Stegoceras (detailed example)

[Morose]

In 1987, J. K. Rigby and colleagues suggested that pachycephalosaur domes were heat-exchange organs used for thermoregulation, based on their internal "radiating structures" (trabeculae). This idea was supported by a few other writers in the mid-1990s. In 1998, Goodwin and colleagues considered the lack of sinuses in the skull of Stegoceras and the "honeycomb"-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 Stegoceras 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 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 Sharpey's fibers 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 bovids) 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.

[Morose]

Restoration of a pair with different coloured domes

In 2008, Eric Snively and Adam Cox tested the performance of 2D and 3D pachycephalosaur skulls through finite element analysis, 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 keratin, a strong material that can withstand much energy without being permanently damaged (like the osteoderms of crocodilians), and therefore incorporated keratin into their test formula. In 2011, American palaeontologists Kevin Padian and John R. Horner proposed that "bizarre structures" in dinosaurs in general (including domes, frills, horns, and crests) were primarily used for species recognition, and dismissed other explanations as unsupported by evidence. Among other studies, these authors cited Goodwin et al.'s 2004 paper on pachycephalosaur domes as support of this idea, and they pointed out that such structures did not appear to be sexually dimorphic. In a response to Padian and Horner the same year, Rob J. Knell and Scott D. Sampson argued that species recognition was not unlikely as a secondary function for "bizarre structures" in dinosaurs, but that sexual selection (used in display or combat to compete for mates) was a more likely explanation, due to the high cost of developing them, and because such structures appear to be highly variable within species. In 2013, the British palaeontologists David E. Hone and Darren Naish criticized the "species recognition hypothesis", and argued that no extant animals use such structures primarily for species recognition, and that Padian and Horner had ignored the possibility of mutual sexual selection (where both sexes are ornamented).

[Morose]

CT cross-sections of the skulls of UALVP 2 (left) and a white-bellied duiker (right)

In 2011, Snively and Jessica M. Theodor conducted a finite element analysis by simulating head-impacts with CT scanned skulls of S. validum (UALVP 2), Prenocephale prenes and several extant head-butting artiodactyls. They found that the correlations between head-striking and skull morphologies found in the living animals also existed in the studied pachycephalosaurs. Stegoceras and Prenocephale both had skull shapes similar to the bighorn sheep with cancellous bone protecting the brain. They also shared similarities in the distribution of compact and cancellous regions with the bighorn sheep, white-bellied duiker and the giraffe. The white-bellied duiker was found to be the closest morphological analogue to Stegoceras; this head-butting species has a dome which is smaller but similarly rounded. Stegoceras was better capable of dissipating force than artiodactyls that butt heads at high forces, but the less vascularized domes of older pachycephalosaurs, and possibly diminished ability to heal from injuries, argued against such combat in older individuals. The study also tested the effects of a keratinous covering of the dome, and found it to aid in performance. Though Stegoceras lacked the pneumatic sinuses that are found below the point of impact in the skulls of head-striking artiodactyls, it instead had vascular struts which could have similarly acted as braces, as well as conduits to feed the development of a keratin covering.

[Morose]

Two S. validum domes with lesions (arrows) shown from above

In 2012, Schott and Evans suggested that the regularity in squamosal ornamentation throughout the ontogeny of Stegoceras was consistent with species recognition, but the change from flat to domed frontoparietals in late age suggests that the function of this feature changed through ontogeny, and was perhaps sexually selected, possibly for intra-specific combat. In 2012, Caleb M. Brown and Anthony P. Russell suggested that the stiffened tails were probably not used as defence against flank-butting, but may have enabled the animals to take a tripodal stance during intra-specific combat, with the tail as support. Brown and Russell found that the tail could thereby help in resisting compressive, tensile, and torsional loading when the animal delivered or received blows with the dome.

