Styracosaurus

Taxobox
name = "Styracosaurus"

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image_width = 250px
image_caption = "Styracosaurus albertensis" skull, American Museum of Natural History.
fossil_range = Late Cretaceous
regnum = Animalia
phylum = Chordata
classis = Sauropsida
superordo = Dinosauria
ordo = Ornithischia
subordo = Cerapoda
infraordo = Ceratopsia
familia = Ceratopsidae
subfamilia = Centrosaurinae
genus = "Styracosaurus"
genus_authority = Lambe, 1913
subdivision_ranks = Species
subdivision =
*"S. albertensis" Lambe, 1913 (type)
*"S. ovatus" Gilmore, 1930

"Styracosaurus" (pronEng|stɪˌrækəˈsɔrəs, meaning "spiked lizard" from Greek "styrax/στυραξ" 'spike at the butt-end of a spear-shaft' and "saurus/σαυρος" 'lizard')cite book|author=Liddell & Scott|year=1980|title=Greek-English Lexicon, Abridged Edition |publisher=Oxford University Press, Oxford, UK|id=ISBN 0-19-910207-4] was a genus of herbivorous ceratopsian dinosaur from the Cretaceous Period (Campanian stage), about 76.5 to 75.0 million years ago. It had four to six long horns extending from its neck frill, a smaller horn on each of its cheeks, and a single horn protruding from its nose, which may have reached dimensions of around 60 centimeters (2 ft) long and 15 centimeters (6 in) wide. The function or functions of the horns and frills have been the subject of debate for many years.

"Styracosaurus" was a large dinosaur, reaching lengths of 5.5 meters (18 ft) and weighing nearly 3 tons. It stood about 1.8 meters (6 ft) tall. "Styracosaurus" possessed four short legs and a bulky body. Its tail was rather short. It also had a beak and flat cheek teeth, indicating that its diet was herbivorous. Like other ceratopsians, this dinosaur may have been a herd animal, traveling in large groups, as suggested by bonebeds.

Named by Lawrence Lambe in 1913, "Styracosaurus" is a member of the Centrosaurinae. Two species, "S. albertensis" and "S. ovatus" are currently assigned to "Styracosaurus". Other species assigned to the genus have since been reassigned elsewhere.

Description

Individuals of the "Styracosaurus" genus were approximately 5.5 m (18 ft) long as adults and weighed around 2.7 tons.Lambert, D. (1993). "The Ultimate Dinosaur Book." Dorling Kindersley: New York, 152–167. ISBN 1-56458-304-X.] The skull was massive, with a large nostril, a tall straight nose horn, and a parietosquasomal frill (a neck frill) crowned with at least four large spikes. Each of the four longest frill spines was comparable in length to the nose horn, at 50 to 55 centimeters long (19.7 to 21.7 in).cite book| last = Dodson| first = P.| authorlink =Peter Dodson | title = The Horned Dinosaurs: A Natural History| publisher = Princeton University Press| year = 1996| location = Princeton| pages = 165–169| isbn = 0-691-05900-4] The nasal horn is estimated at 57 centimeters long (19.7 in) in the type specimen,cite journal |last=Lambe |first=L.M. |year=1913 |title=A new genus and species from the Belly River Formation of Alberta |journal=Ottawa Naturalist |volume=27 |pages=109–116] but the horn is only partially complete. Based on other nasal horn cores from "Styracosaurus" and "Centrosaurus", this horn may have come to a rounded point at around half of that length.cite journal |last=Ryan |first=Michael J. |coauthors=Holmes, Robert; and Russell, A.P. |year=2007 |title=A revision of the late Campanian centrosaurine ceratopsid genus "Styracosaurus" from the Western Interior of North America |journal=Journal of Vertebrate Paleontology |volume=27 |issue=4 |pages=94–962 |doi=10.1671/0272-4634(2007)27 [944:AROTLC] 2.0.CO;2 ] Aside from the large nasal horn and four long frill spikes, the cranial ornamentation was variable. Some individuals had small hook-like projections and knobs at the posterior margin of the frill, similar to but smaller than those in "Centrosaurus". Others had less prominent tabs. Some, like the type individual, had a third pair of long frill spikes. Others had much smaller projections. Second species "S. ovatus" had only four large spikes, but the inner pair curved toward each other, the opposite of what is seen in the much better known "S. albertensis". Other small points are found on the side margins of some but not all specimens. Modest pyramid-shaped brow horns were present in subadults, but were replaced by pits in adults. Like most ceratopsids, "Styracosaurus" had large fenestrae (skull openings) in its frill. The front of the mouth had a toothless beak. The bulky body of "Styracosaurus" resembled that of a rhinoceros. It had powerful shoulders which may have been useful in intraspecies combat. "Styracosaurus" had a relatively short tail. Each toe bore a hooflike ungual which was sheathed in horn.

