Ceratopsids Temporal range: Late Cretaceous, ~ | |
---|---|
Montage of four ceratopsids. Clockwise from top left: Titanoceratops , Styracosaurus , Utahceratops and Triceratops | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | † Ornithischia |
Clade: | † Neornithischia |
Clade: | † Ceratopsia |
Superfamily: | † Ceratopsoidea |
Clade: | † Ceratopsomorpha |
Family: | † Ceratopsidae Marsh, 1888 |
Subgroups | |
| |
Synonyms | |
|
Ceratopsidae (sometimes spelled Ceratopidae) is a family of ceratopsian dinosaurs including Triceratops , Centrosaurus , and Styracosaurus . All known species were quadrupedal herbivores from the Upper Cretaceous. All but one species are known from western North America, which formed the island continent of Laramidia during most of the Late Cretaceous. Ceratopsids are characterized by beaks, rows of shearing teeth in the back of the jaw, elaborate nasal horns, and a thin parietal-squamosal shelf that extends back and up into a frill. The group is divided into two subfamilies—Chasmosaurinae and Centrosaurinae. The chasmosaurines are generally characterized by long, triangular frills and well-developed brow horns. The centrosaurines had well-developed nasal horns or nasal bosses, shorter and more rectangular frills, and elaborate spines on the back of the frill.
These horns and frills show remarkable variation and are the principal means by which the various species have been recognized. Their purpose is not entirely clear. Defense against predators is one possible purpose – although the frills are comparatively fragile in many species – but it is more likely that, as in modern ungulates, they were secondary sexual characteristics used in displays or for intraspecific combat. The massive bosses on the skulls of Pachyrhinosaurus and Achelousaurus resemble those formed by the base of the horns in modern musk oxen, suggesting that they butted heads. Centrosaurines have frequently been found in massive bone beds with few other species present, suggesting that the animals lived in large herds.
Fossil deposits dominated by large numbers of ceratopsids from individual species suggest that these animals were at least somewhat social. [2] However, the exact nature of ceratopsid social behavior has historically been controversial. [3] In 1997, Lehman argued that the aggregations of many individuals preserved in bonebeds originated as local "infestations" and compared them to similar modern occurrences in crocodiles and tortoises. [3] Other authors, such as Scott D. Sampson, interpret these deposits as the remains of large "socially complex" herds. [3]
Modern animals with mating signals as prominent as the horns and frills of ceratopsians tend to form these kinds of large, intricate associations. [4] Sampson found in previous work that the centrosaurine ceratopsids did not achieve fully developed mating signals until nearly fully grown. [5] He finds commonality between the slow growth of mating signals in centrosaurines and the extended adolescence of animals whose social structures are ranked hierarchies founded on age-related differences. [5] In these sorts of groups young males are typically sexually mature for several years before actually beginning to breed, when their mating signals are most fully developed. [6] Females, by contrast do not have such extended adolescence. [6]
Other researchers who support the idea of ceratopsid herding have speculated that these associations were seasonal. [7] This hypothesis portrays ceratopsids as living in small groups near the coasts during the rainy season and inland with the onset of the dry season. [7] Support for the idea that ceratopsids formed herds inland comes from the greater abundance of bonebeds in inland deposits than coastal ones. The migration of ceratopsids away from the coasts may have represented a move to their nesting grounds. [7] Many African herding animals engage in this kind of seasonal herding today. [7] Herds would also have afforded some level of protection from the chief predators of ceratopsids, tyrannosaurids. [8]
Ceratopsids were adapted to processing high-fiber plant material with their highly derived dental batteries and advanced dentition. [9] They may have utilized fermentation to break down plant material with a gut microflora. [9] Mallon et al. (2013) examined herbivore coexistence on the island continent of Laramidia, during the Late Cretaceous. It was concluded that ceratopsids were generally restricted to feeding on vegetation at, or below, the height of 1 meter. [10]
