Brachyrostrans Temporal range: Late Cretaceous, | |
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Mounted cast of a Carnotaurus sastrei skeleton, Chlupáč Museum, Prague | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | Saurischia |
Clade: | Theropoda |
Family: | † Abelisauridae |
Clade: | † Brachyrostra Canale et al., 2008 |
Subgroups | |
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Brachyrostra (meaning "short snouts") is a clade within the theropod dinosaur family Abelisauridae. It includes the famous genera Carnotaurus , Abelisaurus , Aucasaurus as well as their close relatives from the Cretaceous Period of Argentina and Brazil plus Caletodraco from France. [1] The group was first proposed in an analysis conducted by Juan Canale and colleagues in 2008. They found that all South American abelisaurids described up to that point grouped together as a sub-clade of Abelisauridae, which they named based on the relatively unusual shape of their skulls (in comparison with other theropods). They defined the clade Brachyrostra as "all the abelisaurids more closely related to Carnotaurus sastrei than to Majungasaurus crenatissimus." [2]
Brachyrostrans were relatively lightly built compared to other large theropods, ranging in size from 6.1–7.8 m (20–26 ft) [3] and 1400–2000 kg (1.6–2.3 short tons) in weight. [3] [4] They are considered the most derived abelisaurids, with traits like very short, narrow skulls and extremely reduced forearms, even more so than other abelisaurids. [5] [6] Many brachyrostrans had horns or rugosities on the frontal and nasal bones, which have been interpreted as bearing cornified structures or dermal armor. [7]
Studies of the skull anatomy of the most well-known species, Carnotaurus sastrei , lead to debate over what type of prey these animals hunted. Studies by Mazzetta et al. in 1998, 2004, and 2009 suggest that the jaw structure in Carnotaurus was built for swift, rather than strong, bites, with adaptations for mandibular kinesis to assist in swallowing small prey items whole. [8] Surprisingly, it exhibits a form of paracraniokinesis in which the dentary bone articulates against the surangular bone, further jointing the lower jaw and hypothetically allowing this animal a wider array of hunting strategies. [9]
However, in 1998 and 2009, Robert Bakker and Francois Therrien and colleagues contested this finding, stating that Carnotaurus had the exact same skull adaptations (short snout, small teeth, and a fortified occiput) as the Jurassic theropod Allosaurus , which presumably preyed upon large animals by gradual jaw slashing. [10]
Mazzetta et al. 1998–1999 and Phil Currie et al. 2011 found Carnotaurus to be a swift-running predator with semicursorial adaptations such as femoral resistance against bending moments [11] and a hypertrophied caudofemoralis muscle, the primary locomotory muscle in theropods which was located in the tail and pulled the femur backwards. [12] This enlarged caudofemoralis, giving them a speed estimate of 48–56 km/h (30–35 mph), allowed brachyrostrans to be one of the fastest-running large theropod groups yet known. [13] [12]
Within Brachyrostra, there is a slightly more restrictive clade, called Furileusauria ("stiff back lizards"). [14] They represent some of the larger brachyrostrans, with an average length of 7.1 ± 2.1 m (23.3 ± 6.9 ft). [15] The taxon is a stem-based clade and is defined as the most inclusive clade containing Carnotaurus sastrei but not Ilokelesia aguadagrandensis , Skorpiovenator bustingorryi, or Majungasaurus crenatissimus . [14]
Synapomorphies of Furileusauria recovered by Filippi and colleagues include: the presence of a tip in the middle area of the posterior surface of the ventral process of the postorbital, the presence of a knob followed by a deep notch in the postorbital-squamosal contact, the absence of fenestra between the frontal and lacrimal bones, an anterior projection of the distal end of the cervical epiphophyses, a posterior margin of the postzygapophyses which is level with the intervertebral articulation in dorsal vertebrae, a crescent-shaped morphology of the distal tip of the transverse processes in anterior and middle caudal vertebrae, transverse processes of anterior caudal vertebrae that is distally expanded and projected anteriorly, a convex external margin of the transverse processes in anterior caudal vertebrae, and a downturned process on the cnemial crest of the tibia. [14]
A more restrictive clade within Furileusauria is the tribe Carnotaurini. This group is a node-based clade and was first proposed by paleontologists Rodolfo Coria, Luis Chiappe, and Lowell Dingus in 2002, being defined as a clade containing " Carnotaurus sastrei , Aucasaurus garridoi , their most recent common ancestor, and all of its descendants." The tribe Carnotaurini was named in 2002 by Rodolfo Coria et al. in 2002 after their discovery of Aucasaurus garridoi . [6] Their morphological definition of it is by several synapomorphies of the clade, with two ambiguous ones: "the presence of hyposphene–hypantrum articulations in the proximal and middle sections of the caudal series, and cranial processes in the epipophyses of the cervical vertebrae." They defined more ambiguous synapomorphies due to the homologous materials not yet found in all other abelisaurids being: "a very broad coracoid (coracoid maximum width three times the distance across the scapular glenoid area), a humerus with a large and hemispherical head, an extremely short ulna and radius (ulna to humerus ratio 1:3 or less), and frontal prominences (swells or horns) that are located laterally on the skull roof." [6]
