Chilesaurus | |
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Cast of the holotype skeleton | |
Scientific classification ![]() | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | Saurischia |
Clade: | Theropoda (?) |
Genus: | † Chilesaurus Novas et al. 2015 |
Species: | †C. diegosuarezi |
Binomial name | |
†Chilesaurus diegosuarezi Novas et al. 2015 | |
Chilesaurus is an extinct genus of herbivorous dinosaur. While its exact classification is uncertain, many researchers believe it is a theropod, with a minority of academics suggesting that it may be an ornithischian. The type and only known species so far is Chilesaurus diegosuarezi. [1] Chilesaurus lived between 148-147 million years ago (Mya) in the Late Jurassic period of Chile. [2] Showing a combination of traits from theropods, ornithischians, and sauropodomorphs, this genus has far-reaching implications for the evolution of dinosaurs, such as whether the traditional saurischian-ornithischian split is superior or inferior to the proposed group Ornithoscelida. [3] This however, has been contested by several other authors, who believe that the weight of evidence supports its membership within Theropoda, and possibly as a member of Tetanurae. [4] [5]
Chilesaurus was first discovered on 4 February 2004 by a seven-year-old named Diego Suárez. He and his parents, the geologists Manuel Suárez and Rita de la Cruz, were searching for decorative stones in the Aysén Region, and uncovered a vertebra and rib from what would later be named Chilesaurus. More specimens were found that were assigned to various dinosaur species in 2008 but were later recognized as belonging to additional individuals of Chilesaurus. [6] One reason this realization took time was that Chilesaurus has such a bizarre combination of traits, coupled with the fact that the remains were discovered alongsife a few bones from an unrelated diplodocid sauropod. [7]
In 2015, the type species C. diegosuarezi was named and described by Fernando Emilio Novas, Leonardo Salgado, Manuel Suárez, Federico Lisandro Agnolín, Martín Dario Ezcurra, Nicolás Chimento, Rita de la Cruz, Marcelo Pablo Isasi, Alexander Omar Vargas, and David Rubilar-Rogers. The generic name is derived from Chile (the country where it was discovered) and the latinized Greek suffix "saurus", meaning "lizard". The specific name honors Diego Suárez. [1]
The holotype, SNGM-1935, was found in a layer of the Toqui Formation dating from the late Tithonian. It consists of an articulated, rather complete skeleton with skull of a juvenile individual, lacking the feet and most of the tail. Four other partial skeletons (specimens SNGM-1937, SNGM-1936, SNGM-1938, and SNGM-1888) and several single bones (specimens SNGM-1889, SNGM-1895, SNGM-1901, SNGM-1894, SNGM-1898, SNGM-1900, and SNGM-1903) are the paratypes. They represent juvenile and adult individuals. [1]
The holotype of Chilesaurus is relatively complete. It includes a partially complete skull and mandible, a complete series of neck and back vertebrae, most of the ribcage, a complete arm and leg, most of the hips, shoulders, and a few of the front-most tail vertebrae. However, the lack of a complete tail makes a full estimation of its size imprecise. The holotype itself, which represents a skeletally immature individual, is estimated to have been about 1.6 m (5.2 ft) long in life. There are several paratypes representing at least four individuals at varying stages of ontogenetic maturity. The largest of these, believed to be an adult, was used to infer that the full length of an adult Chilesaurus would have been about 3.2 m (10.5 ft). [1]
Chilesaurus can be distinguished from all other dinosaurs by a combination of unique features. It has a very short premaxillary bone, a uniquely quadrilaterally-shaped coracoid bone with thickened outer edges, a hand with only two clawed fingers (the third finger being highly reduced) with very shortened phalanges, and several features of the hip and ankle bones that are distinct from all other theropods. However, one of the most distinctive unusual features of Chilesaurus is its spatula-shaped, elongated teeth, which protrude forwards out of their sockets. Such dentition would be unique among theropods, and is typical for a herbivore, indicating Chilesaurus was a plant-eater. If Chilesaurus was a theropod, it would be only one of a handful of non-coelurosaurian theropods to evolve herbivory, alongside Limusaurus , Berthasaura , and possibly others. [1] Another apparent adaptation for herbivory is the backward-pointing pubic bone in the pelvis, which has been interpreted as an adaptation for developing the more complex digestive apparatus necessaru to digest plants. Such a pelvic arrangement is typical for ornithischians, which has led some authors to suggest that Chilesaurus may be a member of that group. [8]
