Zupaysaurus

Zupaysaurus (/ˌzuːpeɪˈsɔːrəs/; "ZOO-pay-SAWR-us") is a genus of early theropod dinosaur living during the Norian stage of the Late Triassic in what is now Argentina. Fossils of the dinosaur were found in the Los Colorados Formation of the Ischigualasto-Villa Unión Basin in northwestern Argentina. Although a full skeleton has not yet been discovered, Zupaysaurus can be considered a bipedal predator, up to 4 metres (13 ft) long. It may have had two parallel crests running the length of its snout.

Zupaysaurus
Temporal range: Norian
~221–206 Ma
Zupaysaurus
Zupaysaurus in resting pose
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Order: Saurischia
Suborder: Theropoda
Clade: Neotheropoda
Genus: Zupaysaurus
Arcucci & Coria 2003
Species:
Z. rougieri
Binomial name
Zupaysaurus rougieri
Arcucci & Coria 2003

Description

Zupaysaurus skull
Skull diagram

Zupaysaurus was a medium-sized theropod. An adult skull, measured approximately 450 mm (18 in) in length, suggesting a body length of approximately 4 m (13 ft) from its snout to the tip of its tail. Other estimates suggest that Zupaysaurus was at best 5.2 m (17 ft) long and weighed 200 kg (441 lb) at most.[1] Like all theropods, Zupaysaurus walked only on its hindlegs, leaving its forelimbs free to grasp its prey. The length of the neck bones recovered suggests that this genus has a rather long neck. Like the coelophysoids, Zupaysaurus has a kink in its snout, between the premaxillary and maxillary bones of the upper jaw. It is estimated that Zupaysaurus had 24 teeth and an intermandibular hinge is present in the lower jaw.[2] Only one specimen of Zupaysaurus is known to science. The holotype specimen was designated PULR-076, which consists of a nearly complete skull which was very well preserved, the right shoulder girdle, the lower right leg and ankle, and twelve vertebrae from the neck, back, and hips. Additional material of a smaller individual found at the same site may or may not belong to Zupaysaurus.

As Zupaysaurus was originally described, the head bore two thin parallel crests on top of the skull, similar to theropods like Dilophosaurus and Coelophysis kayentakatae. These crests are thought to have been formed by the nasal bones solely, unlike those of many other theropods which also incorporated the lacrimal bones. Crests on the skull were pervasive among theropods and may have been used for communicative purposes such as species or gender recognition.[3] However, more recent analysis of the skull has cast doubt on the presence of these crests in Zupaysaurus. An unpublished abstract presented at a recent conference indicated the structures initially identified as crests were in fact the lacrimal bones displaced upwards during the process of fossilization.[4] Other cranial ornamentation included a rugose laterally-projecting lacrimal ridge on the top of the skull.

A diagnosis is a statement of the anatomical features of an organism (or group) that collectively distinguish it from all other organisms. Some, but not all, of the features in a diagnosis are also autapomorphies. An autapomorphy is a distinctive anatomical feature that is unique to a given organism or group. According to Ezcurra (2006) and Ezcurra and Novas (2006), Zupaysaurus can be distinguished based on the following characteristics: the maxillary fenestra is within the antorbital fossa (according to Ezcurra, 2006), the rostral process of the lacrimal is ventrally bowed (according to Ezcurra, 2006), the ventral process of the squamosal is kinked (according to Ezcurra, 2006), wide contact between squamosal and quadratojugal (according to Ezcurra, 2006). The maxillary-jugal ventral margin describes an obtuse angle in lateral view (according to Ezcurra and Novas, 2006), a notch on the dorsal margin of the ascending process of the maxilla, relating to horizontal ramus of the lacrimal is rostrally tapering onto the forked caudal tip of the ascending process of the maxilla; (according to Ezcurra and Novas, 2006), a lacrimal with a highly pneumatized antorbital recess (according to Ezcurra and Novas, 2006), a short and square-shaped retroarticular process of the mandible (according to Ezcurra and Novas, 2006), the cnemial crest is poorly developed (according to Ezcurra and Novas, 2006).

