~{{fossil range|183|174}} Uncertain allocation of the layers"},"image":{"wt":"Berberosaurus life restoration 2019.jpg"},"image_caption":{"wt":"Life [[paleoart|restoration]] and size comparison"},"genus":{"wt":"Berberosaurus"},"species":{"wt":"liassicus"},"authority":{"wt":"Allain ''et al.'', 2007"},"display_parents":{"wt":"3"},"synonyms":{"wt":"*''Berbesaurus'' Torices ''et al.'', 2012 (''[[lapsus calami]]''){{Cite journal | last1 = Torices | first1 = A. | doi = 10.4202/app.2012.0121 | title = Theropod dinosaurs from the Upper Cretaceous of the South Pyrenees Basin of Spain | journal = Acta Palaeontologica Polonica | year = 2013 | url = http://www.app.pan.pl/article/item/article/item/app20120121.html?pdf=39 | doi-access = free}}"}},"i":0}}]}" id="mwBA">
Berberosaurus Temporal range: Toarcian ~ | |
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Life restoration and size comparison | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | Saurischia |
Clade: | Theropoda |
Clade: | Averostra |
Clade: | † Ceratosauria |
Genus: | † Berberosaurus |
Species: | †B. liassicus |
Binomial name | |
†Berberosaurus liassicus Allain et al., 2007 | |
Synonyms | |
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Berberosaurus (meaning "Berber lizard", in reference to the Berbers of Morocco) is a genus of neotheropod dinosaur, possibly a ceratosaur, from the Toarcian-age (Lower Jurassic) "Toundoute Continental Series" (Azilal Formation) found in the Central High Atlas of Toundoute, Ouarzazate, Morocco. The type species of the genus Berberosaurus is B. liassicus, in reference to the Lias epoch. Berberosaurus might be the oldest known ceratosaur, and is based on partial postcranial remains. This genus represents the oldest formally identified theropod from the North of Africa, as well one of the few from the region in the Early Jurassic. [2]
The remains of Berberosaurus were discovered during a series of expeditions to the High Atlas beginning in the early 2000s, when in 6 years, where made to dig in the local redbeds. It is based on an associated partial postcranial skeleton of a subadult individual cataloged in the Muséum d'Histoire Naturelle de Marrakech; bones from this skeleton include MHNM-Pt9 a neck vertebra;MHNM-Pt23, an anterior part of the sacrum; MHNM-Pt22, a metacarpal; MHNM-Pt19, a femur; MHNM-Pt21, proximal end of the left tibia; MHNM-Pt20, both fibulae; MHNM-Pt16, part of another femur, has been assigned to the genus as well. Its remains were found alongside those of Tazoudasaurus and an indeterminate large-bodied theropod within bone beds in mudflow deposits. Later tectonic activity has affected the bones. [2] Recent papers have quoted that new material of this genus was recovered on the same area, namely the axis, a postorbital, the cranium and teeth, that are currently being studied. [3] Berberosaurus is characterized by the following features: the cervical vertebra is highly pneumatic, with short cervical centra and holes in the neural arch, with low and short neural spine, unlike Elaphrosaurus and Ceratosaurus . It has anteroposteriorly short centra and neural spine; the sacral series markedly arched; the central sacrum narrow transversely; the metacarpal with a very grooved proximal end; anterior femoral trochanter reaches proximally to the midpoint of the femoral head, unlike Ceratosaurus ; large femoral trochanteric platform; tibia with subtriangular distal profile; presence of an oblique ridge that covers proximally the medial sulcus of the fibula. [2]
Ronan Allain and colleagues, who described Berberosaurus, performed a phylogenetic analysis and found their new genus to be the most basal known abelisauroid, more derived than Elaphrosaurus , Ceratosaurus , and Spinostropheus , but less so than Xenotarsosaurus and abelisaurians. Berberosaurus is distinguished from other theropods by anatomical details found in its vertebrae, metacarpals, and hindlimb bones. Its assignment as an abelisauroid pushes back the record of this group and shows that it had diversified by the Early Jurassic. [2] However, Carrano and Sampson (2008) found it to be a basal ceratosaur outside Neoceratosauria instead. [4] Subsequently, the analysis of Xu et al. (2009) recovered it as a dilophosaurid in unresolved polytomy with Dilophosaurus wetherilli , "Dilophosaurus" sinensis, Dracovenator and Cryolophosaurus . [5] The phylogenetic analysis performed by Ezcurra, Agnolin and Novas (2010) recovered Berberosaurus in unresolved polytomy with Ceratosaurus and Abelisauroidea, [6] while the 2018 description of the basal ceratosaur Saltriovenator places Berberosaurus as the sister taxon to that genus. [7] Same year, a paper found Berberosaurus in different positions: considered as a basal ceratosaurian, a neoceratosaurian or a basal abelisauroid. [8] The paper which described Berberosaurus was the last one to find a monophyletic Ceratosauria that contained both Coelophysoids and Neoceratosaurs, most papers have since excluded Coelophysoidea from Ceratosauria and instead have favored the idea that Tetanurae and Ceratosauria are both part of a monophyletic group called Averostra, a classification which was already gaining popularity at around that time. [9]
The classification of Berberosaurus based on Delcourt et al. (2018):
Ceratosauroidea |
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Berberosaurus, like other ceratosaurians, was a bipedal carnivore. [10] It was of moderate size; its estimated femur length of 50.5 centimetres (19.9 inches) [2] is comparable to femur lengths given by Gregory S. Paul for animals like Elaphrosaurus (52.9 cm (20.8 in)) [11] and Dilophosaurus (55.0 cm (21.7 in)). [12] Its remains were found with those of the early sauropod Tazoudasaurus . It has been estimated to be approximately 5 m (16 ft) long and 220–300 kg (490–660 lb) in body mass. [13] [14]
The "Toundoute Continental Series", unlike other members of the Azilal Formation, due to the presence of volcanic material of coeval age. The Azilal Formation recovers a Terrestrial progradation that happened in the Central High Atlas Basin towards the Toarcian, where the older Pliensbachian Carbonate Platform retreated to the east. The lithology of this unit at Tundoute is divided in 5 units from D to H, (A-C represent the units of the underlying marine dolomite, C representing a transition to a terrestrial environment). [15] These layers have been referred to a Pliensbachian-Toarcian age, based on outdated data, as the underliying Carbonates where quoted to be a continuation with less thickess of the ones found in Todhra, that reach the Late Pliensbachian, being eveidence of a latter progradation than the one seen in the N, for example at Béni-Mellal. [16] The section including the bones of Berberosaurus was likely deposited on a channel/floodplain type fluvial system, with sand-filled channels abundant in plant roots (mostly located in fine limestone, probably from the channel margins), developed in the near E-W direction of transit. These layers also recover the presence of thick (up to 6 m) gypsium facies, which suggest the presence of a local Chott, indicating a succession of humid and dry seasons. [15] Based on mesofossils, local vegetation was apparently dominated by ferns, cycads and conifers. [15] Apart from Berberosaurus, Tazoudasaurus and a large-bodied theropod of uncertain affinities, a possible Coelophysidae with juvenile & adult specimens and a small sauropod have been recovered from Acforcid, E of Demnate, as well Gravisaurian ( Tazoudasaurus? [17] ) remains at the E of Azilal village, Medium-Sized Sauropod remains at Mizaguène Hill (SW Azilal) and indeterminate Dinosaur remains from other locations around Azilal and Demnate. [18]
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.
Masiakasaurus is a genus of small predatory noasaurid theropod dinosaurs from the Late Cretaceous of Madagascar. In Malagasy, masiaka means "vicious"; thus, the genus name means "vicious lizard". The type species, Masiakasaurus knopfleri, was named after the musician Mark Knopfler, whose music inspired the expedition crew. It was named in 2001 by Scott D. Sampson, Matthew Carrano, and Catherine A. Forster. Unlike most theropods, the front teeth of M. knopfleri projected forward instead of straight down. This unique dentition suggests that they had a specialized diet, perhaps including fish and other small prey. Other bones of the skeleton indicate that Masiakasaurus were bipedal, with much shorter forelimbs than hindlimbs. M. knopfleri was a small theropod, reaching 1.8–2.1 m (5.9–6.9 ft) long and weighing 20 kg (44 lb).
