Titanoceratops (meaning "titanic horned face") is a controversial genus of herbivorousceratopsiandinosaur. It was a giant chasmosaurineceratopsian that lived in the Late Cretaceous period (Campanian stage, about 75 million years ago[1]) 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 Greektitans. 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.
The holotype specimen is OMNH10165, a partial skeleton including a mostly complete skull and jaws, and much of the skeleton. It was found in either the upper Fruitland Formation or the lower Kirtland Formation. The original quarry is lost, so it is not known which formation the fossil was excavated from. The formations are both late Campanian in age. The skull is incomplete, but as currently reconstructed it measures 2.65 metres (8.7ft) long, making it a candidate for the longest skull of any land animal. With an estimated weight of 6.55 tonnes (6.45 long tons; 7.22 short tons) and length of 6.8 metres (22.3ft), Titanoceratops was comparable in size with the largest ceratopsians, Torosaurus and Triceratops, and was likely the largest animal in its ecosystem, if not in North America, at the time.
Description
Life restorationSkeletal reconstruction, holotype material in white, with human for scale
The skull measures 1.2m (3.9ft) from the tip of the snout to the quadrate, and the restored frill extends its length up to 2.65m (8.7ft)[2] making it a candidate for the longest skull of any land animal. Titanoceratops was as large as the later triceratopsins Triceratops and Torosaurus, with an estimated weight of 6.55 tonnes (7.22 short tons)[2] and a mounted skeleton measuring 6.8 metres (22.3ft) long and 2.5 metres (8.2ft) tall at the back.[3] In 2016 Gregory S. Paul gave a lower estimation of 6.5 meters (21.3ft) and 4.5 tonnes (4.9 short tons). Tom Holtz (2012) noted that it is extremely similar to its closely related contemporaries Eotriceratops and Ojoceratops, which may all be synonymous.[4] The holotype skeleton of Titanoceratops consists of a partial skull with jaws, syncervical, cervical, dorsal, and sacral vertebrae, caudal vertebrae, ribs, humeri, a right radius, femora, tibiae, a right fibula, both ilia, both ischia, and ossified tendons.[2] In total, the amount of material assigned to Titanoceratops means it is quite well known, along with genera like Triceratops, Vagaceratops, Pentaceratops, Chasmosaurus, Centrosaurus, Styracosaurus, and Anchiceratops.[5]
History of study
Beds of the upper Fruitland Formation and lower Kirtland Formation in the vicinity of the Titanoceratops holotype quarryJ. Willis Stovall holds up the humerus, 1941
The holotype of Titanoceratops was collected from the upper Fruitland Formation or the lower Kirtland Formation in July 1941, by a field crew consisting J. Willis Stovall, his student Wann Langston Jr., and Donald E. Savage.[3] The precise location of the quarry is no longer known. The holotype specimen consists of most of the fore and hindlimbs, some vertebrae, a fairly complete skull with only one small section of the frill, and partial lower jaws.[2] The bones, being preserved in a fine-grained shale, were crushed and fragile, and so the skeleton was initially considered unsuitable for mounting. Later, however, the fossils were prepared and the skeleton put on display at the Oklahoma Museum of Natural History.
In 1998, the specimen was described by Thomas Lehman as an aberrant and unusually large individual of Pentaceratops sternbergii, previously described from the same area.[3] The specimen was later reinterpreted as a member of the Triceratopsini, the group including Triceratops, by Nicholas R. Longrich and given the name Titanoceratops ouranos in 2011. The name Titanoceratops is derived from the GreekTitan, a mythical race of giants, keras (κέρας), meaning "horn", and ops (ὤψ), "face". The species name ouranos, refers to Uranus, the father of the Titan race.[2] Longrich's re-interpretation would have major implications for the evolutionary history and biogeography of chasmosaurine dinosaurs. Previously, the origins of Triceratops were poorly known. Until the Longrich's re-interpretation of Titanoceratops, Eotriceratops was thought to be the oldest known triceratopsin, and only dated to 68million years old, from the uppermost region of the Horseshoe Canyon Formation. No Campanian triceratopsins were known, so it appeared as if the group evolved in the Maastrichtian. If Titanoceratops is a member of this group, it would demonstrate that they evolved millions of years earlier than previously thought, and it would imply a five million year long gap in the fossil record and ghost lineage leading to Eotriceratops.[2] However, several subsequent studies have cast doubt on the hypothesis that Titanoceratops is a triceratopsin.[6]
OMNH 10165 is a particularly large chasmosaurine fossil, which Lehman originally assigned to the genus Pentaceratops, believing that it was a particularly large and old specimen. A 2011 study by Longrich disagreed with this interpretation, concluding that it was actually a distinct genus, which he named Titanoceratops. Longrich interpreted the specimen as sharing more characteristics with Triceratops and Torosaurus than with Pentaceratops, and he named a new group, Triceratopsini, to contain all of them.[2] Longrich used the following features to distinguish the specimen from other chasmosaurines: the possession of thin squamosals (Triceratops); an unsealed parietal fenestrae (Triceratops); an epijugal resembling a hornlike structure (Triceratops); a narrow median bar of the parietal (Triceratops, Torosaurus); a narial strut oriented vertically with a narrow base (Triceratops, Torosaurus); an enlarged epoccipital on the rear end of the squamosal (Triceratops, Torosaurus, Eotriceratops); an extremely enlarged premaxillary fossa (Triceratops, Torosaurus, Eotriceratops); and in lacking a narial process of the premaxilla that is dorsally inflected (Triceratops, Torosaurus, Eotriceratops).[2]
Front view of the mount
Lehman ignored Longrich's reclassification in his own subsequent publications.[7] As part of a 2020 study by Fowler and Freedman Fowler, the authors critically re-evaluated the evidence that Titanoceratops was a distinct genus. They agreed with Lehman's original assessment, that the features in the specimen that appeared unique were likely due simply to advanced age and unusually large size. Pending a full re-evaluation of the specimen by other researchers, Fowler and Freedman Fowler opted to consider OMNH 10165 simply a large Pentaceratops.[6]
Paleoecology
The fossilized leaf of Sabalites, a Cretaceous palm, documents the wet, warm climate that prevailed in Late Cretaceous New Mexico
Titanoceratops is known from OMNH10165, a skeleton from the lowermost Fruitland or uppermost Kirtland Formation. The Fruitland Formation is about 100 metres (330ft) thick, and consists of sandstones, mudstones, and abundant coals deposited in a coastal floodplain. Fossil trees are abundant in the area from which the holotype was collected, suggesting a wet, well-forested environment.
