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Stegoceras

Stegoceras is a genus of pachycephalosaurid (dome-headed) dinosaur that lived in what is now North America during the Late Cretaceous period, about 77.5 to 74 million years ago (mya). The first specimens from Alberta, Canada, were described in 1902, and the type species Stegoceras validum was based on these remains. The generic name means "horn roof", and the specific name means "strong". Several other species have been placed in the genus over the years, but these have since been moved to other genera or deemed junior synonyms. Currently only S. validum and S. novomexicanum, named in 2011 from fossils found in New Mexico, remain. The validity of the latter species has also been debated, and it may not even belong to the genus Stegoceras.

History of discovery
of S. validum, shown from the right and underside The first known remains of Stegoceras were collected by palaeontologist Lawrence Lambe from the Belly River Group, in the Red Deer River district of Alberta, Canada. These remains consisted of two partial skull domes (specimens CMN 515 and CMN 1423 in the Canadian Museum of Nature) from two animals of different sizes collected in 1898, and a third partial dome (CMN 1594) collected in 1901. Based on these specimens, Lambe described and named the new monotypic genus and species Stegoceras validus in 1902. The generic name Stegoceras comes from the Greek stegè/στέγη, meaning "roof" and keras/κέρας meaning "horn". The specific name validus means "strong" in Latin, possibly in reference to the thick skull-roof. In 1903, palaeontologist Franz Nopcsa von Felső-Szilvás suggested that the fragmentary domes of Stegoceras were in fact frontal and nasal bones, and that the animal would therefore have had a single, unpaired horn. Lambe was sympathetic to this idea of a new type of "unicorn dinosaur" in a 1903 review of Nopscsa's paper. At this time, there was still uncertainty over which group of dinosaur Stegoceras belonged to, with both ceratopsians (horned dinosaurs) and stegosaurs (plated dinosaurs) as contenders. Hatcher doubted whether the Stegoceras specimens belonged to the same species and whether they were dinosaurs at all, and suggested the domes consisted of the frontal, occipital, and parietal bones of the skull. In 1924, palaeontologist Charles W. Gilmore described a complete skull of S. validus with associated postcranial remains, by then the most complete remains of a dome-headed dinosaur. It was discovered in the Belly River Group by the American palaeontologist George F. Sternberg in 1926, and catalogued as specimen UALVP 2 in the University of Alberta Laboratory for Vertebrate Palaeontology. This find confirmed Hatcher's interpretation of the domes as consisting of the frontoparietal area of the skull. UALVP 2 was found with small, disarticulated bony elements, then thought to be gastralia (abdominal ribs), which are not known in other ornithischian dinosaurs (one of the two main groups of dinosaurs). Gilmore pointed out that the teeth of S. validus were very similar to those of the species Troodon formosus (named in 1856 and by then only known from isolated teeth), and described a skull dome discovered close to the locality where Troodon was found. Therefore, Gilmore considered Stegoceras an invalid junior synonym of Troodon, thereby renaming S. validus into T. validus, and suggested that even the two species might be the same. Furthermore, he found S. brevis to be identical to S. validus, and therefore a junior synonym of the latter. He also placed these species in the new family Troodontidae (since Lambe had not selected a type genus for his Psalisauridae), which he considered closest to the ornithopod dinosaurs. In 1945, after examining casts of T. formosus and S. validus teeth, palaeontologist Charles M. Sternberg demonstrated differences between the two, and instead suggested that Troodon was a theropod dinosaur, and that the dome-headed dinosaurs should be placed in their own family. Though Stegoceras was the first member of this family to be named, Sternberg named the group Pachycephalosauridae after the second genus, as he found that name (meaning "thick head lizard") more descriptive. He also considered T. sternbergi and T. edmontonensis members of Stegoceras, found S. brevis valid, and named a new species, S. lambei, based on a specimen formerly referred to S. validus. The split from Troodon was supported by Russell in 1948, who described a theropod dentary with teeth almost identical to those of T. formosus. of juvenile, flat-headed AMNH 5450 (formerly considered the distinct genus "Ornatotholus"), with sections on the right In 1953, palaeontologist Birger Bohlin