Acrotholus Temporal range: Late Cretaceous, | |
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Holotype dome on display at the Royal Ontario Museum | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | † Ornithischia |
Clade: | † Neornithischia |
Clade: | † Pachycephalosauria |
Family: | † Pachycephalosauridae |
Genus: | † Acrotholus Evans et al., 2013 |
Type species | |
†Acrotholus audeti Evans et al., 2013 |
Acrotholus (Greek for "highest dome"- akros meaning highest and tholos meaning dome) is an extinct genus of pachycephalosaur dinosaur that lived during the Santonian of the late Cretaceous, in the Milk River Formation of Canada. The type species, A. audeti, was named after Roy Audet allowing access to his ranch leading to the discovery of the species. The discovery of this specimen lead to several new revelations in the fossil records questioning the preservation of small-bodied organisms along with the evolution of early pachycephalosaurs. The iconic cranial dome found on Acrotholus makes it one of the earliest indisputable known members of the pachycephalosaur family. [1]
Like others of its clade, Acrotholus was a bipedal herbivore characterized by a dome-shaped head. The dome had often been associated with intra-species combat though exact method of contact have been debated. [2]
The holotype was found in the Deadhorse Coulee Member of the Milk River Formation in Southern Alberta, Canada in 2008. This formation has been known to expose organisms from the Late Santonian. The fossil consists of a nearly complete frontoparietal dome along with the anterior half of the frontoparietal dome. Pachycephalosaurs in general are unique in dinosaur fossil records due to their relatively small size in relation to most dinosaurs in their time period, 40 kg or less. It has been suggested due to smaller bone being susceptible to destruction by carnivores and weathering, preservation bias exist for smaller species. Pachycephalosaurs are unique in this regard due to their recognizable head dome which is resistant towards pre-depositional destruction. Among known fossil specimens, approximately 66% are known from only cranial remains. Acrotholus' discovery further implicates the diversity of small bodied dinosaurs which may not have survived the fossil record, suggesting a much more complex ecology in the late Mesozoic era. [1] A similar conclusion have been made with Asian species of pachycephalosaurs, with their ontogeny under debate due to their flat domed shapes. [3]
Unlike most known pachycephalosaurs, A. audeti's dome on the skull is oval in shape having a maximum thickness of 55 millimetres (2.2 in) above the cerebral fossa. The lack of tesserae or tubercles indicate the specimen was past the juvenile stage of its development. Computed tomography (CT) scans reveal a low vascularity, high density, and fully fused internally fused frontal-frontal and frontoparietal sutures. Peripheral bones are high and well developed on the dome indicating the peripherals were incorporated within the dome. The incorporation of supraorbitals in the dome are similar if not greater than more derived pachycephalosaur. Compared to Stegoceras validum and later Campanian pachycephalosaurids, the dorsally convex frontonasal boss is short and not separated with grooves from the anterior supraorbital lobe. This region is approximately 50% of the thickness to the cerebral fossa (55 mm). Orbital fosse are only slightly concave and pierced by small foramina. Three depression can be seen on the ventral surface of the frontoparietal dome, the orbital cavity, endocranial fossa, and temporal fossa. Comparisons of the skull finds A. audeti to be distinctly different from other pachycephalosaurs in relation to dome. Most notably, the supraorbital region and incorporation of the supraorbital into the dome identify the specimen from other pachycephalosaurs. Very little soft tissue covering have been suggested over the dome due to the structural anatomy and hypothesized use. [2]
A. audeti was recovered as the sister taxon to Prenocephale prenes . Acrotholus is the earliest known specimen of pachycephalosaur with derived traits from the fossil records, predating later known species of flat headed species in Asia. The discovery of this species further suggests pachycephalosaur head features were well established by the Santonian predating later species thought to have primitive traits such as Homalocephale and Stegoceras validum . [1] [4]