A fossil tree stump from the Coal Creek area where Titanoceratops was found, showing the extensive coal beds of the area
The Kirtland Formation, which conformably overlays the Fruitland, is approximately 600 metres (2,000ft) thick, and made up of sandstone, siltstone, mudstone, and shale. Both formations are late Campanian in age. The Fossil Forest Member of the Fruitland is 74.11±0.62million years old, and the Hunter Wash Member of the Kirtland is between 73.37±0.18 and 73.04±0.25million years in age. The two members combined make up the Hunter Wash local fauna. Therefore, Titanoceratops dates between 74 and 73 million years ago.[2] The age Titanoceratops lived in is called the Kirtlandian land-vertebrate age, and it is characterized by the appearance of Pentaceratops sternbergii.[8]
Titanoceratops supports the idea that late Cretaceous dinosaur faunas were highly endemic, with distinct species found in the Southern Great Plains of New Mexico, and the Northern Great Plains of Montana and Canada. Despite extensive sampling to the north in the Dinosaur Park Formation and Two Medicine Formation, triceratopsins are unknown there. This implies that the triceratopsins originally evolved in the south, then spread north in the Maastrichtian.[2]
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'.
Torosaurus is a genus of herbivorous chasmosaurine ceratopsian dinosaur that lived during the late Maastrichtian age of the Late Cretaceous period, between 68 and 66 million years ago, though it is possible that the species range might extend to as far back as 69 million years ago. Fossils have been discovered across the Western Interior of North America, from as far north as Saskatchewan to as far south as Texas.
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. The Chasmosaurs were similar, in overall build and weight, to a white rhinoceros or an Indian rhinoceros; just like rhinos, and all other ceratopsians, they were purely herbivorous, needing to consume around 54 kilograms, or 120 lbs., of plant matter each day.
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.
The Kirtland Formation is a sedimentary geological formation.
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.
Coahuilaceratops is a genus of ceratopsian dinosaur. It is a chasmosaurine ceratopsian which lived during the Late Cretaceous period in what is now southern Coahuila in northern Mexico. It is known from the holotype CPC 276, a partial skeleton of an adult individual which includes several skull elements. Another specimen, CPS 277, may represent a juvenile Coahuilaceratops. All specimens of Coahuilaceratops were collected from a single location in the middle strata of the Cerro del Pueblo Formation, which dates to between 72.5 and 71.4 million years ago.
Ojoceratops is a genus of ceratopsian dinosaur which lived in what is now New Mexico, United States. Ojoceratops fossils have been recovered from strata of the Ojo Alamo Formation, dating to the late Cretaceous period. The type species is Ojoceratops fowleri.
Tatankaceratops is a controversial genus of herbivorous ceratopsian dinosaur. It is a small chasmosaurine ceratopsian which lived during the Late Cretaceous period in what is now South Dakota. It is known from a single partial skull which was collected from the Hell Creek Formation, dating to 66 million years ago. Tatankaceratops was described by Christopher J. Ott and Peter L. Larson in 2010 and the type species is Tatankaceratops sacrisonorum. Tatankaceratops is known from one specimen housed at the Black Hills Institute, BHI 6226.
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.
Triceratopsini is a tribe of herbivorous chasmosaurine dinosaurs that lived between the late Campanian to the late Maastrichtian stages of the Cretaceous period, between 74.73 and 66 million years ago. Fossils of these animals have been found in western North America, in particular West Canada, Western and Midwestern United States, which was once part of the ancient continent of Laramidia. The tribe was named by Nicholas R. Longrich in 2011 for the description of Titanoceratops, which he defined as "all species closer to Triceratops horridus than to Anchiceratops ornatus or Arrhinoceratops brachyops". Triceratopsins were the largest of the chasmosaurines; suggesting that gigantism had evolved in the Ceratopsidae once. In addition there is an evolutionary trend in the solidification of the frills, the most extreme being in Triceratops.
Dinosaur paleobiogeography is the study of dinosaur geographic distribution, based on evidence in the fossil record.
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.
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.
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.
References
↑ Fowler, D. W. 2017. Revised geochronology, correlation, and dinosaur stratigraphic ranges of the Santonian-Maastrichtian (Late Cretaceous) formations of the Western Interior of North America. PLoS ONE 12(11): e0188426.
1 2 3 4 5 Sullivan, R.M.; Lucas, S.G. (2006). "The Kirtlandian Land-Vertebrate "Age"-Faunal Composition, Temporal Position, and Biostratigraphic Correlation in the Nonmarine Upper Cretaceous of Western North America". In Lucas, S.G.; Sullivan, R.M. (eds.). Late Cretaceous vertebrates from the Western Interior(PDF). Vol.35. New Mexico Museum of Natural History and Science Bulletin. pp.7–23. Archived from the original(PDF) on 2009-01-05.
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