Stegoceras - Knowledge

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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.

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itself lacked spikes, and those of the parietosquamosal shelf were in an "ineffective" position, but found it compatible with intra-specific competition. Galton imagined the domes were bashed together, while the vertebral column was held in a horizontal position. This could either be done while facing each other while dealing blows, or while charging each other with lowered heads (analogous to modern sheep and goats). He also noted that the rigidity of the back would have been useful when using the head for this purpose. In 1978, Sues agreed with Galton that the anatomy of pachycephalosaurs was consistent with transmitting dome-to-dome impact stress, based on tests with

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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

2956: 1562: 1826: 2892:(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. 774: 1511: 3042: 1844: 2915:. They also hypothesized that this model of dome growth, with dramatic changes from juvenile to adult, was the common developmental trajectory of pachycephalosaurs. These researchers noted that though Williamson and Carr's observation that the supratemporal fenestrae closed with age was generally correct, there was still a high degree of individual variation in the size of these fenestrae, regardless of the size of the frontoparietal, and this feature may therefore have been independent of ontogeny. 2820: 1835: 138: 1019: 1815: 1028: 3247: 163: 3344: 3030: 1806: 765: 6354: 2868: 1010: 6377: 2028:) and the flat-headed Homalocephalidae (originally spelled Homalocephaleridae). Wall and Galton did not find suborder status for the pachycephalosaurs justified in 1979. By the 1980s, the affinities of the pachycephalosaurs within Ornithischia were unresolved. The main competing views were that the group was closest to either ornithopods or ceratopsians, the latter view due to similarities between the skeleton of 1181: 2097: 2698: 5951: 3150: 1693: 1797: 611: 1742:(space), and the two rows in the premaxilla were separated by a toothless gap at the front. The teeth in the front part of the upper jaw (premaxilla) and front lower jaw were similar; these had taller, more pointed and recurved crowns, and a "heel" at the back. The front teeth in the lower jaw were larger than those of the upper jaw. The front edges of the crowns bore eight 2050:

broad and flattened bar along the postorbital and squamosal bones, and the squamosal bones being deep plates on the occiput. In 1986, Sereno had divided the pachycephalosaurs into different groups based on the extent of the doming of their skulls (grouped in now invalid taxa such as "Tholocephalidae" and "Domocephalinae"), and in 2000 he considered the "partially" domed

3238:, 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. 3624: 2005: 5939: 1685:. The maxilla had a series of foramina that corresponded with each tooth position there, and these functioned as passages for erupting replacement teeth. The mandible articulated with the skull below the back of the orbit. The tooth-bearing part of the lower jaw was long, with the part behind being rather short. Though not preserved, the presence of a 3234:(or "advanced"), 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 1607:(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. 1730:(placed in sockets). It had marginal rows of relatively small teeth, and the rows did not form a straight cutting edge. The teeth were set obliquely along the length of the jaws, and overlapped each other slightly from front to back. On each side, the most complete specimen (UALVP 2) had three teeth in the premaxilla, sixteen in the 3325:

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. In 2023, Horner and colleagues stated that since the dome

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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

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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,

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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.

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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

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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

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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

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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

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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.

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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

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developed early in life, this indicates they were used for visual communication, so that juveniles could recognise other juveniles and adults other adults. They did not rule out that these features could have been used for other purposes, including head-butting, but did not consider trauma seen in

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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

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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

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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

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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

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held its skull vertically, perpendicular to the neck, force would be transmitted from the skull, with little chance of it being dislocated, and the dome could therefore be used as a battering-ram. He believed it was unlikely to have been used mainly as defence against predators, because the dome

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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

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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.

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and other pachycephalosaurs enabled them to take a tripodal stance (first suggested by Maryańska and Osmólska in 1974), Goodwin et al. suggested these structures could have protected the tail against flank-butting, or that the tail itself could have been used as a weapon. In 2004, Goodwin and

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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

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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

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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

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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

718:

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

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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.

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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. Dyer and colleagues found in 2023 that

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colleagues studied the cranial histology of pachycephalosaurs, and found that the vascularity (including the trabeculae) of the domes decreased with age, which they found inconsistent with a function in either head-butting or heat-exchange. They also suggested that a dense layer of

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dinosaur about 2 to 2.5 metres (6.6 to 8.2 ft) long, and weighed around 10 to 40 kilograms (22 to 88 lb). The skull was roughly triangular with a short snout, and had a thick, broad, and relatively smooth dome on the top. The back of the skull had a thick "shelf" over the

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near the surface of the dome indicated that it had an external covering in life, which makes it impossible to know the shape of the dome in a living animal. These researchers instead concluded that the domes were mainly for species recognition and communication (as in some African

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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

2899:. 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 1770:

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. The skull of

531:(or round "outgrowths") and nodes (or "knobs"), many in rows, and the largest formed small horns on the shelf. The teeth were small and serrated. The skull is thought to have been flat in juvenile animals and to have grown into a dome with age. It had a rigid 2922:

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.

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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).

1411:; they agreed that some features used to diagnose the species were indicative of a sub-adult stage, but presented additional diagnostic features in the holotype that distinguish the species. They also pointed out some adult features, which may indicate 3308:(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 4345:

Evans, David C.; Brown, Caleb M.; You, Hailu; Campione, Nicolás E. (October 2021). "Description and revised diagnosis of Asia's first recorded pachycephalosaurid, Sinocephale bexelli gen. nov., from the Upper Cretaceous of Inner Mongolia, China".

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Bohlin, B., 1953. Fossil reptiles from Mongolia and Kansu. Reports from the Scientific Expedition to the North-western Provinces of China under Leadership of Dr. Sven Hedin. VI. Vertebrate Palaeontology 6. The Sino-Swedish Expedition Publication

1746:(serrations), and the back edge bore nine to eleven. The teeth in the back of the upper (maxilla) and lower jaw were triangular in side view and compressed in front view. They had long roots that were oval in section, and the crowns had a marked 3414:, the genus would have had a broad geographic distribution. 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. 2714:

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

3274:"-like network of vascular bone in the dome ill-suited for head-butting, and pointed out that the bones adjacent to the dome risked fracture during such contact. Building on the idea that the ossified tendons that stiffened the tails of 2015:

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

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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

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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.

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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

2803:) 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 2046:, based on similar cranial features, such as the "shelf"-structure above the occiput. He conceded that the evidence for this grouping was not overwhelming, but the validity of the group was supported by Sues and Galton in 1987. 490:

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

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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

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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

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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 1918, Lambe referred another dome (CMN 138) to

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in 1955. In 1970 and 1971, Galton elaborated on this idea, and argued that if the dome was simply ornamental, it would have been less dense, and that the structure was ideal for resisting force. Galton suggested that when

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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

5864:

Weishampel, D. B.; Barrett, P. M.; Coria, R. A.; Le Loeuff, J.; Xu Xing; Z. X.; Sahni, A.; Gomani M.P., Elizabeth; Noto, C. R. (2004). "Dinosaur Distribution". In Weishampel, D. B.; Dodson, P.; Osmólska, H. (eds.).

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thereby help in resisting compressive, tensile, and torsional loading when the animal delivered or received blows with the dome. A 2013 study by Joseph E. Peterson and colleagues identified lesions in skulls of

4624:

Wick, Steven L.; Lehman, Thomas M. (19 September 2024). "A rare 'flat-headed' pachycephalosaur (Dinosauria: Pachycephalosauridae) from West Texas, USA, with morphometric and heterochronic considerations".

2684:; 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. 891:

dinosaurs. Because the skull seemed so specialized compared to the rather "primitive"-looking skeleton, Nopcsa doubted whether these parts actually belonged together, and suggested the skull belonged to a

2749:

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

3075:, and found that they could withstand considerable impact; greater vaulting of the domes allowed for higher forces of impact. They also considered it likely that pachycephalosaur domes were covered in 1419:

changes between related taxa) in the species. They conceded that the paratypes and other assigned specimens differed from the holotype in having more highly domed skulls, instead referring to them as

3006:

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

4959:

Sereno, P. C., 2000. The fossil record, systematics and evolution of pachycephalosaurs and ceratopsians from Asia. 480–516 in Benton, M.J., M.A. Shishkin, D.M. Unwin & E.N. Kurochkin (eds.),

1921:(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 3660: 3418:

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

1870:(articular processes) of successive dorsal (back) vertebrae appears to have prevented sideways movement of the vertebral column, which made it very rigid, and it was further strengthened by 3335:

specimens as evidence for this. They also suggested that features in some pachycephalosaurid skulls indicate the dome would have supported a greater, keratinous structure than just a cap.

3288:) and that use in sexual display was only secondary. They further speculated that the external covering of the domes was brightly coloured in life, or may have changed colour seasonally. 5448:

Galton, P. M. (1971). "A Primitive Dome-Headed Dinosaur (Ornithischia: Pachycephalosauridae) from the Lower Cretaceous of England and the Function of the Dome of Pachycephalosaurids".

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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

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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

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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

1882:(which articulates with the first neck vertebra) indicates that the neck was held in a curved posture, like the "S"- or "U"-shape of most dinosaur necks. Based on their position in 3059:

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

4488:"Re-evaluation of pachycephalosaurids from the Fruitland-Kirtland transition (Kirtlandian, late Campanian), San Juan Basin, New Mexico, with a description of a new species of 941:, and therefore a junior synonym. By this time, the dome-headed dinosaurs were either considered relatives of ornithopods or of ankylosaurs. In 1945, after examining casts of 3068:

by bobbing and presenting their heads to show the size of their domes (intimidation), and thereafter delivered blows to each other, until one opponent signalled submission.

2066:

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

824:

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

653:) 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 3087:), and therefore incorporated keratin into their test formula. In 2011, Snively and Jessica M. Theodor conducted a finite element analysis by simulating head-impacts with 6541: 1661:

was thick, heavily sculpted, and had a convex profile. It formed a boss (shield) on the middle top of the skull together with the frontal bone. The lower front of the

1348: 4132:

Wall, W. P.; Galton, P. M. (1979). "Notes on pachycephalosaurid dinosaurs (Reptilia: Ornithischia) from North America, with comments on their status as ornithopods".

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but not to the extent to which the authors of the study outright referred it to that species. Nevertheless, the authors of the study considered that the holotype of

6457: 2807:

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

1231:(dubious name, without distinguishing characters) due to its incompleteness, and noted its holotype specimen appeared to be lost. In 2003, Thomas E. Williamson and 5639:

Padian, K.; Horner, J. R. (2011). "The evolution of 'bizarre structures' in dinosaurs: biomechanics, sexual selection, social selection or species recognition?".

542:

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

1654:

formed the lower margin of the orbit, extending far forwards and down towards the jaw joint. It was ornamented with ridges and nodes in a radiating arrangement.

546:

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

5983: 2081:

by Evans and colleagues found that some flat-headed pachycephalosaur genera were more closely related to "fully" domed taxa than to the "incompletely" domed

2741:

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,

6511: 3103:. They found that the correlations between head-striking and skull morphologies found in the living animals also existed in the studied pachycephalosaurs. 896:, the skeleton to an ornithopod, and the supposed gastralia (belly ribs) to a fish. This claim was rebutted by Gilmore and Loris S. Russell in the 1930s. 566:

was most likely herbivorous, and it probably had a good sense of smell. The function of the dome has been debated, and competing theories include use in

5732:

Horner, John R.; Goodwin, Mark B.; Evans, David C. (2023). "A new pachycephalosaurid from the Hell Creek Formation, Garfield County, Montana, U.S.A.".

