Difference between revisions of "Triceratops"

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Several varieties of Triceratops are known, from a large number of [[fossil]] remains. They were among the last of the dinosaurs.
 
Several varieties of Triceratops are known, from a large number of [[fossil]] remains. They were among the last of the dinosaurs.
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Triceratops (IPA: /tɹaɪ'sɛɹətɒps/) was a herbivorous genus of ceratopsid dinosaur that lived during the late Maastrichtian (end of the Late Cretaceous Period) around 68 to 65 million years ago (mya) in what is now North America; it was one of the last dinosaurs to appear before the great Cretaceous-Tertiary extinction event.[1] Bearing a large bony frill and three horns on its large four-legged body, and conjuring similarities with the modern rhinoceros, Triceratops is one of the most recognizable of all dinosaurs. The name Triceratops, which literally means "three-horned face", is derived from the Greek tri/τρι- meaning "three", ceras/κέρας meaning "horn", and -ops/ωψ meaning "face".[2] Though it shared the landscape with, and was preyed upon by, the fearsome Tyrannosaurus, it is unclear whether the two battled the way they are commonly depicted in movies and children's dinosaur books.
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Although no complete skeleton has been found,[3] Triceratops is well-known from numerous partial specimens collected since the introduction of the genus in 1887. The function of their frills and three distinctive facial horns has long inspired debate. Although traditionally viewed as defensive weapons against predators, the latest theories suggest these features were primarily used in display for courtship and dominance, much like the antlers and horns of modern reindeer, mountain goats or rhinoceros beetles.[4]
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Triceratops is the best-known of the ceratopsids, though the genus' exact placement within the group has been a point of contention amongst paleontologists. Two species, T. horridus and T. prorsus, are considered valid, although many other species have been named.
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Contents [hide]
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1 Description
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2 Classification
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2.1 Use in phylogenetics
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2.2 Origins
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3 Discoveries and species
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3.1 Number of species
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3.2 Valid species
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3.3 Doubtful species
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3.4 Misassignments
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4 Paleobiology
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4.1 Dentition and diet
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4.2 Functions of the horns and frill
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5 Depiction in recent popular media
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6 References
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7 External links
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[edit] Description
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Triceratops compared in size with a humanIndividual Triceratops have been estimated to reach about 8 m (26 ft) long, 3 m (10 ft) tall,[4] and weigh 6.1 tonnes (13,400 lb).[5] The most distinctive feature is their large skull, one of the largest of all land animals. It could grow to be over 2 m (6.6 ft) in length,[4] and could reach almost a third of the length of the entire animal.[3] It bore a single horn on the snout, above the nostrils, and a pair of horns approximately 1 m (3.3 ft) long, above the eyes. To the rear of the skull was a relatively short bony frill. Most other frilled dinosaurs had large fenestrae in their frills, while the frills of Triceratops were noticeably solid.
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Triceratops species possessed a sturdy build, with robust limbs, and short five-hoofed hands and four-hoofed feet.[6] Although certainly quadrupedal, the posture of these dinosaurs has long been the subject of some debate. Originally, it was believed that the front legs of the animal had to be sprawling at angles from the thorax, in order to better bear the weight of the head.[4] This stance can be seen in paintings by Charles Knight and Rudolph Zallinger. Ichnological evidence in Triceratops trackways, however, and recent reconstructions of skeletons (both physical and digital), seem to show that Triceratops maintained an upright stance during normal locomotion, with the elbows slightly bowed out, in an intermediate state between fully upright and fully sprawling (as in the modern rhinoceros).[7][8] This conclusion does not preclude a sprawling gait for confrontations or feeding.
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[edit] Classification
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Triceratops is the best known genus of the Ceratopsidae, a family of large North American horned dinosaurs. The exact location of Triceratops among the ceratopsians has been debated over the years. Confusion stemmed mainly from the combination of short, solid frills (similar to that of Centrosaurinae), and the long brow horns (more akin to Ceratopsinae, also known as Chasmosaurinae). In the first overview of horned dinosaurs, R. S. Lull hypothesized two lineages, one of Monoclonius and Centrosaurus leading to Triceratops, the other with Ceratops and Torosaurus, making Triceratops a centrosaurine as the group is understood today.[9] Later revisions supported this view, formally describing the first, short-frilled group as Centrosaurinae (including Triceratops), and the second, long-frilled group as Chasmosaurinae.[10][11]
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In 1949, C. M. Sternberg was the first to question this and favoured instead that Triceratops was more closely related to Arrhinoceratops and Chasmosaurus based on skull and horn features, making Triceratops a ceratopsine (chasmosaurine of his usage) genus.[12] However, he was largely ignored with John Ostrom,[13] and later David Norman both placing Triceratops within Centrosaurinae.[14]
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Subsequent discoveries and analyses upheld Sternberg's view on the position of Triceratops, with Lehman defining both subfamilies in 1990 and diagnosing Triceratops as ceratopsine (chasmosaurine of his usage) on the basis of several morphological features. In fact, it fit well into the ceratopsine subfamily, apart from its one feature of a shortened frill.[15] Further research by Peter Dodson, both a 1990 cladistic analysis[16] and a 1993 study using RFTRA (resistant-fit theta-rho analysis),[17] a morphometric technique which systematically measures similarities in skull shape, reinforce Triceratops' placement in the ceratopsine subfamily.
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[edit] Use in phylogenetics
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In phylogenetic taxonomy, the genus has been used as a reference point in the definition of Dinosauria; Dinosaurs have been designated as all descendants of the most recent common ancestor of Triceratops and Neornithes (i.e. modern birds).[18] Furthermore, the bird-hipped dinosaurs, Ornithischia, have been designated as all dinosaurs with a more recent common ancestor to Triceratops than modern birds.[19]
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[edit] Origins
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For many years the origins of Triceratops have been largely obscure. In 1922, the newly discovered Protoceratops was seen as its ancestor by Henry Fairfield Osborn,[20] but many decades passed before additional findings came to light. However, recent years have been fruitful for the discovery of several dinosaurs related to ancestors of Triceratops. Zuniceratops, the earliest known ceratopsian with brow horns, was described in the late 1990s, and Yinlong, the first known Jurassic ceratopsian, in 2005.
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These new finds have been vital in illustrating the origins of horned dinosaurs in general, suggesting an Asian origin in the Jurassic, and the appearance of truly horned ceratopsians by the beginning of the late Cretaceous in North America.[6] As Triceratops is increasingly shown to be a member of the long-frilled Ceratopsinae subfamily, a likely ancestor may have resembled Chasmosaurus, which thrived some 5 million years earlier.