The relatively dome shape of the holotype indicates the specimen was an adult or nearing its completion of its adult transition. Consolidation between flat headed and domed pachycephalosaurs have been greatly debated. Fossils records from more numerous North American specimen suggest a flat domed juvenile stages. Juvenile Pachycephalosaurs wyomingensis have characteristics of nodes on the squamosal nearly identical to holotype fossils of Dracorex hogwartsia and Stygimoloch spinifer. A spike like node on the posteroventral region of the jugal match with that of D. hogwartsia. Such similarities with confirmed juvenile P. wyonmingensis and flat headed species from North America support their recognition as P. wyonmingensis and the ontogenic relationship in growth stages. [6] Comparison of bone development in flat domed taxa from Asia have found patterns indicating active growth at the time of death. [3] Closer analysis of related taxa, Stegoceras novomexicanium found holotype specimen to be juvenile in characteristics, with visible frontal-parietal sutures, small rounded tubercles on the dome, and vascularized internal skull bones. [7] Another Asian taxa, Homalocephale calathocercos was also found exhibit juvenile characteristics in the fossil holotype though it has been proposed some species of pachycephalosaur exhibit pedomorphism. [8]
The most notable physiology of A. audeti is the fused frontal and parietal skull forming a thickened dome structure. The lack of living comparative morphology have led to various theories on the function of the structure. Two accepted hypotheses have been proposed to explain the appearance of the cranial dome. One of the hypotheses suggest the dome plays a visual role, either for sexual display or species recognition. However, such theories have been countered due to the relatively large energy investment required and the morphological changes that occur in the species life span. The more widely accepted and popular hypothesis explains the dome being used as a weapon. In modern species with analogous structures, such head ornaments have been used between intra-species combat. Analysis on dome of the more widely known specimen Pachycephalosaurus wyomingensis have some pathologies in the dome mirroring those of modern-day mammals exhibiting intra-species contact. In P. wyomingensis and other examined pachycephalosaur skulls, chronic osteomyelitis was observed, a pathology often associated with combat. Several skull specimens show various lesions with healing suggesting survival after contact. Flat headed individuals saw little cranial damage supporting their juvenile stage ontogeny. Comparison with extinct and extent species that were known to engage in infra-specific combat have found the formation of lesions to be consistent with their use. [2] [9] [10] Morphological comparisons with modern ungulates practicing head butting have found similar dome shaped structures used in combat. The cancellous region found in head striking artiodactyls have been considered to be covering the domes of pachycephalosaurs to aid in protection against head impacts. In addition, tubular struts in the dome of the related taxa, Stegoceras, were comparable to pneumatized frontal sinuses found in some head-striking mammals. [9] Modern archosaurs such as Ostriches and crocodiles exhibit similar ranges of pathologies due to intra-species combat. [10]
Analysis of pachycephalosaurid skulls by Mallon and Evans, suggest many species frequently inhabited coastal areas contrary to their terrestrial lifestyle. In the same study, it was concluded that North American pachycephalosaurids lived in habitats near floodplains and coastal plains. Hindlimb proportions have been found to be similar to other modern animals such as moose and various wading birds that inhabit wetland areas. [11] The Milk River Formation, where A. audeti is found, is home to many other extinct species in the late Cretaceous including other dinosaurs like Saurornitholestes and mammals like Alphadon . [1]
Marginocephalia is a clade of ornithischian dinosaurs that is characterized by a bony shelf or margin at the back of the skull. These fringes were likely used for display. There are two clades included in Marginocephalia: the thick-skulled Pachycephalosauria and the horned Ceratopsia. All members of Marginocephalia were primarily herbivores. They basally used gastroliths to aid in digestion of tough plant matter until they convergently evolved tooth batteries in Neoceratopsia and Pachycephalosauria. Marginocephalia first evolved in the Jurassic Period and became more common in the Cretaceous. They are basally small facultative quadrupeds while derived members of the group are large obligate quadrupeds. Primitive marginocephalians are found in Asia, but the group migrated upwards into North America.
