Stegoceras is a genus of pachycephalosaurid (dome-headed) dinosaur that lived in what is now North America during the Late Cretaceous period, about 77.5 to 74 million years ago (mya). The first specimens from Alberta, Canada, were described in 1902, and the type species Stegoceras validum was based on these remains. The generic name means "horn roof", and the specific name means "strong". Several other species have been assigned to the genus, but these have since been moved to other genera or deemed junior synonyms. The remaining assigned species, S. novomexicanum, named in 2011 from fossils found in New Mexico, has also been debated, and may not belong to Stegoceras.
Stegoceras was a small, bipedal dinosaur about long, and weighed around . 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 occiput, and it had a thick ridge over the eyes. Much of the skull was ornamented by tubercles (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 vertebral column, and a stiffened tail. The pelvic region was broad, perhaps due to an extended gut.
Originally known only from skull domes, Stegoceras was one of the first known pachycephalosaurs, and the incompleteness of these initial remains led to many theories about the affinities of this group. A complete Stegoceras skull with associated parts of the skeleton was described in 1924, which shed more light on these animals. Pachycephalosaurs are today grouped with the horned ceratopsians in the group Marginocephalia. Stegoceras itself has been considered basal (or "primitive") compared to other pachycephalosaurs. Stegoceras 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 intra-specific combat (head or flank-butting), sexual display, or species recognition. S. validum is known from the Dinosaur Park Formation and the Oldman Formation, whereas S. novomexicanum is known from the Fruitland and Kirtland Formation.
History of discovery
The first known remains of Stegoceras were collected by palaeontologist Lawrence Lambe from the Belly River Group, in the Red Deer River district of Alberta, Canada. These remains consisted of two partial skull domes (specimens CMN 515 and CMN 1423 in the Canadian Museum of Nature) from two animals of different sizes collected in 1898, and a third partial dome (CMN 1594) collected in 1901. Based on these specimens, Lambe described and named the new monotypic genus and species Stegoceras validus in 1902. The generic name Stegoceras comes from the Greek stegè/στέγη, meaning "roof" and keras/κέρας meaning "horn". The specific name validus means "strong" in Latin, possibly in reference to the thick skull-roof. Because the species was based on multiple specimens (a syntype series), CMN 515 was designated as the lectotype specimen by palaeontologist 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, 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 assigned 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).
Confusion with Troodon
In 1924, palaeontologist Charles W. Gilmore described a complete skull of S. validus with associated postcranial remains, by then the most complete remains of a dome-headed dinosaur. It was discovered in the Belly River Group by the American palaeontologist George F. Sternberg in 1926, and catalogued as specimen UALVP 2 in the University of Alberta Laboratory for Vertebrate Palaeontology. This find confirmed Hatcher's interpretation of the domes as consisting of the frontoparietal area of the skull. UALVP 2 was found with small, disarticulated bony elements, then thought to be gastralia (abdominal ribs), which are not known in other ornithischian dinosaurs (one of the two main groups of dinosaurs). Gilmore pointed out that the teeth of S. validus were very similar to those of the species Troodon formosus (named in 1856 and by then only known from isolated teeth), and described a skull dome discovered close to the locality where Troodon was found. Therefore, Gilmore considered Stegoceras an invalid junior synonym of Troodon, thereby renaming S. validus into T. validus, and suggested that even the two species might be the same. Furthermore, he found S. brevis to be identical to S. validus, and therefore a junior synonym of the latter. He also placed these species in the new family Troodontidae (since Lambe had not selected a type genus for his Psalisauridae), which he considered closest to the ornithopod dinosaurs. 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 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, 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 assigned 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.
Later taxonomic developments
In 1953, palaeontologist Birger Bohlin named Troodon bexelli based on a parietal bone from China.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 37:1–113 In 1964, palaeontologist Oskar Kuhn considered this as an unequivocal species of Stegoceras; S. bexelli.Kuhn, O., 1964, Fossilium Catalogus I: Animalia Pars 105. Ornithischia (Supplementum I), IJsel Pers, Deventer, 80 pp In 1974, palaeontologists Teresa Maryańska and Halszka Osmólska concluded that the "gastralia" of Stegoceras were , after identifying such structures in the tail of the pachycephalosaur Homalocephale. In 1979, palaeontologists 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 Brown, who found the holotype specimen (specimen AMNH 5450 in the American Museum of Natural History) in Alberta. This study also named the new genus and species Gravitholus albertae, based on specimen TMP 1972.027.0001 found in the Oldman Formation of Alberta. Its generic name means "heavy dome", and it was thought distinct from Stegoceras by features including its wide dome and small braincase. 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.
In 2000, palaeontologist Robert M. Sullivan assigned S. edmontonensis and S. brevis to the genus Prenocephale (until then only known from the Mongolian species P. prenes), and found it more likely that S. bexelli belonged to Prenocephale than to Stegoceras, but considered it a nomen dubium (dubious name, without distinguishing characters) due to its incompleteness, and noted its holotype specimen appeared to be lost. In 2003, Thomas E. Williamson and Thomas Carr considered Ornatotholus a nomen dubium, or perhaps a juvenile Stegoceras. In a 2003 revision of Stegoceras, Sullivan agreed that Ornatotholus was a junior synonym of Stegoceras, moved S. lambei to the new genus Colepiocephale, and S. sternbergi to the new genus Hanssuesia, named after Sues. He stated that the genus Stegoceras had become a wastebasket taxon for small to medium-sized North American pachycephalosaurs until that point. By this time, dozens of specimens had been assigned to S. validum, including many domes too incomplete to be identified as Stegoceras with certainty. UALVP 2 is still the most complete specimen of Stegoceras, upon which most scientific understanding of the genus is based. S. brevis was moved to the new genus Foraminacephale in 2016 by Ryan K. Schott Schott and David C. Evans, and S. bexelli to Sinocephale in 2021 by Evans and colleagues.
In 2023, palaeontologist Aaron D. Dyer and colleagues analysed sutures and individual elements in the skulls of Gravitholus and Hanssuesia, the former based on synchrotron data, and found no significant distinction between them and Stegoceras validum. They considered both as junior synonyms, with Gravitholus representing the end-stage in the growth of Stegoceras. Supposed distinguishing traits of Hanssuesia were found to be based on misinterpretations.