In their description of the abelisaurid Llukalkan , Federico Gianechini and colleagues performed a phylogenetic analysis to test the affinities of the new taxon. The simplified strict consensus tree of the analysis is shown below. [16] Similar results have been recovered by other analyses including Coria and colleagues (2002), [6] Canale and colleagues (2008), [2] and Cerroni and colleagues (2020). [17]
Abelisauridae |
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Brachyrostrans were initially known exclusively from South America. Members of the group have been unearthed from the Anacleto Formation, the Bajo de la Carpa Formation, the Candeleros Formation, the Huincul Formation, and possibly the Sir Fernandez field of the Allen Formation to the southeast. [18] [5] [19] A single named taxon, Pycnonemosaurus is also known from the Cachoeira do Bom Jardim Formation in Mato Grosso, Brazil. [20] The description of the French taxon Caletodraco , was the first definitive evidence of the clade from outside South America. The Albian taxon Genusaurus, which also hails from France, may also represent a European member of this clade. If it is truly a brachyrostran, it would represent the oldest member of the clade. [21] The enigmatic taxon Dahalokely , from Madagascar may also belong to Brachyrostra, although this remains uncertain.
Giganotosaurus is a genus of theropod dinosaur that lived in what is now Argentina, during the early Cenomanian age of the Late Cretaceous period, approximately 99.6 to 95 million years ago. The holotype specimen was discovered in the Candeleros Formation of Patagonia in 1993 and is almost 70% complete. The animal was named Giganotosaurus carolinii in 1995; the genus name translates to "giant southern lizard", and the specific name honors the discoverer, Ruben Carolini. A dentary bone, a tooth, and some tracks, discovered before the holotype, were later assigned to this animal. The genus attracted much interest and became part of a scientific debate about the maximum sizes of theropod dinosaurs.
Carnotaurus is a genus of theropod dinosaur that lived in South America during the Late Cretaceous period, probably sometime between 72 and 69 million years ago. The only species is Carnotaurus sastrei. Known from a single well-preserved skeleton, it is one of the best-understood theropods from the Southern Hemisphere. The skeleton, found in 1984, was uncovered in the Chubut Province of Argentina from rocks of the La Colonia Formation. Carnotaurus is a derived member of the Abelisauridae, a group of large theropods that occupied the large predatorial niche in the southern landmasses of Gondwana during the late Cretaceous. Within the Abelisauridae, the genus is often considered a member of the Brachyrostra, a clade of short-snouted forms restricted to South America.
Ceratosaurs are members of the clade Ceratosauria, a group of dinosaurs defined as all theropods sharing a more recent common ancestor with Ceratosaurus than with birds. The oldest known ceratosaur, Saltriovenator, dates to the earliest part of the Jurassic, around 199 million years ago. Ceratosauria includes three major clades: Ceratosauridae, Noasauridae, and Abelisauridae, found primarily 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.
Abelisaurus is a genus of predatory abelisaurid theropod dinosaur alive during the Late Cretaceous Period (Campanian) of what is now South America. It was a bipedal carnivore that probably reached about 7.4 metres in length, although this is uncertain as it is known from only one partial skull.
Xenotarsosaurus is a genus of abelisaurid theropod dinosaur that lived during the Late Cretaceous of Argentina.
Abelisauridae is a family of ceratosaurian theropod dinosaurs. Abelisaurids thrived during the Cretaceous period, on the ancient southern supercontinent of Gondwana, and today their fossil remains are found on the modern continents of Africa and South America, as well as on the Indian subcontinent and the island of Madagascar. Isolated teeth were found in the Late Jurassic of Portugal, and the Late Cretaceous genera Tarascosaurus, Arcovenator and Caletodraco have been described in France. Abelisaurids possibly first appeared during the Jurassic period based on fossil records, and some genera survived until the end of the Mesozoic era, around 66 million years ago.
Aucasaurus is a genus of medium-sized abelisaurid theropod dinosaur from Argentina that lived during the Late Cretaceous of the Anacleto Formation. It was smaller than the related Carnotaurus, although more derived in some ways, such as its extremely reduced arms and almost total lack of fingers. The type skeleton is complete to the thirteenth caudal vertebra, and so is relatively well understood, and is the most complete abelisaurid yet described. However, the skull is damaged, causing some paleontologists to speculate that it was involved in a fight prior to death.