named Troodon bexelli based on a parietal bone from China. In 1964, palaeontologist Oskar Kuhn considered this as an unequivocal species of Stegoceras; S. bexelli. In 1974, palaeontologists Teresa Maryańska and Halszka Osmólska concluded that the "gastralia" of Stegoceras were , after identifying such structures in the tail of the pachycephalosaur Homalocephale. In 1983, Galton and Hans-Dieter Sues moved S. browni to its own genus, Ornatotholus (ornatus is Latin for "adorned" and tholus for "dome"), and considered it the first known American member of a group of "flat-headed" pachycephalosaurs, previously known from Asia. In a 1987 review of the pachycephalosaurs, Sues and Galton emended the specific name validus to validum, which has subsequently been used in the scientific literature. These authors synonymized S. brevis, S. sternbergi, and S. lambei with S. validum, found that S. bexelli differed from Stegoceras in several features, and considered it an indeterminate pachycephalosaur. 21st century developments In 2000, palaeontologist Robert M. Sullivan referred S. edmontonensis and S. brevis to the genus Prenocephale (until then only known from the Mongolian species P. prenes), and found it more likely that S. bexelli belonged to Prenocephale than to Stegoceras, but considered it a nomen dubium (dubious name, without distinguishing characters) due to its incompleteness, and noted its holotype specimen appeared to be lost. In 2003, Thomas E. Williamson and Thomas Carr considered Ornatotholus a nomen dubium, or perhaps a juvenile Stegoceras. In a 2003 revision of Stegoceras, Sullivan agreed that Ornatotholus was a junior synonym of Stegoceras, moved S. lambei to the new genus Colepiocephale, and S. sternbergi to Hanssuesia. He stated that the genus Stegoceras had become a wastebasket taxon for small to medium-sized North American pachycephalosaurs until that point. By this time, dozens of specimens had been referred to S. validum, including many domes too incomplete to be identified as Stegoceras with certainty. UALVP 2 is still the most complete specimen of Stegoceras, upon which most scientific understanding of the genus is based. S. brevis was moved to the new genus Foraminacephale in 2016 by Ryan K. Schott Schott and David C. Evans, In 2023, Aaron D. Dyer and colleagues analysed sutures and individual elements in the skulls of the pachycephalosaurs Gravitholus and Hanssuesia, and found no significant distinction between them and Stegoceras validum. They considered both as junior synonyms, with Gravitholus representing the end-stage in the growth of Stegoceras. In 2002, Williamson and Carr described a dome (specimen NMMNH P-33983 in the New Mexico Museum of Natural History and Science) from the San Juan Basin, New Mexico, which they considered a juvenile pachycephalosaur of uncertain species (though perhaps Sphaerotholus goodwini). In 2006, Sullivan and Spencer G. Lucas considered it a juvenile S. validum, which would expand the range of the species considerably. In 2011, palaeontologist Steven E. Jasinski and Sullivan considered the specimen an adult, and made it the holotype of the new species Stegoceras novomexicanum, with two other specimens (SMP VP-2555 and SMP VP-2790) as paratypes. A 2011 phylogenetic analysis by Watabe and colleagues did not place the two Stegoceras species close to each other. In 2016, palaeontologists Williamson and Stephen L. Brusatte restudied the holotype of S. novomexicanum and found that the paratypes did not belong to the same taxon as the holotype, and that all the involved specimens were juveniles. Furthermore, they were unable to determine whether the holotype specimen represented the distinct species S. novomexicanum, or if it was a juvenile of either S. validum or Sphaerotholus goodwini, or another previously known pachycephalosaur. Dyer and colleagues found that the S. novomexicanum holotype could be an immature Sphaerotholus goodwini, because the proposed unique traits of S. novomexicanum disappeared through ontogeny in S. validum.'' ==Description==
Description