5519:"Common Functional Correlates of Head-Strike Behavior in the Pachycephalosaur Stegoceras validum (Ornithischia, Dinosauria) and Combative Artiodactyls" 1322:

in 2021 by Evans and colleagues. In 2023, Aaron D. Dyer and colleagues analysed sutures and individual elements in the skulls of the pachycephalosaurs

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models. The impact would be absorbed through the neck and body, and neck ligaments and muscles would prevent injuries by glancing blows (as in modern

6444: 4939:

gen. et sp. n., a new flat-headed pachycephalosaur (Ornlthlschia, Dinosauria) from the Upper Cretaceous of Mongolia – Acta Palaeontologica Polonica"

4911: 3266:). This idea was supported by a few other writers in the mid-1990s. In 1998, Goodwin and colleagues considered the lack of sinuses in the skull of 1426:(difficult to identify), but found it likely they all belonged to the same taxon (with the assigned specimens being adults), due to the restricted 3957:"Homology and Architecture of the Caudal Basket of Pachycephalosauria (Dinosauria: Ornithischia): The First Occurrence of Myorhabdoi in Tetrapoda" 907:

in their 1943 review of the dome-headed dinosaurs, by then known from 46 skulls. From these specimens, Brown and Schlaikjer named the new species

1610:

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

1762:, and both sides were divided vertically by a ridge. Each edge had about seven or eight denticles, with the front edge usually having the most. 6516: 5392:"Cranial Histology of Pachycephalosaurs (Ornithischia: Marginocephalia) Reveals Transitory Structures Inconsistent with Head-Butting Behavior" 6521: 6370: 5919: 5882: 5848: 5813: 5234:

Chapman, R. E.; Galton, Pe. M.; Sepkoski, J. J.; Wall, W. P. (1981). "A Morphometric Study of the Cranium of the Pachycephalosaurid Dinosaur

5042: 4817: 4701: 4668: 3637: 2852:

In 1998, Goodwin and colleagues instead proposed that the inflation of the dome was an ontogenetic feature that changed with age, based on a

2074:(or "primitive") than the "fully-domed" members of the subfamily Pachycephalosaurinae, elaborating on conclusions reached by Sereno in 1986. 1894:" in the tail, consisting of parallel rows, with the extremities of each tendon contacting the next successively. Such structures are called 5668:"Bizarre structures in dinosaurs: species recognition or sexual selection? A response to Padian and Horner: Bizarre structures in dinosaurs" 3158: 711: 5759:

Mallon, J. C.; Evans, D. C. (2014). "Taphonomy and habitat preference of North American pachycephalosaurids (Dinosauria, Ornithischia)".

3204: 3195: 1537:

individual known to date. Its length is estimated to have been about 2 to 2.5 metres (6.6 to 8.2 ft), comparable to the size of a

5025:

Maryańska, T.; Chapman, R. E.; Weishampel, D. B. (2004). "Pachycephalosauria". In Weishampel, D. B.; Dodson, P.; Osmólska, H. (eds.).

4460:

Williamson, T. E.; Carr, T. D. (2002). "A juvenile pachycephalosaur (Dinosauria: Pachycephalosauridae) from the Fruitland Formation".

2009: 1108: 5701:"The 'species recognition hypothesis' does not explain the presence and evolution of exaggerated structures in non-avialan dinosaurs" 2058:). The dome-headed/flat-headed division of the pachycephalosaurs was abandoned in the following years, as flat heads were considered 4017:

Brown, B.; E. M., Schlaikjer (1943). "A study of the troödont dinosaurs, with the description of a new genus and four new species".

1960:, and became wider towards the hind part. The broadness of the pelvic region may have accommodated a rear extension of the gut. The 1925:

was mainly thin and plate-like. The humerus had a slender shaft, was slightly twisted along its length, and was slightly bowed. The

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at their bases. The denticles here were compressed and directed towards the top of the crowns. Both the outer and inner side of the

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for small to medium-sized North American pachycephalosaurs until that point. By this time, dozens of specimens had been referred to

4322:

gen. nov. and the diversity of pachycephalosaurid dinosaurs (Ornithischia: Cerapoda) in the Belly River Group of Alberta, Canada".

4840:

Schott, Ryan K.; Evans, David C.; Goodwin, Mark B.; Horner, John R.; Brown, Caleb Marshall; Longrich, Nicholas R. (29 June 2011).

3115:

protecting the brain. They also shared similarities in the distribution of compact and cancellous regions with the bighorn sheep,

6526: 5976: 3016:

their foreheads and horns form a broad contact surface, unlike the narrow surface of pachycephalosaur domes. Because the dome of

2918:

A 2012 study by Schott and Evans found that the number and shape of the individual nodes on the squamosal shelf of the examined

2415: 684:

means "strong" in Latin, possibly in reference to the thick skull-roof. Because the species was based on multiple specimens (a

5576:

Moore, Bryan R. S.; Roloson, Mathew J.; Currie, Philip J.; Ryan, Michael J.; Patterson, R. Timothy; Mallon, Jordan C. (2022).

4487: 2795:

had a keen sense of smell. The researchers found that the dinosaur could have had either a scroll-shaped turbinate (like in a

851:(named in 1856 and by then only known from isolated teeth), and described a skull dome discovered close to the locality where 748:). By this time, he considered these animals as members of Stegosauria (then composed of both families of armoured dinosaurs, 4398:"Problematic putative pachycephalosaurids: Synchrotron µCT imaging shines new light on the anatomy and taxonomic validity of 3171:

domes with lesions (arrows, left) shown from above, and distribution of observed pachycephalosaurid skull lesions shown on a

4555:"Pachycephalosaurs (Dinosauria: Ornithischia) from the Upper Cretaceous (upper Campanian) of New Mexico: A reassessment of 4167:

Galton, P. M.; Sues, H.-D. (1983). "New data on pachycephalosaurid dinosaurs (Reptilia: Ornithischia) from North America".

6536: 2935:

has been used as a model for experimentation in various studies. The dome has mainly been interpreted as a weapon used in

1996:

are not completely known, they were most likely like other pachycephalosaurs in having five-fingered hands and four toes.

3899: 3304:

argued that species recognition was not unlikely as a secondary function for "bizarre structures" in dinosaurs, but that

5328:"Distributions of Cranial Pathologies Provide Evidence for Head-Butting in Dome-Headed Dinosaurs (Pachycephalosauridae)" 1927: 1896: 1888: 1872: 1748: 1724: 1628: 1620: 1596: 1232: 2955: 2085:, which suggests they represent juveniles of domed taxa, and that flat heads do not indicate taxonomic affinities. The 5969: 2271: 1164:

in several features, and considered it an indeterminate pachycephalosaur. In 1998, Goodwin and colleagues considered

5187:"Breathing life into dinosaurs: tackling challenges of soft-tissue restoration and nasal airflow in extinct species" 1689:

is indicated by facets at the front of the lower jaw. Like other pachycephalosaurs, it would have had a small beak.

162: 6380: 4521: 3231: 1561: 5185:

Bourke, J. M.; Porter, Wm. R.; Ridgely, R. C.; Lyson, T. R.; Schachner, E. R.; Bell, P. R.; Witmer, L. M. (2014).

2745:

was an indiscriminate bulk-feeder that cropped large amounts of vegetation, while the teeth and narrow rostrum of

1565:

CT images of skull UALVP 2 in multiple views, showing relative surface densities of bone (blue: low, yellow: high)

6531: 2532: 2063: 969:

after the second genus, as he found that name (meaning "thick head lizard") more descriptive. He also considered

650: 5418: 4293: 4280:

Williamson, T. E.; Carr, T. D. (2003). "A new genus of derived pachycephalosaurian from western North America".

1825: 1371:, Steven E. Jasinski and Sullivan considered the specimen an adult, and made it the holotype of the new species 6074: 4978:"The oldest North American pachycephalosaurid and the hidden diversity of small-bodied ornithischian dinosaurs" 3375: 2652:

stage of the Late Cretaceous period (about 84 million years ago). This is before the supposedly more primitive

2126: 1743: 3794: 3316:

In 2012, Schott and Evans suggested that the regularity in squamosal ornamentation throughout the ontogeny of

773: 6353: 4256:(Lambe) new comb. (Dinosauria: Ornithischia: Pachycephalosauria) from the Upper Cretaceous of North America" 3449: 3446: 3371: 3348: 3072: 2432: 1938: 1638:. The occiput sloped backwards and down, and the occipital condyle was deflected in the same direction. The 583: 509:, remain. The validity of the latter species has also been debated, and it may not even belong to the genus 1465:, but different in some aspects. They considered it a possible representative of a new southern species of 997:

was supported by Russell in 1948, who described a theropod dentary with teeth almost identical to those of

6397: 5268:

Schott, R. K.; Evans, D. C. (2012). "Squamosal Ontogeny and Variation in the Pachycephalosaurian Dinosaur

4522:"A new pachycephalosaurid from the Baynshire Formation (Cenomanian-late Santonian), Gobi Desert, Mongolia" 2515: 2247: 1615: 1611: 1510: 904: 5943: 4912:"Agonistic behavior in pachycephalosaurs (Ornithischia, Dinosauria); a new look at head-butting behavior" 1738:

of the lower jaw. The teeth in the premaxilla were separated from those behind in the maxilla by a short

6483: 6101: 3041: 2977:

and other pachycephalosaurs were used for butting heads was first suggested by American palaeontologist

2936: 2819: 2719:, and tooth replacement happened in backwards progression in sequential threes. The occipital region of 2177: 2160: 2078: 2071: 1843: 1618:

on the top back of the skull was very reduced in size, due to the thickening of the frontoparietal. The

1483: 1407:, or another previously known pachycephalosaur. In 2016, Jasinski and Sullivan defended the validity of 829: 567: 559: 4211:(Ornithischia: Pachycephalosauria) with comments on cranial display structures in agonistic behavior". 3071:

In 2008, Eric Snively and Adam Cox tested the performance of 2D and 3D pachycephalosaur skulls through

2054:

a transition between the supposedly "primitive" flat-headed and advanced "fully" domed genera (such as

5099:

Hudgins, Michael Naylor; Currie, Philip J.; Sullivan, Corwin (16 October 2021). "Dental assessment of

3688: 887:(since Lambe had not selected a type genus for his Psalisauridae), which he considered closest to the 6248: 6196: 5768: 5593: 5530: 5406: 5339: 5281: 5151: 4989: 4857: 4846:(Dinosauria: Pachycephalosauria): A Quantitative Model of Pachycephalosaur Dome Growth and Variation" 4762: 4570: 4355: 4220: 4176: 4141: 3968: 3856: 3116: 3048: 2669: 2473: 2295: 1642:

formed the lower front margin of the orbit, and its surface had rows of node-like ornamentation. The

1368: 966: 950: 662: 292: 150: 3844: 2969:

skulls (left, arrows), with possible combat orientations (right), according to Peterson et al., 2013

2042:

supported the relationship between pachycephalosaurs and ceratopsians, and united them in the group

1525:

is one of the most completely known North American pachycephalosaurs, and one of the few known from

5911: 3399: 3280: 2944: 2665: 2456: 2367: 1674: 1387: 1078: 1074: 825: 595: 575: 137: 2672:). The late occurrence of pachycephalosaurs compared to the related ceratopsians indicates a long 1834: 6299: 6164: 6143: 6113: 5992: 5457: 5430: 5422: 5305: 5297: 5247: 5216: 5167: 5124: 4778: 4735: 4371: 4297: 4088: 4057: 3872: 3798: 3403: 3367: 3235: 2774:

and other pachycephalosaurs had a good sense of smell (olfaction), based on the study of cranial

2233: 2139: 2112: 1852:

Bones of UALVP 2 including those of the limbs, pelvis, vertebrae, ribs, and ossified tail tendons

921: 817: 782: 689: 599: 459: 363: 268: 255: 157: 5840: 5834: 4809: 4803: 3743:"Anatomy and classification of the North American Pachycephalosauria (Dinosauria: Ornithischia)" 1814: 1018: 6488: 5874: 5805: 5799: 5034: 3246: 3079:, a strong material that can withstand much energy without being permanently damaged (like the 1027: 883:, and therefore a junior synonym of the latter. He also placed these species in the new family 6470: 6462: 5915: 5878: 5844: 5830: 5809: 5621: 5558: 5367: 5208: 5081: 5038: 5007: 4885: 4813: 4723: 4697: 4693: 4686: 4664: 4638: 4602: 4434: 4247: 3996: 3880: 3608: 3571: 3343: 2999: 2895:

In 2011, Schott and colleagues made a more comprehensive analysis of cranial dome ontogeny in

2836: 2731:

In 2021, the Canadian palaeontologist Michael N. Hudgins and colleagues examined the teeth of

2716: 2384: 1879: 1573:

was roughly triangular in shape when viewed from the side, with a relatively short snout. The

1553:, but it is disputed whether the known specimens (incomplete skulls) are adults or juveniles. 1284: 1104: 756:), in a new family he called Psalisauridae (named for the vaulted or dome-shaped skull roof). 693: 638: 6475: 5795: 2839:

in the degree of doming, and hypothesized that flat-headed specimens such as AMNH 5450 (

2831:

Several explanations have historically been proposed for the variation seen in the skulls of

1085:

were ossified tendons, after identifying such structures in the tail of the pachycephalosaur

5903: 5866: 5776: 5741: 5712: 5679: 5648: 5611: 5601: 5548: 5538: 5414: 5357: 5347: 5289: 5198: 5159: 5116: 5071: 5026: 4997: 4875: 4865: 4770: 4630: 4578: 4413: 4363: 4327: 4289: 4228: 4184: 4149: 4026: 3986: 3976: 3864: 3790: 3700: 3629: 3379: 3305: 3301: 3259: 2657: 2024:

split Pachycephalosauria into two families; the dome-headed Pachycephalosauridae (including

1859: 1696: 1582: 1112: 1041: 847: 649:. These remains consisted of two partial skull domes (specimens CMN 515 and CMN 1423 in the 587: 532: 474: 6435: 5955: 4207:

Goodwin, M. B.; Buchholtz, E. A.; Johnson, R. E. (1998). "Cranial anatomy and diagnosis of

1140:, which has subsequently been used in the scientific literature. These authors synonymized 841:

dinosaurs (one of the two main groups of dinosaurs). Gilmore pointed out that the teeth of

6278: 6059: 6043: 4799: 3583: 3535: 3363: 3112: 3029: 2779: 2751: 2681: 2319: 2043: 1686: 1682: 1643: 1587: 1454: 1305: 961:

dinosaur, and that the dome-headed dinosaurs should be placed in their own family. Though

551: 492: 471: 48: 5904: 5475:

Sues, H. D. (1978). "Functional morphology of the dome in pachycephalosaurid dinosaurs".