Discovery

Discovered in May 1997 by Santiago Reuil ("Vultur"), part of the crew of Guillermo Rougier. It was later described by Arcucci and Coria and published in 2003. The name Zupaysaurus is composed of the Quechua word supay meaning "devil" and the Greek word sauros (σαυρος) meaning "lizard"; thus "devil lizard". In Incan mythology, supay was both the god of death and ruler of the ukhu pacha, the Incan underworld. The type species was named Z. rougieri in the honor of Guillermo Rougier, the scientist who led the expedition which discovered and collected the holotype (original specimen) PULR-076. Zupaysaurus was first described and named in the scientific journal Ameghiniana by Argentine paleontologists Andrea Arcucci and Rodolfo Coria in 2003.[2]

Classification

Zupaysaurus was classified as the earliest known tetanuran theropod due to several features of its skull, dentition, and hindlimb. However, several features typical of more basal theropods were also noted by the original authors.[2] Analyses by Carano (2005), Tykoski (2005), and Ezcurra and Novas (2005) have classified Zupaysaurus as a coelophysoid related to Segisaurus and probably Liliensternus, though more basal than Coelophysis.[4][5][6] Yates (2006) found Zupaysaurus to form a group with Dilophosaurus and Dracovenator, placing it in a monophyletic Dilophosauridae.[7] But later studies found Zupaysaurus to be a sister taxon sister to a clade containing dilophosaurids, ceratosaurs and tetanurans.[8][9]

Below is a cladogram based on the phylogenetic analysis conducted by Sues et al. in 2011, showing the relationships of Zupaysaurus:[10]

Theropoda 
 Herrerasauridae 

Staurikosaurus

Herrerasaurus

Chindesaurus

Eoraptor

Daemonosaurus

Tawa

 Neotheropoda 
 Coelophysidae 

Coelophysis

Liliensternus

Zupaysaurus

Cryolophosaurus

Dilophosaurus

Jurassic theropods (which include ceratosaurs and tetanurans)

Paleoecology

Zupaysaurus was discovered in red siliciclastic sediments at the "Quebrada de los Jachaleros" locality within the Los Colorados Formation of the La Rioja province in Argentina. This formation has been shown by magnetostratigraphy to date to the Norian stage of the Late Triassic period, approximately 228 to 208 million years ago.[11] but has also been assigned to the slightly younger Rhaetian stage, which was approximately 208 to 201 million years ago.[12] Both specimens assigned to this genus are housed in the collection of the National University of La Rioja in La Rioja, Argentina.[2]

The Los Colorados Formation was interpreted as an ancient floodplain and it was home to several types of early sauropodomorph dinosaurs (including Riojasaurus, Coloradisaurus , and Lessemsaurus), all of which shared the same paleoenvironment with Zupaysaurus.[11] It is recognized as one of the earliest known faunal assemblages dominated by dinosaurs, which were 43% of the number of tetrapod species currently known. The non-dinosaurs that inhabited this locality included pseudosuchians, therapsids like Cynodontia, other early reptiles, and possible archosaurs.

References

  1. ^ "ZUPAYSAURUS". Dinochecker.com. Retrieved 5 May 2013.
  2. ^ a b c d Arcucci, A.B. & Coria R.A. 2003. A new Triassic carnivorous dinosaur from Argentina. Ameghiniana 40(2):217-228.
  3. ^ Currie, P.J. & Zhao X. 1993. A new carnosaur (Dinosauria, Theropoda) from the Jurassic of Xinjiang, People's Republic of China. Canadian Journal of Earth Sciences 30: 2037-2081.
  4. ^ a b Ezcurra, M.D. & Novas, F.E. 2005. Phylogenetic relationships of the Triassic theropod *Zupaysaurus rougieri* from NW Argentina. Presented in August 2005 during the II Latin American Congress of Vertebrate Paleontology Archived May 4, 2006, at the Wayback Machine in Rio de Janeiro, Brazil. This analysis will be published in peer-reviewed print form later in 2006. A summary of the talk can be seen here.
  5. ^ Carrano, M.T., Hutchinson, J.R., & Sampson, S.D. 2005. New information on Segisaurus halli, a small theropod dinosaur from the Early Jurassic of Arizona. Journal of Vertebrate Paleontology 25(4): 835-849.
  6. ^ Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Dissertation. University of Texas at Austin. 553 pp.
  7. ^ Yates, A.M., 2006 (for 2005). "A new theropod dinosaur from the Early Jurassic of South Africa and its implications for the early evolution of theropods." Palaeontologia Africana, 41: 105-122.
  8. ^ Smith N.D., Makovicky P. J., Hammer W. R. & Currie P. J. 2007 Osteology of Cryolophosaurus ellioti (Dinosauria: Theropoda) from the Early Jurassic of Antarctica and implications for early theropod evolution. Zool. J. Linn. Soc. 151, 377–421.
  9. ^ Nesbitt, S. J., Smith, N. D., Irmis, R. B., Turner, A. H., Downs, A., & M. A. Norell. 2009. A complete skeleton of a Late Triassic saurischian and the early evolution of dinosaurs. Science 326:1530-1533.
  10. ^ Hans-Dieter Sues, Sterling J. Nesbitt, David S. Berman and Amy C. Henrici (2011). "A late-surviving basal theropod dinosaur from the latest Triassic of North America". Proceedings of the Royal Society B 278 (1723): 3459–3464
  11. ^ a b Weishampel, D.B., Barrett, P.M., Coria, R.A., Le Loueff, J., Xu X., Zhao X., Sahni, A., Gomani, E.M.P., & Noto, C.R. 2004. Dinosaur distribution. In: Weishampel, D.B., Dodson, P., & Osmólska, H. (Eds.). The Dinosauria (2nd Edition). Berkeley: University of California Press. Pp. 517-606.
  12. ^ Heckert, A.B. & Lucas, S.G. 1998. Global correlation of the Triassic theropod record. Gaia 15: 63-74. [not printed until 2000]

External links

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.