Stegoceras is one of the most completely known North American pachycephalosaurs, and one of the few known from postcranial remains; S. validum specimen UALVP 2 is the most complete Stegoceras individual known to date. Its length is estimated to have been about , comparable to the size of a goat. Stegoceras was small to medium in size compared to other pachycephalosaurs. S. novomexicanum appears to have been smaller than S. validum, but it is disputed whether the known specimens (incomplete skulls) are adults or juveniles. Skull and dentition The skull of Stegoceras was roughly triangular in shape when viewed from the side, with a relatively short snout. The frontal and parietal bones were very thick and formed an elevated dome. The suture between these two elements was obliterated (only faintly visible in some specimens), and they are collectively termed the "frontoparietal". The frontoparietal dome was broad and had a relatively smooth surface, with only the sides being rugose (wrinkled). It was narrowed above and between the orbita (eye sockets). The frontoparietal narrowed at the back, was wedged between the squamosal bones, and ended in a depression above the at the back of the skull. The parietal and squamosal bones formed a thick shelf over the occiput termed the parietosquamosal shelf, whose extent varied between specimens. The squamosal was large, not part of the dome, and the back part was swollen. It was ornamented by irregularly spaced tubercles (or round outgrowths), and a row of nodes (knobs) extended along its upper edges, ending in a pointed tubercle (or small horn) on each side at the back of the skull. An inner row of smaller tubercles ran parallel with the larger one. Except for the upper surface of the dome, much of the skull was ornamented with nodes, many arranged in rows. of S. validum, with integument speculatively based on other small ornithischian dinosaurs The scapula (shoulder blade) was longer than the humerus (upper arm bone); its blade was slender and narrow, and slightly twisted, following the contour of the ribs. The scapula did not expand at the upper end but was very expanded at the base. The coracoid was mainly thin and plate-like. The humerus had a slender shaft, was slightly twisted along its length, and was slightly bowed. The (where the deltoid and pectoral muscles attached) was weakly developed. The ends of the ulna were expanded, and ridges extended along the shaft. The radius was more robust than the ulna, which is unusual. When seen from above, the pelvic girdle was very broad for a bipedal archosaur, and became wider towards the hind part. The broadness of the pelvic region may have accommodated a rear extension of the gut. The ilium was elongated and the ischium was long and slender. Though the pubis is not known, it was probably reduced in size like that of Homalocephale. The femur (thigh bone) was slender and inwards curved, the tibia was slender and twisted, and the fibula was slender and wide at the upper end. The metatarsus of the foot appears to have been narrow, and the single known ungual (claw bone) of a toe was slender and slightly curved. ==Classification==
Classification
During the 1970s, more pachycephalosaur genera were described from Asian fossils, which provided more information about the group. In 1974, Maryańska and Osmólska concluded that pachycephalosaurs are distinct enough to warrant their own suborder within Ornithischia, Pachycephalosauria. In 1978, the Chinese palaeontologist Dong Zhiming split Pachycephalosauria into two families; the dome-headed Pachycephalosauridae (including Stegoceras) and the flat-headed Homalocephalidae (originally spelled Homalocephaleridae). Wall and Galton did not find suborder status for the pachycephalosaurs justified in 1979. The dome-headed/flat-headed division of the pachycephalosaurs was abandoned in the following years, as flat heads were considered paedomorphic (juvenile-like) or derived ("advanced") traits in most revisions, but not a sexually dimorphic trait. In 2006, Sullivan argued against the idea that the extent of doming was useful in determining taxonomic affinities between pachycephalosaurs. In 2003, Sullivan found Stegoceras itself to be more basal (or "primitive") than the "fully-domed" members of the subfamily Pachycephalosaurinae, elaborating on conclusions reached by Sereno in 1986. The cladogram below shows the placement of Stegoceras within Pachycephalosauridae according to Schott and colleagues, 2016: }} The biogeography and early evolutionary history of pachycephalosaurs is poorly understood, and can only be clarified by new discoveries. Pachycephalosaurs appear abruptly in the fossil record, and are present in both North America and Asia, so it is unknown when they first originated, and from which direction they dispersed. The oldest known members of the group (such as Acrotholus) are "fully domed" and known from the Santonian stage of the Late Cretaceous period (about 84 million years ago). This is before the supposedly more primitive Stegoceras from the Middle Campanian (77 million years ago) and Homalocephale from the Early Maastrichtian (70 million years ago), so the doming of the skull may be a homoplastic trait (a form of convergent evolution). The late occurrence of pachycephalosaurs compared to the related ceratopsians indicates a long ghost lineage (inferred, but missing from the fossil record) spanning 66 million years, from the Late Jurassic to the Cretaceous. Since pachycephalosaurs were mainly small, this may be due to taphonomic bias; smaller animals are less likely to be preserved through fossilisation. More delicate bones are also less likely to be preserved, which is why pachycephalosaurs are mainly known from their robust skulls. ==Palaeobiology==
