Ceratosaurs are members of a group of theropod dinosaurs defined as all theropods sharing a more recent common ancestry with Ceratosaurus than with birds. Ceratosaurs are believed to have diverged from the rest of Theropoda by the early Jurassic, however, the oldest confirmed discovered specimens date to the Late Jurassic. According to the majority of the latest research, Ceratosauria includes the Late Jurassic to Late Cretaceous theropods Ceratosaurus, Elaphrosaurus, and Abelisaurus, found primarily (though not exclusively) in the Southern Hemisphere. Originally, Ceratosauria included the above dinosaurs plus the Late Triassic to Early Jurassic Coelophysoidea and Dilophosauridae, implying a much earlier divergence of ceratosaurs from other theropods. However, most recent studies have shown that coelophysoids and dilophosaurids do not form a natural group with other ceratosaurs, and are excluded from this group.
Ceratosauria derives its names from the type species, Ceratosaurus nasicornis, described by O.C. Marsh in 1884. A moderately large predator from the Late Jurassic, Ceratosaurus nasicornis, was the first ceratosaur to be discovered. Ceratosaurs are generally moderately large in size, with some exceptions like the larger Carnotaurus and the significantly smaller noasaurs. The major defining characteristics of Ceratosauria include a robust skull with increased ornamentation or height and a shortening of the arms. Both of these characteristics are generally accentuated in later members of the group, such as the abelisaurs, whereas more basal species such as C. nasicornis appear more similar to other basal theropods. The highly fragmented nature of the ceratosaur fossil record, means that the characteristics, relationships, and early history of Ceratosauria remain mysterious and highly debated.
|Skeletal reconstruction of Ceratosaurus nasicornis|
Ceratosauria was first described by O.C. Marsh in the American Journal of Science in 1884. Writing about the newly discovered C. nasicornis, he noted the similarities between the firmly united metatarsals of C. nasicornis and those of Archaeopteryx. Since C. nasicornis was the only other dinosaur discovered at the time to share this trait, Marsh concluded that Ceratosauria must be placed very near Archaeopteryx and its related groups.
However, the idea of the Ceratosauria was soon contested by Marsh's rival, Edward Drinker Cope. Cope argued that the taxon was invalid. The idea of the Ceratosauria would regain some support more than thirty years later when Gilmore argued in its favor in 1920. Despite Gilmore’s support, few species were added to the group following World War I, and little emphasis was placed on it. In fact, the scientific communities most common interaction with Ceratosauria throughout much of the 20th century was the disputation of its existence, performed by the likes of Romer, Lapparent, Lavocat, Colbert, and Charig amongst others.
Ceratosauria’s fortune changed in 1986 when Jacques Gauthier, in an attempt to clarify the evolution of birds, grouped the majority of theropods into either Ceratosauria or Tetanurae. In Ceratosauria, he placed the ceratosaurs and coelophysoids. Gauthier’s paper brought Ceratosauria’s use back in vogue, and by the early 1990s, Abelisaridae had also been included under Ceratosauria. The triumvirate model of ceratosaurs, coelophysoids, and abelisaurids would go unchallenged until the early 2000s. Beginning at the turn of the millennium, a large number of paleontologists began excluding coelophysoids from Ceratosauria. This view is now widely held thanks to several similarities between Ceratosauria and Tetanurae not found in coelophysoids.
Most paleontologists have postulated that Ceratosauria split off from other theropods in the late Triassic or early Jurassic. Despite this, no ceratosaurs have been discovered prior the middle Jurassic, and even in the middle Jurassic, species are sparse. Many scientists, such as Carrano and Sampson, have postulated the lack of specimens is due to a poor fossil record, rather than an indictment on the abundance of ceratosaurs at the time. A similarly large gap of specimens exist in the lower Cretaceous, particularly for Abelisaridae. Several recent discoveries of possible ceratosaurs have begun reshaping the phylogeny of Ceratosauria by filling in some of these gaps. Specifically, this has allowed paleontologists to begin altering the sub-groups inside Ceratosauria, in an attempt to find their most basal members.
Currently, most paleontologists agree that Ceratosauria divides into two sub groups, Ceratosauridae and Abelisauroidea, with some variance as to which taxa are placed into basal polotomy. Abelisauroidea is further divided into the Abelisauridae and Noasauridae, with Abelisauridae, including Carnotaurinae. Recently, Rauhut and Carrano have placed Elaphrosaurinae inside Noasauridae while simultaneously moving the previous noasaurs into Noasaurinae. Into their new Noasauridae, they have uniquely included Deltadromeus and Limusaurus.