[Morose]

A 2013 study by Joseph E. Peterson and colleagues identified lesions in skulls of Stegoceras and other pachycephalosaurs, which were interpreted as infections caused by trauma. Lesions were found on 22% of sampled pachycephalosaur skulls (a frequency consistent across genera), but were absent from flat-headed specimens (which have been interpreted as juveniles or females), which is consistent with use in intra-specific combat (for territory or mates). The distribution of lesions in these animals tended to concentrate at the top of the dome, which supports head-butting behaviour. Flank-butting would probably result in fewer injuries, which would instead be concentrated on the sides of the dome. These observations were made while comparing the lesions with those on the skulls and flanks of modern sheep skeletons. The researchers noted that modern head-butting animals use their weapons for both combat and display, and that pachycephalosaurs could therefore also have used their domes for both. Displaying a weapon and willingness to use it can be enough to settle disputes in some animals.

[Morose]

Nasal passages

In 1989, Emily B. Griffin found that Stegoceras and other pachycephalosaurs had a good sense of smell (olfaction), based on the study of cranial endocasts that showed large olfactory bulbs in the brain. In 2014, Jason M. Bourke and colleagues found that Stegoceras would have needed cartilaginous nasal turbinates in the front of the nasal passages for airflow to reach the olfactory region. Evidence for the presence of this structure is a bony ridge to which it could have attached. The size of the olfactory region also indicates that Stegoceras had a keen sense of smell. The researchers found that the dinosaur could have had either a scroll-shaped turbinate (like in a turkey) or a branched one (as in an ostrich) as both could have directed air to the olfactory region. The blood vessel system in the passages also suggest that the turbinates served to cool down warm arterial blood from the body that was heading to the brain. The skull of S. validum specimen UALVP 2 was suited for a study of this kind due to its exceptional preservation; it has ossified soft tissue in the nasal cavity, which would otherwise be cartilaginous and therefore not preserved through mineralization.

[Morose]

Palaeoecology

The Dinosaur Park Formation in Alberta, Canada, where S. validumwas was first discovered

S. validum is known from the late Late Cretaceous Belly River Group (the Canadian equivalent to the Judith River Group in the US), and specimens have been recovered from the Dinosaur Park Formation (late Campanian, 76.5 to 75 mya) in Dinosaur Provincial Park (including the lectotype specimen), and the Oldman Formation (middle Campanian, 77.5 to 76.5 mya) of Alberta, Canada.

[Morose]

The pachycephalosaurs Hanssuesia and Foraminacephale are also known from both formations. S. novomexicanum is known from the Fruitland (late Campanian, about 75 mya) and lower Kirtland Formation (late Campanian, about 74 mya) of New Mexico, and if this species correctly belongs in Stegoceras, the genus would have had a broad geographic distribution. 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. It has traditionally been suggested that pachycehalosaurs inhabited mountain environments; wear of their skulls was supposedly a result of them having been rolled by water from upland areas, and comparisons with bighorn sheep reinforced the theory. In 2014, Jordan C. Mallon and Evans disputed this idea, as the wear and original locations of the skulls is not consistent with having been transported in such a way, and they instead proposed that North American pachycephalosaurs inhabited alluvial (associated with water) and coastal plain environments.

[Morose]

The Dinosaur Park Formation is interpreted as a low-relief setting of rivers and floodplains that became more swampy and influenced by marine conditions over time as the Western Interior Seaway transgressed westward. The climate was warmer than present-day Alberta, without frost, but with wetter and drier seasons. Coniferswere apparently the dominant canopy plants, with an understory of ferns, tree ferns, and angiosperms. Dinosaur Park is known for its diverse community of herbivores. As well as Stegoceras, the formation has also yielded fossils of the ceratopsians Centrosaurus, Styracosaurus and Chasmosaurus, the hadrosauridsProsaurolophus, Lambeosaurus, Gryposaurus, Corythosaurus, and Parasaurolophus, and the ankylosaurs Edmontonia and Euoplocephalus. Theropods present include the tyrannosaurids Gorgosaurus and Daspletosaurus. Other dinosaurs known from the Oldman Formation include the hadrosaur Brachylophosaurus, the ceratopsians Coronosaurus and Albertaceratops, ornithomimids, therizinosaurs and possibly ankylosaurs. Theropods included troodontids, oviraptorosaurs, the dromaeosaurid Saurornitholestes and possibly an albertosaurine tyrannosaur.