Ceratopsians probably had the "low mass-specific metabolic rat[e]" typical of large bodied animals. [9]
According to Scott D. Sampson, if ceratopsids were to have sexual dimorphism modern ecological analogues suggest it would be in their mating signals like horns and frills. [11] No convincing evidence for sexual dimorphism in body size or mating signals is known in ceratopsids, although was present in the more primitive ceratopsian Protoceratops andrewsi whose sexes were distinguishable based on frill and nasal prominence size. [11] This is consistent with other known tetrapod groups where midsized animals tended to exhibit markedly more sexual dimorphism than larger ones. [12] However, if there were sexually dimorphic traits, they may have been soft tissue variations like colorations or dewlaps that would not have been preserved as fossils. [12]
Scott D. Sampson has compared the evolution of ceratopsids to that of some mammal groups: both were rapid from a geological perspective and precipitated the simultaneous evolution of large body size, derived feeding structures, and "varied hornlike organs." [3] The earliest ceratopsids, including members of both Centrosaurinae and Chasmosaurinae are known from the early Campanian stage, though the fossil record for early ceratopsids is poor. [13] All but one of the named species of ceratopsid is known from Western North America, which formed the island continent of Laramidia during the Late Cretaceous, separated from the island continent of Appalachia to the east by the Western Interior Seaway. The latitudinal range of ceratopsians across Laramidia extends from Alaska to Mexico. The only named ceratopsid outside of Laramidia is Sinoceratops , a centrosaurine from the late Campanian of China. [1] An indeterminate tooth of a ceratopsid is known from Mississippi dating to the late Maastrichtian, a few million years prior to the close of the Cretaceous, indicating that ceratopsids dispersed into eastern North America corresponding to the closure of the Western Interior Seaway at the end of the Cretaceous. [14]
The chief predators of ceratopsids were tyrannosaurids. [8]
There is evidence for an aggressive interaction between a Triceratops and a Tyrannosaurus in the form of partially healed tyrannosaur tooth marks on a Triceratops brow horn and squamosal (a bone of the neck frill); the bitten horn is also broken, with new bone growth after the break. It is not known what the exact nature of the interaction was, though: either animal could have been the aggressor. [15] Since the Triceratops wounds healed, it is most likely that the Triceratops survived the encounter and managed to overcome the Tyrannosaurus. Paleontologist Peter Dodson estimates that in a battle against a bull Tyrannosaurus, the Triceratops had the upper hand and would successfully defend itself by inflicting fatal wounds to the Tyrannosaurus using its sharp horns. [16]
The clade Ceratopsidae was in 1998 defined by Paul Sereno as the group including the last common ancestor of Pachyrhinosaurus and Triceratops; and all its descendants. [17] In 2004, it was by Peter Dodson defined to include Triceratops , Centrosaurus , and all descendants of their most recent common ancestor. [18] Ceratopsidae was given an official definition in the PhyloCode by Daniel Madzia and colleagues in 2021 as "the smallest clade containing Centrosaurus apertus , Ceratops montanus , Chasmosaurus belli , and Triceratops horridus ". [19] This definition ensures that the type species of Ceratopsidae, Ceratops montanus, is included in the clade's definition. [19]
Triceratops is a genus of chasmosaurine ceratopsian dinosaur that lived during the late Maastrichtian age of the Late Cretaceous period, about 68 to 66 million years ago in what is now western North America. It was one of the last-known non-avian dinosaurs and lived until the Cretaceous–Paleogene extinction event 66 million years ago. The name Triceratops, which means 'three-horned face', is derived from the Greek words trí- meaning 'three', kéras meaning 'horn', and ṓps meaning 'face'.
Ceratopsia or Ceratopia is a group of herbivorous, beaked dinosaurs that thrived in what are now North America, Europe, and Asia, during the Cretaceous Period, although ancestral forms lived earlier, in the Jurassic. The earliest known ceratopsian, Yinlong downsi, lived between 161.2 and 155.7 million years ago. The last ceratopsian species, Triceratops prorsus, became extinct during the Cretaceous–Paleogene extinction event, 66 million years ago.
