Bravoceratops Temporal range: Late Cretaceous, | |
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Hypothetical restoration of the skeleton with the holotype material in white | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | † Ornithischia |
Clade: | † Neornithischia |
Clade: | † Ceratopsia |
Family: | † Ceratopsidae |
Subfamily: | † Chasmosaurinae |
Genus: | † Bravoceratops Wick & Lehman, 2013 |
Type species | |
†Bravoceratops polyphemus Wick & Lehman, 2013 |
Bravoceratops is a genus of large chasmosaurine ceratopsid dinosaur that lived approximately 70 million years ago, and is known from the Late Cretaceous Javelina Formation in what is now Texas, United States.
Bravoceratops is only known from the holotype specimen TMM 46015-1; it is housed in the collection of the Texas Memorial Museum, Austin, Texas. The skull consists of a number of fragments, altogether including: the braincase , parts of each brow horn, the rear end of the left postorbital , assorted parts of the nasal area and horn, the jugals , quadrates and quadratojugals from each side, a section of the parietal and epiparietals , multiple parts of the dentary, and some of the right squamosal . It was recovered from the lowermost rocks of the Javelina Formation, in Big Bend National Park; ceratopsid fossils are uncommon in this formation, giving the discovery of Bravoceratops importance in clarifying the groups' diversity. The specimen was found in sandy conglomerate sediment at the Hippiewalk geologic locality. Signs of erosion are presented in the larger bone fragments, which were found disorganized over an area of ten square metres; the site is thought to have been a lag deposit of a river, only most durable extremities able to survive preservation. The animal would have lived during the late Campanian or early Maastrichtian stage of the Cretaceous period, approximately 70 million years ago. [1]
The genus name means "wild horn-face", and is derived from the Mexican name for the Rio Grande, "Rio Bravo del Norte" (wild river of the north), and the Greek words "keras" (κέρας) meaning "horn" and "ops" (ὤψ) referring to the "face". The specific name polyphemus, refers to the giant cyclops Polyphemus confronted by Odysseus in the Greek epic poem, The Odyssey. This is in reference to the presumption that it had a single central horn on the surface of the frill, recalling the placement of a Cyclops' eye. The genus was described and named by Steven L. Wick and Thomas M. Lehman in 2013 and the type species is Bravoceratops polyphemus. [1]
Two autapomorphies (unique derived traits) were originally proposed. Firstly, the median parietal bar (which runs down the middle of the frill between its large holes) at mid-length splays out to the rear like a fan and its rear edge is not notched or embayed. Secondly, the upper surface of the bar is, at the midline, hollowed out by a symmetrical depression. It is this hollow in the form of an inverted tear that occasioned the specific name as it resembled the single eye of a cyclops, thus the allusion with Polyphemus. The authors assumed it formed the base of an epiparietal, or small horn, unpreserved in the fossil. [1] In 2020, paleontologists Denver W. Fowler and Elizabeth A. Freedman Fowler suggested that the parietal median bar of the specimen had been reconstructed upside down, and that the genus therefore did not have any distinguishing features among chasmosaurines, making it a nomen dubium , or invalid genus. [2] However, a number of authors considered the genus valid (see classification below) and it was included as part of a southwest clade of chasmosaurines by Dalman et al., (2021).
Unlike Fowler and Freedman-Folwer, Sebastian G. Dalman and colleagues, in a 2021 paper naming the related genus Sierraceratops , treated Bravoceratops as valid, noting both distinct traits and those unique to it and Sierraceratops. A flange in front of its quadrate near the jugal bones is present, large as in Sierraceratops while small or not present in other chasmosaurs. The epijugal, a small horn on the jugals, is longer than tall and sharp on its tip, as in Sierraceratops and unlike other derived chasmosaurines. The two brow horns are compressed, being taller than wide rather than conical, a trait shared with Sierraceratops and the distantly related Judiceratops. The cross section of the lower bar of its parietal bar has a D-shape shared with Sierraceratops and similar but not identical to that of Coahuilaceratops , but unlike other chasmosaurines. Bravoceratops is distinguishable from Coahuilaceratops in lacking enormous brow horns; it is distinguishable from Sierraceratops in having larger brow horns and a smaller epijugal horn. [3]
Bravoceratops is assigned to the Chasmosaurinae. It is an "intermediate" form of chasmosaurine; it, along with genera like Pentaceratops and Anchiceratops , was distinctly more derived ("advanced") than forms like Chasmosaurus but less so than taxa like Triceratops . Its position in a phylogenetic analysis performed by Wick and Lehman (2013), who first named the taxon. It was found that Bravoceratops may be the sister taxon of Coahuilaceratops (a relationship later supported by Dalman et al., 2021). The first analysis, with unmodified characters from a previous study, found the pair to be relatively basal . The second, with two added characters, instead found Bravoceratops to be a relatively derived taxon. Both positions were considered to be somewhat unintuitive; several of its derived characters would be odd for a taxon in the basal position, but the genus also possessed primitive characters would require evolutionary reversals if the more derived position was correct. Regardless, the more derived placement was considered to be somewhat more likely. The first tree is reproduced on the left, and the second on the right: [1]
Future studies found trees showing the derived position, with Bravoceratops found related to Arrhinoceratops and Triceratopsini. However, the fragmentary nature of Bravoceratops caused it to generally be removed from phylogenetic analyses, as it was a wildcard taxon (meaning it was unstable and caused poorer results). [4] [5] [6] Fowler and Freedman Fowler (2020) recovered Bravoceratops in the more basal position, and, similar to Wick and Lehman (2013), found it close to Pentaceratops, and once again sister to Coahuilaceratops. However, they also decided to remove it from their analysis due to their unsubstantiated conclusion the taxon was invalid. [2] However, Dalman et al. (2021) convincingly validated the genus. They demonstrated that Bravoceratops shared a number morphological traits with other chasmosaurines from the same time and region. As a result, Bravoceratops was determined to be part of a clade endemic to the Southwestern United states. Sister taxa include Coahuilaceratops from Mexico and Sierraceratops from New Mexico. [3]
Chasmosaurus is a genus of ceratopsid dinosaur from the Late Cretaceous Period in North America. Its given name means 'opening lizard', referring to the large openings (fenestrae) in its frill. With a length of 4.3–4.8 metres (14.1–15.7 ft) and a weight of 1.5–2 tonnes —or anywhere from 2,200 to nearly 5,000 lbs., give or take—Chasmosaurus was of a slightly smaller to ‘average’ size, especially when compared to larger ceratopsians.