Palaeobiology
Feeding mechanics It is uncertain what pachycephalosaurs ate; having very small, ridged teeth they could not have chewed tough, fibrous plants as effectively as other dinosaurs of the same period. It is assumed that their sharp, serrated teeth were ideally suited for a mixed diet of leaves, seeds, fruit and insects. Sues and Galton suggested in 1987 that Stegoceras may have had an entirely herbivorous diet, as the tooth crowns were similar to those of iguanid lizards. The premaxillary teeth show wear facets from contact with the predentary bone, and the maxillary teeth have double wear facets similar to those seen in other ornithischian dinosaurs. Every third maxillary tooth of UALVP 2 are erupting replacement teeth, and tooth replacement happened in backwards progression in sequential threes. The occipital region of Stegoceras was well-demarcated for muscle-attachment and it is believed that the jaw movement of Stegoceras and other pachycephalosaurs was mostly limited to up-and-down motions with only a slight capability for jaw rotation. This is based on the structure of the jaw and dental microwear and wear facets of the teeth indicate that the bite-force was used more for shearing than for crushing. In 2021, palaeontologist Michael N. Hudgins and colleagues examined the teeth of Stegoceras and Thescelosaurus and found that while both had heterodont teeth, they could be statistically distinguished from each other. Due to its broad rostrum and more uniform teeth, Stegoceras was an indiscriminate bulk-feeder that cropped large amounts of vegetation, while the teeth and narrow rostrum of Thescelosaurus indicates it was a selective feeder. Pachycephalosaurs and Thescelosaurids occur in the same North American formations, and it appears that their coexistence was made possible by them occupying different ecomorphospaces (though Stegoceras and Thescelosaurus themselves were not contemporaries). In 2026, palaeontologists Louis-Philippe Bateman and Hans C.E. Larsson examined the feeding performance in pachycephalosaurs using Stegoceras as a model, and found that the presence of their domes limited the space and movement of their jaw muscles. Compared with other orhithischians, the feeding performance of Stegoceras was closer to basal ornithiscians and ornithopods than would be expected of a derived taxon from the Late Cretaceous. They concluded that the dome constrained pachycephalosaurs to a more plesiomorphic (ancestral) dentition with a low-fiber herbivorous niche, and that it was an example of a sociosecual display structure constraining evolution of feeding traits. Nasal passages In 1989, Emily B. Griffin found that Stegoceras and other pachycephalosaurs had a good sense of smell (olfaction), based on the study of cranial endocasts that showed large olfactory bulbs in the brain. In 2014, Jason M. Bourke and colleagues found that Stegoceras would have needed cartilaginous nasal turbinates in the front of the nasal passages for airflow to reach the olfactory region. Evidence for the presence of this structure is a bony ridge to which it could have attached. The size of the olfactory region also indicates that Stegoceras had a keen sense of smell. The researchers found that the dinosaur could have had either a scroll-shaped turbinate (like in a turkey) or a branched one (as in an ostrich) as both could have directed air to the olfactory region. The blood vessel system in the passages also suggest that the turbinates served to cool down warm arterial blood from the body that was heading to the brain. The skull of S. validum specimen UALVP 2 was suited for a study of this kind due to its exceptional preservation; it has ossified soft tissue in the nasal cavity, which would otherwise be cartilaginous and therefore not preserved through mineralization. Ontogenetic changes Several explanations have historically been proposed for the variation seen in the skulls of Stegoceras and other pachycephalosaurs. Brown and Schlaikjer suggested that there was sexual dimorphism in the degree of doming, and hypothesized that flat-headed specimens such as AMNH 5450 (Ornatotholus) represented the female morph of Stegoceras. This idea was supported by a 1981 morphometric study by Champan and colleagues, which found that males had larger and thicker domes. After