2668:(70 million years ago), so the doming of the skull may be a homoplastic trait (a form of 1359:, which they considered a juvenile pachycephalosaur of uncertain species (though perhaps 5772: 5597: 5534: 5410: 5343: 5285: 5155: 4993: 4861: 4766: 4574: 4359: 4224: 4180: 4145: 3972: 3860: 2710:

may have had an entirely herbivorous diet, as the tooth crowns were similar to those of

1980:(thigh bone) was slender and inwards curved, the tibia was slender and twisted, and the 1928: 1805: 764: 6209: 6177: 5616: 5577: 5553: 5518: 5362: 5327: 4880: 4841: 4656: 3991: 3956: 3600: 3565: 3550: 3511: 3296: 3065: 2940: 2737: 1934: 1647: 1591: 1352: 1265: 860: 804: 753: 685: 642: 634: 571: 288: 239: 5582:(Ornithischia: Pachycephalosauridae) and implications for the head-butting hypothesis" 5391: 2867: 2723:

was well-demarcated for muscle-attachment and it is believed that the jaw movement of

6505: 6327: 6292: 6264: 6185: 5867: 5684: 5667: 5652: 5142:

Giffin, E. B. (1989). "Pachycephalosaur Paleoneurolagy (Archosauria: Ornithischia)".

5128: 5027: 4934: 4375: 3742: 3604: 3592: 3555: 3529: 3496: 3465: 3423: 3255: 3011:

which would have risked damaging the spinal cord on impact. Modern bighorn sheep and

2992: 2978: 2808: 2796: 2677: 2673: 2034: 1950: 1873: 1708: 1639: 1578: 1427: 1376: 1087: 1051: 749: 670: 535:, and a stiffened tail. The pelvic region was broad, perhaps due to an extended gut. 467: 88: 5434: 5309: 4301: 3802: 3222:

Bryan R. S. Moore and colleagues examined and reconstructed the limb musculature of

1180: 1009: 6313: 6086: 6034: 5220: 5163: 4774: 4739: 4232: 4044:

Sternberg, C. M. (1945). "Pachycephalosauridae Proposed for Dome-Headed Dinosaurs,

3588: 3577: 3517: 3502: 3490: 3309: 3292: 3100: 3084: 2965: 2853: 2848: 2787: 2697: 2676:(inferred, but missing from the fossil record) spanning 66 million years, from the 2638: 2096: 2059: 2021: 1969: 1961: 1903: 1867: 1759: 1755: 1670: 1574: 1541:. The weight has been estimated to be about 10 to 40 kilograms (22 to 88 lb). 1461:

based on morphometric analyses. It was a juvenile, very comparable to juveniles of

1412: 1227: 1205: 1092: 900: 884: 875:, and suggested that even the two species might be the same. Furthermore, he found 838: 724: 704:

on the midline of the head, and noted their similarity to the nasal horn-core of a

631: 547: 482: 300: 223: 5745: 5120: 4634: 4583: 4554: 3254:

In 1987, J. Keith Rigby and colleagues suggested that pachycephalosaur domes were

1889: 1692: 5606: 5543: 5496:"Structural Mechanics of Pachycephalosaur Crania Permitted Head-butting Behavior" 5352: 5293: 4870: 3981: 3123:. The white-bellied duiker was found to be the closest morphological analogue to 2680:

to the Cretaceous. Since pachycephalosaurs were mainly small, this may be due to

6429: 6334: 6320: 6306: 6285: 6271: 6237: 6216: 5899: 3559: 3523: 3507: 3227: 3149: 3035:

CT images showing the internal density of bone in UALVP 2 (blue: low, red: high)

2988: 2889: 2343: 2039: 1621: 1526: 1324: 1318: 706: 701: 63: 31: 5950: 3929:, an orthopodous dinosaur from the Belly River Cretaceous of Alberta, Canada". 3382:(middle Campanian, 77.5 to 76.5 mya) of Alberta, Canada. The pachycephalosaurs 3127:; this head-butting species has a dome which is smaller but similarly rounded. 2995:). Sues also suggested that the animals could have butted each other's flanks. 1897: 1796: 1634:) was shortened and distanced from the regions below the orbits and around the 1590:(eye sockets). The frontoparietal narrowed at the back, was wedged between the 6257: 6223: 5060:"Evolutionary Trends in the Jaw Adductor Mechanics of Ornithischian Dinosaurs" 4976:

Evans, D. C.; Schott, R. K.; Larson, D. W.; Brown, C. M.; Ryan, M. J. (2013).

3868: 3619: 3596: 3544: 3540: 3481: 3469: 3434: 3263: 3088: 3080: 2900: 2644: 2020:

within Ornithischia, Pachycephalosauria. In 1978, the Chinese palaeontologist

1985: 1954: 1725: 1719: 1704: 1662: 1658: 1651: 1629: 1375:, with two other specimens (SMP VP-2555 and SMP VP-2790) as paratypes. A 2011 1356: 1275: 1062: 1044:

skull (left), and 1924 illustrations of same skull with labelled bones (right)

888: 834: 728: 610: 527:, and it had a thick ridge over the eyes. Much of the skull was ornamented by 506: 108: 73: 6420: 4642: 3484:. Dinosaur Park is known for its diverse community of herbivores. As well as 1749: 1347:

In 2002, Williamson and Carr described a dome (specimen NMMNH P-33983 in the

5495: 3704: 3477: 3407: 3271: 3137: 3132:

keratinous covering of the dome, and found it to aid in performance. Though

3002:

pointed out that the dorsal vertebrae from the back of the pachycephalosaur

2775: 2649: 2086: 1957: 1678: 1604: 1597: 1187:

of juvenile, flat-headed AMNH 5450 (formerly considered the distinct genus "

1184: 790: 654: 615: 528: 174: 113: 57: 17: 5625: 5562: 5371: 5212: 5085: 5011: 4889: 4724:"A taxonomic review of the Pachycephalosauridae (Dinosauria: Ornithischia)" 4529:

Fossil Record 3. New Mexico Museum of Natural History and Science, Bulletin

4503:

Fossil Record 3. New Mexico Museum of Natural History and Science, Bulletin

4367: 4000: 3884: 2835:

and other pachycephalosaurs. Brown and Schlaikjer suggested that there was

1949:

was more robust than the ulna, which is unusual. When seen from above, the

1669:(hole) was present in the suture between the premaxillae, leading into the 965:

was the first member of this family to be named, Sternberg named the group

5938: 4439:

from the Upper Cretaceous Fruitland Formation, San Juan Basin, New Mexico"

4318:

Schott, R. K.; Evans, D. C. (2016). "Cranial variation and systematics of

2004: 1992:(claw bone) of a toe was slender and slightly curved. Though the limbs of 1886:, the ossified tendons found with UALVP 2 would have formed an intricate " 1650:

were fused and formed a thick ridge above the orbit. The relatively large

6414: 6025: 6016: 4753:

Peczkis, J. (1995). "Implications of Body-Mass Estimates for Dinosaurs".

3419: 3213:

Pelvic, hind limb, pectoral, and brachial musculature reconstructions of

2904: 2017: 1922: 1907: 1739: 1416: 1103:

by Galton in 1971. The specific name honours Barnum Brown, who found the

958: 893: 463: 207: 194: 103: 98: 83: 78: 68: 5301: 3569:. Other dinosaurs known from the Oldman Formation include the hadrosaur 3410:, about 74 mya) of New Mexico, and if this species correctly belongs in 2062:(juvenile-like) or derived traits in most revisions, but not a sexually 899:

Gilmore's classification was supported by the American palaeontologists

6449: 5461: 5426: 5251: 5171: 5002: 4977: 4782: 4418: 4397: 4117:

Fossilium Catalogus I: Animalia Pars 105. Ornithischia (Supplementum I)

4092: 4061: 3876: 3461: 3453: 3120: 3076: 2879:

In 2003, Williamson and Carr published a hypothetical growth series of

2800: 2711: 1965: 1918: 1914: 1910:(four-limbed) animals, and may have functioned in stiffening the tail. 1735: 1731: 1681:

sense organ). The maxilla was short and deep, and probably contained a

1666: 795: 786: 524: 519: 478: 323: 118: 93: 5780: 5717: 5700: 4331: 3429:

The Dinosaur Park Formation is interpreted as a low-relief setting of

2093:

within Pachycephalosauridae according to Schott and colleagues, 2016:

6010: 5961: 5203: 5186: 5076: 5059: 4613:. New Mexico Museum of Natural History and Science. pp. 107–116. 3661:"New genera and species from the Belly River Series (mid-Cretaceous)" 3285: 1989: 1981: 1946: 1635: 953:

demonstrated differences between the two, and instead suggested that

646: 477:, about 77.5 to 74 million years ago (mya). The first specimens from 184: 6391: 4188: 4153: 4030: 3665:

Geological Survey of Canada, Contributions to Canadian Palaeontology

3230:

dinosaurs, but the pelvic and hindlimb musculature was instead more

5326:

Peterson, J. E.; Dischler, C.; Longrich, N. R.; Dodson, P. (2013).

5033:(2nd ed.). Berkeley: University of California Press. pp. 4663:. Jefferson, North Carolina: McFarland & Co. pp. 834–838. 1363:). In 2006, Sullivan and Spencer G. Lucas considered it a juvenile 3457: 3442: 3438: 3430: 3342: 3245: 3012: 2954: 2866: 2818: 2696: 2095: 2003: 1977: 1691: 1614:(opening) behind the eye was narrow and sloped backwards, and the 1560: 1509: 1481:

The description also included the holotype of the dubious species

1430:

interval and geographic range. Dyer and colleagues found that the

1299:, upon which most scientific understanding of the genus is based. 1179: 800: 609: 455: 3904:

typifying a new family referred provisionally to the Stegosauria"

2782:

in the brain. In 2014, Jason M. Bourke and colleagues found that

1545:

was small to medium in size compared to other pachycephalosaurs.

3473: 1942: 1665:(front bone of the upper jaw) was rugose and thickened. A small 1538: 6395: 6003: 5965: 5107:(Ornithischia: Thescelosauridae): paleoecological inferences". 2008:

Reconstructed skeleton showing ossified tail tendons in place,

1367:, which would expand the range of the species considerably. In 1295:

with certainty. UALVP 2 is still the most complete specimen of

785:'s 1924 skeletal reconstruction of UALVP 2 (left), showing the 3687:

Hatcher, J.B.; Lull, R.S.; Marsh, O.C.; Osborn, H. F. (1907).

1988:

of the foot appears to have been narrow, and the single known

1420: 744:, based on specimen CMN 1423 (which he originally included in 731:(plated dinosaurs) as contenders. Hatcher doubted whether the 3488:, the formation has also yielded fossils of the ceratopsians 2931:

The function of pachycephalosaur domes has been debated, and

1473:

since the study concluded it was very dissimilar from other

1099:, based on a flattened dome, formerly described as a female 5477:

Neues Jahrbuch für Geologie und Paläontologie - Monatshefte

5419:

10.1666/0094-8373(2004)030<0253:chopom>2.0.co;2

4294:

10.1671/0272-4634(2002)022[0779:ANGODP]2.0.CO;2

1972:

is not known, it was probably reduced in size like that of

4548: 4546: 4544: 4542: 4402:

from the Belly River Group (Campanian) of Alberta, Canada"

2642:

dispersed. The oldest known members of the group (such as

1878:. Though the neck vertebrae are not known, the downturned 1291:, including many domes too incomplete to be identified as 5873:(2nd ed.). University of California Press. pp. 5833:. In Currie, Philip J.; Koppelhus, Eva Bundgaard (eds.). 5798:. In Currie, Philip J.; Koppelhus, Eva Bundgaard (eds.). 5272:

Lambe, 1902, from the Dinosaur Park Formation, Alberta".