Pentaceratops is a genus of herbivorous ceratopsid dinosaur from the late Cretaceous Period of what is now North America. Fossils of this animal were first discovered in 1921, but the genus was named in 1923 when its type species, Pentaceratops sternbergii, was described. Pentaceratops lived around 76–73 million years ago, its remains having been mostly found in the Kirtland Formation in the San Juan Basin in New Mexico. About a dozen skulls and skeletons have been uncovered, so anatomical understanding of Pentaceratops is fairly complete. One exceptionally large specimen later became its own genus, Titanoceratops, due to its more derived morphology, similarities to Triceratops, and lack of unique characteristics shared with Pentaceratops.
Medusaceratops is an extinct genus of centrosaurine ceratopsian dinosaur known from the Late Cretaceous Judith River Formation of Montana, northern United States. It contains a single species, Medusaceratops lokii.
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.
Eotriceratops is a genus of herbivorous ceratopsian dinosaurs which lived in the area of North America during the late Cretaceous period. The only named species is Eotriceratops xerinsularis.
The Aguja Formation is a geological formation in North America, exposed in Texas, United States and Chihuahua and Coahuila in Mexico, whose strata date back to the Late Cretaceous. Dinosaur remains are among the fossils that have been recovered from the formation. Fossil palms have also been unearthed here.
Coahuilaceratops is a genus of chasmosaurine ceratopsian dinosaur that lived during the early Maastrichtian age of the Late Cretaceous epoch, about 71.5 to 70.5 million years ago in what is now northern Mexico. It contains a single species, Coahuilaceratops magnacuerna.
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.
Titanoceratops is a controversial genus of herbivorous ceratopsian dinosaur. It was a giant chasmosaurine ceratopsian that lived in the Late Cretaceous period in what is now New Mexico. Titanoceratops was named for its large size, being one of the largest known horned dinosaurs and the type species was named T. ouranos, after Uranus (Ouranos), the father of the Greek titans. It was named in 2011 by Nicholas R. Longrich for a specimen previously referred to Pentaceratops. Longrich believed that unique features found in the skull reveal it to have been a close relative of Triceratops, classified within the subgroup Triceratopsini. However, other researchers have expressed skepticism, and believe "Titanoceratops" to simply be an unusually large, old specimen of Pentaceratops.
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.
Regaliceratops is a monospecific genus of chasmosaurine ceratopsid dinosaur from Alberta, Canada that lived during the Late Cretaceous in what is now the St. Mary River Formation. The type and only species, Regaliceratops peterhewsi, is known only from an adult individual with a nearly complete skull lacking the lower jaw, which was nicknamed "Hellboy". Regaliceratops was named in 2015 by Caleb M. Brown and Donald M. Henderson. Regaliceratops has an estimated length of 5 metres (16 ft) and body mass of 2 metric tons. The skull of Regaliceratops displays features more similar to centrosaurines, which suggests convergent evolution in display morphology in ceratopsids.
Spiclypeus is an extinct genus of chasmosaurine ceratopsian dinosaur known from the Late Cretaceous Judith River Formation of Montana, United States.
Yehuecauhceratops is a genus of horned centrosaurine ceratopsid dinosaur from the Late Cretaceous of Coahuila, Mexico. It contains a single species, Y. mudei, described from two partial specimens by Rivera-Sylva et al. in 2016 and formally named by Rivera-Sylva et al. in 2017. It was a small centrosaurine with a body length of 3 metres (9.8 ft), making it smaller than Agujaceratops and Coahuilaceratops, the other two ceratopsids in its environment; the three may have been ecologically segregated. A ridge bearing a single roughened projection near the bottom of the squamosal bone, which probably supported a small horn, allows Yehuecauhceratops to be distinguished from other centrosaurines. Its affinities to nasutoceratopsin centrosaurines, such as Avaceratops and Nasutoceratops, are supported by various morphological similarities to the former.
Terminocavus is a genus of ceratopsid dinosaur from the late Cretaceous Period of what is now North America. The genus contains a single species, the type species Terminocavus sealeyi, known from a parietal and some other associated fragments. The holotype specimen was discovered in the Kirtland Formation of New Mexico in 1997, and was later described and named in a 2020 study. It was similar in anatomy to Pentaceratops and Anchiceratops, which it was closely related to, but had a distinctive heart-shaped upper frill with very narrow notch. It has been hypothesized to form an anagenetic series with several other chasmosaur species.
Sierraceratops is a genus of chasmosaurine ceratopsian from the Late Cretaceous Hall Lake Formation of New Mexico, United States. The genus contains a single species, Sierraceratops turneri, known from a partial skeleton discovered in 1997.
Bisticeratops is a genus of chasmosaurine ceratopsian from outcrops of the Campanian age Kirtland Formation found in the Bisti/De-Na-Zin Wilderness in northwestern New Mexico, United States. The type and only species is B. froeseorum, known from a nearly complete skull.