other flat-headed pachycephalosaurs were discovered, the degree of doming was proposed to be a feature with taxonomic importance, and AMNH 5450 was therefore considered a distinct taxon from 1979 onwards. In 1998, Goodwin and colleagues instead proposed that the inflation of the dome was an ontogenetic feature that changed with age, based on a histological study of an S. validum skull that showed the dome consisted of vascular, fast-growing bone, consistent with an increase in doming through age. These authors found that the supposedly distinct features of Ornatotholus could easily be the results of ontogeny. A 2012 study by Schott and Evans found that the number and shape of the individual nodes on the squamosal shelf of the examined S. validum skulls varied considerably, and that this variability does not seem to correlate with ontogenic changes, but was due to individual variation. These researchers found no correlation between the width of supratemporal fenestrae and the size of the squamosal. Dome function The function of pachycephalosaur domes has been debated, and Stegoceras has been used as a model for experimentation in various studies. The dome has mainly been interpreted as a weapon used in intra-specific combat, a sexual display structure, or a means for species recognition. In 1997, palaeontologist Kenneth Carpenter pointed out that the dorsal vertebrae from the back of the pachycephalosaur Homalocephale show that the back curved downwards just before the neck (which was not preserved), and unless the neck curved upwards, the head would point to the ground. He therefore inferred that the necks of Stegoceras and other pachycephalosaurs were held in a curved posture (as is the norm in dinosaurs), and that they would therefore not have been able to align their head, neck, and body horizontally straight, which would be needed to transmit stress. Their necks would have to be held below the level of the back, which would have risked damaging the spinal cord on impact. Modern bighorn sheep and bison overcome this problem by having strong ligaments from the neck to the tall neural spines over the shoulders (which absorb the force of impact), but such features are not known in pachycephalosaurs. These animals also absorb the force of impact through sinus chambers at the base of their horns, and their foreheads and horns form a broad contact surface, unlike the narrow surface of pachycephalosaur domes. Because the dome of Stegoceras was rounded, it would have given a very small area for potential impact, and the domes would have glanced off each other (unless the impact was perfectly centred). Combating pachycephalosaurs would have had difficulty seeing each other while their heads were lowered, due to the bony ridges above the eyes. In 2011, Snively and Jessica M. Theodor conducted a finite element analysis by simulating head-impacts with CT scanned skulls of S. validum (UALVP 2), Prenocephale prenes and several extant head-butting artiodactyls. They found that the correlations between head-striking and skull morphologies found in the living animals also existed in the studied pachycephalosaurs. Stegoceras and Prenocephale both had skull shapes similar to the bighorn sheep with cancellous bone protecting the brain. They also shared similarities in the distribution of compact and cancellous regions with the bighorn sheep, white-bellied duiker and the giraffe. The white-bellied duiker was found to be the closest morphological analogue to Stegoceras; this head-butting species has a dome which is smaller but similarly rounded. Stegoceras was better capable of dissipating force than artiodactyls that butt heads at high forces, but the less vascularized domes of older pachycephalosaurs, and possibly diminished ability to heal from injuries, argued against such combat in older individuals. The study also tested the effects of a keratinous covering of the dome, and found it to aid in performance. Though Stegoceras lacked the pneumatic sinuses that are found below the point of impact in the skulls of head-striking artiodactyls, it instead had vascular struts which could have similarly acted as braces, as well as conduits to feed the development of a keratin covering. In 2012, Caleb M. Brown and Anthony P. Russell suggested that the stiffened tails were probably not used as defence against flank-butting, but may have enabled the animals to take a tripodal stance during intra-specific combat, with the tail as support. Brown and Russell found that the tail could thereby help in resisting compressive, tensile, and torsional loading when the animal delivered or received blows with the dome. Bryan R. S. Moore and colleagues examined and reconstructed the