Pachycephalosauria is a clade of ornithischian dinosaurs. Along with Ceratopsia, it makes up the clade Marginocephalia. With the exception of two species, most pachycephalosaurs lived during the Late Cretaceous Period, dating between about 85.8 and 66 million years ago. They are exclusive to the Northern Hemisphere, all of them being found in North America and Asia. They were all bipedal, herbivorous/omnivorous animals with thick skulls. Skulls can be domed, flat, or wedge-shaped depending on the species, and are all heavily ossified. The domes were often surrounded by nodes and/or spikes. Partial skeletons have been found of several pachycephalosaur species, but to date no complete skeletons have been discovered. Often isolated skull fragments are the only bones that are found.
Pachycephalosaurus is a genus of pachycephalosaurid ornithischian dinosaur. The type species, P. wyomingensis, is the only known species, but some researchers argue that Stygimoloch might be a second species, P. spinifer, or a juvenile specimen of P. wyomingensis. It lived during the Maastrichtian age of the Late Cretaceous period in what is now western North America. Remains have been excavated in Montana, South Dakota, Wyoming, and Alberta. The species is known mainly from a single skull, plus a few extremely thick skull roofs. More complete fossils would come to be found in the following years.
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 placed in the genus over the years, but these have since been moved to other genera or deemed junior synonyms. Currently only S. validum and S. novomexicanum, named in 2011 from fossils found in New Mexico, remain. The validity of the latter species has also been debated.
Homalocephale is a genus of pachycephalosaurid dinosaur that lived during the Late Cretaceous period of what is now the Nemegt Formation, Mongolia, about 70 million years ago. The genus was described in 1974 by Halszka Osmólska and Teresa Maryańska, and consists of a single species, H. calathocercos. Though Homalocephale has been regarded as a synonym of Prenocephale, juvenile specimens of the latter indicate that they were distinct. Homalocephale was 1.8 m (5.9 ft) long and possibly a omnivore.
Prenocephale is a genus of small pachycephalosaurid dinosaur from the Late Cretaceous Nemegt Formation of Mongolia. It was similar in many ways to its close relative, Homalocephale.
Yaverlandia is a genus of maniraptoran dinosaur. Known from a partial fossil skull found in Lower Cretaceous strata of the Wessex Formation on the Isle of Wight. it was described as the earliest known member of the pachycephalosaurid family, but research by Darren Naish shows it to have actually been a theropod, seemingly a maniraptoran. The type species is Y. bitholus.
Tylocephale is a genus of pachycephalosaurid dinosaur, a group of dome-headed, herbivorous ornithischians, that lived during the Late Campanian stage of the Late Cretaceous in what is now Mongolia. It is known from a partial skull and associated mandible that were unearthed in 1971 by a Polish-Mongolian Expedition to the Barun Goyot Formation of the Gobi Desert. The specimen was described in 1974 by Polish paleontologists Teresa Maryańska and Halszka Osmólska as a new genus and species.
Colepiocephale is a genus of pachycephalosaurid dinosaur from Late Cretaceous deposits of Alberta, Canada. It was collected from the Foremost Formation. The type species, C. lambei, was originally described by Sternberg, and later renamed by Sullivan in 2003. C. lambei is a domed pachycephalosaur characterized principally by the lack of a lateral and posteriosquamosal shelf, a steeply down-turned parietal, and the presence of two incipient nodes tucked under the posterior margin of the parietosquamosal border.
Goyocephale is an extinct genus of pachycephalosaurian ornithischian that lived in Mongolia during the Late Cretaceous about 76 million years ago. It was first described in 1982 by Altangerel Perle, Teresa Maryańska and Halszka Osmólska for a disarticulated skeleton with most of a skull, part of the forelimb and hindlimb, some of the pelvic girdle, and some vertebrae. Perle et al. named the remains Goyocephale lattimorei, from the Mongolian гоё (goyo), meaning "decorated", and the Ancient Greek κεφαλή (kephale), for head. The species name honours Owen Lattimore.
