Riojasaurus

Riojasaurus (meaning "Rioja lizard") was a herbivorous sauropodomorph dinosaur named after La Rioja Province in Argentina where it was found in the Los Colorados Formation in the Ischigualasto-Villa Unión Basin by José Bonaparte. It lived during the Late Triassic (Norian stage) and grew to about 10 metres (33 ft) long.[1] Riojasaurus is the only known riojasaurid to live in South America.

Riojasaurus
Temporal range: Norian
~227–213 Ma
Riojasaurus skull
Riojasaurus skull cast
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Order: Saurischia
Suborder: Sauropodomorpha
Family: Riojasauridae
Genus: Riojasaurus
Bonaparte 1969
Type species
Riojasaurus incertus

Description

Riojasaurus Scale
Size comparison

Riojasaurus had a heavy body, bulky legs, and a long neck and tail. Its leg bones were dense and massive for an early sauropodomorph.[1] By contrast, its vertebrae were lightened by hollow cavities, and unlike most early sauropodomorphs, Riojasaurus had four sacral vertebrae instead of three.[1] It has been thought it probably moved slowly on all fours and was unable to rear up on its back legs.[1][2] The nearly equal length of the fore and hindlimbs[1] has also been interpreted as suggestive of an obligatorily quadrupedal gait. However, in 2016, Scott Hartman found the hand anatomy, relatively straight back and largely immobile shoulder girdle of Riojasaurus supported it being a biped.[3]

No skull was found with the first skeleton of Riojasaurus,[4] although a well-preserved skull attributed to Riojasaurus was found later.[5] The teeth of Riojasaurus were leaf shaped and serrated. The upper jaw contained 5 teeth at the front, with 24 more behind them in a row that ended under the eyes.

Comparisons between the scleral rings of Riojasaurus and modern birds and reptiles suggest that it may have been cathemeral, active throughout the day at short intervals.[6]

Classification

Many scientists think that Riojasaurus was closely related to Melanorosaurus,[1] known from the Triassic-Early Jurassic period. However, studies at Bristol University, England, suggest that it is unique in some key ways, such as the longer bones in its neck. It is certainly quite different from other sauropodomorphs found in the Los Colorados Formation of Argentina.[7]

Due to their size and limb anatomy Riojasaurus and the possibly related Melanorosaurus have been considered close relatives of the earliest sauropods.[1]

References

  1. ^ a b c d e f g "Riojasaurus." In: Dodson, Peter & Britt, Brooks & Carpenter, Kenneth & Forster, Catherine A. & Gillette, David D. & Norell, Mark A. & Olshevsky, George & Parrish, J. Michael & Weishampel, David B. The Age of Dinosaurs. Publications International, LTD. p. 41. ISBN 0-7853-0443-6.
  2. ^ Van Heerden, J. and Galton, P.M. (1997). "The affinities of Melanorosaurus a Late Triassic prosauropod dinosaur from South Africa". Neues Jahrbuch für Geologie und Paläontologie Monatshefte. (1):39-55
  3. ^ Hartman, Scott. "La Rioja's ponderous biped".
  4. ^ Bonaparte, J.F. (1969). Dos nuevas "faunas" de reptiles triásicos de Argentina. Ameghiniana 10(1): 89-102.
  5. ^ Bonaparte, J.F. & Pumares, J.A. (1995). Notas sobre el primer craneo de Riojasaurus incertus (Dinosauria, Prosauropoda, Melanorosauridae) del Triasico Superios de La Rioja, Argentina. Ameghiniana 32(4): 341-349.
  6. ^ Schmitz, L.; Motani, R. (2011). "Nocturnality in Dinosaurs Inferred from Scleral Ring and Orbit Morphology". Science. 332 (6030): 705–8. Bibcode:2011Sci...332..705S. doi:10.1126/science.1200043. PMID 21493820.
  7. ^ Moody, Richard. Dinofile. pg 20. Octopus Publishing Group Ltd., 2006
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.

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.