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[edit] Discoveries and species
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Triceratops skull, showing horns and frill (neck armour?), Oxford University Museum of Natural HistoryThe first named specimen now attributed to Triceratops is a pair of brow horns attached to a skull roof, found near Denver, Colorado in the spring of 1887.[21] This specimen was sent to Othniel Charles Marsh, who believed that the formation from which it came dated from the Pliocene, and that the bones belonged to a particularly large and unusual bison, which he named Bison alticornis.[22][21] He realized that there were horned dinosaurs by the next year, which saw his publication of the genus Ceratops from fragmentary remains,[23] but he still believed B. alticornis to be a Pliocene mammal. It took a third and much more complete skull to change his mind. The specimen, collected in 1888 by John Bell Hatcher from the Lance Formation of Wyoming, was initially described as another species of Ceratops,[24] but after reflection Marsh changed his mind and gave it the generic name Triceratops, accepting his Bison alticornis as another species of Ceratops[25] (it would later be added to Triceratops[9]). The sturdy nature of the animal's skull has ensured that many examples have been preserved as fossils, allowing variations between species and individuals to be studied. Triceratops remains have subsequently been found in the American states of Montana and South Dakota (in addition to Colorado and Wyoming), and in the Canadian provinces of Saskatchewan and Alberta.
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[edit] Number of species
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Within the first decades after Triceratops was described, various skulls were collected, which varied to a lesser or greater degree from the original Triceratops, named T. horridus by Marsh (from the Latin horridus; "rough, rugose", suggesting the roughened texture of those bones belonging to the type specimen, later identified as an aged individual). This variation is unsurprising, given that Triceratops skulls are large three-dimensional objects, coming from individuals of different ages and both sexes, and which were subjected to different amounts and directions of pressure during fossilization.[4] Discoverers would name these as separate species (listed below), and came up with several phylogenetic schemes for how they were related to each other.
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In the first attempt to understand the many species, Lull found two groups, although he did not say how he distinguished them: one composed of T. horridus, T. prorsus, and T. brevicornus; the other of T. elatus and T. calicornis. Two species (T. serratus and T. flabellatus) stood apart from these groups.[9] By 1933, and his revision of the landmark 1907 Hatcher-Marsh-Lull monograph of all known ceratopsians, he retained his two groups and two unaffiliated species, with a third lineage of T. obtusus and T. hatcheri that was characterized by a very small nasal horn.[11] T. horridus-T. prorsus-T. brevicornus was now thought to be the most conservative lineage, with an increase in skull size and a decrease in nasal horn size, and T.-elatus-T. calicornis was defined by large brow horns and small nasal horn.[11] C. M. Sternberg made one modification, adding T. eurycephalus and suggesting that it linked the second and third lineages closer together than they were to the T. horridus lineage.[12] This pattern was followed until the major studies of the 1980s and 1990s.
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With time, however, the idea that the differing skulls might be representative of individual variation within one (or two) species gained popularity. In 1986, Ostrom and Wellnhofer published a paper in which they proposed that there was only one species, Triceratops horridus.[26] Part of their rationale was that generally there are only one or two species of any large animal in a region (modern examples being the elephant and the giraffe in modern Africa). To their findings, Lehman added the old Lull-Sternberg lineages combined with maturity and sexual dimorphism, suggesting that the T. horridus-T. prorsus-T. brevicornus lineage was composed of females, the T.calicornis-T.elatus lineage was made up of males, and the T. obtusus-T. hatcheri lineage was of pathologic old males.[15] His reasoning was that males had taller, more erect horns and larger skulls, and females had smaller skulls with shorter, forward-facing horns.
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Triceratops model, Royal Belgian Institute of Natural Sciences, BrusselsThese findings, however, were contested a few years later by Catherine Forster, who reanalysed Triceratops material more comprehensively and concluded that the remains fell into two species, T. horridus and T. prorsus, although the distinctive skull of T. (now tentatively Diceratops) hatcheri differed enough to warrant a separate genus.[27] She found that T. horridus and several other species belonged together, and T. prorsus and T. brevicornus stood alone, and since there were many more specimens in the first group, she suggested that this meant the two groups were two species. It is still possible to interpret this reasoning as describing a single species with sexual dimorphism.[4][28]
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[edit] Valid species
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T. horridus (Marsh, 1889) (originally Ceratops) (type species)
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T. prorsus (Marsh, 1890)
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[edit] Doubtful species
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The following species are considered nomina dubia ("dubious names"), and are based on remains that are too poor or incomplete to be distinguished from pre-existing Triceratops species.
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T. albertensis (C. M. Sternberg, 1949)
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T. alticornis (Marsh, 1887 (originally Bison))
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T. eurycephalus (Schlaikjer, 1935)
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T. galeus (Marsh, 1889)
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T. ingens (Lull, 1915)
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T. maximus (Brown, 1933)
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T. sulcatus (Marsh, 1890)
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[edit] Misassignments
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T. brevicornus (Hatcher, 1905) (=T. prorsus)
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T. calicornus (Marsh, 1898) (=T. horridus)
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T. elatus (Marsh, 1891) (=T. horridus)
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T. flabellatus (Marsh, 1889) (=T. horridus)
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T. hatcheri (Lull, 1907) (=Diceratops hatcheri)
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T. mortuarius (Cope, 1874) (nomen dubium; originally Polyonax; =Polyonax mortuarius)
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T. obtusus (Marsh, 1898) (=T. horridus)
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T. serratus (Marsh, 1890) (=T. horridus)
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T. sylvestris (Cope, 1872) (nomen dubium; originally Agathaumas sylvestris)
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[edit] Paleobiology
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Although Triceratops are commonly portrayed as herding animals, there is currently no solid evidence that they lived in herds. Unlike other horned dinosaurs, some of which are known from sites preserving dozens or hundreds of individuals, all Triceratops finds known at present preserve only solitary individuals.[4] However, these remains are very common; for example, Bruce Erickson, a paleontologist of the Science Museum of Minnesota, has reported having seen 200 specimens of T. prorsus in the Hell Creek Formation of Montana.[29] Similarly, Barnum Brown claimed to have seen over 500 skulls in the field.[30] Because Triceratops teeth, horn fragments, frill fragments, and other skull fragments are such abundant fossils in the Lancian faunal stage of the late Maastrichtian (late Cretaceous, 68 to 65 mya) Period of western North America, it is regarded as among the dominant herbivores of the time, if not the most dominant herbivore; in 1986, Robert Bakker estimated it as making up 5/6ths of the large dinosaur fauna at the end of the Cretaceous.[31]