Various limb positions have been proposed for "Styracosaurus" and ceratopsids in general, including forelegs which were held underneath the body, or, alternately, held in a sprawling position. The most recent work has put forward an intermediate crouched position as most likely.cite web |url=http://palaeo-electronica.org/2007_1/step/index.html |title=Forelimb stance and step cycle in "Chasmosaurus irvinensis" (Dinosauria:Neoceratopsia |accessdate=2007-05-28 |author=Thompson, Stefan |coauthors= and Holmes, Robert |year=2007 |month=April |format=HTML |publisher=Palaeontologia Electronica ] Paleontologists Gregory Paul and Per Christiansen of the Zoological Museum of the University of Copenhagen in Denmark proposed that large ceratopsians such as "Styracosaurus" were able to run at speeds exceeding that of an elephant, based on possible ceratopsian trackways which did not exhibit signs of sprawling forelimbs. [cite journal| last = Paul| first = Gregory| authorlink =| coauthors = Per Christiansen| title = Forelimb posture in neoceratopsian dinosaurs: implications for gait and locomotion | journal = Paleobiology| volume = 26| issue = 3| pages = 450–465| publisher = BioOne|month=September | year=2000| url = http://www.bioone.org/perlserv/?request=get-document&doi=10.1666%2F0094-8373(2000)026%3C0450%3AFPINDI%3E2.0.CO%3B2 |id =doi| id = 10.1666/0094-8373(2000)026%3C0450:FPINDI%3E2.0.CO;2 | label = 10.1666/0094-8373(2000)026<0450:FPINDI>2.0.CO;2 | accessdate = 2007-02-25| doi = 10.1666/0094-8373(2000)026<0450:FPINDI>2.0.CO;2| format = subscription required| year = 2000 ]