Marginocephalia is a clade of ornithischian dinosaurs that is characterized by a bony shelf or margin at the back of the skull. These fringes were likely used for display. This clade was officially defined in the PhyloCode by Daniel Madzia and colleagues in 2021 as "the smallest clade containing Ceratops montanus, Pachycephalosaurus wyomingensis, and Triceratops horridus". There are two clades included in Marginocephalia: the thick-skulled Pachycephalosauria and the horned Ceratopsia. All members of Marginocephalia were primarily herbivores. They basally used gastroliths to aid in digestion of tough plant matter until they convergently evolved tooth batteries in Neoceratopsia and Pachycephalosauria. Marginocephalia first evolved in the Jurassic Period and became more common in the Cretaceous. They are basally small facultative quadrupeds while derived members of the group are large obligate quadrupeds. Primitive marginocephalians are found in Asia, but the group migrated upwards into North America.
Styracosaurus is an extinct genus of herbivorous ceratopsian dinosaur from the Late Cretaceous of North America. It had four to six long parietal spikes extending from its neck frill, a smaller jugal horn on each of its cheeks, and a single horn protruding from its nose, which may have been up to 60 centimeters long and 15 centimeters wide. The function or functions of the horns and frills have been debated for many years.
Einiosaurus is a genus of herbivorous centrosaurine ceratopsian dinosaur from the Upper Cretaceous of northwestern Montana. The name means 'bison lizard', in a combination of Blackfeet Indian eini and Latinized Ancient Greek sauros; the specific name (procurvicornis) means 'with a forward-curving horn' in Latin. Einiosaurus is medium-sized with an estimated body length at 4.5 metres (15 ft).
Achelousaurus is a genus of centrosaurine ceratopsid dinosaur that lived during the Late Cretaceous Period of what is now North America, about 74.2 million years ago. The first fossils of Achelousaurus were collected in Montana in 1987, by a team led by Jack Horner, with more finds made in 1989. In 1995, Achelousaurus horneri was described and named by Scott D. Sampson; the generic name means "Achelous lizard", in reference to the Greek deity Achelous, and the specific name refers to Horner. The genus is known from a few specimens consisting mainly of skull material from individuals, ranging from juveniles to adults.
Centrosaurus is a genus of centrosaurine ceratopsian dinosaur from Campanian age of Late Cretaceous Canada. Their remains have been found in the Dinosaur Park Formation, dating from 76.5 to 75.5 million years ago.
Anchiceratops is an extinct genus of chasmosaurine ceratopsid dinosaur that lived approximately 72 to 71 million years ago during the latter part of the Cretaceous Period in what is now Alberta, Canada. Anchiceratops was a medium-sized, heavily built, ground-dwelling, quadrupedal herbivore that could grow up to an estimated 4.3 metres (14 ft) long. Its skull featured two long brow horns and a short horn on the nose. The skull frill was elongated and rectangular, its edges adorned by coarse triangular projections. About a dozen skulls of the genus have been found.
Arrhinoceratops is a genus of herbivorous ceratopsian dinosaur. The name was coined as its original describer concluded it was special because the nose-horn was not a separate bone, however further analysis revealed this was based on a misunderstanding. It lived during the latest Campanian/earliest Maastrichtian stage of the Late Cretaceous, predating its famous relative Triceratops by a few million years, although it was contemporary with Anchiceratops. Its remains have been found in Canada.
Monoclonius is an extinct genus of herbivorous ceratopsian dinosaur found in the Late Cretaceous layers of the Judith River Formation in Montana, United States, and the uppermost rock layers of the Dinosaur Park Formation in Alberta, Canada dated to between 75 and 74.6 million years ago.
Brachyceratops is a dubious genus of ceratopsian dinosaur known only from partial juvenile specimens dating to the late Cretaceous Period of Montana, United States.