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Triceratops was one of the last ceratopsians to appear before the Cretaceous-Tertiary extinction event. The related Diceratops and Torosaurus, and the more distantly related diminutive Leptoceratops, were also present, though their remains have been rarely encountered.[4]
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[edit] Dentition and diet
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Triceratops were herbivorous, and because of their low head, their primary food was probably low growth, although they may have been able to knock down taller plants with their horns, beak, and bulk.[32][6] The jaws were tipped with a deep, narrow beak, believed to have been better at grasping and plucking than biting.[33]
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Triceratops teeth were arranged in groups called batteries, of 36 to 40 tooth columns with 3 to 5 stacked teeth per column, depending on the size of the animal.[6] This gives a range of 432 to 800 teeth, of which only a fraction were in use at any given time (tooth replacement was continuous and occurred throughout the life of the animal).[6] They functioned by shearing in a vertical to near-vertical orientation.[6] The great size and numerous teeth of Triceratops suggests that they ate large volumes of fibrous plant material,[6] with some suggesting palms and cycads,[34][35] and others suggesting ferns, which then grew in prairies.[36]
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[edit] Functions of the horns and frill
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Triceratops head from the front
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Triceratops head from the side
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Juvenile and adult skulls — the juvenile skull is about the size of an adult human headThere has been much speculation over the functions of Triceratops' head adornments. The two main theories have revolved around use in combat, or display in courtship, with the latter thought now to be the most likely primary function.[6]
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Early on, Lull postulated that the frills may have served as anchor points for the jaw muscles to aid chewing by allowing increased size and thus power for the muscles.[37] This has been put forward by other authors over the years, but later studies do not find evidence of large muscle attachments on the frill bones.[38]
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Triceratops were long thought to have possibly used their horns and frills in combat with predators such as Tyrannosaurus, the idea being discussed first by C. H. Sternberg in 1917 and 70 years later by Robert Bakker.[39][40]There is evidence that Tyrannosaurus did prey upon them, as a Triceratops pelvis has been found with tyrannosaur toothmarks and subsequent healing, indicating the wound was made while the animal was alive.[41]
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In 2005, a BBC documentary, The Truth About Killer Dinosaurs, tested how Triceratops might have defended themselves against large predators like Tyrannosaurus. To see if Triceratops could have charged other dinosaurs, as would a modern-day rhinoceros, an artificial Triceratops skull was made and propelled into simulated Tyrannosaurus skin at 24 km/h (15 mph). The brow horns penetrated the skin, but the blunt nose horn and the beak could not, and the front of the skull broke. The conclusion drawn was that it would have been impossible for Triceratops to have defended themselves in this way—instead they probably stood their ground when attacked by large predators, using their horns for goring if the predator came close enough.
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In addition to combat with predators using horns, Triceratops are classically shown engaging each other in combat with horns locked. While studies show that such activity would be feasible, if unlike that of present-day horned animals,[42] there is no evidence that they actually did so. Additionally, although pitting, holes, lesions, and other damage on Triceratops skulls (and the skulls of other ceratopsids) are often attributed to horn damage in combat, a recent study finds 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.[43]
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The large frill also may have helped to increase body area to regulate body temperature.[44] A similar theory has been proposed regarding the plates of Stegosaurus,[45] although this use alone would not account for the bizarre and extravagant variation seen in different members of the Ceratopsidae.[6] This observation is highly suggestive of what is now believed to be the primary function, display.
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The theory of their use in sexual display was first proposed by Davitashvili in 1961 and has gained increasing acceptance since.[46][38][15] Evidence that visual display was important, either in courtship or in other social behaviour, 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 behaviour.[47] A recent study of the smallest Triceratops skull, ascertained to be a juvenile, shows the frill and horns developed at a very early age, predating sexual development and thus probably important for visual communication and species recognition in general.[48] The large eyes and shortened features, a hallmark of "cute" baby mammals, also suggest that the parent Triceratops may have cared for its young.
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[edit] Depiction in recent popular media
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Main article: Triceratops in popular culture
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The distinctive appearance of Triceratops has led to them being frequently depicted in various forms of popular culture including films, computer games and documentaries. In more recent media, they have notably appeared in the film Jurassic Park and its sequels, as well as in three major dinosaur documentaries: Walking with Dinosaurs, The Truth About Killer Dinosaurs and Prehistoric Park.
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They are famously known as "three-horns" (and are so named in The Land Before Time animated film and its numerous sequels) due to the three prominent horns on their head and nose, which have become almost synonymous with the dinosaurs. The shorthand "Trike" is another common informal name, and is also the name of the Triceratops character in the children's book series and television cartoon series Harry and His Bucket Full of Dinosaurs.
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A recurring theme in children's dinosaur books is a climactic showdown or battle between Triceratops and T. rex.[49][50][51][52] As such these two dinosaurs are often depicted and thought of as natural enemies. A memorable but anachronistic battle with Ceratosaurus substituting for T. rex is featured in the 1966 movie One Million Years B.C.
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Triceratops appears in video games either derived directly from the Jurassic Park series or similarly themed, namely the 1997 PC games Jurassic Park: Chaos Island and Turok: Dinosaur Hunter, and the 2000 PC and Playstation game Dino Crisis 2. As well, it is a popular creature used in games designed by Nintendo, including Diddy Kong Racing and Starfox Adventures. Triceratops (the species are not identified) is also the official state fossil of South Dakota,[53] and the official state dinosaur of Wyoming.[54]
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[edit] References
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^ Lehman T.M. (1987). "Late Maastrichtian paleoenvironments and dinosaur biogeography in the Western Interior of North America". Paleogeography, Paleoacclimatology and Paleoecology 60: pp. 189–217. 
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^ Liddell & Scott (1980). Greek-English Lexicon, Abridged Edition. Oxford University Press, Oxford, UK. ISBN 0-19-910207-4. 
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^ a b Lambert, D. (1993). The Ultimate Dinosaur Book. Dorling Kindersley, New York, pp. 110—129. ISBN 1-56458-304-X. 
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^ a b c d e f g h Dodson, P. (1996). The Horned Dinosaurs. Princeton University Press, Princeton, New Jersey. ISBN 0-691-02882-6. 
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^ Alexander, R.M. (1985). Mechanics of posture and gait of some large dinosaurs. Zoological Journal of the Linnean Society, 83, 1–25.
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^ a b c d e f g h i 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.
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^ Christiansen, P., and Paul, G.S. (2001). Limb bone scaling, limb proportions, and bone strength in neoceratopsian dinosaurs. Gaia 16:13–29.