Classification

"Styracosaurus" is a member of the Centrosaurinae, a subfamily of large North American horned dinosaurs characterized by their "prominent nasal horns, subordinate brow horns, short squamosals in a short frill, a tall, deep face relative to the ceratopines, and a projection into the rear of the nasal fenestra."cite web| last =Tweet| first =J.| authorlink =| coauthors =| title =Centrosaurinae| work =Thescelosaurus| publisher =Qwest.net| year =2007| url =http://www.users.qwest.net/~jstweet1/centrosaurinae.htm| format =| doi =| accessdate =2007-04-22] Other members of the clade include "Centrosaurus" (from which the group takes its name),cite book |last=Dodson |first=P. |year=1990 |chapter=On the status of the ceratopsids "Monoclonius" and "Centrosaurus" |editor=Carpenter, K.; and Currie, P.J. (eds.) |title=Dinosaur Systematics: Perspectives and Approaches |publisher=Cambridge University Press |location=Cambridge |pages=231–243 |isbn=0-521-36672-0] cite journal| last =Ryan| first =M.J.| authorlink =| coauthors =A.P. Russell| title =New centrosaurine ceratopsids from the late Campanian of Alberta and Montana and a review of contemporaneous and regional patterns of centrosaurine evolution| journal =Journal of Vertebrate Paleontology | volume =23| issue =3| pages =| publisher =| year =2003 ] "Pachyrhinosaurus",cite journal| last =Ryan| first =M.J.| authorlink =| coauthors =A.P. Russell| title =A new centrosaurine ceratopsid from the Oldman Formation of Alberta and its implications for centrosaurine taxonomy and systematics| journal =Canadian Journal of Earth Sciences | volume =42| issue =| pages =1369–1387| publisher =| year =2005| doi =10.1139/e05-029] "Avaceratops", "Einiosaurus", "Albertaceratops",cite journal| last =Ryan| first =M.J.| authorlink =| coauthors =| title =A new basal centrosaurine ceratopsid from the Oldman Formation, southeastern Alberta| journal =Journal of Paleontology| volume =81| issue =2| pages =376–396 | publisher =| url =| doi =10.1666/0022-3360(2007)81 [376:ANBCCF] 2.0.CO;2| id =| accessdate =| year =2007 ] "Achelousaurus", "Brachyceratops",Dodson, P., Forster, C. A, and Sampson, S. D. (2004) "Ceratopsidae". In: Weishampel, D. B., Dodson, P., and Osmólska, H. (eds.), "The Dinosauria" (second edition). University of California Press:Berkeley, pp. 494–513. ISBN 0-520-24209-2.] and "Monoclonius", although these last two are dubious. Because of the variation between species and even individual specimens of centrosaurines, there has been much debate over which genera and species are valid, particularly whether "Centrosaurus" and/or "Monoclonius" are valid genera, undiagnosable, or possibly members of the opposite sex. In 1996, Peter Dodson found enough variation between "Centrosaurus", "Styracosaurus", and "Monoclonius" to warrant separate genera, and that "Styracosaurus" resembled "Centrosaurus" more closely than either resembled "Monoclonius". Dodson also believed one species of "Monoclonius", "M. nasicornis", may actually have been a female "Styracosaurus".Dodson, P. (1996). "The Horned Dinosaurs: A Natural History". Princeton University Press: Princeton, New Jersey, p. 197–199. ISBN 0-691-02882-6.] His assessments have been partially followed, with other researchers not accepting "Monoclonius nasicornis" as a female "Styracosaurus", or "Monoclonius" as a valid genus.cite book |last=Ryan |first=Michael J. |coauthors=and Evans, David C. |editors=Currie, Phillip J., and Koppelhus, Eva |title=Dinosaur Provincial Park: A Spectacular Ancient Ecosystem Revealed |chapter=Ornithischian Dinosaurs |year=2005 |publisher=Indiana University Press |location=Bloomington |pages=312–348 |isbn=0-253-34595-2 ] While sexual dimorphism has been proposed for an earlier ceratopsian "Protoceratops", [cite journal | last = Dodson | first = P. | title = Quantitative aspects of relative growth and sexual dimorphism in "Protoceratops" | journal = Journal of Paleontology | volume = 50 | pages = 929–940 | doi =] there is no firm evidence for sexual dimorphism in any ceratopsids.Forster, C. A. (1990). The cranial morphology and systematics of "Triceratops", with a preliminary analysis of ceratopsian phylogeny. Ph.D. Dissertation. University of Pennsylvania, Philadelphia. 227 pp.] cite journal|last=Lehman|first=T. M.|year=1998|title=A gigantic skull and skeleton of the horned dinosaur "Pentaceratops sternbergi" from New Mexico|journal=Journal of Paleontology|volume=72|issue=5|pages=894–906] cite journal |last=Sampson |first=S. D. |coauthors=Ryan, M.J.; and Tanke, D.H. |year=1997 |title=Craniofacial ontogeny in centrosaurine dinosaurs (Ornithischia: Ceratopsidae): taphonomic and behavioral phylogenetic implications |journal=Zoological Journal of the Linnean Society |volume=121 |pages=293–337]