4728:

New Mexico Museum of Natural History and Science Bulletin

4443:

New Mexico Museum of Natural History and Science Bulletin

4260:

New Mexico Museum of Natural History and Science Bulletin

4127: 4125: 3795:

10.1671/0272-4634(2003)23[181:rotdsl]2.0.co;2

1309:

in 2016 by Ryan K. Schott Schott and David C. Evans, and

915:(both from Alberta), as well as moving the large species 4603:"The validity of the Late Cretaceous pachycephalosaurid 4596: 4594: 3826:

Centralblatt für Mineralogie, Geologie und Paläontologie

3111:

both had skull shapes similar to the bighorn sheep with

3047:

CT cross-sections of the skulls of UALVP 2 (left) and a

1945:

were expanded, and ridges extended along the shaft. The

1941:

muscles attached) was weakly developed. The ends of the

1657:

The nasal openings were large and faced frontwards. The

1332:, and found no significant distinction between them and 3370:

in the US), and specimens have been recovered from the

562:(or "primitive") compared to other pachycephalosaurs. 4520:

Watabe, M.; Tsogtbaatar, K.; Sullivan, R. M. (2011).

3931:

Department of Geology, University of Alberta Bulletin

4609:. In Sullivan, Robert M.; Lucas, Spencer G. (eds.). 2811:

and therefore not preserved through mineralization.

1866:

is incompletely known. The articulation between the

345: 332: 307: 6404: 6247: 6195: 6163: 6112: 6085: 6058: 5263: 5261: 3955:Brown, C. M.; Russell, A. P.; Farke, A. A. (2012). 3250:

Restoration of a pair with different coloured domes

1906:; the feature is unique to pachycephalosaurs among 1734:(both part of the upper jaw), and seventeen in the 828:in 1926, and catalogued as specimen UALVP 2 in the 4685: 4481: 4479: 4477: 4475: 4396:Dyer, Aaron; Powers, Mark; Currie, Philip (2023). 4275: 4273: 3777:Sullivan, R. M. (2003). "Revision of the dinosaur 3682: 3680: 3678: 3366:Belly River Group (the Canadian equivalent to the 3262:, based on their internal "radiating structures" ( 1477:specimens and therefore probably not referable to 1209:(until then only known from the Mongolian species 3693:Monographs of the United States Geological Survey 1581:were very thick and formed an elevated dome. The 2871:CT images of TMP 84.5.1, a juvenile or subadult 1779:appears to have had a smaller frontal boss than 1349:New Mexico Museum of Natural History and Science 1336:. They considered both as junior synonyms, with 5898:Eberth, D. A. (1997). "Judith River Wedge". In 4717: 4715: 4713: 4202: 4200: 4198: 1783:, and seems to have been more gracile overall. 1379:by Watabe and colleagues did not place the two 837:(abdominal ribs), which are not known in other 793:, and comparison between its teeth and that of 4391: 4389: 4387: 4385: 1491:where it was also found to be very similar to 1061:based on a parietal bone from China. In 1964, 5977: 5829:Braman, Dennis R.; Koppelhus, Eva B. (2005). 5321: 5319: 4971: 4969: 4313: 4311: 3781:Lambe (Ornithischia, Pachycephalosauridae)". 3736: 3734: 3595:and possibly ankylosaurs. Theropods included 3456:was warmer than present-day Alberta, without 1065:considered this as an unequivocal species of 710:specimen. In 1903, Hungarian palaeontologist 8: 5385: 5383: 5381: 3950: 3948: 3946: 3944: 3732: 3730: 3728: 3726: 3724: 3722: 3720: 3718: 3716: 3714: 3612:and possibly an albertosaurine tyrannosaur. 3378:(including the lectotype specimen), and the 1499:was probably an adult specimen of the genus 1340:representing the end-stage in the growth of 4961:The Age of Dinosaurs in Russia and Mongolia 4794: 4792: 4553:Williamson, T. E.; Brusatte, S. L. (2016). 4406:Vertebrate Anatomy Morphology Palaeontology 3908:Transactions of the Royal Society of Canada 1984:was slender and wide at the upper end. The 989:, based on a specimen formerly referred to 925:, along with two other species. They found 919:(which was named in 1931) to the new genus 6392: 6201: 6169: 6160: 6091: 6064: 6055: 6000: 5984: 5970: 5962: 5666:Knell, R. J.; Sampson, S. (January 2011). 4012: 4010: 3772: 3770: 3768: 3766: 3764: 3762: 3760: 845:were very similar to those of the species 481:, Canada, were described in 1902, and the 136: 38: 5716: 5683: 5615: 5605: 5552: 5542: 5361: 5351: 5202: 5103:(Ornithischia: Pachycephalosauridae) and 5075: 5001: 4963:. Cambridge University Press, Cambridge. 4879: 4869: 4835: 4833: 4831: 4829: 4601:Jasinski, S. E.; Sullivan, R. M. (2016). 4582: 4486:Jasinski, S. E.; Sullivan, R. M. (2011). 4417: 4324:Zoological Journal of the Linnean Society 3990: 3980: 3654: 3652: 3402:(late Campanian, about 75 mya) and lower 2864:could easily be the results of ontogeny. 2100:Partial skull of a juvenile, flat-headed 855:was found. Therefore, Gilmore considered 680:/κέρας meaning "horn". The specific name 30:For the stegosaur of a similar name, see 4692:. New York: Dorling Kindersley. p. 3741:Sues, H. D. & Galton, P. M. (1987). 2973:The hypothesis that the domed skulls of 2827:skulls, according to Schott et al., 2011 1438:, because the proposed unique traits of 714:suggested that the fragmentary domes of 538:Originally known only from skull domes, 6542:Multispecific non-avian dinosaur genera 4905: 4903: 4901: 4899: 4808:. Princeton University Press. pp. 4075:Russell, L. S. (1948). "The dentary of 3648: 2648:) are "fully domed" and known from the 1073:. In 1974, the Polish palaeontologists 378: 5910:. San Diego: Academic Press. pp. 5390:Goodwin, M. B.; Horner, J. R. (2004). 4910:Carpenter, Kenneth (1 December 1997). 4805:The Princeton Field Guide to Dinosaurs 4433:Sullivan, R. M.; Lucas, S. G. (2006). 2998:In 1997, the American palaeontologist 1594:, and ended in a depression above the 1533:specimen UALVP 2 is the most complete 816:In 1924, the American palaeontologist 505:, named in 2011 from fossils found in 5839:. Indiana University Press. pp. 5804:. Indiana University Press. pp. 4252:(Brown and Schlaikjer) new comb. and 3638:Timeline of pachycephalosaur research 3460:, but with wetter and drier seasons. 3394:are also known from both formations. 2770:In 1989, Emily B. Griffin found that 2762:themselves were not contemporaries). 1917:(shoulder blade) was longer than the 1195:In 2000, Robert M. Sullivan referred 149:skeletons based on specimen UALVP 2, 27:Genus of pachycephalosaurid dinosaurs 7: 5517:Snively, E.; Theodor, J. M. (2011). 3374:(late Campanian, 76.5 to 75 mya) in 2847:. This idea was supported by a 1981 2038:. In 1986, American palaeontologist 1399:, or if it was a juvenile of either 1091:. In 1979, William Patrick Wall and 949:teeth, the American palaeontologist 622:, shown from the right and underside 4607:(Dinosauria: Pachycephalosauridae)" 3291:In 2011, American palaeontologists 2507: 2448: 2407: 2400: 2359: 2335: 2311: 2287: 2263: 2239: 2152: 2145: 2118: 2108: 6512:Pachycephalosaurs of North America 5734:Journal of Vertebrate Paleontology 5699:Hone, D. W. E.; Naish, D. (2013). 5274:Journal of Vertebrate Paleontology 5144:Journal of Vertebrate Paleontology 4755:Journal of Vertebrate Paleontology 4348:Canadian Journal of Earth Sciences 4282:Journal of Vertebrate Paleontology 4213:Journal of Vertebrate Paleontology 4169:Canadian Journal of Earth Sciences 4134:Canadian Journal of Earth Sciences 4079:, a genus of theropod dinosaurs". 3783:Journal of Vertebrate Paleontology 2843:) represented the female morph of 2010:National Museum of Natural History 1902:, and are otherwise only known in 1774:can be distinguished from that of 1549:appears to have been smaller than 1111:) in Alberta. In 1983, Galton and 1109:American Museum of Natural History 1081:concluded that the "gastralia" of 25: 4435:"The pachycephalosaurid dinosaur 2907:) from flat to domed, supporting 1968:was long and slender. Though the 1213:), and found it more likely that 688:), CMN 515 was designated as the 6376: 6375: 6352: 5949: 5937: 5685:10.1111/j.1469-7998.2010.00758.x 5653:10.1111/j.1469-7998.2010.00719.x 4933:Perle, A.; Osmólska, H. (1982). 3622: 3554:. Theropods present include the 3326:and associated ornamentation of 3203: 3194: 3157: 3148: 3099:and several extant head-butting 3040: 3028: 2786:would have needed cartilaginous 2032:and the "primitive" ceratopsian 1842: 1833: 1824: 1813: 1804: 1795: 1673:, and possibly connected to the 1442:disappeared through ontogeny in 1026: 1017: 1008: 985:valid, and named a new species, 772: 763: 161: 2416:Pachycephalosaurus wyomingensis 1457:was described, and assigned to 5164:10.1080/02724634.1989.10011739 4775:10.1080/02724634.1995.10011575 4462:New Mexico: New Mexico Geology 4233:10.1080/02724634.1998.10011064 2799:) or a branched one (as in an 1434:holotype could be an immature 1191:"), with sections on the right 1172:, therefore a junior synonym. 820:described a complete skull of 712:Franz Nopcsa von Felső-Szilvás 1: 6517:Fossil taxa described in 1902 5746:10.1080/02724634.2023.2190369 5578:"The appendicular myology of 5494:Snively, E.; Cox, A. (2008). 5121:10.1016/j.cretres.2021.105058 4943:Acta Palaeontologica Polonica 4635:10.1016/j.geobios.2024.08.006 4584:10.1016/j.cretres.2016.01.012 4119:, IJsel Pers, Deventer, 80 pp 3747:Palaeontographica Abteilung A 3464:were apparently the dominant 2701:Mandible and teeth of UALVP 2 2089:below shows the placement of 1383:species close to each other. 803:which Gilmore argued was the 355:Jasinski & Sullivan, 2011 6522:Taxa named by Lawrence Lambe 5607:10.1371/journal.pone.0268144 5544:10.1371/journal.pone.0021422 5353:10.1371/journal.pone.0068620 5294:10.1080/02724634.2012.679878 4871:10.1371/journal.pone.0021092 4611:Fossil Record 5: Bulletin 74 4246:Sullivan, Robert M. (2000). 3982:10.1371/journal.pone.0030212 3445:conditions over time as the 3422:(associated with water) and 1487:in its morphometric analysis 1040:UALVP 2, the first complete 590:, whereas the controversial 426:Brown & Schlaikjer, 1943 4661:Dinosaurs: The Encyclopedia 3925:Gilmore, C. W. (1924). "On 2660:(77 million years ago) and 2272:Homalocephale calathocercos 1303:was moved to the new genus 1279:. He stated that the genus 1127:is Latin for "adorned" and 1107:(specimen AMNH 5450 in the 740:, and named a new species, 676:/στέγη, meaning "roof" and 630:were collected by Canadian 626:The first known remains of 558:itself has been considered 6558: 5500:Palaeontologia Electronica 4688:The Ultimate Dinosaur Book 4250:Prenocephale edmontonensis 3351:in Alberta, Canada, where 2943:structure, or a means for 1390:restudied the holotype of 466:that lived in what is now 29: 6365: 6350: 6204: 6172: 6159: 6140: 6094: 6067: 6054: 5999: 5906:Encyclopedia of Dinosaurs 5794:Eberth, David A. (2005). 3869:10.1126/science.18.445.60 3816:Nopcsa, F. (1903). "Über 3330:and the ornamentation of 3242:Other suggested functions 2533:Sphaerotholus buchholtzae 2529: 2512: 2505: 2470: 2453: 2446: 2429: 2412: 2405: 2398: 2381: 2364: 2357: 2340: 2333: 2316: 2309: 2292: 2285: 2268: 2261: 2244: 2237: 2208: 2191: 2174: 2157: 2150: 2143: 2123: 2116: 1415:(difference in timing of 1176:21st century developments 651:Canadian Museum of Nature 570:(head or flank-butting), 369: 362: 329: 322: 306: 299: 158:Scientific classification 156: 144: 135: 41: 5902:; Padian, Kevin (eds.). 5836:Dinosaur Provincial Park 5831:"Campanian palynomorphs" 5801:Dinosaur Provincial Park 5105:Thescelosaurus neglectus 5058:Nabavizadeh, A. (2016). 4722:Sullivan, R. M. (2006). 4605:Stegoceras novomexicanum 4557:Stegoceras novomexicanum 3845:"Recent Zoopaleontology" 3376:Dinosaur Provincial Park 2212:Stegoceras novomexicanum 2127:Wannanosaurus yansiensis 1386:In 2016, Williamson and 1373:Stegoceras novomexicanum 1255:was a junior synonym of 1247:. In a 2003 revision of 1243:, or perhaps a juvenile 6527:Paleontology in Alberta 5450:Journal of Paleontology 5240:Journal of Paleontology 4081:Journal of Paleontology 4050:Journal of Paleontology 3705:10.5962/bhl.title.60500 3447:Western Interior Seaway 3372:Dinosaur Park Formation 3362:is known from the late 3349:Dinosaur Park Formation 3073:finite element analysis 2433:Alaskacephale gangloffi 1449:In 2024, a specimen of 1251:, Sullivan agreed that 727:(horned dinosaurs) and 723:belonged to, with both 584:Dinosaur Park Formation 417:Wall & Galton, 1979 408:Galton & Sues, 1983 400:Wall & Galton, 1979 4937:Goyocephale lattimorei 4916:Rocky Mountain Geology 4494:Texascephale langstoni 4492:and a reassessment of 4368:10.1139/cjes-2020-0190 4320:Foraminacephale brevis 4048:, n. sp., Described". 