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^ Chapman, R.E., Snyder, R.A., Jabo, S., and Andersen, A. (2001). On a new posture for the horned dinosaur Triceratops. Journal of Vertebrate Paleontology 21 (Supplement to Number 3), Abstracts of Papers, 61st Annual Meeting:39A–40A.
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^ a b c Hatcher, J. B., Marsh, O. C. and Lull, R. S. (1907) The Ceratopsia. Government Printing Office, Washington, D.C. ISBN 0405127138.
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^ Lambe, L.M. (1915). On Eoceratops canadensis, gen. nov., with remarks on other genera of Cretaceous horned dinosaurs. Canada Department of Mines Geological Survey Museum Bulletin 12:1–49.
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^ a b c Lull, R. S. (1933) A revision of the Ceratopsia or horned dinosaurs. Memoirs of the Peabody Museum of Natural History 3(3):1–175.
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^ a b Sternberg, C. M. (1949). The Edmonton fauna and description of a new Triceratops from the Upper Edmonton member; phylogeny of the Ceratopsidae. National Museum of Canada Bulletin 113:33–46.
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^ Ostrom, J. H. (1966). Functional morphology and evolution of the ceratopsian dinosaurs. Evolution 20:220–227.
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^ Norman, David (1985). The Illustrated Encyclopaedia of Dinosaurs. London: Salamander Books. ISBN 0-517-468905. 
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^ a b c Lehman, T. M. (1990). The ceratopsian subfamily Chasmosaurinae: sexual dimorphism and systematics. in: Carpenter, K., and Currie, P. J. (eds.). Dinosaur Systematics: Perspectives and Approaches, Cambridge University Press, Cambridge, pp. 211–229. ISBN 0-521-36672-0.
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^ Dodson, P., and Currie, P. J. (1990). Neoceratopsia. 593–618. in Weishampel, D. B., Dodson, P., & Osmólska, H. (eds.). The Dinosauria University of California Press:Berkeley, pp. 593-618. ISBN 0-520-06727-4.
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^ Dodson, P. (1993). Comparative Craniology of the Ceratopsia. In: American Journal of Science 293, pp. 200–234.
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^ Gauthier, J. A. (1986). Saurischian monophyly and the origin of birds. The Origin of Birds and the Evolution of Flight, K. Padian (ed.), Memoirs of the California Academy of Sciences 8:1–55.
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^ Sereno, P. C. (1998). A rationale for phylogenetic definitions, with application to the higher-level taxonomy of Dinosauria. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 210(1):41–83.
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^ Dodson, P. (1996). The Horned Dinosaurs. Princeton University Press:Princeton, New Jersey, p. 244. ISBN 0-691-02882-6.
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^ a b Carpenter, K. (2006). "Bison" alticornis and O.C. Marsh's early views on ceratopsians. In: Carpenter, K. (ed.). Horns and Beaks: Ceratopsian and Ornithopod Dinosaurs Indiana University Press:Bloomington and Indianapolis, pp. 349–364. ISBN 0-253-34817-X.
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^ Marsh, O.C. (1887). Notice of new fossil mammals. American Journal of Science 34:323–331.
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^ Marsh, O.C. (1888). A new family of horned Dinosauria, from the Cretaceous. American Journal of Science 36:477–478.
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^ Marsh, O.C. (1889a). Notice of new American Dinosauria. American Journal of Science 37:331–336.
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^ Marsh, O.C. (1889b). Notice of gigantic horned Dinosauria from the Cretaceous. American Journal of Science 38:173–175.
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^ Ostrom, J. H., and Wellnhofer, P. (1986). The Munich specimen of Triceratops with a revision of the genus. Zitteliana 14:111–158.
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^ Forster, C.A. (1996). Species resolution in Triceratops: cladistic and morphometric approaches. Journal of Vertebrate Paleontology 16(2):259–270.
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^ Lehman, T. M. (1998). A gigantic skull and skeleton of the horned dinosaur Pentaceratops sternbergi from New Mexico. Journal of Paleontology 72(5):894–906.
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^ Erickson, B.R. (1966). Mounted skeleton of Triceratops prorsus in the Science Museum. Scientific Publications of the Science Museum 1:1–16.
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^ Dodson, P. (1996). The Horned Dinosaurs. Princeton University Press:Princeton, New Jersey, p. 79. ISBN 0-691-02882-6.
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^ Bakker, R.T. (1986). The Dinosaur Heresies: New Theories Unlocking The Mystery of the Dinosaurs and Their Extinction William Morrow:New York, p. 438. ISBN 0140100555.
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^ Tait J., and Brown, B. (1928). How the Ceratopsia carried and used their head. Transactions of the Royal Society of Canada. 22:13–23.
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^ Ostrom, J. H. (1966). "Functional morphology and evolution of the ceratopsian dinosaurs". Evolution 20 (3): pp. 290–308. 
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^ Ostrom, J. H. (1964). A functional analysis of jaw mechanics in the dinosaur Triceratops. Postilla, Yale Peabody Museum 88:1–35.
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^ Weishampel, D. B. (1984). Evolution of jaw mechanisms in ornithopod dinosaurs. Advances in Anatomy, Embryology, and Cell Biology 87:1-110.
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^ 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.
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^ Lull, R. S. (1908). The cranial musculature and the origin of the frill in the ceratopsian dinosaurs. American Journal of Science 4(25):387–399.
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^ a b 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.
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^ Sternberg, C. H. (1917). Hunting Dinosaurs in the Badlands of the Red Deer River, Alberta, Canada. Published by the author, San Diego, California, 261 pp.
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^ Bakker, R. T. (1986). The Dinosaur Heresies: New Theories Unlocking The Mystery of the Dinosaurs and Their Extinction William Morrow:New York. ISBN 0140100555.
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^ Fowler, D. W., and Sullivan, R. M. (2006). A ceratopsid pelvis with toothmarks from the Upper Cretaceous Kirtland Formation, New Mexico: evidence of late Campanian tyrannosaurid feeding behavior. New Mexico Museum of Natural History and Science Bulletin 35:127–130.
 +
^ Farke, A. A. (2004). Horn Use in Triceratops (Dinosauria: Ceratopsidae): Testing Behavioral Hypotheses Using Scale Models. Palaeo-electronica.
 +
^ 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.
 +
^ Wheeler, P.E. (1978). "Elaborate CNS cooling structures in large dinosaurs". Nature (275): 441–443. 
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^ Farlow, J. O., Thompson, C. V., and Rosner, D. E. (1976). "Plates of the dinosaur Stegosaurus: Forced convection heat loss fins?". Science 192: pp. 1123–1125. 
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^ Davitashvili L (1961). The Theory of sexual selection. Izdatel'stvo Akademia nauk SSSR, p. 538. 
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^ Farlow, J. O., and Dodson, P. (1975). "The behavioral significance of frill and horn morphology in ceratopsian dinosaurs". Evolution 29 (2): pp. 353–361. 
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^ Goodwin, M. B., Clemens, W. A., Horner, J. R., and Padian, K. (2006). "The smallest known Triceratops skull: new observations on ceratopsid cranial anatomy and ontogeny.". Journal of Vertebrate Paleontology 26 (1): pp. 103–112. 
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^ Geis, D. (1982). How and Why Book of Dinosaurs. Allan Publishers. ISBN 0-824150015.
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^ Watson, J. W. (1981). The Giant Golden Book of Dinosaurs. Random House Children's Books. ISBN 0-307137643.
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^ Pistorius A. (1958) What dinosaur is it? Follett Pub. Co., Chicago.
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^ Wise, W. (1963) In The Time of The Dinosaurs. Scholastic. ISBN 0-590323512.
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^ State of South Dakota. Signs and Symbols of South Dakota. Retrieved on 2007-01-20.
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^ State of Wyoming.