The oldest known ceratosaur currently described is Berberosaurus liassicus at 185 Ma. Some paleontologists believe that Berberosaurus liassicus, is the most basal ceratosaur currently discovered as well. The placement of B. liassicus has been a topic for debate, with some considering it an abelisaur, while others classifying as a basal polotomy.
A different conclusion was reached in a 2017 paper on Limusaurus ontogeny. The phylogenetics from the analysis with composite Limusaurus codings is shown below. Unlike other analyses, Noasauridae was placed more basal than Ceratosaurus, with the latter being within Abelisauridae by definition.
Some of the defining characteristics of Ceratosauria include an increase in height and ornamentation of the skull, as well as a shortening of the forelimbs. Likewise, ceratosaurs fused their ilium, ischium, and pubis together, as well as the astralagus and calcaneum. For less derived members of the group, such as C. nasicornis, traits such as raising of the skull and shortening of the forelimbs were not as noticeable. The skull of C. nasicornis was rather similar to the basal theropod mold, with a distinguishing nasal crest to go along with lacrimal crests similar to the contemporary Allosaurus. C. nasicornis had larger teeth than Allosaurus, and some paleontologists postulate that it would have had a difficult time attacking larger prey. Abelisaurids, however, carried many of these defining traits to their extremes. Most abelisaurids had largely shortened forelimbs, with Carnotaurus having shrunk them further than any large theropod. After analyzing the features of the newly discovered Rugops primus, Paul Sereno has postulated that many of these abelisaurid features may lend themselves to scavenging. Despite the huge reduction in size, no taxa in Ceratosauria ever lost a digit or any critical elements of the forelimb. Some joint variation has also been observed in Ceratosauria, and it has been postulated that they may have had better shoulder mobility than other large theropods.
Ceratosaurs appeared to have had a global population that diverged by the early Jurassic. However, they appear to have largely disappeared from Laurasia in the Cretaceous, with those few specimens that have been discovered having been possibly reintroduced from Gondwana. No confirmed specimens of ceratosaurs in North America during the Cretaceous have been found.
Abelisaurids in particular had great success in Gondwana, particularly in the Cretaceous. Some Gondwana specimens have recently been found and dated to Late Jurassic, and possibly even the Middle Jurassic, greatly extending the Abelisaurid timeline. Some paleontologists have postulated that a large desert may have kept abelisaurids locked in southern Gondwana until the late Jurassic. Whether correlation or causation, it has been largely observed that late Cretaceous ceratosaurs were found less in areas dominated by basal tetanurans (Africa) or coelosaurs (North America and Asia). The below phylogeny follows a simplified cladogram of Hendrickx et al. (2015), limited to Ceratosauria.
Ceratosaurs were theropods and thus carnivores, with one exception, Limusaurus inextricabilis was an herbivore with a toothless beak. Ceratosaurus has been argued to have eaten a large amount of fish and other aquatic creatures, though this has been disputed by many paleontologists. Tooth marks on large animals such as Allosaurus indicate that Ceratosaurus likely utilized scavenging often. The interesting jaws of the abelisaurids have drawn mixed dietary predictions. One study on Carnotaurus found that its bite, thanks to its shortened skull, was suited for hunting small prey, thanks to a quick, but relatively weak bite. On the other hand, other groups of paleontologists have found that the bite of Carnotaurus was relatively powerful, and more adept at hunting and wounding large prey. Others have postulated its skull was built for scavenging. The debate over the eating habits of ceratosaurs is quite active, particularly recently with the increase in abelisaur discoveries.
Abelisauroidea is a clade of theropod dinosaurs within the Ceratosauria. Some well-known dinosaurs of this group include the abelisaurids Abelisaurus, Carnotaurus, Thanos and Majungasaurus.