Origins

The evolutionary origins of "Styracosaurus" were not understood for many years because fossil evidence for early ceratopsians was sparse. The discovery of "Protoceratops", in 1922, shed light on early ceratopsid relationships,Dodson, P. (1996). "The Horned Dinosaurs: A Natural History". Princeton University Press: Princeton, New Jersey, p. 244. ISBN 0-691-02882-6.] but several decades passed before additional finds filled in more of the blanks. Fresh discoveries in the late 1990s and 2000s, including "Zuniceratops", the earliest known ceratopsian with brow horns, and "Yinlong", the first known Jurassic ceratopsian, indicate what dinosaurs related to the ancestors of "Styracosaurus" may have looked like. These new discoveries have been important in illustrating the origins of horned dinosaurs in general, and suggest that the group originated during the Jurassic in Asia, and the appearance of truly horned ceratopsians by the beginning of the late Cretaceous in North America.Dodson, P., Forster, C. A, and Sampson, S. D. (2004) "Ceratopsidae". In: Weishampel, D. B., Dodson, P., and Osmólska, H. (eds.), "The Dinosauria" (second edition). University of California Press:Berkeley, pp. 494–513. ISBN 0-520-24209-2.]

Discoveries and species

The first fossil remains of "Styracosaurus" were collected in Alberta, Canada by C.M. Sternberg (from an area now known as the Dinosaur Provincial Park, in a formation now called the Dinosaur Park Formation) and named by Lawrence Lambe in 1913. This quarry was revisited in 1935 by a Royal Ontario Museum crew who found the missing lower jaws and most of the skeleton. These fossils indicate that "S. albertensis" was around 5.5 to 5.8 meters in length and stood about 1.65 meters high at the hips. An unusual feature of this first skull is that the smallest frill spike on the left side is partially overlapped at its base by the next spike. It appears that the frill suffered a break at this point in life and was shortened by about 6 centimeters (2 in). The correct shape of this area is unknown because the corresponding area of the right side of the frill was not recovered.

Barnum Brown and Erich Maren Schlaikjer, working for the American Museum of Natural History in New York, collected a nearly complete articulated skeleton with a partial skull in 1915. These fossils were also found in the Dinosaur Park Formation, near Steveville, Alberta. Brown and Schlaikjer compared the finds, and, though they allowed that both specimens were from the same general locality and geological formation, they considered the specimen sufficiently distinct from the holotype to warrant erecting a new species, and described the fossils as "Styracosaurus parksi", named in honor of William Parks.cite journal| last = Brown| first = Barnum| authorlink =| coauthors = Erich Maren Schlaikjer| title = The skeleton of "Styracosaurus" with the description of a new species| journal = American Museum novitates| volume = no. 955| pages = 12| publisher = New York City : The American Museum of Natural History| year = 1937| url = http://digitallibrary.amnh.org/dspace/handle/2246/2191| accessdate =2007-03-29] Among the differences between the specimens cited by Brown and Schlaikjer were a cheekbone quite different from that of "S. albertensis", and smaller tail vertebrae. "S. parksi" also had a more robust jaw, a shorter dentary, and the frill differed in shape from that of the type species. However, much of the skull consisted of plaster reconstruction, and the original 1937 paper did not illustrate the actual skull bones. It is now accepted as a specimen of "S. albertensis".

In the summer of 2006, Darren Tanke of the Royal Tyrrell Museum of Palaeontology in Drumheller, Alberta relocated the long lost "S. parksi" site. Pieces of the skull, evidently abandoned by the 1915 crew, were found in the quarry. These were collected and it is hoped more pieces will be found, perhaps enough to warrant a redescription of the skull and test whether "S. albertensis" and "S. parksi" are the same. The Tyrrell Museum has also collected several partial "Styracosaurus" skulls. At least one confirmed bonebed (bonebed 42) in Dinosaur Provincial Park has also been explored (other proposed "Styracosaurus" bonebeds instead have fossils from a mix of animals, and nondiagnostic ceratopsian remains). Bonebed 42 is known to contain numerous pieces of skulls such as horncores, jaws and frill pieces.