Revision as of 13:50, 21 March 2007

Triceratops was a large North American ceratopsian (horned) dinosaur. It had a large bony frill protecting its neck and three horns - one over each eye and a smaller one on its beaked snout.

Triceratops was herbivorous, up to 26 feet long, and could weigh up to 6 tons. The skull of Triceratops, which is approximately a third of the entire animal's length, is among the largest known for all land animals.

Several varieties of Triceratops are known, from a large number of fossil remains. They were among the last of the dinosaurs. Triceratops (IPA: /tɹaɪ'sɛɹətɒps/) was a herbivorous genus of ceratopsid dinosaur that lived during the late Maastrichtian (end of the Late Cretaceous Period) around 68 to 65 million years ago (mya) in what is now North America; it was one of the last dinosaurs to appear before the great Cretaceous-Tertiary extinction event.[1] Bearing a large bony frill and three horns on its large four-legged body, and conjuring similarities with the modern rhinoceros, Triceratops is one of the most recognizable of all dinosaurs. The name Triceratops, which literally means "three-horned face", is derived from the Greek tri/τρι- meaning "three", ceras/κέρας meaning "horn", and -ops/ωψ meaning "face".[2] Though it shared the landscape with, and was preyed upon by, the fearsome Tyrannosaurus, it is unclear whether the two battled the way they are commonly depicted in movies and children's dinosaur books.