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 and Arcovenator have been described in France. Abelisaurids first appear in the fossil record of the early middle Jurassic period, and at least three genera survived until the end of the Mesozoic era 66 million years ago.
Deltadromeus is a genus of theropod dinosaur from the Aoufous Formation of Morocco.
Elaphrosaurus is a genus of ceratosaurian theropod dinosaur that lived approximately 154 to 150 million years ago during the Late Jurassic Period in what is now Tanzania in Africa. Elaphrosaurus was a medium-sized but lightly built member of the group that could grow up to 6.2 m (20 ft) long. Morphologically, this dinosaur is significant in two ways. Firstly, it has a relatively long body but is very shallow-chested for a theropod of its size. Secondly, it has very short hindlimbs in comparison with its body. Phylogenetic analyses indicate that this genus is likely a ceratosaur. Earlier suggestions that it is a late surviving coelophysoid have been examined but generally dismissed. Elaphrosaurus is currently believed to be a very close relative of Limusaurus, an unusual beaked ceratosaurian which may have been either herbivorous or omnivorous.
Tazoudasaurus is a genus of gravisaurian, probably a vulcanodontid sauropod dinosaurs hailing from the late Early Jurassic (Toarcian), that was recovered in the "Toundoute Continental Series" located in the High Atlas Mountains of Morocco in North Africa. Along with Patagosaurus, Volkheimeria, Bagualia and Perijasaurus represents one of the few sauropods named from this stage on Gondwana, as well the only one from Africa.
Sarcosaurus is a genus of basal neotheropod dinosaur, roughly 3.5 metres (11 ft) long. It lived in what is now England and maybe Ireland and Scotland during the Hettangian-Sinemurian stages of the Early Jurassic, about 199-196 million years ago. Sarcosaurus is one of the earliest known Jurassic theropods, and one of only a handful of theropod genera from this time period. Along with Dracoraptor hanigani it is one of the two described neotheropods from the lowermost Jurassic of the United Kingdom.
Genusaurus is a genus of abelisaurid 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.
Noasaurus is a genus of ceratosaurian theropod dinosaur genus from the late Campanian-Maastrichtian of Argentina. The type and only species is N. leali.
Noasauridae is an extinct family of theropod dinosaurs belonging to the group Ceratosauria. They were closely related to the short-armed abelisaurids, although most noasaurids had much more traditional body types generally similar to other theropods. Their heads, on the other hand, had unusual adaptations depending on the subfamily. 'Traditional' noasaurids, sometimes grouped in the subfamily Noasaurinae, had sharp teeth which splayed outwards from a downturned lower jaw.
Limusaurus is a genus of theropod dinosaur that lived in what is now China during the Late Jurassic, around 161 to 157 million years ago. The type and only species Limusaurus inextricabilis was described in 2009 from specimens found in the Upper Shishugou Formation in the Junggar Basin of China. The genus name consists of the Latin words for "mud" and "lizard", and the species name means "impossible to extricate", both referring to these specimens possibly dying after being mired. Limusaurus was a small, slender animal, about 1.7 m in length and 15 kg (33 lb) in weight, which had a long neck and legs but very small forelimbs. It underwent a drastic morphological transformation as it aged: while juveniles were toothed, these teeth were completely lost and replaced by a beak with age. Several of these features were convergently similar to the later ornithomimid theropods as well as the earlier non-dinosaurian shuvosaurids.
Austrocheirus is an extinct genus of theropod dinosaur that lived during the Late Cretaceous period. It was named and described by Martin Ezcurra, Federico Agnolin and Fernando Novas in 2010. It contains the type species Austrocheirus isasii. The generic name means "southern hand". The specific epithet honours discoverer and preparator Marcelo Pablo Isasi.
Eoabelisaurus is a genus of abelisaurid theropod dinosaur from the Lower Jurassic Cañadón Asfalto Formation of the Cañadón Asfalto Basin in Argentina, South America. The generic name combines a Greek ἠώς, (eos), "dawn", with the name Abelisaurus, in reference to the fact it represents an early relative of the latter. Only one species is currently recognized, E. mefi; the specific name honours the MEF, the Museo Paleontológico "Egidio Feruglio", where discoverer Diego Pol is active. It is characterized by reduced forelimb proportions that show primitive characteristics of the Abelisauridae family.