A third species, "S. ovatus", from the Two Medicine Formation of Montana, was described by Gilmore in 1930. The fossil material is limited, with the best being a portion of the parietal bone of the frill, but one unusual feature is that the pair of spikes closest to the midline converge towards the midline, rather than away from it as in "S. albertensis". There also may only have been two sets of spikes on each side of the frill, instead of three. The spikes are much shorter than in "S. albertensis", with the longest only 295 millimeters (11.6 in) long.cite journal |last=Gilmore |first=Charles W. |authorlink=Charles W. Gilmore |year=1930 |title=On dinosaurian reptiles from the Two Medicine Formation of Montana |journal=Proceedings of the United States National Museum |volume=77 |issue=16 |pages=1–39 ] Recent review of styracosaur skull remains by Ryan, Holmes, and Russell finds it to be a distinct species.

Several other species which were assigned to "Styracosaurus" have since been assigned to other genera. "S. sphenocerus", described by Edward Drinker Cope in 1890 as a species of "Monoclonius" and based on a nasal bone with a broken "Styracosaurus"-like straight nose horn, was attributed to "Styracosaurus" in 1915.cite journal|last=Lambe|first=L. M.|year=1915|title=On "Eoceratops canadensis", gen. nov., with remarks on other genera of Cretaceous horned dinosaurs|journal=Canada Geological Survey Bulletin, Geological Series|volume=12|issue=24|pages=1–49] "S. makeli", mentioned informally by amateur paleontologists Stephen and Sylvia Czerkas in 1990 in a caption to an illustration, is an early name for "Einiosaurus".cite book|chapter=Einiosaurus |last=Glut |first=Donald F. |title=Dinosaurs: The Encyclopedia |year=1997 |publisher=McFarland & Co |location=Jefferson, North Carolina|pages=396–398 |isbn=0-89950-917-7] "S. borealis" is an early informal name for "S. parksi".cite book|chapter=Styracosaurus |last=Glut |first=Donald F. |title=Dinosaurs: The Encyclopedia |year=1997 |publisher=McFarland & Co |location=Jefferson, North Carolina|pages=865–868 |isbn=0-89950-917-7]

Paleobiology

"Styracosaurus" and other horned dinosaurs are often depicted in popular culture as herding animals. A bonebed composed of "Styracosaurus" remains is known from the Dinosaur Park Formation of Alberta, about halfway up the formation. This bonebed is associated with different types of river deposits.cite book |last=Eberth |first=David A.|coauthors=and Getty, Michael A. |editors=Currie, Phillip J., and Koppelhus, Eva |title=Dinosaur Provincial Park: A Spectacular Ancient Ecosystem Revealed |chapter=Ceratopsian bonebeds: occurrence, origins, and significance |year=2005 |publisher=Indiana University Press |location=Bloomington |pages=501–536 |isbn=0-253-34595-2 ] [Although this article mentioned two bonebeds, including BB 156, the recent review by Ryan "et al." only accepted BB 42.] The mass deaths may have been a result of otherwise non-herding animals congregating around a waterhole in a period of drought, with evidence suggesting the environment may have been seasonal and semiarid.cite journal|last=Rogers|first=R. R.|year=1990|title=Taphonomy of three dinosaur bone beds in the Upper Cretaceous Two Medicine Formation, northwestern Montana: Evidence for drought-related mortality|journal=Palaios|volume=5|pages=394–41|doi=10.2307/3514834]

"Styracosaurus" is known from higher in the formation than the closely-related "Centrosaurus", suggesting that "Styracosaurus" displaced "Centrosaurus" as the environment changed over time.cite book |last=Ryan |first=Michael J. |coauthors=and Evans, David C. |editors=Currie, Phillip J., and Koppelhus, Eva |title=Dinosaur Provincial Park: A Spectacular Ancient Ecosystem Revealed |chapter=Ornithischian Dinosaurs |year=2005 |publisher=Indiana University Press |location=Bloomington |pages=312–348 |isbn=0-253-34595-2 ]

Dentition and diet

"Styracosaurus" were herbivorous dinosaurs; they probably fed mostly on low growth because of the head's position. They may, however, have been able to knock down taller plants with their horns, beak, and bulk. [Tait J., and Brown, B. (1928). How the Ceratopsia carried and used their head. "Transactions of the Royal Society of Canada." 22:13–23.] The jaws were tipped with a deep, narrow beak, believed to have been better at grasping and plucking than biting.cite journal|author=Ostrom, J. H.|year=1966| title=Functional morphology and evolution of the ceratopsian dinosaurs| journal=Evolution| volume=20| issue=3| pages=290–308| doi=10.2307/2406631]