Agujaceratops is a genus of horned dinosaur from the Late Cretaceous (Campanian) of west Texas. It is a chasmosaurine (long-frilled) ceratopsian. Two species are known, Agujaceratops mariscalensis, and A. mavericus.
Chasmosaurinae is a subfamily of ceratopsid dinosaurs. They were one of the most successful groups of herbivores of their time. Chasmosaurines appeared in the early Campanian, and became extinct, along with all other non-avian dinosaurs, during the Cretaceous–Paleogene extinction event. Broadly, the most distinguishing features of chasmosaurines are prominent brow horns and long frills lacking long spines; centrosaurines generally had short brow horns and relatively shorter frills, and often had long spines projecting from their frills.
Centrosaurinae is a subfamily of ceratopsid, a group of large quadrupedal ornithischian dinosaur. Centrosaurine fossil remains are known primarily from the northern region of Laramidia but isolated taxa have been found in China and Utah as well.
Kosmoceratops is a genus of ceratopsid dinosaur that lived in North America about 76–75.9 million years ago during the Late Cretaceous period. Specimens were discovered in Utah in the Kaiparowits Formation of the Grand Staircase–Escalante National Monument in 2006 and 2007, including an adult skull and postcranial skeleton and partial subadults. In 2010, the adult was made the holotype of the new genus and species Kosmoceratops richardsoni; the generic name means "ornate horned face", and the specific name honors Scott Richardson, who found the specimens. The find was part of a spate of ceratopsian discoveries in the early 21st century, and Kosmoceratops was considered significant due to its elaborate skull ornamentation.
Vagaceratops is a genus of herbivorous ceratopsian dinosaur. It is a chasmosaurine ceratopsian which lived during the Late Cretaceous period in what is now Alberta. Its fossils have been recovered from the Upper Dinosaur Park Formation. It is sometimes included in the genus Chasmosaurus as Chasmosaurus irvinensis instead of being recognized as its own genus.
Nasutoceratops is genus of ceratopsid dinosaur that lived in North America during the Late Cretaceous period, about 76.0–75.5 million years ago. The first known specimens were discovered in Utah in the Kaiparowits Formation of the Grand Staircase–Escalante National Monument (GSENM) from 2006 onwards, including a subadult skull with a partial postcranial skeleton and rare skin impressions and two other partial skulls. In 2013, the subadult was made the holotype of the new genus and species Nasutoceratops titusi; the generic name means "large-nosed horned face", and the specific name honors the paleontologist Alan L. Titus for his work at the GSENM. The dinosaur was noted for its large nose in news reports, and later featured in Jurassic World films.
Coronosaurus is a genus of centrosaurine ceratopsian dinosaurs which lived in the Late Cretaceous, in the middle Campanian stage. Its remains, two bone beds, were discovered by Phillip J. Currie in the Oldman Formation of Alberta, Canada, and its type and only species, Coronosaurus brinkmani, was first described in 2005, as a new species within the genus Centrosaurus. Later studies questioned the presence of a direct relationship, and in 2012 it was named as a separate genus. Coronosaurus means "crowned lizard", coming from "corona", Latin for crown, and "sauros", Greek for lizard; this name refers to the unique, crown-like shape of the horns on the top of its frill.
This timeline of ceratopsian research is a chronological listing of events in the history of paleontology focused on the ceratopsians, a group of herbivorous marginocephalian dinosaurs that evolved parrot-like beaks, bony frills, and, later, spectacular horns. The first scientifically documented ceratopsian fossils were described by Edward Drinker Cope starting in the 1870s; however, the remains were poorly preserved and their true nature was not recognized. Over the next several decades, Cope named several such genera and species. Cope's hated rival, Othniel Charles Marsh, also described ceratopsian remains. In 1887, Marsh mistook a Triceratops horn for one belonging to a new species of prehistoric Bison. Marsh also named the eponymous genus Ceratops in 1888. The next year, he named the most famous ceratopsian, Triceratops horridus. It was the discovery of Triceratops that illuminated the ceratopsian body plan, and he formally named the Ceratopsia in 1890.