Stegoceras (Greek for "roof horn") was a genus of ornithischian plant eating pachycephalosaurid dinosaur that was found in North America in the Late Cretaceous period 74-65 million years ago. It has an estimated length of 1.53 m (5 ft) and was 50 cm (19⅝ in) high at the hips and weighed 13 kg (28⅝ lb) which is quite small for a dinosaur. It was named by Lawrence Lambe in 1902.
Stegoceras' name means "roof horn" from the Greek words stegos/στέγος- "roof" or "covered" and kéras/κέρας -"horn".
Stegoceras sported a 5.1 centimeter (2 inch)-thick dome. It was initially proposed that male Stegoceras (and individuals of other pachycephalosaurid species) would ram each other headlong, not unlike contemporary bighorn sheepor musk oxen. It was later suggested that they engaged in flank-butting rather than ramming, a widely evidenced theory. Foremost, the rounded shape of the skull roof would lessen the contacted surface area during head-butting, resulting in glancing blows.
Second, pachycephalosaurs would not have been able to align their head, neck, and body in a perfect horizontal line (which would be needed to transmit stress) -- it was more likely that they carried their neck in an "S"- or "U"-shaped curve (Stegoceras seemed to carry their spine in a less extreme curve, due to their thick neck muscles).
Lastly, the relatively large width of most pachycephalosaurs would have served to protect vital organs from harm during flank-butting. When a partial skeleton of Stegoceras was first discovered, it was thought to have gastralia, or belly ribs, not typically found in other ornithischian dinosaurs. They were subsequently found to be ossified tendons.
Description[]
Size of S. validum compared to a human
Stegoceras had an estimated length of up to 2 metres (6.6 ft) and weighed 10 kilograms (22 lb) – 40 kilograms (88 lb). It had a relatively large brain that was encased in a dome of 3 inches (7.6 cm) of thick bone divided into two parts. The dome had a fairly smooth surface, but was irregularly pitted by foramina which gives an entrance to channels within the bone. Stegoceras (at least S. validum) can be distinguished by having a prominent parietosquamosal shelf with open supratemporal fosse, incipient doming of the frontopariental and minute node in clusters on postorbital and squamosals. Stegoceras had rounded eye sockets that faced forward, which suggests it had good vision and was thus capable of binocular vision. The teeth are small and curved with serrated edges. The head was supported by an "S"- or "U"-shaped neck. When a partial skeleton of Stegoceras was first discovered, it was thought to have gastralia, or belly ribs, not typically found in other ornithischian dinosaurs. They were subsequently found to be ossified tendons. The legs were more than three times the length of the arms.
Discovery[]
Partial dome CMN 515, lectotype of S. validum, shown from the right and underside
The first known remains of Stegoceras were collected by Canadian palaeontologist Lawrence Lambe from the Belly River Group, in the Red Deer River district of Alberta, Canada. These remains consisted of two partial skull domes (specimens CMN 515 and CMN 1423 in the Canadian Museum of Nature) from two animals of different sizes collected in 1898, and a third partial dome (CMN 1594) collected in 1901. Based on these specimens, Lambe described and named the new monotypic genus and species Stegoceras validus in 1902. The generic name Stegoceras comes from the Greek stegè/στέγη, meaning "roof" and keras/κέρας meaning "horn". The specific name validus means "strong" in Latin, possibly in reference to the thick skull-roof. Because the species was based on multiple specimens (a syntype series), CMN 515 was designated as the lectotype specimen by John Bell Hatcher in 1907.
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 nasal bones on the midline of the head, and noted their similarity to the nasal horn-core of a Triceratops specimen. In 1903, Hungarian palaeontologist Franz Nopcsa von Felső-Szilvás suggested that the fragmentary domes of Stegoceras were in fact frontal and nasal bones, and that the animal would therefore have had a single, unpaired horn. Lambe was sympathetic to this idea of a new type of "unicorn dinosaur" in a 1903 review of Nopscsa's paper. At this time, there was still uncertainty over which group of dinosaur Stegoceras belonged to, with both ceratopsians (horned dinosaurs) and stegosaurs (plated dinosaurs) as contenders. Hatcher doubted whether the Stegoceras specimens belonged to the same species and whether they were dinosaurs at all, and suggested the domes consisted of the frontal, occipital, and parietal bones of the skull. In 1918, Lambe referred another dome (CMN 138) to S. validus, and named a new species, S. brevis, based on specimen CMN 1423 (which he originally included in S. validus). By this time, he considered these animals as members of Stegosauria (then composed of both families of armoured dinosaurs, Stegosauridae and Ankylosauridae), in a new family he called Psalisauridae (named for the vaulted or dome-shaped skull roof).
