Their World. Indiana University Press. pp. 162–242. ISBN 0-253-34870-6.
7. Gilmore, C.W. (1920). "Osteology of the carnivorous Dinosauria in the United States National Museum, with special reference to the genera Antrodemus (Allosaurus) and Ceratosaurus". Bulletin of the United States National Museum 110: 1–154. doi:10.5479/si.03629236.110.i.
8. Tykoski, R.S.; Rowe, T. (2004). "Ceratosauria". In Weishampel, D.B.; Dodson, P.; Osmólska, H. The Dinosauria: Second Edition. University of California Press. pp. 47–70. ISBN 0-520-24209-2.
9. Norman, D.B. (1985). "Carnosaurs". The Illustrated Encyclopedia of Dinosaurs. Salamander Books Ltd. pp. 62–67. ISBN 0-517-46890-5.
10. Mortimer, Michael. "C.dentisulcatus". Retrieved 2 March 2013.
11. Paul, Gregory S. (1988). "Ceratosaurs". Predatory Dinosaurs of the World. Simon & Schuster. pp. 274–279. ISBN 0-671-61946-2.
12. Madsen, J.H.; Welles, S.P. (2000). Ceratosaurus (Dinosauria, Theropoda): A Revised Osteology. Utah Geological Survey. pp. 1–80. ISBN 1-55791-380-3.
13. Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria, Saurischia). Ph.D. dissertation, Univ. Bristol [U.K.]. 440 pp.
14. Madsen JH, Welles SP. Ceratosaurus (Dinosauria, Therapoda), a Revised Osteology. Miscellaneous Publication. Utah Geological Survey. ISBN 1-55791-380-3
15. Britt, B. B.; Chure, D. J.; Holtz, T. R., Jr.; Miles, C. A.; Stadtman, K. L. (2000). "A reanalysis of the phylogenetic affinties of Ceratosaurus (Theropoda, Dinosauria) based on new specimens from Utah, Colorado, and Wyoming". Journal of Vertebrate Paleontology 20 (suppl.): 32A. doi:10.1080/02724634.2000.10010765.
16. Rauhut, O. W. M. (2003). The Interrelationships and Evolution of Basal Theropod Dinosaurs. Special Papers in Palaeontology (Wiley). p. 25.
17. Carrano, Matthew T.; Sampson, Scott D. (January 2008). "The Phylogeny of Ceratosauria (Dinosauria: Theropoda)". Journal of Systematic Palaeontology 6: 192. doi:10.1017/S1477201907002246. ISSN 1477-2019.
18. http://dml.cmnh.org/2000May/msg00490.html
19. Rauhut, Oliver W. M. (2011). "Theropod dinosaurs from the Late Jurassic of Tendaguru (Tanzania)". Special Papers in Palaeontology 86: 195–239. doi:10.1111/j.1475-4983.2011.01084.x.
20. Foster, J. (2007). "Appendix." Jurassic West: The Dinosaurs of the Morrison Formation and Their World. Indiana University Press. pp. 327-329.
21. Diego Pol & Oliver W. M. Rauhut (2012). "A Middle Jurassic abelisaurid from Patagonia and the early diversification of theropod dinosaurs". Proceedings of the Royal Society B: Biological Sciences 279 (1804): 3170–5. doi:10.1098/rspb.2012.0660. PMC 3385738. PMID 22628475.
22. Bakker RT, Bir G (2004). "Dinosaur Crime Scene Investigations". In Currie PJ, Koppelhus EB, Shugar MA, Wright JL. Feathered Dragons. Indiana University Press. pp. 301–342. ISBN 0-253-34373-9.
23. Bakker, Robert T.; Bir, Gary (2004). "Dinosaur crime scene investigations: theropod behavior at Como Bluff, Wyoming, and the evolution of birdness". In Currie, Philip J.; Koppelhus, Eva B.; Shugar, Martin A.; and Wright, Joanna L. (eds.). Feathered Dragons: Studies on the Transition from Dinosaurs to Birds. Bloomington and Indianapolis: Indiana University Press. pp. 301–342. ISBN 978-0-253-34373-4.
24. Chure, Daniel J. (2000). "Prey bone utilization by predatory dinosaurs in the Late Jurassic of North America, with comments on prey bone use by dinosaurs throughout the Mesozoic" (PDF). Gaia 15: 227–232. ISSN 0871-5424.
25. Glut, D.F. (1997). "Ceratosaurus". Dinosaurs: The Encyclopedia. McFarland & Company. pp. 266–270. ISBN 0-89950-917-7.
26. Rothschild, B., Tanke, D. H., and Ford, T. L., 2001, Theropod stress fractures and tendon avulsions as a clue to activity: In: Mesozoic Vertebrate Life, edited by Tanke, D. H., and Carpenter, K., Indiana University Press, p. 331-336.
27. Molnar, R. E., 2001, Theropod paleopathology: a literature survey: In: Mesozoic Vertebrate Life, edited by Tanke, D. H., and Carpenter, K., Indiana University Press, p. 337-363.
28. Glut, Donald F.; Brett-Surman, Michael K. (1997). "Dinosaurs and the media". The Complete Dinosaur. Bloomington and Indianapolis: Indiana University Press. pp. 675–706. ISBN 0-253-33349-0.

Further reading