Abelisauroids flourished in the Southern hemisphere during the Cretaceous period, but their origins can be traced back to at least the Middle Jurassic, when they had a more global distribution (the earliest known abelisauroid remains come from Australian and South American deposits dated to about 170 million years ago). By the Cretaceous period, abelisauroids had apparently become extinct in Asia and North America, possibly due to competition from tyrannosauroids. However, advanced abelisauroids of the family Abelisauridae persisted in the southern continents until the Cretaceous–Paleogene extinction event 66 million years ago.Averostra
Averostra, or "bird snouts", is a clade that includes most theropod dinosaurs that have a promaxillary fenestra (fenestra promaxillaris), an extra opening in the front outer side of the maxilla, the bone that makes up the upper jaw. Two groups of averostrans, the Ceratosauria and the Orionides, survived into the Cretaceous period. When the Cretaceous–Paleogene extinction event occurred, ceratosaurians and two groups of orionideans within the clade Coelurosauria, the Tyrannosauroidea and Maniraptoriformes, were still extant. Only one subgroup of maniraptoriformes, Aves, survived the extinction event and persisted to the present day.Avetheropoda
Avetheropoda, or "bird theropods", is a clade that includes carnosaurians and coelurosaurs to the exclusion of other dinosaurs.Cerapoda
Cerapoda ("ceratopsians and ornithopods") is a clade of the dinosaur order Ornithischia.Ceratosauridae
Ceratosauridae is a family of theropod dinosaurs belonging to the infraorder Ceratosauria. The family's type genus, Ceratosaurus, was first found in Jurassic rocks from North America. Ceratosauridae is made up of the genera Ceratosaurus, found in North America, Tanzania, and Portugal, and Genyodectes, from the Early Cretaceous of Argentina. Unnamed probable ceratosaurids are known from limited material in the Middle Jurassic of Madagascar, the Late Jurassic of Switzerland, and the Late Jurassic or possibly Early Cretaceous of Uruguay.Ceratosaurus
Ceratosaurus (from Greek κέρας/κέρατος, keras/keratos meaning "horn" and σαῦρος/sauros meaning "lizard") was a carnivorous theropod dinosaur in the Late Jurassic period (Kimmeridgian to Tithonian). This genus was first described in 1884 by American paleontologist Othniel Charles Marsh based on a nearly complete skeleton discovered in Garden Park, Colorado, in rocks belonging to the Morrison Formation. The type species is Ceratosaurus nasicornis.
The Garden Park specimen remains the most complete skeleton known from the genus, and only a handful of additional specimens have been described since. Two additional species, Ceratosaurus dentisulcatus and Ceratosaurus magnicornis, have been described in 2000 from two fragmentary skeletons from the Cleveland-Lloyd Quarry of Utah and from the vicinity of Fruita, Colorado. The validity of these additional species has been questioned, however, and all three skeletons possibly represent different growth stages of the same species. In 1999, the discovery of the first juvenile specimen was reported. Since 2000, a partial specimen was excavated and described from the Lourinhã Formation of Portugal, providing evidence for the presence of the genus outside of North America. Fragmentary remains have also been reported from Tanzania, Uruguay, and Switzerland, although their assignment to Ceratosaurus is currently not accepted by most paleontologists.
Ceratosaurus was a medium-sized theropod. The original specimen is estimated to be 5.3 m (17 ft) or 5.69 m (18.7 ft) long, while the specimen described as C. dentisulcatus was larger, at around 7 m (23 ft) long. Ceratosaurus was characterized by deep jaws that supported proportionally very long, blade-like teeth, a prominent, ridge-like horn on the midline of the snout, and a pair of horns over the eyes. The forelimbs were very short, but remained fully functional; the hand had four fingers. The tail was deep from top to bottom. A row of small osteoderms (skin bones) was present down the middle of the neck, back, and tail. Additional osteoderms were present at unknown positions on the animal's body.
Ceratosaurus gives its name to the Ceratosauria, a clade of theropod dinosaurs that diverged early from the evolutionary lineage leading to modern birds. Within the Ceratosauria, some paleontologists proposed it to be most closely related to Genyodectes from Argentina, which shares the strongly elongated teeth. The geologically older genus Proceratosaurus from England, although originally described as a presumed antecedent of Ceratosaurus, was later found to be unrelated. Ceratosaurus shared its habitat with other large theropod genera including Torvosaurus and Allosaurus, and it has been suggested that these theropods occupied different ecological niches to reduce competition. Ceratosaurus may have preyed upon plant-eating dinosaurs, although some paleontologists suggested that it hunted aquatic prey such as fish. The nasal horn was probably not used as a weapon as was originally suggested by Marsh, but more likely was used solely for display.Coelophysoidea
Coelophysoidea were common dinosaurs of the Late Triassic and Early Jurassic periods. They were widespread geographically, probably living on all continents. Coelophysoids were all slender, carnivorous forms with a superficial similarity to the coelurosaurs, with which they were formerly classified, and some species had delicate cranial crests. Sizes range from about 1 to 6 m in length. It is unknown what kind of external covering coelophysoids had, and various artists have portrayed them as either scaly or feathered. Some species may have lived in packs, as inferred from sites where numerous individuals have been found together.