In 1924, the American palaeontologist Charles W. Gilmore described a complete skull of S. validus with associated postcranial remains, by then the most complete remains of a dome-headed dinosaur. It was discovered in the Belly River Group by the American palaeontologist George F. Sternberg in 1926, and catalogued as specimen UALVP 2 in the University of Alberta Laboratory for Vertebrate Palaeontology. This find confirmed Hatcher's interpretation of the domes as consisting of the frontoparietal area of the skull. UALVP 2 was found with small, disarticulated bony elements, then thought to be gastralia (abdominal ribs), which are not known in other ornithischian dinosaurs (one of the two main groups of dinosaurs). Gilmore pointed out that the teeth of S. validus were very similar to those of the species Troodon formosus (named in 1856 and by then only known from isolated teeth), and described a skull dome discovered close to the locality where Troodon was found. Therefore, Gilmore considered Stegoceras an invalid junior synonym of Troodon, thereby renaming S. validus into T. validus, and suggested that even the two species might be the same. Furthermore, he found S. brevis to be identical to S. validus, and therefore a junior synonym of the latter. He also placed these species in the new family Troodontidae (since Lambe had not selected a type genus for his Psalisauridae), which he considered closest to the ornithopod dinosaurs. 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 nodosaur, 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.
Gilmore's classification was supported by the American palaeontologists Barnum Brown and Erich Maren Schlaikjer 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 T. sternbergi and T. edmontonensis (both from Alberta), as well as moving the large species T. wyomingensis (which was named in 1931) to the new genus Pachycephalosaurus, along with two other species. They found T. validus distinct from T. formosus, but considered S. brevis the female form of T. validus, 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 T. formosus and S. validus teeth, the American palaeontologist Charles M. Sternberg demonstrated differences between the two, and instead suggested that Troodon was a theropod dinosaur, and that the dome-headed dinosaurs should be placed in their own family. Though Stegoceras was the first member of this family to be named, Sternberg named the group Pachycephalosauridae after the second genus, as he found that name (meaning "thick head lizard") more descriptive. He also considered T. sternbergi and T. edmontonensis members of Stegoceras, found S. brevis valid, and named a new species, S. lambei, based on a specimen formerly referred to S. validus. The split from Troodon was supported by Russell in 1948, who described a theropod dentary with teeth almost identical to those of T. formosus.
In 1953, Birger Bohlin named Troodon bexelli based on a parietal bone from China. In 1964, Oskar Kuhn considered this as an unequivocal species of Stegoceras; S. bexelli. In 1974, the Polish palaeontologists Teresa Maryańska and Halszka Osmólska concluded that the "gastralia" of Stegoceras were ossified tendons, after identifying such structures in the tail of the pachycephalosaur Homalocephale. In 1979, William Patrick Wall and Peter Galton named the new species Stegoceras browni, based on a flattened dome, formerly described as a female S. validus by Galton in 1971. The specific name honours Barnum Brown, who found the holotype specimen (specimen AMNH 5450 in the American Museum of Natural History) in Alberta. In 1983, Galton and Hans-Dieter Sues moved S. browni to its own genus, Ornatotholus (ornatus is Latin for "adorned" and tholus for "dome"), and considered it the first known American member of a group of "flat-headed" pachycephalosaurs, previously known from Asia. In a 1987 review of the pachycephalosaurs, Sues and Galton emended the specific name validus to validum, which has subsequently been used in the scientific literature. These authors synonymized S. brevis, S. sternbergi, and S. lambei with S. validum, found that S. bexelli differed from Stegoceras in several features, and considered it an indeterminate pachycephalosaur. In 1998, Goodwin and colleagues considered Ornatotholus a juvenile S. validum, therefore a junior synonym.