Arcovenator is an extinct genus of abelisaurid theropod dinosaurs hailing from the Late Cretaceous of France. The type and only described species is Arcovenator escotae.
Fosterovenator is a genus of ceratosaur dinosaur known from the Late Jurassic Morrison Formation of Wyoming. The holotype is YPM VP 058267A, B, and C, a tibia with an articulated astragalus. An additional specimen is known, the paratype YPM VP 058267D, a fibula of a larger individual.
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.
Afromimus is a genus of theropod dinosaur from the Early Cretaceous Elrhaz Formation of Niger. It contains a single species, A. tenerensis, named in 2017 by Paul Sereno from parts of the right leg, vertebrae, and ribs found in the Ténéré Desert. It was originally classified as an ornithomimosaurian, but subsequently it was argued to be an abelisauroid.
The Azilal Formation, also known as Toundoute Continental Series and Wazzant Formation, is a geological unit in the Azilal, Béni-Mellal, Imilchil, Zaouiat Ahansal, Ouarzazate, Tinerhir and Errachidia areas of the High Atlas of Morocco, part of the Tafraout Group that covers the Early Toarcian to Middle Aalenian stages of the Jurassic Period. While there have been atributions of it´s lowermost layers to the Latest Pliensbachian, the current oldest properly measured are part of the Earliest Toarcian regresion ("MRST10"), part of the Lower-Middle Palymorphum biozone. The Azilal Formation consists mainly of claystones rich in continental plant debris and laminated microbial facies. It is a continental deposit which overlies marine dolomites of equivalent age to the Rotzo Formation of Italy, mostly part of the Aganane Formation. The formation is the continental part of a massive Siliciclastic-Carbonate platform, best assigned to an alluvial environment occasionally interrupted by shallow marine incursions and marks a dramatic decrease of the carbonate productivity under increasing terrigenous sedimentation. Dinosaur remains, such the sauropod Tazoudasaurus and the basal ceratosaur Berberosaurus are known from the unit, along with several undescribed genera. The toarcian High Atlas is divided in 5 units: the continental layers with paralic deposits belong to the Azilal, along the shoreface layers of the Tagoudite Formation and Tafraout Formation & Aguerd-nˈTazoult Formation, connected with the offshore Ait Athmane Formation and the deeper shelf deposits of the Agoudim 1 Formation.
Dornraptor is an extinct genus of averostran theropod dinosaur from the Early Jurassic of Charmouth, Dorset, England. The genus contains a single species, D. normani, known from a fragmentary knee joint and femur that were initially described by Sir Richard Owen as belonging to the early armored dinosaur Scelidosaurus. Dornraptor lived in what is now England, along other theropods like Dracoraptor and Sarcosaurus.
The Tafraout Group is a geological group of formations of Toarcian-Aalenian age in the Azilal, Béni-Mellal, Imilchil, Zaouiat Ahansal, Ouarzazate, Tinerhir and Errachidia areas of the High Atlas of Morocco. The Group represents the remnants of a local massive Siliciclastic-Carbonate platform, best assigned to succession W-E of alluvial environment occasionally interrupted by shallow marine incursions and inner platform to open marine settings, and marks a dramatic decrease of the carbonate productivity under increasing terrigenous sedimentation. Fossils include large reef biotas with richness in "lithiotid" bivalves and coral mounts, but also by remains of vertebrates such as the sauropod Tazoudasaurus and the basal ceratosaur Berberosaurus, along with several undescribed genera. While there have been attributions of its lowermost layers to the Latest Pliensbachian, the current oldest properly measured are part of the Earliest Toarcian regression ("MRST10"), part of the Lower-Middle Palymorphum biozone. This group is composed of the following units, which extend from west to east: the Azilal Formation ; the Amezraï Formation ; the Aguerd-nˈTazoult Formation ; the Tagoudite Formation & the Trafraout Formation. They are connected with the offshore Ait Athmane Formation and the deeper shelf deposits of the Agoudim 1 Formation. Overall, this group represents a mixed carbonate-siliciclastic system of several hundred meters thick, dominated by deposits of shallow marine platforms linked to a nearby hinterland dominated by conglomerates. The strata of the group extend towards the central High Atlas, covering different anticlines and topographic features along the mountain range.