limb musculature of Stegoceras in 3D in 2022, using the very complete UALVP 2 specimen as basis. They found that the musculature of the forelimbs was conservative, particularly compared to those of early bipedal saurischian dinosaurs, but the pelvic and hindlimb musculature was instead more derived, due to peculiarities of the skeleton. These areas had large muscles, and combined with the wide pelvis and stout hind limbs (and possibly enlarged ligaments), this resulted in a strong, stable pelvic structure that would have helped during head-butting between individuals. Since the skull domes of pachycephalosaurs grew with positive allometry, and may have been used in combat, these researchers suggested it may have been the case for the hindlimb muscles as well, if they were used to propel the body forwards during head-butting. They cautioned that while UALVP 2 is very complete for a pachycephalosaur, their study was limited by it missing large portions of its vertebral column and outer limb elements. Other suggested functions In 1987, J. Keith Rigby and colleagues suggested that pachycephalosaur domes were heat-exchange organs used for thermoregulation, based on their internal "radiating structures" (trabeculae). This idea was supported by a few other writers in the mid-1990s. In 2011, palaeontologists Kevin Padian and John R. Horner proposed that "bizarre structures" in dinosaurs in general (including domes, frills, horns, and crests) were primarily used for species recognition, and dismissed other explanations as unsupported by evidence. Among other studies, these authors cited Goodwin et al.'s 2004 paper on pachycephalosaur domes as support of this idea, and they pointed out that such structures did not appear to be sexually dimorphic. In a response to Padian and Horner the same year, Rob J. Knell and Scott D. Sampson argued that species recognition was not unlikely as a secondary function for "bizarre structures" in dinosaurs, but that sexual selection (used in display or combat to compete for mates) was a more likely explanation, due to the high cost of developing them, and because such structures appear to be highly variable within species. In 2013, the British palaeontologists David E. Hone and Darren Naish criticized the "species recognition hypothesis", and argued that no extant animals use such structures primarily for species recognition, and that Padian and Horner had ignored the possibility of mutual sexual selection (where both sexes are ornamented). In 2012, Schott and Evans suggested that the regularity in squamosal ornamentation throughout the ontogeny of Stegoceras was consistent with species recognition, but the change from flat to domed frontoparietals in late age suggests that the function of this feature changed through ontogeny, and was perhaps sexually selected, possibly for intra-specific combat. ==Palaeoenvironment==
Palaeoenvironment
in Alberta, Canada, where S. validum was first discovered S. validum is known from the late Late Cretaceous Belly River Group (the Canadian equivalent to the Judith River Group in the US), and specimens have been recovered from the Dinosaur Park Formation (late Campanian, 76.5 to 75 mya) in Dinosaur Provincial Park (including the lectotype specimen), and the Oldman Formation (middle Campanian, 77.5 to 76.5 mya) of Alberta, Canada. The pachycephalosaurs Hanssuesia (if not a synonym of Stegoceras) and Foraminacephale are also known from both formations. that lived alongside S. validum The Dinosaur Park Formation is interpreted as a low-relief setting of rivers and floodplains that became more swampy and influenced by marine conditions over time as the Western Interior Seaway transgressed westward. The climate was warmer than present-day Alberta, without frost, but with wetter and drier seasons. Conifers were apparently the dominant canopy plants, with an understory of ferns, tree ferns, and angiosperms. Dinosaur Park is known for its diverse community of herbivores. As well as Stegoceras, the formation has also yielded fossils of the ceratopsians Centrosaurus, Styracosaurus and Chasmosaurus, the hadrosaurids Prosaurolophus, Lambeosaurus, Gryposaurus, Corythosaurus, and Parasaurolophus, and the ankylosaurs Edmontonia and Euoplocephalus. Theropods present include the tyrannosaurids Gorgosaurus and Daspletosaurus. Other dinosaurs known from the Oldman Formation include the hadrosaur Brachylophosaurus, the ceratopsians Coronosaurus and Albertaceratops, ornithomimids, therizinosaurs and possibly ankylosaurs. Theropods included troodontids, oviraptorosaurs, the dromaeosaurid Saurornitholestes and possibly an albertosaurine tyrannosaur. ==See also==
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