Although no complete skeleton has been found,[3] Triceratops is well-known from numerous partial specimens collected since the introduction of the genus in 1887. The function of their frills and three distinctive facial horns has long inspired debate. Although traditionally viewed as defensive weapons against predators, the latest theories suggest these features were primarily used in display for courtship and dominance, much like the antlers and horns of modern reindeer, mountain goats or rhinoceros beetles.[4]

Triceratops is the best-known of the ceratopsids, though the genus' exact placement within the group has been a point of contention amongst paleontologists. Two species, T. horridus and T. prorsus, are considered valid, although many other species have been named.

Contents [hide] 1 Description 2 Classification 2.1 Use in phylogenetics 2.2 Origins 3 Discoveries and species 3.1 Number of species 3.2 Valid species 3.3 Doubtful species 3.4 Misassignments 4 Paleobiology 4.1 Dentition and diet 4.2 Functions of the horns and frill 5 Depiction in recent popular media 6 References 7 External links


[edit] Description

Triceratops compared in size with a humanIndividual Triceratops have been estimated to reach about 8 m (26 ft) long, 3 m (10 ft) tall,[4] and weigh 6.1 tonnes (13,400 lb).[5] The most distinctive feature is their large skull, one of the largest of all land animals. It could grow to be over 2 m (6.6 ft) in length,[4] and could reach almost a third of the length of the entire animal.[3] It bore a single horn on the snout, above the nostrils, and a pair of horns approximately 1 m (3.3 ft) long, above the eyes. To the rear of the skull was a relatively short bony frill. Most other frilled dinosaurs had large fenestrae in their frills, while the frills of Triceratops were noticeably solid.

Triceratops species possessed a sturdy build, with robust limbs, and short five-hoofed hands and four-hoofed feet.[6] Although certainly quadrupedal, the posture of these dinosaurs has long been the subject of some debate. Originally, it was believed that the front legs of the animal had to be sprawling at angles from the thorax, in order to better bear the weight of the head.[4] This stance can be seen in paintings by Charles Knight and Rudolph Zallinger. Ichnological evidence in Triceratops trackways, however, and recent reconstructions of skeletons (both physical and digital), seem to show that Triceratops maintained an upright stance during normal locomotion, with the elbows slightly bowed out, in an intermediate state between fully upright and fully sprawling (as in the modern rhinoceros).[7][8] This conclusion does not preclude a sprawling gait for confrontations or feeding.


[edit] Classification Triceratops is the best known genus of the Ceratopsidae, a family of large North American horned dinosaurs. The exact location of Triceratops among the ceratopsians has been debated over the years. Confusion stemmed mainly from the combination of short, solid frills (similar to that of Centrosaurinae), and the long brow horns (more akin to Ceratopsinae, also known as Chasmosaurinae). In the first overview of horned dinosaurs, R. S. Lull hypothesized two lineages, one of Monoclonius and Centrosaurus leading to Triceratops, the other with Ceratops and Torosaurus, making Triceratops a centrosaurine as the group is understood today.[9] Later revisions supported this view, formally describing the first, short-frilled group as Centrosaurinae (including Triceratops), and the second, long-frilled group as Chasmosaurinae.[10][11]

In 1949, C. M. Sternberg was the first to question this and favoured instead that Triceratops was more closely related to Arrhinoceratops and Chasmosaurus based on skull and horn features, making Triceratops a ceratopsine (chasmosaurine of his usage) genus.[12] However, he was largely ignored with John Ostrom,[13] and later David Norman both placing Triceratops within Centrosaurinae.[14]

Subsequent discoveries and analyses upheld Sternberg's view on the position of Triceratops, with Lehman defining both subfamilies in 1990 and diagnosing Triceratops as ceratopsine (chasmosaurine of his usage) on the basis of several morphological features. In fact, it fit well into the ceratopsine subfamily, apart from its one feature of a shortened frill.[15] Further research by Peter Dodson, both a 1990 cladistic analysis[16] and a 1993 study using RFTRA (resistant-fit theta-rho analysis),[17] a morphometric technique which systematically measures similarities in skull shape, reinforce Triceratops' placement in the ceratopsine subfamily.


[edit] Use in phylogenetics In phylogenetic taxonomy, the genus has been used as a reference point in the definition of Dinosauria; Dinosaurs have been designated as all descendants of the most recent common ancestor of Triceratops and Neornithes (i.e. modern birds).[18] Furthermore, the bird-hipped dinosaurs, Ornithischia, have been designated as all dinosaurs with a more recent common ancestor to Triceratops than modern birds.[19]


[edit] Origins For many years the origins of Triceratops have been largely obscure. In 1922, the newly discovered Protoceratops was seen as its ancestor by Henry Fairfield Osborn,[20] but many decades passed before additional findings came to light. However, recent years have been fruitful for the discovery of several dinosaurs related to ancestors of Triceratops. Zuniceratops, the earliest known ceratopsian with brow horns, was described in the late 1990s, and Yinlong, the first known Jurassic ceratopsian, in 2005.

These new finds have been vital in illustrating the origins of horned dinosaurs in general, suggesting an Asian origin in the Jurassic, and the appearance of truly horned ceratopsians by the beginning of the late Cretaceous in North America.[6] As Triceratops is increasingly shown to be a member of the long-frilled Ceratopsinae subfamily, a likely ancestor may have resembled Chasmosaurus, which thrived some 5 million years earlier.