Ceratopsid teeth, including those of "Styracosaurus", were arranged in groups called batteries. Older teeth on top were continually replaced by the teeth underneath them; this occurred throughout the life of the animal. Unlike hadrosaurids, which also had dental batteries, ceratopsid teeth sliced but did not grind. Some scientists have suggested ceratopsids like "Styracosaurus" ate palms and cycads, [Weishampel, D. B. (1984). Evolution of jaw mechanisms in ornithopod dinosaurs. "Advances in Anatomy, Embryology, and Cell Biology" 87:1–110.] while others have suggested ferns. [Coe, M. J., Dilcher, D. L., Farlow, J. O., Jarzen, D. M., and Russell, D. A. (1987). Dinosaurs and land plants. In: Friis, E. M., Chaloner, W. G., and Crane, P. R. (eds.) "The Origins of Angiosperms and their Biological Consequences" Cambridge University Press, pp. 225–258. ISBN 0-521-32357-6.] Dodson has proposed that Late Cretaceous ceratopsians may have knocked down angiosperm trees and then sheared off leaves and twigs.Dodson, P. (1996). "The Horned Dinosaurs: A Natural History". Princeton University Press: Princeton, New Jersey, p. 266. ISBN 0-691-02882-6.]

Horns and frill

The large nasal horns and frills of "Styracosaurus" are among the most distinctive facial adornments of all dinosaurs. Their function has been the subject of debate ever since the first horned dinosaurs were discovered.

Early in the 20th century, paleontologist R. S. Lull put forth the hypothesis that the frills of ceratopsian dinosaurs acted as anchor points for their jaw muscles.cite journal| last =Lull| first=R.S. | authorlink=R. S. Lull | title =The cranial musculature and the origin of the frill in the ceratopsian dinosaurs| journal =American Journal of Science| volume =4| issue =25| pages =387–399| year=1908 ] He later noted that for "Styracosaurus", the spikes would have given it a formidable appearance.cite journal |last=Lull |first=R.S. |authorlink=R. S. Lull |year=1933 |title=A revision of the Ceratopsia or horned dinosaurs |journal=Memoirs of the Peabody Museum of Natural History |volume=3 |issue=3 |pages=1–175] In 1996, Dodson supported the idea of muscle attachments in part and created detailed diagrams of possible muscle attachments in the frills of "Styracosaurus" and "Chasmosaurus", but did not subscribe to the idea that they completely filled in the fenestrae.Dodson, P. (1996). "The Horned Dinosaurs: A Natural History". Princeton University Press: Princeton, New Jersey, p. 269. ISBN 0-691-02882-6.] C.A. Forster, however, found no evidence of large muscle attachments on the frill bones.Forster, C. A. (1990). The cranial morphology and systematics of "Triceratops", with a preliminary analysis of ceratopsian phylogeny. Ph.D. Dissertation. University of Pennsylvania, Philadelphia. 227 pp. OCLC 61500040 ]

It was long believed ceratopsians like "Styracosaurus" used their frills and horns in defence against the large predatory dinosaurs of the time. Although pitting, holes, lesions, and other damage on ceratopsid skulls are often attributed to horn damage in combat, a 2006 study found no evidence for horn thrust injuries causing these forms of damage (for example, there is no evidence of infection or healing). Instead, non-pathological bone resorption, or unknown bone diseases, are suggested as causes.Tanke, D. H, and Farke, A. A. (2006). Bone resorption, bone lesions, and extracranial fenestrae in ceratopsid dinosaurs: a preliminary assessment. in: Carpenter, K. (ed.). "Horns and Beaks: Ceratopsian and Ornithopod Dinosaurs" Indiana University Press: Bloomington. pp. 319–347. ISBN 0-253-34817-X.]