Ilokelesia is an extinct genus of abelisaurid theropod, preserved in the layers of the earliest Late Cretaceous of the Huincul Formation, Neuquén Group, located near Plaza Huincul, Neuquén Province, Argentina. The specimen, consisting of very fragmentary elements of the skull and the axial and appendicular skeleton, was described by Rodolfo Coria and Leonardo Salgado in late 1998.
Quilmesaurus is a genus of carnivorous abelisaurid theropod dinosaur from the Patagonian Upper Cretaceous of Argentina. It was a member of Abelisauridae, closely related to genera such as Carnotaurus. The only known remains of this genus are leg bones which share certain similarities to a variety of abelisaurids. However, these bones lack unique features, which may render Quilmesaurus a nomen vanum.
Ekrixinatosaurus is a genus of abelisaurid theropod which lived approximately 100 to 97 million years ago during the Late Cretaceous period. Its fossils have been found in Argentina. Only one species is currently recognized, Ekrixinatosaurus novasi, from which the specific name honors of Dr. Fernando Novas for his contributions to the study of abelisaurid theropods, while the genus name refers to the dynamiting of the holotype specimen. It was a large abelisaur, measuring between 6.5 and 8 m in length and weighing 800 kg (1,800 lb).
Genusaurus is a genus of abelisauroid theropod from the Early Cretaceous. Its fossils were found in France. Genusaurus is believed to have lived during the Albian stage, around 112-100 million years ago.
Pycnonemosaurus is a genus of carnivorous theropod dinosaur that belonged to the family Abelisauridae. It was found in the Upper Cretaceous red conglomerate sandstones of the Cachoeira do Bom Jardim Formation, Mato Grosso, Brazil, and it lived during the late Campanian to early Maastrichtian stage of the Late Cretaceous.
Skorpiovenator is a genus of abelisaurid theropod dinosaur from the Late Cretaceous Huincul Formation of Argentina. It is one of the most complete and informative abelisaurids yet known, described from a nearly complete and articulated skeleton.
Bonapartenykus is a monospecific genus of alvarezsauroid dinosaur from Argentina that lived during the Late Cretaceous (Campanian-Maastrichtian) in what is now the upper Allen Formation of the Río Negro Province. The type and only species, Bonapartenykus ultimus, is known from a nearly articulated but partial skeleton that was found in close association to two incomplete eggs and several clusters of eggshells belonging to the oogenus Arriagadoolithus. Bonapartenykus was named in 2012 by Federico L. Agnolin, Jaime E. Powell, Fernando E. Novas and Martin Kundrát. Bonapartenykus has an estimated length of 2.5 m (8.2 ft) and weight of 72 kg (159 lb), making it the largest member of the clade Alvarezsauroidea.
Arcovenator is an extinct genus of abelisaurid theropod dinosaurs hailing from the Late Cretaceous of France and possibly Spain. The type and only described species is Arcovenator escotae.
Majungasaurinae is a subfamily of large carnivorous theropods from the Upper Cretaceous, found in Madagascar, India, and France. It is a subgroup within the theropod family Abelisauridae, a Gondwanan clade known for their thick and often horned skulls and vestigial arms. The two subfamilies of Abelisauridae are Carnotaurinae, best known from the South American Carnotaurus, and Majungasaurinae, consisting of Madagascar’s Majungasaurus and its closest relatives. Their ancestors emerged in the Middle Jurassic, and the clade lasted until the Upper Cretaceous.
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.
Tralkasaurus is a genus of abelisaurid dinosaur from the Huincul Formation from Río Negro Province in Argentina. The type and only species is Tralkasaurus cuyi, named in 2020 by Mauricio Cerroni and colleagues based on an incomplete skeleton. A medium-sized abelisaurid, Tralkasaurus exhibits a conflicting blend of characteristics found among the early-diverging abelisauroids with others that characterize the highly specialized clade Brachyrostra, and thus its position within the clade is poorly-resolved.
Teratopodus is an ichnogenus of titanosaurian sauropod footprint. It includes a single species, T. malarguensis, known from prints found in the Late Cretaceous Anacleto Formation of Argentina. The Teratopodus tracks represent some of the best sauropod pes tracks currently known from South America.
Elemgasem is an extinct genus of brachyrostran abelisaurid from the Late Cretaceous Portezuelo Formation of Patagonia, Argentina. The genus contains a single species, Elemgasem nubilis. The cladistic position of Elemgasem within Brachyrostra is uncertain, given that phylogenetic analyses recover it as either a sister taxon to Furileusauria or in several positions within this clade.
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