Examples of coelophysoids include Coelophysis, Procompsognathus and Liliensternus. Most dinosaurs formerly referred to as being in the dubious taxon "Podokesauridae" are now classified as coelophysoids.Dinosaur vision
Dinosaur vision was, in general, better than the vision of most other reptiles, although vision varied between dinosaur species. Coelurosaurs, for example, had good stereoscopic or binocular vision, whereas large carnosaurs had poor binocular vision, comparable to that of modern alligators.Dinosauriformes
Dinosauriformes is a clade of archosaurian reptiles that include the dinosaurs and their most immediate relatives. All dinosauriformes are distinguished by several features, such as shortened forelimbs and a partially to fully perforated acetabulum, the hole in the hip socket traditionally used to define dinosaurs. The oldest known member is Asilisaurus, dating to about 245 million years ago in the Anisian age of the middle Triassic period.Echinodon
Echinodon (pronounced eh-KY-no-don) meaning "hedgehog tooth" in reference to the spines on its teeth (Ancient Greek: εχινος, romanized: echinos, lit. 'hedgehog', + ὀδών, odṓn, 'tooth'), occasionally known as Saurechinodon, is a genus of small European dinosaur of the early Cretaceous Period (Berriasian age), 140 million years ago.Gojirasaurus
Gojirasaurus (meaning "Godzilla Lizard") is a dubious genus of coelophysoid theropod dinosaur named after the giant monster movie character Gojira (the Japanese name for the monster Godzilla).Jingshanosaurus
Jingshanosaurus (meaning "Jingshan lizard") is a genus of sauropodomorph dinosaurs from the early Jurassic period.Lophostropheus
Lophostropheus (pron.:" LOAF-oh-STRO-fee-us") is an extinct genus of coelophysoid theropod dinosaur that lived approximately 200 million years ago during the boundary between the Late Triassic Period and the Early Jurassic Period, in what is now Normandy, France. Lophostropheus is one of the few dinosaurs that may have survived the Triassic–Jurassic extinction event.
Lophostropheus was a small to medium-sized, moderately-built, ground-dwelling, bipedal carnivore, that could grow up to 3 m (9.8 ft) long. Over the years it had been incorrectly classified as Halticosaurus and Liliensternus, but was later recognized as a new genus and was reassigned to Lophostropheus in 2007.Neotheropoda
Neotheropoda (meaning "new theropods") is a clade that includes coelophysoids and more advanced theropod dinosaurs, and the only group of theropods who survived the Triassic–Jurassic extinction event. Yet all of the neotheropods became extinct during the early Jurassic period except for Averostra.Orionides
Orionides is a clade of tetanuran theropod dinosaurs from the Middle Jurassic to the Present. The clade includes most theropod dinosaurs, including birds.Orodrominae
Orodrominae is a subfamily of parksosaurid dinosaurs from the Cretaceous of North America and Asia.Tetanurae
Tetanurae (/ˌtɛtəˈnjuːriː/ or "stiff tails") is a clade that includes most theropod dinosaurs, including megalosauroids, allosauroids, tyrannosauroids, ornithomimosaurs, maniraptorans, and birds. Tetanurans are defined as all theropods more closely related to modern birds than to Ceratosaurus and contain the majority of predatory dinosaur diversity. Tetanurae likely diverged from its sister group, Ceratosauria, during the late Triassic. Tetanurae first appeared in the fossil record by the Early Jurassic about 190 mya and by the Middle Jurassic had become globally distributed.The group was named by Jacques Gauthier in 1986 and originally had two main subgroups: Carnosauria and Coelurosauria, the clade containing birds and related dinosaurs such as compsognathids, tyrannosaurids, ornithomimosaurs, and maniraptorans. The original Carnosauria was a polyphyletic group including any large carnivorous theropod. Many of Gauthier's carnosaurs, such as tyrannosaurids, have since been re-classified as coelurosaurs or primitive tetanurans. Carnosauria has been reclassified as a group containing allosaurids that split from the Coelurosauria at the Neotetanurae/Avetheropoda node. Members of Spinosauroidea are believed to represent basal tetanurans.Tetanuran evolution was characterized by parallel diversification of multiple lineages, repeatedly attaining large body size and similar locomotor morphology. Cryolophosaurus has been claimed as the first true member of the group, but