3900:"The Cretaceous genus 3356: 3251: 2970: 2876: 2828: 2702: 2516:Sphaerotholus goodwini 2320:Foraminacephale brevis 2248:Goyocephale lattimorei 2105: 2012: 1964:was elongated and the 1712: 1616:supratemporal fenestra 1612:infratemporal fenestra 1566: 1519: 1436:Sphaerotholus goodwini 1405:Sphaerotholus goodwini 1361:Sphaerotholus goodwini 1225:, but considered it a 1192: 1095:named the new species 905:Erich Maren Schlaikjer 623: 614:Partial dome CMN 515, 6484:Paleobiology Database 5064:The Anatomical Record 4982:Nature Communications 4842:"Cranial Ontogeny in 3898:Lambe, L. M. (1918). 3659:Lambe, L. M. (1902). 3386:(if not a synonym of 3346: 3249: 2958: 2937:intra-specific combat 2870: 2822: 2700: 2344:Amtocephale gobienses 2178:Colepiocephale lambei 2161:Hanssuesia sternbergi 2099: 2079:phylogenetic analysis 2007: 1953:was very broad for a 1722:(differentiated) and 1707:based on other small 1695: 1564: 1513: 1484:Texacephale langstoni 1377:phylogenetic analysis 1183: 830:University of Alberta 645:district of Alberta, 613: 568:intra-specific combat 431:Texacephale langstoni 422:Hanssuesia sternbergi 6537:Ornithischian genera 6249:Pachycephalosaurinae 6197:Pachycephalosauridae 5946:at Wikimedia Commons 4684:Lambert, D. (1993). 4400:Gravitholus albertae 4209:Stygimoloch spinifer 4019:Bulletin of the AMNH 3843:Lambe, L.M. (1903). 3355:was first discovered 3117:white-bellied duiker 3049:white-bellied duiker 2670:convergent evolution 2474:Stygimoloch spinifer 2296:Tylocephale gilmorei 2234:Pachycephalosaurinae 2140:Pachycephalosauridae 1787:Postcranial skeleton 1718:had teeth that were 967:Pachycephalosauridae 951:Charles M. Sternberg 606:History of discovery 413:Gravitholus albertae 269:Pachycephalosauridae 151:Royal Tyrrell Museum 5773:2014Letha..47..567M 5598:2022PLoSO..1768144M 5535:2011PLoSO...621422S 5411:2004Pbio...30..253G 5344:2013PLoSO...868620P 5286:2012JVPal..32..903S 5156:1989JVPal...9...67G 5109:Cretaceous Research 4994:2013NatCo...4.1828E 4862:2011PLoSO...621092S 4767:1995JVPal..14..520P 4575:2016CrRes..62...29W 4563:Cretaceous Research 4360:2021CaJES..58..981E 4225:1998JVPal..18..363G 4181:1983CaJES..20..462G 4146:1979CaJES..16.1176W 3973:2012PLoSO...730212B 3861:1903Sci....18...60L 3575:, the ceratopsians 3175:skull model (right) 3097:Prenocephale prenes 2945:species recognition 2815:Ontogenetic changes 2666:Early Maastrichtian 2457:Dracorex hogwartsia 2368:Prenocephale prenes 2104:, UCMZ(VP) 2008.001 1929:deltopectoral crest 1557:Skull and dentition 1518:compared to a human 1388:Stephen L. Brusatte 937:the female form of 879:to be identical to 867:, thereby renaming 826:George F. Sternberg 665:. The generic name 576:species recognition 405:Ornatotholus browni 6300:Pachycephalosaurus 6165:Pachycephalosauria 6144:Pachycephalosauria 6114:Pachycephalosauria 5993:Pachycephalosauria 5705:Journal of Zoology 5672:Journal of Zoology 5641:Journal of Zoology 5580:Stegoceras validum 5270:Stegoceras validum 5101:Stegoceras validum 5003:10.1038/ncomms2749 4844:Stegoceras validum 4437:Stegoceras validum 4419:10.18435/vamp29388 3441:and influenced by 3404:Kirtland Formation 3398:is known from the 3368:Judith River Group 3357: 3332:Pachycephalosaurus 3252: 3236:positive allometry 2971: 2877: 2829: 2778:that showed large 2703: 2195:Stegoceras validum 2113:Pachycephalosauria 2106: 2070:itself to be more 2056:Pachycephalosaurus 2013: 1713: 1567: 1520: 1334:Stegoceras validum 1193: 1119:to its own genus, 922:Pachycephalosaurus 818:Charles W. Gilmore 690:lectotype specimen 659:Stegoceras validus 657:genus and species 624: 600:Kirtland Formation 594:is known from the 582:is known from the 487:Stegoceras validum 460:pachycephalosaurid 381:Stegoceras validus 312:Stegoceras validum 256:Pachycephalosauria 145:Two reconstructed 6499: 6498: 6471:Open Tree of Life 6398:Taxon identifiers 6389: 6388: 6361: 6360: 6348: 6347: 6344: 6343: 6155: 6154: 6151: 6150: 6136: 6135: 6132: 6131: 5942:Media related to 5921:978-0-12-226810-6 5900:Currie, Philip J. 5884:978-0-520-24209-8 5850:978-0-253-34595-0 5815:978-0-253-34595-0 5781:10.1111/let.12082 5718:10.1111/jzo.12035 5197:(11): 2148–2186. 5191:Anatomical Record 5044:978-0-520-24209-8 4819:978-0-691-13720-9 4703:978-1-56458-304-8 4670:978-0-89950-917-4 4332:10.1111/zoj.12465 4046:Stegoceras lambei 3609:Saurornitholestes 3572:Brachylophosaurus 3437:that became more 3339:Palaeoenvironment 3000:Kenneth Carpenter 2837:sexual dimorphism 2823:Growth series of 2717:replacement teeth 2693:Feeding mechanics 2634: 2633: 2625: 2624: 2616: 2615: 2607: 2606: 2598: 2597: 2589: 2588: 2580: 2579: 2571: 2570: 2562: 2561: 2553: 2552: 2544: 2543: 2494: 2493: 2485: 2484: 2385:Acrotholus audeti 2223: 2222: 1880:occipital condyle 1471:S. novomexicanum, 1285:wastebasket taxon 1263:to the new genus 1105:holotype specimen 1097:Stegoceras browni 993:. The split from 933:, but considered 694:John Bell Hatcher 639:Belly River Group 495:. Currently only 447: 446: 440: 427: 418: 409: 401: 397:Stegoceras browni 393: 385: 375: 356: 342: 295: 16:(Redirected from 6549: 6532:Campanian genera 6492: 6491: 6479: 6478: 6466: 6465: 6453: 6452: 6440: 6439: 6438: 6425: 6424: 6423: 6393: 6379: 6378: 6356: 6202: 6170: 6161: 6092: 6065: 6056: 6049: 6048: 6001: 5986: 5979: 5972: 5963: 5954:Data related to 5953: 5941: 5926: 5925: 5909: 5895: 5889: 5888: 5872: 5861: 5855: 5854: 5826: 5820: 5819: 5791: 5785: 5784: 5756: 5750: 5749: 5729: 5723: 5722: 5720: 5696: 5690: 5689: 5687: 5663: 5657: 5656: 5636: 5630: 5629: 5619: 5609: 5573: 5567: 5566: 5556: 5546: 5514: 5508: 5507: 5491: 5485: 5484: 5472: 5466: 5465: 5445: 5439: 5438: 5396: 5387: 5376: 5375: 5365: 5355: 5323: 5314: 5313: 5265: 5256: 5255: 5231: 5225: 5224: 5206: 5204:10.1002/ar.23046 5182: 5176: 5175: 5139: 5133: 5132: 5096: 5090: 5089: 5079: 5077:10.1002/ar.23306 5055: 5049: 5048: 5032: 5022: 5016: 5015: 5005: 4973: 4964: 4957: 4951: 4950: 4930: 4924: 4923: 4907: 4894: 4893: 4883: 4873: 4837: 4824: 4823: 4796: 4787: 4786: 4750: 4744: 4743: 4719: 4708: 4707: 4691: 4681: 4675: 4674: 4653: 4647: 4646: 4621: 4615: 4614: 4598: 4589: 4588: 4586: 4550: 4537: 4536: 4526: 4517: 4511: 4510: 4500: 4483: 4470: 4469: 4457: 4451: 4450: 4430: 4424: 4423: 4421: 4393: 4380: 4379: 4342: 4336: 4335: 4315: 4306: 4305: 4277: 4268: 4267: 4243: 4237: 4236: 4204: 4193: 4192: 4164: 4158: 4157: 4140:(6): 1176–1186. 4129: 4120: 4115:Kuhn, O., 1964, 4113: 4107: 4103: 4097: 4096: 4072: 4066: 4065: 4041: 4035: 4034: 4014: 4005: 4004: 3994: 3984: 3952: 3939: 3938: 3922: 3916: 3915: 3895: 3889: 3888: 3840: 3834: 3833: 3813: 3807: 3806: 3774: 3755: 3754: 3738: 3709: 3708: 3689:"The Ceratopsia" 3684: 3673: 3672: 3656: 3632: 3630:Dinosaurs portal 3627: 3626: 3625: 3468:plants, with an 3396:S. novomexicanum 3380:Oldman Formation 3306:sexual selection 3302:Scott D. Sampson 3281:Sharpey's fibers 3260:thermoregulation 3258:organs used for 3207: 3198: 3161: 3152: 3044: 3032: 2903:(in contrast to 2788:nasal turbinates 2658:Middle Campanian 2508: 2449: 2408: 2401: 2360: 2336: 2312: 2288: 2264: 2240: 2153: 2146: 2119: 2109: 1930: 1899: 1891: 1875: 1874:ossified tendons 1860:vertebral column 1846: 1837: 1828: 1817: 1808: 1799: 1772:S. novomexicanum 1751: 1727: 1697:Life restoration 1675:Jacobson's organ 1631: 1623: 1599: 1547:S. novomexicanum 1440:S. novomexicanum 1432:S. novomexicanum 1424:S. novomexicanum 1409:S. novomexicanum 1397:S. novomexicanum 1392:S. novomexicanum 1197:S. edmontonensis 1113:Hans-Dieter Sues 1079:Halszka Osmólska 1075:Teresa Maryańska 1042:pachycephalosaur 1030: 1021: 1012: 975:T. edmontonensis 913:T. edmontonensis 848:Troodon formosus 776: 767: 592:S. novomexicanum 588:Oldman Formation 533:vertebral column 502:S. novomexicanum 434: 425: 416: 407: 399: 391: 383: 373: 354: 350:S. novomexicanum 347: 340: 334: 309: 287: 280: 267: 254: 238: 222: 166: 165: 140: 130: 60: 47:Temporal range: 39: 21: 6557: 6556: 6552: 6551: 6550: 6548: 6547: 6546: 6502: 6501: 6500: 6495: 6487: 6482: 6474: 6469: 6461: 6456: 6448: 6443: 6434: 6433: 6428: 6419: 6418: 6413: 6400: 6390: 6385: 6357: 6340: 6279:Foraminacephale 6243: 6191: 6147: 6128: 6108: 6081: 6075:Avemetatarsalia 6060:Avemetatarsalia 6050: 6044:Marginocephalia 6006: 6005: 5995: 5990: 5934: 5929: 5922: 5897: 5896: 5892: 5885: 5863: 5862: 5858: 5851: 5828: 5827: 5823: 5816: 5793: 5792: 5788: 5758: 5757: 5753: 5731: 5730: 5726: 5698: 5697: 5693: 5665: 5664: 5660: 5638: 5637: 5633: 5592:(9): e0268144. 5575: 5574: 5570: 5516: 5515: 5511: 5493: 5492: 5488: 5474: 5473: 5469: 5447: 5446: 5442: 5394: 5389: 5388: 5379: 5325: 5324: 5317: 5267: 5266: 5259: 5233: 5232: 5228: 5184: 5183: 5179: 5141: 5140: 5136: 5098: 5097: 5093: 5057: 5056: 5052: 5045: 5024: 5023: 5019: 4975: 4974: 4967: 4958: 4954: 4932: 4931: 4927: 4909: 4908: 4897: 4839: 4838: 4827: 4820: 4798: 4797: 4790: 4752: 4751: 4747: 4721: 4720: 4711: 4704: 4683: 4682: 4678: 4671: 4655: 4654: 4650: 4623: 4622: 4618: 4600: 4599: 4592: 4552: 4551: 4540: 4524: 4519: 4518: 4514: 4498: 4485: 4484: 4473: 4459: 4458: 4454: 4432: 4431: 4427: 4395: 4394: 4383: 4354:(10): 981–992. 4344: 4343: 4339: 4317: 4316: 4309: 4279: 4278: 4271: 4245: 4244: 4240: 4206: 4205: 4196: 4189:10.1139/e83-043 4166: 4165: 4161: 4154:10.1139/e79-104 4131: 4130: 4123: 4114: 4110: 4104: 4100: 4074: 4073: 4069: 4043: 4042: 4038: 4016: 4015: 4008: 3954: 3953: 3942: 3927:Troodon validus 3924: 3923: 3919: 3897: 3896: 3892: 3842: 3841: 3837: 3815: 3814: 3810: 3776: 3775: 3758: 3740: 3739: 3712: 3686: 3685: 3676: 3658: 3657: 3650: 3646: 3628: 3623: 3621: 3618: 3601:oviraptorosaurs 3584:Albertaceratops 3536:Parasaurolophus 3392:Foraminacephale 3364:Late Cretaceous 3341: 3244: 3220: 3219: 3218: 3217: 3210: 3209: 3208: 3200: 3199: 3179: 3178: 3177: 3176: 3164: 3163: 3162: 3154: 3153: 3113:cancellous bone 3056: 3055: 3054: 3053: 3052: 3045: 3037: 3036: 3033: 2953: 2929: 2875:, with sections 2817: 2780:olfactory bulbs 2768: 2752:ecomorphospaces 2695: 2690: 2682:taphonomic bias 2635: 2626: 2617: 2608: 2599: 2590: 2581: 2572: 2563: 2554: 2545: 2495: 2486: 2224: 2044:Marginocephalia 2002: 1932: 1901: 1893: 1877: 1856: 1855: 1854: 1853: 1849: 1848: 1847: 1839: 1838: 1830: 1829: 1820: 1819: 1818: 1810: 1809: 1801: 1800: 1789: 1753: 1729: 1687:predentary bone 1648:palpebral bones 1633: 1625: 1601: 1592:squamosal bones 1559: 1508: 1455:Aguja Formation 1306:Foraminacephale 1178: 1048: 1047: 1046: 1045: 1033: 1032: 1031: 1023: 1022: 1014: 1013: 917:T. wyomingensis 814: 813: 812: 811: 789:of the tail as 779: 778: 777: 769: 768: 669:comes from the 632:palaeontologist 608: 552:Marginocephalia 493:junior synonyms 472:Late Cretaceous 443: 389:Troodon validus 377: 376: 318: 315: 286: 278: 265: 252: 236: 220: 160: 131: 129: 128: 127: 126: 121: 116: 111: 106: 101: 96: 91: 86: 81: 76: 71: 66: 55: 54: 52: 49:Late Cretaceous 45: 35: 28: 23: 22: 15: 12: 11: 5: 6555: 6553: 6545: 6544: 6539: 6534: 6529: 6524: 6519: 6514: 6504: 6503: 6497: 6496: 6494: 6493: 6480: 6467: 6454: 6441: 6426: 6410: 6408: 6402: 6401: 6396: 6387: 6386: 6384: 6383: 6373: 6366: 6363: 6362: 6359: 6358: 6351: 6349: 6346: 6345: 6342: 6341: 6339: 6338: 6331: 6324: 6317: 6310: 6303: 6296: 6289: 6282: 6275: 6268: 6261: 6253: 6251: 6245: 6244: 6242: 6241: 6234: 6227: 6220: 6213: 6210:Colepiocephale 6205: 6199: 6193: 6192: 6190: 6189: 6182: 6178:Ferganocephale 6173: 6167: 6157: 6156: 6153: 6152: 6149: 6148: 6141: 6138: 6137: 6134: 6133: 6130: 6129: 6127: 6126: 6125: 6124: 6118: 6116: 6110: 6109: 6107: 6106: 6105: 6104: 6095: 6089: 6083: 6082: 6080: 6079: 6078: 6077: 6068: 6062: 6052: 6051: 6047: 6046: 6037: 6028: 6019: 6013: 6004: 5997: 5996: 5991: 5989: 5988: 5981: 5974: 5966: 5960: 5959: 5958:at Wikispecies 5947: 5933: 5932:External links 5930: 5928: 5927: 5920: 5890: 5883: 5869:The Dinosauria 5856: 5849: 5821: 5814: 5786: 5767:(4): 567–578. 