Classification[]
Reconstructed skeleton showing ossified tail tendons in place, National Museum of Natural History
During the 1970s, more pachycephalosaur genera were described from Asian fossils, which provided more information about the group. In 1974, Maryańska and Osmólska concluded that pachycephalosaurs are distinct enough to warrant their own suborder within Ornithischia, Pachycephalosauria. In 1978, the Chinese palaeontologist Dong Zhiming split Pachycephalosauria into two families; the dome-headed Pachycephalosauridae (including Stegoceras) and the flat-headed Homalocephalidae (originally spelled Homalocephaleridae). Wall and Galton did not find suborder status for the pachycephalosaurs justified in 1979. 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 Stegoceras and the "primitive" ceratopsian Protoceratops. In 1986, American palaeontologist Paul Sereno supported the relationship between pachycephalosaurs and ceratopsians, and united them in the group Marginocephalia, 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.
By the early 21st century, few pachycephalosaur genera were known from postcranial remains, and many taxa were only known from domes, which made classification within the group difficult. Pachycephalosaurs are thus mainly defined by cranial features, such as the flat to domed frontoparietal, the broad and flattened bar along the postorbital and squamosal bones, and the squamosal bones being deep plates on the occiput. 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 Stegoceras a transition between the supposedly "primitive" flat-headed and advanced "fully" domed genera (such as Pachycephalosaurus). The dome-headed/flat-headed division of the pachycephalosaurs was abandoned in the following years, as flat heads were considered paedomorphic (juvenile-like) or derived traits in most revisions, but not a sexually dimorphic trait. In 2006, Sullivan argued against the idea that the extent of doming was useful in determining taxonomic affinities between pachycephalosaurs. In 2003, Sullivan found Stegoceras itself to be more basal (or "primitive") than the "fully-domed" members of the subfamily Pachycephalosaurinae, elaborating on conclusions reached by Sereno in 1986.
A 2013 phylogenetic analysis by Evans and colleagues found that some flat-headed pachycephalosaur genera were more closely related to "fully" domed taxa than to the "incompletely" domed Stegoceras, which suggests they represent juveniles of domed taxa, and that flat heads do not indicate taxonomic affinities. The cladogram below shows the placement of Stegoceras within Pachycephalosauridae according to Schott and colleagues, 2016:
Partial skull of a juvenile, flat-headed S. validum, UCMZ(VP) 2008.001
| Pachycephalosauria |
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The biogeography and early evolutionary history of pachycephalosaurs is poorly understood, and can only be clarified by new discoveries. Pachycephalosaurs appear abruptly in the fossil record, and are present in both North America and Asia, so it is unknown when they first originated, and from which direction they dispersed. The oldest known members of the group (such as Acrotholus) are "fully domed" and known from the Santonian stage of the Late Cretaceous period (about 84 million years ago). This is before the supposedly more primitive Stegoceras from the Middle Campanian (77 million years ago) and Homalocephale from the Early Maastrichtian (70 million years ago), so the doming of the skull may be a homoplastic trait (a form of convergent evolution). The late occurrence of pachycephalosaurs compared to the related ceratopsians indicates a long ghost lineage (inferred, but missing from the fossil record) spanning 66 million years, from the Late Jurassic to the Cretaceous. Since pachycephalosaurs were mainly small, this may be due to taphonomic bias; smaller animals are less likely to be preserved through fossilisation. More delicate bones are also less likely to be preserved, which is why pachycephalosaurs are mainly known from their robust skulls.
JPInstitute.com Description[]
Stegoceras was one of the smallest members of the same family that includes the more well known Pachycephalosaurus. It had a thick dome on its head, but instead of a ring of bumps or spikes, it had a small ridge of bone at the base of its dome. These were small, swift-running hunters who may have also eaten plants.
Discovered early in the 20th century, many subsequent dinosaurs were assigned to this genus and later reassigned. The genus is known from a complete skull, a number of partial skulls, and other skeletal material. To date, there is only one valid species for this genus. The skeletal remains of this dinosaur seem to support the theory of head butting for this family, which adds fuel to the debate over whether this activity would have been possible without injury to the spine.
Appearance in other media[]
Jurassic Park[]
- Stegoceras appears in Jurassic World: Alive as a Epic.
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Read more Stegoceras on Jurassic Park Wiki |