Hanssuesia is a genus of pachycephalosaurid dinosaurs from the late Cretaceous period. It lived in what is now Alberta and Montana, and contains the single species Hanssuesia sternbergi.
Sphaerotholus is a genus of pachycephalosaurid dinosaur from the Upper Cretaceous of the western United States and Canada. To date, five species have been described: the type species, S. goodwini, from the Den-na-zin Member of the Kirtland Formation of San Juan County, New Mexico, USA; S. buchholtzae, from the Hell Creek Formation of western Carter County, Montana, USA and the Frenchman Formation of Saskatchewan, Canada; S. edmontonensis, from the Horseshoe Canyon Formation of Alberta, Canada; S. lyonsi, from the Dinosaur Park Formation (Campanian) of Alberta, Canada; and S. triregnum from the Hell Creek Formation of Garfield County, Montana, USA.
Alaskacephale is an extinct genus of pachycephalosaurid, a group of dome-headed, herbivorous ornithischian dinosaurs, that lived during the Campanian stage of the Late Cretaceous period in what is now northern Alaska. The genus is one of the few known Arctic dinosaurs and was found in the Prince Creek Formation, which preserves a menagerie of fossils. The only known specimen, a squamosal bone, was found in 1999 and later described in 2005. However, Alaskacephale was not formally named until the next year.
Texacephale is a pachycephalosaurid dinosaur from the Campanian stage of the Late Cretaceous. Its fossils come from the Aguja Formation of Big Bend National Park, in Texas, and were described in 2010 by Longrich, Sankey and Tanke. The generic name means Texas + "head" in reference to its place of discovery, and the specific name langstoni honors Wann Langston.
Foraminacephale is a genus of pachycephalosaurid dinosaur from Late Cretaceous deposits of Canada.
Amtocephale is a genus of pachycephalosaurid dinosaur from early Late Cretaceous deposits of southern Gobi Desert, Mongolia.
This timeline of pachycephalosaur research is a chronological listing of events in the history of paleontology focused on the pachycephalosaurs, a group of dome-skulled herbivorous marginocephalian dinosaurs. One of the first major events related to the history of pachycephalosaur research actually regards the discovery of an unrelated dinosaur called Troodon, reported from the western United States by Joseph Leidy in 1856. The type specimen of Troodon was simply an unusual tooth, but the close resemblance between Troodon teeth and pachycephalosaur teeth would cause taxonomic confusion for over a century. This was resolved by Phil Currie in 1987, who realized that Troodon belonged to a group of bird-like carnivores then known as saurornithoidids, but since renamed Troodontidae after Troodon itself. The first scientifically documented true pachycephalosaur remains were discovered in Early Cretaceous rocks from England and named Stenopelix not long after Troodon was named in America. Other notable early finds include the well-known pachycephalosaur Stegoceras validum.
Triopticus is a genus of archosauriform reptile from the Late Triassic of Texas, United States. It contains a single species, Triopticus primus, described in 2016 by Stocker et al. It has an unusually domed head reminiscent of the later pachycephalosaurian dinosaurs in an example of convergent evolution.
Sinocephale is a genus of pachycephalosaurid dinosaur that lived in Inner Mongolia, China during the Cretaceous period. The only species, Sinocephale bexelli, was originally named as a species of the genus Troodon in 1953, and later transferred to the genus Stegoceras. After decades of being considered dubious, it was re-evaluated in 2021 and recognized as a valid taxon, being given a unique generic name. The original holotype was lost, with modern research conducted using rediscovered plaster casts. Scant material makes for limited knowledge of its life appearance, but it is distinguished by an embayment on the back of the domed skull, which would give it a heart shape as seen from above. It is potentially the oldest known pachycephalosaurid and falls within the subset of the family called Pachycephalosaurinae, related to animals such as Pachycephalosaurus and Prenocephale. The geologic context of the species has been historically unclear but it is currently thought to originate in rocks belonging to the Ulansuhai Formation.