[edit] Discoveries and species

Triceratops skull, showing horns and frill (neck armour?), Oxford University Museum of Natural HistoryThe first named specimen now attributed to Triceratops is a pair of brow horns attached to a skull roof, found near Denver, Colorado in the spring of 1887.[21] This specimen was sent to Othniel Charles Marsh, who believed that the formation from which it came dated from the Pliocene, and that the bones belonged to a particularly large and unusual bison, which he named Bison alticornis.[22][21] He realized that there were horned dinosaurs by the next year, which saw his publication of the genus Ceratops from fragmentary remains,[23] but he still believed B. alticornis to be a Pliocene mammal. It took a third and much more complete skull to change his mind. The specimen, collected in 1888 by John Bell Hatcher from the Lance Formation of Wyoming, was initially described as another species of Ceratops,[24] but after reflection Marsh changed his mind and gave it the generic name Triceratops, accepting his Bison alticornis as another species of Ceratops[25] (it would later be added to Triceratops[9]). The sturdy nature of the animal's skull has ensured that many examples have been preserved as fossils, allowing variations between species and individuals to be studied. Triceratops remains have subsequently been found in the American states of Montana and South Dakota (in addition to Colorado and Wyoming), and in the Canadian provinces of Saskatchewan and Alberta.


[edit] Number of species Within the first decades after Triceratops was described, various skulls were collected, which varied to a lesser or greater degree from the original Triceratops, named T. horridus by Marsh (from the Latin horridus; "rough, rugose", suggesting the roughened texture of those bones belonging to the type specimen, later identified as an aged individual). This variation is unsurprising, given that Triceratops skulls are large three-dimensional objects, coming from individuals of different ages and both sexes, and which were subjected to different amounts and directions of pressure during fossilization.[4] Discoverers would name these as separate species (listed below), and came up with several phylogenetic schemes for how they were related to each other.

In the first attempt to understand the many species, Lull found two groups, although he did not say how he distinguished them: one composed of T. horridus, T. prorsus, and T. brevicornus; the other of T. elatus and T. calicornis. Two species (T. serratus and T. flabellatus) stood apart from these groups.[9] By 1933, and his revision of the landmark 1907 Hatcher-Marsh-Lull monograph of all known ceratopsians, he retained his two groups and two unaffiliated species, with a third lineage of T. obtusus and T. hatcheri that was characterized by a very small nasal horn.[11] T. horridus-T. prorsus-T. brevicornus was now thought to be the most conservative lineage, with an increase in skull size and a decrease in nasal horn size, and T.-elatus-T. calicornis was defined by large brow horns and small nasal horn.[11] C. M. Sternberg made one modification, adding T. eurycephalus and suggesting that it linked the second and third lineages closer together than they were to the T. horridus lineage.[12] This pattern was followed until the major studies of the 1980s and 1990s.

With time, however, the idea that the differing skulls might be representative of individual variation within one (or two) species gained popularity. In 1986, Ostrom and Wellnhofer published a paper in which they proposed that there was only one species, Triceratops horridus.[26] Part of their rationale was that generally there are only one or two species of any large animal in a region (modern examples being the elephant and the giraffe in modern Africa). To their findings, Lehman added the old Lull-Sternberg lineages combined with maturity and sexual dimorphism, suggesting that the T. horridus-T. prorsus-T. brevicornus lineage was composed of females, the T.calicornis-T.elatus lineage was made up of males, and the T. obtusus-T. hatcheri lineage was of pathologic old males.[15] His reasoning was that males had taller, more erect horns and larger skulls, and females had smaller skulls with shorter, forward-facing horns.


Triceratops model, Royal Belgian Institute of Natural Sciences, BrusselsThese findings, however, were contested a few years later by Catherine Forster, who reanalysed Triceratops material more comprehensively and concluded that the remains fell into two species, T. horridus and T. prorsus, although the distinctive skull of T. (now tentatively Diceratops) hatcheri differed enough to warrant a separate genus.[27] She found that T. horridus and several other species belonged together, and T. prorsus and T. brevicornus stood alone, and since there were many more specimens in the first group, she suggested that this meant the two groups were two species. It is still possible to interpret this reasoning as describing a single species with sexual dimorphism.[4][28]


[edit] Valid species T. horridus (Marsh, 1889) (originally Ceratops) (type species) T. prorsus (Marsh, 1890)

[edit] Doubtful species The following species are considered nomina dubia ("dubious names"), and are based on remains that are too poor or incomplete to be distinguished from pre-existing Triceratops species.

T. albertensis (C. M. Sternberg, 1949) T. alticornis (Marsh, 1887 (originally Bison)) T. eurycephalus (Schlaikjer, 1935) T. galeus (Marsh, 1889) T. ingens (Lull, 1915) T. maximus (Brown, 1933) T. sulcatus (Marsh, 1890)

[edit] Misassignments T. brevicornus (Hatcher, 1905) (=T. prorsus) T. calicornus (Marsh, 1898) (=T. horridus) T. elatus (Marsh, 1891) (=T. horridus) T. flabellatus (Marsh, 1889) (=T. horridus) T. hatcheri (Lull, 1907) (=Diceratops hatcheri) T. mortuarius (Cope, 1874) (nomen dubium; originally Polyonax; =Polyonax mortuarius) T. obtusus (Marsh, 1898) (=T. horridus) T. serratus (Marsh, 1890) (=T. horridus) T. sylvestris (Cope, 1872) (nomen dubium; originally Agathaumas sylvestris)

[edit] Paleobiology Although Triceratops are commonly portrayed as herding animals, there is currently no solid evidence that they lived in herds. Unlike other horned dinosaurs, some of which are known from sites preserving dozens or hundreds of individuals, all Triceratops finds known at present preserve only solitary individuals.[4] However, these remains are very common; for example, Bruce Erickson, a paleontologist of the Science Museum of Minnesota, has reported having seen 200 specimens of T. prorsus in the Hell Creek Formation of Montana.[29] Similarly, Barnum Brown claimed to have seen over 500 skulls in the field.[30] Because Triceratops teeth, horn fragments, frill fragments, and other skull fragments are such abundant fossils in the Lancian faunal stage of the late Maastrichtian (late Cretaceous, 68 to 65 mya) Period of western North America, it is regarded as among the dominant herbivores of the time, if not the most dominant herbivore; in 1986, Robert Bakker estimated it as making up 5/6ths of the large dinosaur fauna at the end of the Cretaceous.[31]