Eucnemesaurus

Eucnemesaurus (; meaning "good tibia lizard", for its robust tibiae) is a basal sauropodomorph dinosaur genus usually considered to be a synonym of Euskelosaurus. Recent study by Yates (2006), however, indicates that it is valid and the same animal as putative "giant herrerasaurid" Aliwalia.

Eucnemesaurus was named in 1920 by Egbert Cornelis Nicolaas van Hoepen. The type species is Eucnemesaurus fortis. The specific name means "strong" in Latin. It is based on holotype TrM 119, a partial skeleton including vertebrae, part of a pubis, a femur, and two tibiae. The remains were found by Van Hoepen in the late Carnian-early Norian-age Upper Triassic Lower Elliot Formation of the Slabberts district, Orange Free State, South Africa.

Yates assigned the genus to the new family Riojasauridae, with Riojasaurus, usually regarded as a melanorosaurid.

Jingshanosaurus

Jingshanosaurus (meaning "Jingshan lizard") is a genus of sauropodomorph dinosaurs from the early Jurassic period.

La Rioja Province, Argentina

La Rioja (Spanish pronunciation: [la ˈrjoxa]) is one of the provinces of Argentina and is located in the west of the country. Neighboring provinces are from the north clockwise Catamarca, Córdoba, San Luis and San Juan. The dinosaur Riojasaurus is named after the province.

Los Colorados Formation

The Los Colorados Formation is a sedimentary rock formation of the Ischigualasto-Villa Unión Basin, found in the provinces of San Juan and La Rioja in Argentina. The formation dates back to the Norian age of the Late Triassic.

The up to 600 metres (2,000 ft) thick formation comprises sandstones, siltstones, mudstones and conglomerates with gypsum layers deposited in a fluvial to lacustrine environment. The formation is the uppermost stratigraphic unit of the Agua de la Peña Group, overlying the Lagerstätte of the Ischigualasto Formation. Los Colorados Formation is partly covered by the Cretaceous Cerro Rajado Formation, separated by an unconformity.

The formation is known for its fossils of early dinosaurs, including the coelophysoid Zupaysaurus and the "prosauropods" Coloradisaurus, Lessemsaurus, and Riojasaurus. Magnetostratigraphic analysis suggests that the Los Colorados Formation was deposited between 227 and 213 million years ago.

Massospondylidae

Massospondylidae is a family of early massopod dinosaurs that existed in Asia, Africa, South America and Antarctica during the Late Triassic to the Early Jurassic periods. Several dinosaurs have been classified as massospondylids over the years. The largest cladistic analysis of early sauropodomorphs, which was presented by Apaldetti and colleagues in November 2011, found Adeopapposaurus, Coloradisaurus, Glacialisaurus, Massospondylus, Leyesaurus and Lufengosaurus to be massospondylids. This result supports many previous analyses that tested fewer taxa. However, this analysis found the two recently described North American massopods, Sarahsaurus and Seitaad, and the South African Ignavusaurus to nest outside Massospondylidae, as opposed to some provisional proposals. Earlier in 2011, Pradhania, a sauropodomorph from India, was tested for the first time in a large cladistic analysis and was found to be a relatively basal massospondylid. Mussaurus and Xixiposaurus may also be included within Massospondylidae. In 2019, a specimen previously assigned to Massospondylus from South Africa was re-examined and found to belong to a separate genus that was named Ngwevu.

Mussaurus

Mussaurus (meaning "mouse lizard") is a genus of herbivorous sauropodomorph dinosaur that lived in southern Argentina during the Late Triassic, about 215 million years ago. It receives its name from the small size of the skeletons of juvenile and infant individuals, which were once the only known specimens of the genus. However, since Mussaurus is now known from adult specimens, the name is something of a misnomer; adults possibly reached 6 metres (20 ft) in length and weighed more than 1,000 kilograms (2,200 lb). Mussaurus possesses anatomical features suggesting a close, possibly transitional evolutionary relationship with true sauropods.

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.

Plateosauria

Plateosauria is a clade of sauropodomorph dinosaurs which lived during the Late Triassic to the Late Cretaceous. The name Plateosauria was first coined by Gustav Tornier in 1913. The name afterwards fell out of use until the 1980s.