Coelophysidae

Coelophysidae is a family of primitive carnivorous theropod dinosaurs. Most species were relatively small in size. The family flourished in the Late Triassic and Early Jurassic periods, and has been found on numerous continents. Many members of Coelophysidae are characterized by long, slender skulls and light skeletons built for speed. One member, Coelophysis, displays the earliest known furcula in a dinosaur.Under cladistic analysis, Coelophysidae was first defined by Paul Sereno in 1998 as the most recent common ancestor of Coelophysis bauri and Procompsognathus triassicus, and all of that common ancestor's descendants. However, Tykoski (2005) has advocated for the definition to change to include the additional taxa of "Syntarsus" kayentakatae and Segisaurus halli. Coelophysidae is part of the superfamily Coelophysoidea, which in turn is a subset of the larger Neotheropoda clade. As part of Coelophysoidea, Coelophysidae is often placed as sister to the Dilophosauridae family, however, the monophyly of this clade has often been disputed. The older term "Podokesauridae", named 14 years prior to Coelophysidae (which would normally grant it priority), is now usually ignored, since its type specimen was destroyed in a fire and can no longer be compared to new finds.

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.

Daemonosaurus

Daemonosaurus (pron.:"DAY-mow-no-SORE-us") is an extinct genus of theropod dinosaur from the Late Triassic of New Mexico. Fossils have been found from deposits in the Chinle Formation, which is latest Triassic in age. While theropods had diversified into several specialized groups by this time, Daemonosaurus is a basal theropod that lies outside the clade Neotheropoda. Daemonosaurus is unusual among early theropods in that it had a short skull and long protruding teeth.

Dilophosauridae

Dilophosauridae is a family of medium to large sized theropod dinosaurs. The name Dilophosauridae is derived from Greek, with “di” meaning “two,” “lophos” meaning “crest,” “sauros” meaning “lizard,” and “idae” meaning “family”. While the name suggests that all dilophosaurids have two crests, this is not applicable to all dilophosaurids. The Dilophosauridae is anchored by the genus Dilophosaurus, and therefore the name comes from the distinctive two crests of the genus.

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.

Dracovenator

Dracovenator () is a genus of dilophosaurid theropod dinosaur that lived approximately 201 to 199 million years ago during the early part of the Jurassic Period in what is now South Africa. Dracovenator was a medium-sized, moderately-built, ground-dwelling, bipedal carnivore, that could grow up to an estimated 7 m (23.0 ft) long. Its type specimen was based on only a partial skull that was recovered.

Jeholosauridae

Jeholosaurids were herbivorous neornithischian dinosaurs from the Cretaceous Period (Aptian - Santonian, with a possible Campanian record) of Asia. The family was first proposed by Han et al. in 2012. The jeholosaurids were defined as those ornithischians more closely related to Jeholosaurus shangyuanensis than to Hypsilophodon foxii, Iguanodon bernissartensis, Protoceratops andrewsi, Pachycephalosaurus wyomingensis, or Thescelosaurus neglectus. The Jeholosauridae includes the type genus Jeholosaurus and Yueosaurus.

Jingshanosaurus

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

Lepidus praecisio

Lepidus is a genus of extinct coelophysoidean theropod from the Upper Triassic of the United States. It lived in the Otis Chalk localities of the Dockum Group in Texas, around 223 million years ago.

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.

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.

Panguraptor

Panguraptor ("Pangu [a Chinese god] plunderer") is a genus of coelophysid theropod dinosaur known from fossils discovered in Lower Jurassic rocks of southern China. The type and only known species is Panguraptor lufengensis. The generic name refers to the deity Pangu but also to the supercontinent Pangaea for which in a geological context the same characters are used: 盘古. Raptor means "seizer", "robber" in Latin. The specific name is a reference to the Lufeng Formation.The holotype specimen was recovered on 12 October 2007 from the Lufeng Formation of Yunnan, which is noted for sauropodomorph fossils. It was described in 2014 by You Hai-Lu and colleagues.

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.

Xixiposaurus

Xixiposaurus is a genus of prosauropod dinosaur which existed in what is now Lower Lufeng Formation, China during the lower Jurassic period. It was first named by Sekiya Toru in 2010 and the type species is Xixiposaurus suni.

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