Stegoceras
Stegoceras | |
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Two reconstructed S. validum skeletons based on specimen UALVP 2, Royal Tyrrell Museum
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | †Ornithischia |
Clade: | †Pachycephalosauria |
Family: | †Pachycephalosauridae |
Genus: | †Stegoceras Lambe, 1902 |
Type species | |
†Stegoceras validum Lambe, 1902
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Species | |
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Synonyms | |
List
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Stegoceras is a
Stegoceras was a small,
Originally known only from skull domes, Stegoceras was one of the first known pachycephalosaurs, and the incompleteness of these initial remains led to many theories about the affinities of this group. A complete Stegoceras skull with associated parts of the skeleton was described in 1924, which shed more light on these animals. Pachycephalosaurs are today grouped with the horned
History of discovery
The first known remains of Stegoceras were collected by Canadian
As no similar remains had been found in the area before, Lambe was unsure of what kind of dinosaur they were, and whether they represented one species or more; he suggested the domes were "prenasals" situated before the
In 1924, the American palaeontologist
Gilmore's classification was supported by the American palaeontologists
In 1953,
21st century developments
In 2000, 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.[18] In 2003, Thomas E. Williamson and Thomas Carr considered Ornatotholus a nomen dubium, or perhaps a juvenile Stegoceras.[19] 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.[4] S. brevis was moved to the new genus Foraminacephale in 2016 by Ryan K. Schott Schott and David C. Evans,[20] and S. bexelli to Sinocephale in 2021 by Evans and colleagues.[21] 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.[22]
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.[23][24] In 2011, 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.[25] A 2011 phylogenetic analysis by Watabe and colleagues did not place the two Stegoceras species close to each other.[26]
In 2016, Williamson and
Description
Stegoceras is one of the most completely known North American pachycephalosaurs, and one of the few known from
Skull and dentition
The skull of Stegoceras was roughly triangular in shape when viewed from the side, with a relatively short snout. The
The large orbit was shaped like an imperfect ellipse (with the longest axis from front to back), and faced to the side and slightly forward. The
The nasal openings were large and faced frontwards. The
Stegoceras had teeth that were
The skull of Stegoceras can be distinguished from those of other pachycephalosaurs by features such as its pronounced parietosquamosal shelf (though this became smaller with age), the "incipient" doming of its frontopariental (though the doming increased with age), its inflated nasal bones, its ornamentation of tubercles on the sides and back of the squamosal bones, rows of up to six tubercles on the upper side of each squamosal, and up to two nodes on the backwards projection of the parietal. It is also distinct in its lack of nasal ornamentation, and in having a reduced diastema.[4][34] The skull of S. novomexicanum can be distinguished from that of S. validum in features such as the backwards extension of the parietal bone being more reduced and triangular, having larger supratemporal fenestrae (though this may be due to the possible juvenile status of the specimens), and having roughly parallel suture contacts between the squamosal and parietal. It also appears to have had a smaller frontal boss than S. validum,[25][27] and seems to have been more gracile overall.[28]
Postcranial skeleton
The
The
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
By the early 21st century, few pachycephalosaur genera were known from postcranial remains, and many taxa were only known from domes, which made classification within the group difficult. Pachycephalosaurs are thus mainly defined by cranial features, such as the flat to domed frontoparietal, the broad and flattened bar along the postorbital and squamosal bones, and the squamosal bones being deep plates on the occiput.
A 2013
Pachycephalosauria |
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The
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.