The large frill on "Styracosaurus" and related genera also may have helped to increase body area to regulate body temperature, [cite journal|author=Wheeler, P.E.|year=1978|title=Elaborate CNS cooling structures in large dinosaurs|journal=Nature|issue=275|pages=441–443|doi=10.1038/275441a0|volume=275] like the ears of the modern elephant. A similar theory has been proposed regarding the plates of "Stegosaurus", [cite journal|author=Farlow, J. O., Thompson, C. V., and Rosner, D. E.|year=1976| title=Plates of the dinosaur "Stegosaurus": Forced convection heat loss fins?| journal=Science| volume=192| pages=1123&ndash;1125| doi=10.1126/science.192.4244.1123| pmid=17748675] although this use alone would not account for the bizarre and extravagant variation seen in different members of the Ceratopsidae. This observation is highly suggestive of what is now believed to be the primary function, display.

The theory of frill use in sexual display was first proposed in 1961 by Davitashvili. This theory has gained increasing acceptance.cite book|title=The Theory of sexual selection|year=1961|author=Davitashvili L|pages= 538|publisher=Izdatel'stvo Akademii Nauk SSSR] Evidence that visual display was important, either in courtship or in other social behavior, can be seen in the fact that horned dinosaurs differ markedly in their adornments, making each species highly distinctive. Also, modern living creatures with such displays of horns and adornments use them in similar behavior. [cite journal|author=Farlow, J. O., and Dodson, P.|year=1975|title=The behavioral significance of frill and horn morphology in ceratopsian dinosaurs|journal=Evolution|volume=29|issue=2|pages=353–361| doi=10.2307/2407222]

In popular culture

Because of the distinctive frill and horns of "Styracosaurus", depictions of these animals are easily recognizable. The spines, hooks, and horns attached to the head of this dinosaur sparked the imagination of filmmakers during the earliest days of motion pictures, and this has led to its appearance in films ever since. Notable among them are: "The Son of Kong" (1933), where a "Styracosaurus" battles the movie's heroes;cite web| last =Erickson| first =G. | authorlink =| coauthors =| title =DVD Savant Review: "The Son of Kong"| publisher =DVD Talk.com| year =2005| url =http://www.dvdtalk.com/dvdsavant/s1827son.html| accessdate =2007-04-24] "The Valley of Gwangi" (1969), where "Styracosaurus" is pitted against a carnivorous dinosaur;cite web| last =Perez| first =D.| title =The Styracosaurus 1969| publisher =The Seventh Voyage.com| year =2006| url =http://theseventhvoyage.com/Styracosaurus.htm| accessdate =2007-04-23] and Disney's CGI film "Dinosaur" (2000), where an anthropomorphic "Styracosaurus" named Eema has a pet "Ankylosaurus".cite web| last =Kobak| first =S.| title =Dinosaur| publisher =Films on Disc|year=2002 |url=http://www.filmsondisc.com/dvdpages/dinosaur_dvd_review.htm| accessdate =2007-04-23] The genus also appeared in the novel of "Jurassic Park", in the list of dinosaurs present in the park, but was not seen in the film adaptation.

On the small screen, "Styracosaurus" has appeared in many cartoons and anime, including ' ("Bakuryuu Sentai Abaranger"), "Dinozaurs", "Dino-Riders", "Dinosaucers",cite web| last =| first =| authorlink =| coauthors =| title =Dinosaucers Cast and Crew| work =| publisher =TV.com| year =2007| url =http://www.tv.com/dinosaucers/show/9208/cast.html| accessdate =2007-04-28] and "Zoids".cite web| title =Red Horn| work =Zoids Tribute| publisher =Mopock.com| year =2001| url =http://mopock.com/ZOIDS/photos/RedHorn.html| accessdate =2007-04-28] "Styracosaurus" has also been featured in various video games, including the popular "Jurassic Park", "Turok", and ' game franchises. It was also a huntable dinosaur in the dinosaur hunting game "Primal Prey".

"Styracosaurus" has appeared in some dinosaur scale model lines, such as the Carnegie Collection.

References

External links

* [http://www.dinoruss.com/de_4/5a795cb.htm "Styracosaurus", from the Dinosaur Encyclopaedia]