The hind limb of Chilesaurus is less adapted for running than many other groups of dinosaurs. This is inferred from the presence of a small cnemial crest on the front top of the shinbone, and a broad foot with a weight-bearing first toe. These adaptations would have made running for long periods of time much less efficient, and they are generally not present in animals which are known to have been proficcient runners. However, it is possible that Chilesaurus could defend itself relatively strong arms which bore a large claw on the first finger. [1]
In its original description, Chilesaurus was found to be a very basally-branching member of the theropod clade Tetanurae. An abbreviated version of the cladogram produced by the authors of that paper is shown below. [1]
In 2017, Baron and Barrett proposed that Chilesaurus may instead be a basal ornithischian. The publication in which this was proposed also resurrected the previously-discredited "Ornithoscelida hypothesis", which suggested that theropods shared a more recent common ancestor with ornithischians than with sauropodomorphs. [8] [9] Less than a year later in 2018, Müller and colleagues published a reply to Baron & Barrett (2017), arguing that their phylogenetic dataset actually suggested that Chilesaurus was a basal sauropodomorph rather than an ornithischian. [10] Baron and Barrett reached out to the other team of authors to inform them that they had accidentally published a faulty early version of their dataset with many traits scored incorrectly, and that their original results were based on an edited final dataset. They corrected their original publication and supplied the final dataset to Müller and colleagues, who agreed that it supported the placement of Chilesaurus in Ornithischia as had been originally argued by Baron and Barrett in 2017. [10] [3] However, Müller and colleagues also noted that Baron and Barrett did not test the original proposal of Chilesaurus as a theropod, and that its classification remained uncertain. [10] Below is a cladogram illustrating the classification proposed by Baron & Barrett (2017), which places Chilesaurus as the most basal ornithischian. [8]
In a latter re-revision of his own datasets, Baron concluded that Chilesaurus remains enigmatic and could be either a theropod or an ornithischian, while noting that its affinities as a theropod were more generally supported by other authors. [11] In his major work on theropod classification in 2024, Andrea Cau recovered Chilesaurus as a sister taxon of Tetanurae. [4] Fonseca and colleagues conducted a similar work on the classification of ornithischians, which also recovered Chilesaurus as a theropod. [5]
A full osteological description of Chilesaurus was published as part of the PhD thesis of Nicolás Roberto Chimento Ortiz in 2018. In it, he noted the likely theropod affinities of the taxon, highlighting elements such as the structure of the metatarsals, characteristics of the tibia, and the opisthopubic pelvis. [12]
Ornithischia is an extinct clade of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name Ornithischia, or "bird-hipped", reflects this similarity and is derived from the Greek stem ornith- (ὀρνιθ-), meaning "bird", and ischion (ἴσχιον), meaning "hip". However, birds are only distantly related to this group, as birds are theropod dinosaurs.
Eoraptor is a genus of small, lightly built, basal sauropodomorph dinosaur. One of the earliest-known dinosaurs and one of the earliest sauropodomorphs, it lived approximately 231 to 228 million years ago, during the Late Triassic in Western Gondwana, in the region that is now northwestern Argentina. The type and only species, Eoraptor lunensis, was first described in 1993, and is known from an almost complete and well-preserved skeleton and several fragmentary ones. Eoraptor had multiple tooth shapes, which suggests that it was omnivorous.
Ankylosauria is a group of herbivorous dinosaurs of the clade Ornithischia. It includes the great majority of dinosaurs with armor in the form of bony osteoderms, similar to turtles. Ankylosaurs were bulky quadrupeds, with short, powerful limbs. They are known to have first appeared in North Africa during the Middle Jurassic, and persisted until the end of the Late Cretaceous. The two main families of ankylosaurians, Nodosauridae and Ankylosauridae are primarily known from the Northern Hemisphere, but the more basal Parankylosauria are known from southern Gondwana during the Cretaceous.
Guaibasaurus is an extinct genus of basal saurischian dinosaur known from the Late Triassic Caturrita Formation of Rio Grande do Sul, southern Brazil. Most analyses recover it as a sauropodomorph, although there are some suggestions that it was a theropod instead. In 2016 Gregory S. Paul estimated it at 2 meters and 10 kg, whereas in 2020 Molina-Pérez and Larramendi listed it at 3 meters and 35 kg.