Plateosauria is a node-based taxon. In 1998, Paul Sereno defined Plateosauria as the last common ancestor of Plateosaurus engelhardti and Massospondylus carinatus, and its descendants. Peter Galton and Paul Upchurch in 2004 used a different definition: the last common ancestor of Plateosaurus engelhardti and Jingshanosaurus xinwaensis, and its descendants. In their cladistic analysis the Plateosauria belonged to the Prosauropoda, and included the Plateosauridae subgroup. In Galton's and Upchurch's study also Coloradisaurus, Euskelosaurus, Jingshanosaurus, Massospondylus, Mussaurus, Sellosaurus, and Yunnanosaurus proved to be plateosaurians.However, recent cladistic analyses suggest that the Prosauropoda as traditionally defined is paraphyletic to sauropods. Prosauropoda, as currently defined, is a synonym of Plateosauridae as both contain the same taxa by definition.

The following cladogram simplified after an analysis presented by Apaldetti and colleagues in 2011.

The following cladogram simplified after an analysis presented by Blair McPhee and colleagues in 2014.

Riojasauridae

Riojasauridae is a family of sauropod-like dinosaurs from the Upper Triassic. It is known primarily from the genera Riojasaurus and Eucnemesaurus. Sites containing Riojasauridae include the Lower Elliot Formation of Orange Free State, South Africa (where fossils of Eucnemesaurus have been found), and Ischigualasto, in La Rioja Province, Argentina ( where fossils of Riojasaurus have been recovered).

Riojasuchus

Not to be confused with the sauropodomorph dinosaur, Riojasaurus

Riojasuchus is an extinct genus of Late Triassic (Norian) quadrupedal crurotarsan archosaur. Riojasuchus is a member of Ornithosuchidae, a family of facultatively bipedal carnivores that were geographically widespread during the Late Triassic. Two other genera, Ornithosuchus and Venaticosuchus, are currently known. The holotype specimen is PVL 3827. It was found in the Los Colorados Formation of the Ischigualasto-Villa Unión Basin in northwestern Argentina.

Sarahsaurus

Sarahsaurus is a genus of basal sauropodomorph dinosaur which lived during the lower Jurassic period in what is now northeastern Arizona, United States.

Sauropodomorpha

Sauropodomorpha ( SOR-ə-POD-ə-MOR-fə; from Greek, meaning "lizard-footed forms") is an extinct clade of long-necked, herbivorous, saurischian dinosaurs that includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had long necks and tails, were quadrupedal, and became the largest animals to ever walk the Earth. The "prosauropods", which preceded the sauropods, were smaller and were often able to walk on two legs. The sauropodomorphs were the dominant terrestrial herbivores throughout much of the Mesozoic Era, from their origins in the mid-Triassic (approximately 230 Ma) until their decline and extinction at the end of the Cretaceous (approximately 66 Ma).

Xingxiulong

Xingxiulong (meaning "Xingxiu Bridge dragon") is a genus of bipedal sauropodiform from the Early Jurassic of China. It contains a single species, X. chengi, described by Wang et al. in 2017 from three specimens, two adults and an immature individual, that collectively constitute a mostly complete skeleton. Adults of the genus measured 4–5 metres (13–16 ft) long and 1–1.5 metres (3 ft 3 in–4 ft 11 in) tall. Phylogenetic analysis suggests that Xingxiulong is most closely related to its contemporary Jingshanosaurus, although an alternative position outside of both the Sauropodiformes and Massospondylidae is also plausible.

Despite their close relationship, Xingxiulong prominently differs from Jingshanosaurus - and from most basal sauropodomorphs - in having a number of sauropod-like traits. These include a sacrum containing four vertebrae; a pubis with an exceptionally long top portion; and the femur, the first and fifth metatarsals on the foot, and the scapula being wide and robust. These probably represent adaptations to supporting high body weight, in particular a large gut. Unlike sauropods, however, Xingxiulong would still have been bipedal.

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