In 2021, the Canadian palaeontologist Michael N. Hudgins and colleagues examined the teeth of Stegoceras and
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
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
In 2003, Williamson and Carr published a hypothetical growth series of S. validum, showing Ornatotholus as the juvenile stage. They suggested that juveniles were characterized by a flat, thickened frontoparietal roof, with larger supratemporal fenestrae, and studded with closely spaced tubercles and nodes. The parietosquamosal shelf was not reduced in size, and the frontoparietal suture was open. Sub-adults had mound-like domes, with the back part of the parietal and skull-roof being flat. The supratemporal fenestrae showed asymmetry in size, and the closure of the frontoparietal suture was variable. The nodes were stretched or almost obliterated as the dome expanded during growth, with a tesserated surface remaining. The pattern was often obliterated at the highest point (apex) of the dome, the area where maximum expansion occurred. The tubercles on the skull were stretched in different directions, and those at the margin of the parietosquamosal shelf may have been hypertrophied (enlarged) tubercles. The back and sides of sub-adult and adult skulls were ornamented by less modified tubercles. Before being incorporated into the enlarging dome, the skull bones expanded, resulting in junctions between these bones. The adult dome was broad and convex, and incorporated most of the shelf, which was reduced in size and overhung the occiput as a thick "lip". The supratempooral fenestrae were closed, but the suture between the frontoparietal and connected skull bones was not always closed in adults and subadults.[19]
In 2011, Schott and colleagues made a more comprehensive analysis of cranial dome ontogeny in S. validum. The study found that the parietosquamosal shelf conserved the arrangement of ornamentation throughout growth, and that vascularity of the frontoparietal domes decreased with size. It also found that dome shape and size was strongly correlated with growth, and that growth was
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.[45]
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
Combat
The hypothesis that the domed skulls of Stegoceras and other pachycephalosaurs were used for butting heads was first suggested by American palaeontologist
In 1997, the American 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.[35]
Because of the problems he found with the head-butting hypothesis, Carpenter instead suggested the domes were adaptations for flank-butting (as seen in some large African mammals); he imagined that two animals would stand parallel, facing each other or the same direction, and direct blows to the side of the opponent. The relatively large body width of pachycephalosaurs may consequently have served to protect vital organs from harm during flank-butting. It is possible that Stegoceras and similar pachycephalosaurs would have delivered the blows with a movement of the neck from the side and a rotation of the head. The upper sides of the dome have the greatest surface area, and may have been the point of impact. The thickness of the dome would have increased the power behind a blow to the sides, and this would ensure that the opponent felt the force of the impact, without being seriously injured. The bone rim above the orbit may have protected the aggressor's eye when making a blow. Carpenter suggested that the pachycephalosaurs would have first engaged in
In 2008, Eric Snively and Adam Cox tested the performance of 2D and 3D pachycephalosaur skulls through
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.[9] A 2013 study by Joseph E. Peterson and colleagues identified lesions in skulls of Stegoceras and other pachycephalosaurs, which were interpreted as infections caused by trauma. Lesions were found on 22% of sampled pachycephalosaur skulls (a frequency consistent across genera), but were absent from flat-headed specimens (which have been interpreted as juveniles or females), which is consistent with use in intra-specific combat (for territory or mates). The distribution of lesions in these animals tended to concentrate at the top of the dome, which supports head-butting behaviour. Flank-butting would probably result in fewer injuries, which would instead be concentrated on the sides of the dome. These observations were made while comparing the lesions with those on the skulls and flanks of modern sheep skeletons. The researchers noted that modern head-butting animals use their weapons for both combat and display, and that pachycephalosaurs could therefore also have used their domes for both. Displaying a weapon and willingness to use it can be enough to settle disputes in some animals.[46]
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
Other suggested functions
In 1987, J. Keith Rigby and colleagues suggested that pachycephalosaur domes were
In 2011, American palaeontologists
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.[45] Dyer and colleagues found in 2023 that Stegoceras specimens differed in the thickness of the frontonasal boss, and that skulls with the most bone pathologies were those with the tallest bosses, which they considered indication that variation in boss thickness represents intersexual variation.[22]
Palaeoenvironment
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.[4][22] S. novomexicanum is known from the Fruitland (late Campanian, about 75 mya) and lower Kirtland Formation (late Campanian, about 74 mya) of New Mexico, and if this species correctly belongs in Stegoceras, the genus would have had a broad geographic distribution.[25] The presence of similar pachycephalosaurs in both the west and north of North America during the latest Cretaceous shows that they were an important part of the dinosaur faunas there.[27]
It has traditionally been suggested that pachycehalosaurs inhabited mountain environments; wear of their skulls was supposedly a result of them having been rolled by water from upland areas, and comparisons with bighorn sheep reinforced the theory. In 2014, Jordan C. Mallon and Evans disputed this idea, as the wear and original locations of the skulls is not consistent with having been transported in such a way, and they instead proposed that North American pachycephalosaurs inhabited
The Dinosaur Park Formation is interpreted as a low-relief setting of
See also
References
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External links
- Media related to Stegoceras at Wikimedia Commons
- Data related to Stegoceras at Wikispecies