Saurischia is one of the two basic divisions of dinosaurs, classified by their hip structure. Saurischia and Ornithischia were originally called orders by Harry Seeley in 1888 though today most paleontologists classify Saurischia as an unranked clade rather than an order.
Guaibasauridae is a family of basal sauropodomorph dinosaurs, known from fossil remains of late Triassic period formations in Brazil, Argentina and India.
Fernando Emilio Novas is an Argentine paleontologist working for the Comparative Anatomy Department of the Bernardino Rivadavia Natural Sciences Museum in Buenos Aires, Argentina. Novas holds a PhD in Natural sciences.
Phytodinosauria is a group of dinosaurs proposed in 1986, combining the Sauropodomorpha and Ornithischia as sister groups, conceptualized as a superorder of herbivorous dinosaurs excluding the carnivorous Theropoda. This hypothesis has been refuted by modern cladistic analysis, showing such a group to be polyphyletic. Modern studies either combine the Theropoda and Sauropodormorpha in the Saurischia or the Theropoda and Ornithischia in the Ornithoscelida.
Ornithoscelida is a proposed clade that includes various major groupings of dinosaurs. An order Ornithoscelida was originally proposed by Thomas Henry Huxley but later abandoned in favor of Harry Govier Seeley's division of Dinosauria into Saurischia and Ornithischia. The term was revived in 2017 after a new cladistic analysis by Baron et al.
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Zuolong is an extinct genus of tetanuran theropod from the Late Jurassic period of China. The type and only species is Z. salleei. The generic name of Zuolong is in honor of General Zuo Zōngtáng with the Chinese word "long" which means dragon. The specific epithet "salleei" is in honor of Hilmar Sallee, who funded the expedition which led to the specimen's discovery.
Eodromaeus is an extinct genus of probable basal theropod dinosaurs from the Late Triassic of Argentina. Like many other of the earliest-known dinosaurs, it hails from the Carnian-age Ischigualasto Formation, within the Ischigualasto-Villa Unión Basin of northwestern Argentina. Upon its discovery, it was argued to be one of the oldest true theropods, supplanting its contemporary Eoraptor, which was reinterpreted as a basal sauropodomorph.
Daemonosaurus is an extinct genus of possible theropod dinosaur from the Late Triassic of New Mexico. The only known fossil is a skull and neck fragments from deposits of the latest Triassic Chinle Formation at Ghost Ranch. Daemonosaurus was an unusual dinosaur with a short skull and large, fang-like teeth. It lived alongside early neotheropods such as Coelophysis, which would have been among the most common dinosaurs by the end of the Triassic. However, Daemonosaurus retains several plesiomorphic ("primitive") traits of the snout, and it likely lies outside the clade Neotheropoda. It may be considered a late-surviving basal theropod or non-theropod basal saurischian, possibly allied to other early predatory dinosaurs such as herrerasaurids or Tawa.
Buriolestes is a genus of early sauropodomorph dinosaurs from the Late Triassic Santa Maria Formation of the Paraná Basin in southern Brazil. It contains a single species, B. schultzi, named in 2016. The type specimen was found alongside a specimen of the lagerpetid dinosauromorph Ixalerpeton.
The Toqui Formation is a geological formation in the Aysén Region of southern Chile. It has been dated to the Tithonian stage of the Late Jurassic by uranium–lead dating of zircons, providing ages between 148.7 ± 1.4 Ma and 147 ± 1.0 Ma. It consists of a sequence of clastic sedimentary sandstones and conglomerates, interbedded with volcanic tuffs and ignimbrite. The dinosaurs Chilesaurus and indeterminate diplodocids and the mesoeucrocodylian Burkesuchus are known from the formation. The formation was deposited in a fluvio-deltaic environment.
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Nhandumirim is a genus of basal sauropodomorph dinosaur from the Carnian age of Late Triassic Brazil. It is currently considered a saturnaliid sauropodomorph. The type and only species, Nhandumirim waldsangae, is known from a single immature specimen including vertebrae, a chevron, pelvic material, and a hindlimb found in the Santa Maria Formation in Rio Grande do Sul.
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