Triceratops was one of the last ceratopsians to appear before the Cretaceous-Tertiary extinction event. The related Diceratops and Torosaurus, and the more distantly related diminutive Leptoceratops, were also present, though their remains have been rarely encountered.[4]


[edit] Dentition and diet Triceratops were herbivorous, and because of their low head, their primary food was probably low growth, although they may have been able to knock down taller plants with their horns, beak, and bulk.[32][6] The jaws were tipped with a deep, narrow beak, believed to have been better at grasping and plucking than biting.[33]

Triceratops teeth were arranged in groups called batteries, of 36 to 40 tooth columns with 3 to 5 stacked teeth per column, depending on the size of the animal.[6] This gives a range of 432 to 800 teeth, of which only a fraction were in use at any given time (tooth replacement was continuous and occurred throughout the life of the animal).[6] They functioned by shearing in a vertical to near-vertical orientation.[6] The great size and numerous teeth of Triceratops suggests that they ate large volumes of fibrous plant material,[6] with some suggesting palms and cycads,[34][35] and others suggesting ferns, which then grew in prairies.[36]


[edit] Functions of the horns and frill

Triceratops head from the front Triceratops head from the side Juvenile and adult skulls — the juvenile skull is about the size of an adult human headThere has been much speculation over the functions of Triceratops' head adornments. The two main theories have revolved around use in combat, or display in courtship, with the latter thought now to be the most likely primary function.[6]

Early on, Lull postulated that the frills may have served as anchor points for the jaw muscles to aid chewing by allowing increased size and thus power for the muscles.[37] This has been put forward by other authors over the years, but later studies do not find evidence of large muscle attachments on the frill bones.[38]

Triceratops were long thought to have possibly used their horns and frills in combat with predators such as Tyrannosaurus, the idea being discussed first by C. H. Sternberg in 1917 and 70 years later by Robert Bakker.[39][40]There is evidence that Tyrannosaurus did prey upon them, as a Triceratops pelvis has been found with tyrannosaur toothmarks and subsequent healing, indicating the wound was made while the animal was alive.[41]

In 2005, a BBC documentary, The Truth About Killer Dinosaurs, tested how Triceratops might have defended themselves against large predators like Tyrannosaurus. To see if Triceratops could have charged other dinosaurs, as would a modern-day rhinoceros, an artificial Triceratops skull was made and propelled into simulated Tyrannosaurus skin at 24 km/h (15 mph). The brow horns penetrated the skin, but the blunt nose horn and the beak could not, and the front of the skull broke. The conclusion drawn was that it would have been impossible for Triceratops to have defended themselves in this way—instead they probably stood their ground when attacked by large predators, using their horns for goring if the predator came close enough.

In addition to combat with predators using horns, Triceratops are classically shown engaging each other in combat with horns locked. While studies show that such activity would be feasible, if unlike that of present-day horned animals,[42] there is no evidence that they actually did so. Additionally, although pitting, holes, lesions, and other damage on Triceratops skulls (and the skulls of other ceratopsids) are often attributed to horn damage in combat, a recent study finds 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.[43]

The large frill also may have helped to increase body area to regulate body temperature.[44] A similar theory has been proposed regarding the plates of Stegosaurus,[45] although this use alone would not account for the bizarre and extravagant variation seen in different members of the Ceratopsidae.[6] This observation is highly suggestive of what is now believed to be the primary function, display.

The theory of their use in sexual display was first proposed by Davitashvili in 1961 and has gained increasing acceptance since.[46][38][15] Evidence that visual display was important, either in courtship or in other social behaviour, 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 behaviour.[47] A recent study of the smallest Triceratops skull, ascertained to be a juvenile, shows the frill and horns developed at a very early age, predating sexual development and thus probably important for visual communication and species recognition in general.[48] The large eyes and shortened features, a hallmark of "cute" baby mammals, also suggest that the parent Triceratops may have cared for its young.


[edit] Depiction in recent popular media Main article: Triceratops in popular culture The distinctive appearance of Triceratops has led to them being frequently depicted in various forms of popular culture including films, computer games and documentaries. In more recent media, they have notably appeared in the film Jurassic Park and its sequels, as well as in three major dinosaur documentaries: Walking with Dinosaurs, The Truth About Killer Dinosaurs and Prehistoric Park.

They are famously known as "three-horns" (and are so named in The Land Before Time animated film and its numerous sequels) due to the three prominent horns on their head and nose, which have become almost synonymous with the dinosaurs. The shorthand "Trike" is another common informal name, and is also the name of the Triceratops character in the children's book series and television cartoon series Harry and His Bucket Full of Dinosaurs.

A recurring theme in children's dinosaur books is a climactic showdown or battle between Triceratops and T. rex.[49][50][51][52] As such these two dinosaurs are often depicted and thought of as natural enemies. A memorable but anachronistic battle with Ceratosaurus substituting for T. rex is featured in the 1966 movie One Million Years B.C.

Triceratops appears in video games either derived directly from the Jurassic Park series or similarly themed, namely the 1997 PC games Jurassic Park: Chaos Island and Turok: Dinosaur Hunter, and the 2000 PC and Playstation game Dino Crisis 2. As well, it is a popular creature used in games designed by Nintendo, including Diddy Kong Racing and Starfox Adventures. Triceratops (the species are not identified) is also the official state fossil of South Dakota,[53] and the official state dinosaur of Wyoming.[54]


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