5751: 5724: 5711:(3): 172–180. 5691: 5658: 5631: 5568: 5509: 5486: 5467: 5440: 5405:(2): 253–267. 5377: 5315: 5280:(4): 903–913. 5257: 5246:(3): 608–618. 5226: 5177: 5134: 5091: 5070:(3): 271–294. 5050: 5043: 5029:The Dinosauria 5017: 4965: 4952: 4925: 4895: 4825: 4818: 4788: 4761:(4): 520–533. 4745: 4709: 4702: 4676: 4669: 4648: 4616: 4590: 4538: 4512: 4471: 4452: 4425: 4381: 4337: 4307: 4288:(4): 779–801. 4269: 4238: 4219:(2): 363–375. 4194: 4175:(3): 462–472. 4159: 4121: 4108: 4098: 4087:(5): 625–629. 4067: 4056:(5): 534–538. 4036: 4006: 3940: 3917: 3890: 3835: 3822:Stereocephalus 3808: 3789:(1): 181–207. 3756: 3710: 3674: 3647: 3645: 3642: 3641: 3640: 3634: 3633: 3617: 3614: 3593:therizinosaurs 3566:Daspletosaurus 3556:tyrannosaurids 3551:Euoplocephalus 3512:Prosaurolophus 3452:westward. The 3426:environments. 3340: 3337: 3297:John R. Horner 3243: 3240: 3212: 3211: 3202: 3201: 3193: 3192: 3191: 3190: 3189: 3166: 3165: 3156: 3155: 3147: 3146: 3145: 3144: 3143: 3066:threat display 3046: 3039: 3038: 3034: 3027: 3026: 3025: 3024: 3023: 2952: 2949: 2941:sexual display 2928: 2925: 2911:as a juvenile 2816: 2813: 2767: 2766:Nasal passages 2764: 2760:Thescelosaurus 2747:Thescelosaurus 2738:Thescelosaurus 2694: 2691: 2689: 2686: 2632: 2631: 2628: 2627: 2623: 2622: 2619: 2618: 2614: 2613: 2610: 2609: 2605: 2604: 2601: 2600: 2596: 2595: 2592: 2591: 2587: 2586: 2583: 2582: 2578: 2577: 2574: 2573: 2569: 2568: 2565: 2564: 2560: 2559: 2556: 2555: 2551: 2550: 2547: 2546: 2542: 2541: 2538: 2537: 2528: 2525: 2524: 2521: 2520: 2511: 2506: 2504: 2501: 2500: 2497: 2496: 2492: 2491: 2488: 2487: 2483: 2482: 2479: 2478: 2469: 2466: 2465: 2462: 2461: 2452: 2447: 2445: 2442: 2441: 2438: 2437: 2428: 2425: 2424: 2421: 2420: 2411: 2406: 2404: 2399: 2397: 2394: 2393: 2390: 2389: 2380: 2377: 2376: 2373: 2372: 2363: 2358: 2356: 2353: 2352: 2349: 2348: 2339: 2334: 2332: 2329: 2328: 2325: 2324: 2315: 2310: 2308: 2305: 2304: 2301: 2300: 2291: 2286: 2284: 2281: 2280: 2277: 2276: 2267: 2262: 2260: 2257: 2256: 2253: 2252: 2243: 2238: 2236: 2230: 2229: 2226: 2225: 2221: 2220: 2217: 2216: 2207: 2204: 2203: 2200: 2199: 2190: 2187: 2186: 2183: 2182: 2173: 2170: 2169: 2166: 2165: 2156: 2151: 2149: 2144: 2142: 2136: 2135: 2132: 2131: 2122: 2117: 2115: 2107: 2001: 2000:Classification 1998: 1926: 1895: 1887: 1871: 1851: 1850: 1841: 1840: 1832: 1831: 1823: 1822: 1821: 1812: 1811: 1803: 1802: 1794: 1793: 1792: 1791: 1790: 1788: 1785: 1747: 1723: 1627: 1626:(floor of the 1619: 1595: 1579:parietal bones 1558: 1555: 1507: 1504: 1353:San Juan Basin 1266:Colepiocephale 1177: 1174: 1160:differed from 1035: 1034: 1025: 1024: 1016: 1015: 1007: 1006: 1005: 1004: 1003: 929:distinct from 861:junior synonym 805:senior synonym 781: 780: 771: 770: 762: 761: 760: 759: 758: 754:Ankylosauridae 686:syntype series 643:Red Deer River 635:Lawrence Lambe 607: 604: 572:sexual display 462:(dome-headed) 445: 444: 442: 441: 428: 419: 410: 402: 394: 386: 372: 371: 370: 367: 366: 360: 359: 358: 357: 343: 327: 326: 320: 319: 316: 304: 303: 297: 296: 276: 272: 271: 263: 259: 258: 250: 243: 242: 240:Neornithischia 234: 227: 226: 218: 211: 210: 205: 198: 197: 192: 188: 187: 182: 178: 177: 172: 168: 167: 154: 153: 142: 141: 133: 132: 124: 123: 122: 117: 112: 107: 102: 97: 92: 87: 82: 77: 72: 67: 62: 61: 46: 26: 24: 14: 13: 10: 9: 6: 4: 3: 2: 6554: 6543: 6540: 6538: 6535: 6533: 6530: 6528: 6525: 6523: 6520: 6518: 6515: 6513: 6510: 6509: 6507: 6490: 6485: 6481: 6477: 6472: 6468: 6464: 6459: 6455: 6451: 6446: 6442: 6437: 6431: 6427: 6422: 6416: 6412: 6411: 6409: 6407: 6403: 6399: 6394: 6382: 6374: 6372: 6368: 6367: 6364: 6355: 6337: 6336: 6332: 6330: 6329: 6328:Sphaerotholus 6325: 6323: 6322: 6318: 6316: 6315: 6311: 6309: 6308: 6304: 6302: 6301: 6297: 6295: 6294: 6293:Homalocephale 6290: 6288: 6287: 6283: 6281: 6280: 6276: 6274: 6273: 6269: 6267: 6266: 6265:Alaskacephale 6262: 6260: 6259: 6255: 6254: 6252: 6250: 6246: 6240: 6239: 6235: 6233: 6232: 6228: 6226: 6225: 6221: 6219: 6218: 6214: 6212: 6211: 6207: 6206: 6203: 6200: 6198: 6194: 6188: 6187: 6186:Wannanosaurus 6183: 6180: 6179: 6175: 6174: 6171: 6168: 6166: 6162: 6158: 6146: 6145: 6139: 6122: 6121: 6120: 6119: 6117: 6115: 6111: 6103: 6099: 6098: 6097: 6096: 6093: 6090: 6088: 6084: 6076: 6072: 6071: 6070: 6069: 6066: 6063: 6061: 6057: 6053: 6045: 6041: 6038: 6036: 6032: 6029: 6027: 6023: 6020: 6018: 6014: 6012: 6008: 6007: 6002: 5998: 5994: 5987: 5982: 5980: 5975: 5973: 5968: 5967: 5964: 5957: 5952: 5948: 5945: 5940: 5936: 5935: 5931: 5923: 5917: 5913: 5908: 5907: 5901: 5894: 5891: 5886: 5880: 5876: 5871: 5870: 5860: 5857: 5852: 5846: 5842: 5838: 5837: 5832: 5825: 5822: 5817: 5811: 5807: 5803: 5802: 5797: 5796:"The Geology" 5790: 5787: 5782: 5778: 5774: 5770: 5766: 5762: 5755: 5752: 5747: 5743: 5739: 5735: 5728: 5725: 5719: 5714: 5710: 5706: 5702: 5695: 5692: 5686: 5681: 5677: 5673: 5669: 5662: 5659: 5654: 5650: 5646: 5642: 5635: 5632: 5627: 5623: 5618: 5613: 5608: 5603: 5599: 5595: 5591: 5587: 5583: 5581: 5572: 5569: 5564: 5560: 5555: 5550: 5545: 5540: 5536: 5532: 5529:(6): e21422. 5528: 5524: 5520: 5513: 5510: 5505: 5501: 5497: 5490: 5487: 5482: 5478: 5471: 5468: 5463: 5459: 5455: 5451: 5444: 5441: 5436: 5432: 5428: 5424: 5420: 5416: 5412: 5408: 5404: 5400: 5393: 5386: 5384: 5382: 5378: 5373: 5369: 5364: 5359: 5354: 5349: 5345: 5341: 5338:(7): e68620. 5337: 5333: 5329: 5322: 5320: 5316: 5311: 5307: 5303: 5299: 5295: 5291: 5287: 5283: 5279: 5275: 5271: 5264: 5262: 5258: 5253: 5249: 5245: 5241: 5237: 5230: 5227: 5222: 5218: 5214: 5210: 5205: 5200: 5196: 5192: 5188: 5181: 5178: 5173: 5169: 5165: 5161: 5157: 5153: 5149: 5145: 5138: 5135: 5130: 5126: 5122: 5118: 5114: 5110: 5106: 5102: 5095: 5092: 5087: 5083: 5078: 5073: 5069: 5065: 5061: 5054: 5051: 5046: 5040: 5036: 5031: 5030: 5021: 5018: 5013: 5009: 5004: 4999: 4995: 4991: 4987: 4983: 4979: 4972: 4970: 4966: 4962: 4956: 4953: 4948: 4944: 4940: 4938: 4929: 4926: 4921: 4917: 4913: 4906: 4904: 4902: 4900: 4896: 4891: 4887: 4882: 4877: 4872: 4867: 4863: 4859: 4856:(6): e21092. 4855: 4851: 4847: 4845: 4836: 4834: 4832: 4830: 4826: 4821: 4815: 4811: 4807: 4806: 4801: 4795: 4793: 4789: 4784: 4780: 4776: 4772: 4768: 4764: 4760: 4756: 4749: 4746: 4741: 4737: 4733: 4729: 4725: 4718: 4716: 4714: 4710: 4705: 4699: 4695: 4690: 4689: 4680: 4677: 4672: 4666: 4662: 4658: 4652: 4649: 4644: 4640: 4636: 4632: 4628: 4620: 4617: 4612: 4608: 4606: 4597: 4595: 4591: 4585: 4580: 4576: 4572: 4568: 4564: 4560: 4558: 4549: 4547: 4545: 4543: 4539: 4534: 4530: 4523: 4516: 4513: 4508: 4504: 4497: 4495: 4491: 4482: 4480: 4478: 4476: 4472: 4467: 4463: 4456: 4453: 4448: 4444: 4440: 4438: 4429: 4426: 4420: 4415: 4411: 4407: 4403: 4401: 4392: 4390: 4388: 4386: 4382: 4377: 4373: 4369: 4365: 4361: 4357: 4353: 4349: 4341: 4338: 4333: 4329: 4325: 4321: 4314: 4312: 4308: 4303: 4299: 4295: 4291: 4287: 4283: 4276: 4274: 4270: 4265: 4261: 4257: 4255: 4251: 4242: 4239: 4234: 4230: 4226: 4222: 4218: 4214: 4210: 4203: 4201: 4199: 4195: 4190: 4186: 4182: 4178: 4174: 4170: 4163: 4160: 4155: 4151: 4147: 4143: 4139: 4135: 4128: 4126: 4122: 4118: 4112: 4109: 4102: 4099: 4094: 4090: 4086: 4082: 4078: 4071: 4068: 4063: 4059: 4055: 4051: 4047: 4040: 4037: 4032: 4028: 4024: 4020: 4013: 4011: 4007: 4002: 3998: 3993: 3988: 3983: 3978: 3974: 3970: 3967:(1): e30212. 3966: 3962: 3958: 3951: 3949: 3947: 3945: 3941: 3936: 3932: 3928: 3921: 3918: 3913: 3909: 3905: 3903: 3894: 3891: 3886: 3882: 3878: 3874: 3870: 3866: 3862: 3858: 3854: 3850: 3846: 3839: 3836: 3831: 3827: 3823: 3819: 3812: 3809: 3804: 3800: 3796: 3792: 3788: 3784: 3780: 3773: 3771: 3769: 3767: 3765: 3763: 3761: 3757: 3752: 3748: 3744: 3737: 3735: 3733: 3731: 3729: 3727: 3725: 3723: 3721: 3719: 3717: 3715: 3711: 3706: 3702: 3698: 3694: 3690: 3683: 3681: 3679: 3675: 3670: 3666: 3662: 3655: 3653: 3649: 3643: 3639: 3636: 3635: 3631: 3620: 3615: 3613: 3611: 3610: 3606: 3605:dromaeosaurid 3602: 3598: 3594: 3590: 3589:ornithomimids 3586: 3585: 3580: 3579: 3574: 3573: 3568: 3567: 3562: 3561: 3557: 3553: 3552: 3547: 3546: 3542: 3538: 3537: 3532: 3531: 3530:Corythosaurus 3526: 3525: 3520: 3519: 3514: 3513: 3509: 3505: 3504: 3499: 3498: 3497:Styracosaurus 3493: 3492: 3487: 3483: 3479: 3475: 3471: 3467: 3463: 3459: 3455: 3451: 3448: 3444: 3440: 3436: 3432: 3427: 3425: 3424:coastal plain 3421: 3415: 3413: 3409: 3405: 3401: 3397: 3393: 3389: 3385: 3381: 3377: 3373: 3369: 3365: 3361: 3354: 3350: 3345: 3338: 3336: 3333: 3329: 3324: 3319: 3314: 3311: 3307: 3303: 3298: 3294: 3289: 3287: 3282: 3277: 3273: 3269: 3265: 3261: 3257: 3256:heat-exchange 3248: 3241: 3239: 3237: 3233: 3229: 3225: 3216: 3206: 3197: 3188: 3185: 3174: 3170: 3160: 3151: 3142: 3139: 3135: 3130: 3126: 3122: 3118: 3114: 3110: 3106: 3102: 3098: 3094: 3090: 3086: 3082: 3078: 3074: 3069: 3067: 3062: 3050: 3043: 3031: 3022: 3019: 3014: 3009: 3005: 3004:Homalocephale 3001: 2996: 2994: 2993:bighorn sheep 2990: 2985: 2980: 2979:Edwin Colbert 2976: 2968: 2967: 2962: 2957: 2950: 2948: 2946: 2942: 2938: 2934: 2927:Dome function 2926: 2924: 2921: 2916: 2914: 2910: 2906: 2902: 2898: 2893: 2891: 2890:hypertrophied 2886: 2882: 2874: 2869: 2865: 2863: 2859: 2855: 2850: 2846: 2842: 2838: 2834: 2826: 2821: 2814: 2812: 2810: 2809:cartilaginous 2806: 2802: 2798: 2794: 2789: 2785: 2781: 2777: 2773: 2765: 2763: 2761: 2757: 2753: 2748: 2744: 2740: 2739: 2734: 2729: 2726: 2722: 2718: 2713: 2709: 2699: 2692: 2688:Palaeobiology 2687: 2685: 2683: 2679: 2678:Late Jurassic 2675: 2674:ghost lineage 2671: 2667: 2663: 2662:Homalocephale 2659: 2655: 2651: 2647: 2646: 2640: 2630: 2629: 2621: 2620: 2612: 2611: 2603: 2602: 2594: 2593: 2585: 2584: 2576: 2575: 2567: 2566: 2558: 2557: 2549: 2548: 2540: 2539: 2536: 2535: 2534: 2527: 2526: 2523: 2522: 2519: 2518: 2517: 2510: 2509: 2503: 2502: 2499: 2498: 2490: 2489: 2481: 2480: 2477: 2476: 2475: 2468: 2467: 2464: 2463: 2460: 2459: 2458: 2451: 2450: 2444: 2443: 2440: 2439: 2436: 2435: 2434: 2427: 2426: 2423: 2422: 2419: 2418: 2417: 2410: 2409: 2403: 2402: 2396: 2395: 2392: 2391: 2388: 2387: 2386: 2379: 2378: 2375: 2374: 2371: 2370: 2369: 2362: 2361: 2355: 2354: 2351: 2350: 2347: 2346: 