subsequent studies have disagreed on whether it is a dilophosaurid or tetanuran. Arcucci and Coria (2003) classified Zupaysaurus as an early tetanuran, but it was later placed as a sister taxon to the clade containing dilophosaurids, ceratosaurs, and tetanurans.Shared tetanuran features include a ribcage indicating a sophisticated air-sac-ventilated lung system similar to that in modern birds. This character would have been accompanied by an advanced circulatory system. Other tetanuran characterizing features include the absence of the fourth digit of the hand, placement of the maxillary teeth anterior to the orbit, a strap-like scapula, maxillary fenestrae, and stiffened tails. During the Late Jurassic and Early Cretaceous, large spinosaurids and allosaurs flourished but possibly died out in the northern hemisphere before the end of the Cretaceous, and were replaced as apex predators by tyrannosauroid coelurosaurs. At least in South America, carcharodontosaurid allosaurs persisted until the end of the Mesozoic Era, and died out at the same time the non-avian coelurosaurs.Theropoda
Theropoda ( or , from Greek θηρίον "wild beast" and πούς, ποδός "foot") or theropods () are a dinosaur suborder that is characterized by hollow bones and three-toed limbs. They are generally classed as a group of saurischian dinosaurs, although a 2017 paper has instead placed them in the proposed clade Ornithoscelida as the closest relatives of the Ornithischia. Theropods were ancestrally carnivorous, although a number of theropod groups evolved to become herbivores, omnivores, piscivores, and insectivores. Theropods first appeared during the Carnian age of the late Triassic period 231.4 million years ago (Ma) and included the sole large terrestrial carnivores from the Early Jurassic until at least the close of the Cretaceous, about 66 Ma. In the Jurassic, birds evolved from small specialized coelurosaurian theropods, and are today represented by about 10,500 living species.Timeline of ceratosaur research
This timeline of ceratosaur research is a chronological listing of events in the history of paleontology focused on the ceratosaurs, a group of relatively primitive, often horned, predatory theropod dinosaurs that became the apex predators of the southern hemisphere during the Late Cretaceous. The nature and taxonomic composition of the Ceratosauria has been controversial since the group was first distinguished in the late 19th century. In 1884 Othniel Charles Marsh described the new genus and species Ceratosaurus nasicornis from the Late Jurassic Morrison Formation of the western United States. He felt that it belonged in a new family that he called the Ceratosauridae. He created the new taxon Ceratosauria to include both the Ceratosauridae and the ostrich-like ornithomimids. The idea of the Ceratosauria was soon contested, however. Later that same decade both Lydekker and Marsh's hated rival Edward Drinker Cope argued that the taxon was invalid.The idea of the Ceratosauria would regain some support more than thirty years later when Gilmore argued in its favor in 1920. Nevertheless, the validity of Ceratosauria was disputed throughout much of the 20th century by researchers like Romer, Lapparent, Lavocat, Colbert, and Charig. However, in 1986, more than a century after Marsh first coined the name, Jacques Gauthier revived the idea. Three years later, Rowe published a new definition of Ceratosauria, all taxa more closely related to Ceratosaurus than to birds, based on Gauthier's use of the term. This modern use of the term was thought to include the many theropods discovered since the 1880s known as coelophysoids. Ceratosaurus itself had loose joints between bones in the skull whose interpretation has been controversial. Paleontologist Robert T. Bakker has interpreted this condition as an adaptation to swallow prey larger than it would otherwise be able to fit through its jaws.Since the 1980s, major developments in ceratosaur taxonomy have centered on the discovery of the Abelisauridae, a new family of large ceratosaurs that were among the dominant predators of the southern hemisphere during the Cretaceous. One of the most notable of these was Carnotaurus, an unusual horned theropod with a short face. More recent noteworthy non-abelisaur ceratosaur discoveries include the protruding-toothed noasaurid Masiakasaurus knopfleri, named after the lead guitarist from Dire Straits.