2345: 2338: 2337: 2331: 2330: 2327: 2326: 2323: 2322: 2321: 2314: 2313: 2307: 2306: 2303: 2302: 2299: 2298: 2297: 2290: 2289: 2283: 2282: 2279: 2278: 2275: 2274: 2273: 2266: 2265: 2259: 2258: 2255: 2254: 2251: 2250: 2249: 2242: 2241: 2235: 2232: 2231: 2228: 2227: 2219: 2218: 2215: 2214: 2213: 2206: 2205: 2202: 2201: 2198: 2197: 2196: 2189: 2188: 2185: 2184: 2181: 2180: 2179: 2172: 2171: 2168: 2167: 2164: 2163: 2162: 2155: 2154: 2148: 2147: 2141: 2138: 2137: 2134: 2133: 2130: 2129: 2128: 2121: 2120: 2114: 2111: 2110: 2103: 2098: 2094: 2092: 2088: 2084: 2080: 2075: 2073: 2069: 2065: 2061: 2057: 2053: 2047: 2045: 2041: 2037: 2036: 2035:Protoceratops 2031: 2027: 2023: 2019: 2011: 2006: 1999: 1997: 1995: 1991: 1987: 1983: 1979: 1975: 1974:Homalocephale 1971: 1967: 1963: 1959: 1956: 1952: 1951:pelvic girdle 1948: 1944: 1940: 1936: 1931: 1924: 1920: 1916: 1911: 1909: 1905: 1900: 1892: 1890:caudal basket 1885: 1884:Homalocephale 1881: 1876: 1869: 1865: 1861: 1845: 1836: 1827: 1816: 1807: 1798: 1786: 1784: 1782: 1777: 1773: 1768: 1765:The skull of 1763: 1761: 1757: 1752: 1745: 1741: 1737: 1733: 1728: 1721: 1717: 1710: 1709:ornithischian 1706: 1702: 1698: 1694: 1690: 1688: 1684: 1680: 1676: 1672: 1668: 1664: 1660: 1655: 1653: 1649: 1645: 1641: 1640:lacrimal bone 1637: 1632: 1624: 1617: 1613: 1608: 1606: 1600: 1593: 1589: 1584: 1580: 1576: 1572: 1569:The skull of 1563: 1556: 1554: 1552: 1548: 1544: 1540: 1536: 1532: 1528: 1524: 1517: 1512: 1505: 1503: 1502: 1498: 1494: 1490: 1486: 1485: 1480: 1476: 1472: 1468: 1464: 1460: 1456: 1452: 1447: 1445: 1441: 1437: 1433: 1429: 1428:stratigraphic 1425: 1422: 1418: 1414: 1410: 1406: 1402: 1398: 1393: 1389: 1384: 1382: 1378: 1374: 1370: 1366: 1362: 1358: 1354: 1350: 1345: 1343: 1339: 1335: 1331: 1327: 1326: 1321: 1320: 1315: 1312: 1308: 1307: 1302: 1298: 1294: 1290: 1286: 1283:had become a 1282: 1278: 1277: 1272: 1271:S. sternbergi 1268: 1267: 1262: 1258: 1254: 1250: 1246: 1242: 1238: 1234: 1230: 1229: 1224: 1220: 1216: 1212: 1208: 1207: 1203:to the genus 1202: 1198: 1190: 1186: 1182: 1175: 1173: 1171: 1167: 1163: 1159: 1156:, found that 1155: 1151: 1147: 1146:S. sternbergi 1143: 1139: 1135: 1130: 1126: 1122: 1118: 1114: 1110: 1106: 1102: 1098: 1094: 1090: 1089: 1088:Homalocephale 1084: 1080: 1076: 1072: 1068: 1064: 1060: 1057: 1053: 1052:Birger Bohlin 1043: 1039: 1029: 1020: 1011: 1002: 1000: 996: 992: 988: 984: 980: 976: 972: 971:T. sternbergi 968: 964: 960: 956: 952: 948: 944: 940: 936: 932: 928: 924: 923: 918: 914: 910: 909:T. sternbergi 906: 902: 897: 895: 890: 886: 882: 878: 874: 870: 866: 862: 858: 854: 850: 849: 844: 840: 839:ornithischian 836: 831: 827: 823: 819: 810: 806: 802: 798: 797: 792: 788: 784: 775: 766: 757: 755: 751: 750:Stegosauridae 747: 743: 739: 734: 730: 726: 722: 717: 713: 709: 708: 703: 697: 695: 691: 687: 683: 679: 675: 672: 668: 664: 660: 656: 652: 648: 644: 640: 636: 633: 629: 621: 617: 612: 605: 603: 601: 597: 593: 589: 585: 581: 577: 573: 569: 565: 561: 557: 553: 550:in the group 549: 545: 541: 536: 534: 530: 526: 521: 518:was a small, 517: 513: 512: 508: 504: 503: 498: 494: 489: 488: 484: 480: 476: 473: 469: 468:North America 465: 461: 457: 453: 452: 438: 432: 429: 423: 420: 414: 411: 406: 403: 398: 395: 392:Gilmore, 1924 390: 387: 382: 379: 368: 365: 361: 352: 351: 344: 339: 338: 331: 330: 328: 325: 321: 314: 313: 305: 302: 298: 294: 290: 285: 284: 277: 274: 273: 270: 264: 261: 260: 257: 251: 248: 245: 244: 241: 235: 232: 229: 228: 225: 219: 216: 213: 212: 209: 206: 203: 200: 199: 196: 193: 190: 189: 186: 183: 180: 179: 176: 173: 170: 169: 164: 159: 155: 152: 148: 143: 139: 134: 120: 115: 110: 105: 100: 95: 90: 85: 80: 75: 70: 65: 59: 56:77.5–74 50: 44: 40: 37: 33: 19: 6405: 6333: 6326: 6319: 6314:Prenocephale 6312: 6305: 6298: 6291: 6284: 6277: 6270: 6263: 6256: 6236: 6230: 6229: 6222: 6215: 6208: 6184: 6176: 6142: 6102:Ornithischia 6087:Ornithischia 6039: 6035:Ornithischia 6030: 6021: 5905: 5893: 5868: 5859: 5835: 5824: 5800: 5789: 5764: 5760: 5754: 5737: 5733: 5727: 5708: 5704: 5694: 5678:(1): 18–22. 5675: 5671: 5661: 5644: 5640: 5634: 5589: 5585: 5579: 5571: 5526: 5522: 5512: 5503: 5499: 5489: 5480: 5476: 5470: 5456:(1): 40–47. 5453: 5449: 5443: 5402: 5399:Paleobiology 5398: 5335: 5331: 5277: 5273: 5269: 5243: 5239: 5235: 5229: 5194: 5190: 5180: 5150:(1): 67–77. 5147: 5143: 5137: 5112: 5108: 5104: 5100: 5094: 5067: 5063: 5053: 5028: 5020: 4985: 4981: 4960: 4955: 4946: 4942: 4936: 4928: 4919: 4915: 4853: 4849: 4843: 4804: 4758: 4754: 4748: 4731: 4727: 4687: 4679: 4660: 4651: 4626: 4619: 4610: 4604: 4566: 4562: 4556: 4532: 4528: 4515: 4506: 4502: 4493: 4489: 4465: 4461: 4455: 4446: 4442: 4436: 4428: 4409: 4405: 4399: 4351: 4347: 4340: 4323: 4319: 4285: 4281: 4263: 4259: 4253: 4249: 4241: 4216: 4212: 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2857: 2856:study of an 2854:histological 2849:morphometric 2844: 2841:Ornatotholus 2840: 2832: 2830: 2824: 2804: 2792: 2783: 2771: 2769: 2759: 2755: 2746: 2742: 2736: 2732: 2730: 2724: 2720: 2707: 2704: 2661: 2653: 2643: 2639:biogeography 2636: 2531: 2530: 2514: 2513: 2472: 2471: 2455: 2454: 2431: 2430: 2414: 2413: 2383: 2382: 2366: 2365: 2342: 2341: 2318: 2317: 2294: 2293: 2270: 2269: 2246: 2245: 2211: 2210: 2209: 2194: 2193: 2192: 2176: 2175: 2159: 2158: 2125: 2124: 2101: 2090: 2082: 2076: 2067: 2060:paedomorphic 2055: 2051: 2048: 2033: 2029: 2025: 2022:Dong Zhiming 2014: 1993: 1973: 1912: 1904:teleost fish 1883: 1868:zygagophyses 1863: 1857: 1780: 1775: 1771: 1766: 1764: 1756:tooth crowns 1715: 1714: 1700: 1671:nasal cavity 1656: 1609: 1570: 1568: 1550: 1546: 1542: 1534: 1530: 1522: 1521: 1515: 1500: 1496: 1492: 1488: 1482: 1478: 1474: 1470: 1466: 1462: 1458: 1450: 1448: 1443: 1439: 1435: 1431: 1423: 1413:heterochrony 1408: 1404: 1400: 1396: 1391: 1385: 1380: 1372: 1364: 1360: 1346: 1341: 1337: 1333: 1329: 1323: 1317: 1313: 1310: 1304: 1300: 1296: 1292: 1288: 1280: 1274: 1270: 1264: 1260: 1256: 1253:Ornatotholus 1252: 1248: 1244: 1241:nomen dubium 1240: 1237:Ornatotholus 1236: 1228:nomen dubium 1226: 1222: 1219:Prenocephale 1218: 1217:belonged to 1214: 1210: 1206:Prenocephale 1204: 1200: 1196: 1194: 1189:Ornatotholus 1188: 1169: 1166:Ornatotholus 1165: 1161: 1157: 1153: 1149: 1145: 1141: 1137: 1133: 1128: 1124: 1121:Ornatotholus 1120: 1116: 1100: 1096: 1093:Peter Galton 1086: 1082: 1070: 1066: 1058: 1055: 1049: 1037: 998: 994: 990: 986: 982: 978: 974: 970: 962: 954: 946: 942: 938: 934: 930: 926: 920: 916: 912: 908: 901:Barnum Brown 898: 885:Troodontidae 880: 876: 872: 868: 864: 856: 852: 846: 842: 821: 815: 808: 794: 745: 741: 737: 732: 725:ceratopsians 720: 715: 705: 698: 681: 677: 673: 666: 658: 627: 625: 619: 591: 579: 563: 555: 548:ceratopsians 543: 539: 537: 515: 514: 510: 501: 500: 496: 486: 485: 483:type species 450: 449: 448: 436: 430: 421: 412: 404: 396: 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S. 4734:: 347–365. 4657:Glut, D. F. 3914:(4): 23–36. 3855:(445): 60. 3597:troodontids 3560:Gorgosaurus 3541:ankylosaurs 3524:Gryposaurus 3482:angiosperms 3435:floodplains 3228:saurischian 3136:lacked the 3095:(UALVP 2), 2989:plexi-glass 2040:Paul Sereno 1933:(where the 1622:basicranium 1527:postcranial 1506:Description 1501:Stegoceras. 1497:Texacephale 1479:Stegoceras. 1417:ontogenetic 1351:) from the 1338:Gravitholus 1325:Gravitholus 1319:Sinocephale 1235:considered 1233:Thomas Carr 1168:a juvenile 999:T. formosus 977:members of 943:T. formosus 931:T. formosus 859:an invalid 799:(right), a 707:Triceratops 702:nasal bones 511:Stegoceras. 470:during the 384:Lambe, 1902 341:Lambe, 1902 317:Lambe, 1902 32:Stegosaurus 6506:Categories 6436:Stegoceras 6406:Stegoceras 6369:See also: 6258:Acrotholus 6231:Stegoceras 6224:Hanssuesia 6123:see below↓ 6026:Dinosauria 5956:Stegoceras 5944:Stegoceras 5483:: 459–472. 5236:Stegoceras 5115:: 105058. 4949:: 115–127. 4535:: 489–497. 4509:: 202–215. 4490:Stegoceras 4449:: 329–330. 3902:Stegoceras 3832:: 266–267. 3818:Stegoceras 3779:Stegoceras 3644:References 3545:Edmontonia 3539:, and the 3486:Stegoceras 3478:tree ferns 3470:understory 3412:Stegoceras 3388:Stegoceras 3384:Hanssuesia 3360:S. validum 3353:S. validum 3328:Stegoceras 3323:Stegoceras 3318:Stegoceras 3276:Stegoceras 3268:Stegoceras 3264:trabeculae 3224:Stegoceras 3215:S. validum 3184:Stegoceras 3173:S. validum 3169:S. validum 3134:Stegoceras 3129:Stegoceras 3125:Stegoceras 3105:Stegoceras 3093:S. validum 3091:skulls of 3089:CT scanned 3081:osteoderms 3061:Stegoceras 3018:Stegoceras 3008:Stegoceras 2984:Stegoceras 2975:Stegoceras 2961:S. validum 2933:Stegoceras 2920:S. validum 2913:Stegoceras 2901:allometric 2897:S. validum 2883:, showing 2881:S. validum 2873:S. validum 2858:S. validum 2845:Stegoceras 2833:Stegoceras 2825:S. validum 2805:S. validum 2793:Stegoceras 2784:Stegoceras 2772:Stegoceras 2756:Stegoceras 2743:Stegoceras 2733:Stegoceras 2725:Stegoceras 2721:Stegoceras 2708:Stegoceras 2654:Stegoceras 2645:Acrotholus 2102:S. validum 2091:Stegoceras 2083:Stegoceras 2068:Stegoceras 2052:Stegoceras 2030:Stegoceras 2026:Stegoceras 1994:Stegoceras 1986:metatarsus 1898:myorhabdoi 1864:Stegoceras 1781:S. validum 1776:S. validum 1767:Stegoceras 1720:heterodont 1716:Stegoceras 1705:integument 1701:S. validum 1663:premaxilla 1659:nasal bone 1652:jugal bone 1644:prefrontal 1571:Stegoceras 1551:S. validum 1543:Stegoceras 1535:Stegoceras 1531:S. validum 1523:Stegoceras 1516:S. validum 1493:S. validum 1475:Stegoceras 1469:, but not 1467:Stegoceras 1463:S. validum 1459:Stegoceras 1451:Stegoceras 1444:S. validum 1401:S. validum 1381:Stegoceras 1365:S. validum 1357:New Mexico 1342:Stegoceras 1330:Hanssuesia 1297:Stegoceras 1293:Stegoceras 1289:S. validum 1281:Stegoceras 1276:Hanssuesia 1257:Stegoceras 1249:Stegoceras 1245:Stegoceras 1223:Stegoceras 1215:S. bexelli 1170:S. validum 1162:Stegoceras 1158:S. bexelli 1154:S. validum 1101:S. validus 1083:Stegoceras 1071:S. bexelli 1067:Stegoceras 1063:Oskar Kuhn 1038:S. validum 991:S. validus 979:Stegoceras 963:Stegoceras 947:S. validus 939:T. validus 927:T. validus 889:ornithopod 881:S. validus 873:T. validus 869:S. validus 857:Stegoceras 843:S. validus 822:S. validus 809:Stegoceras 746:S. validus 738:S. validus 733:Stegoceras 729:stegosaurs 721:Stegoceras 716:Stegoceras 667:Stegoceras 628:Stegoceras 620:S. validum 580:S. validum 564:Stegoceras 556:Stegoceras 544:Stegoceras 540:Stegoceras 516:Stegoceras 507:New Mexico 497:S. validum 451:Stegoceras 337:S. validum 283:Stegoceras 208:Dinosauria 147:S. validum 43:Stegoceras 6009:Kingdom: 5129:239253658 4643:0016-6995 4569:: 29–43. 4376:244227050 4266:: 177–90. 4254:P. brevis 3408:Campanian 3400:Fruitland 3272:honeycomb 3270:and the " 3138:pneumatic 2905:isometric 2776:endocasts 2664:from the 2656:from the 2650:Santonian 2087:cladogram 2064:dimorphic 1958:archosaur 1744:denticles 1726:thecodont 1711:dinosaurs 1679:olfactory 1630:braincase 1605:tubercles 1529:remains; 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