Peteinosaurus

Peteinosaurus (/pɛˌtaɪnəˈsɔːrəs/ peh-TY-nə-SOR-əs;[1] meaning "winged lizard") was a prehistoric genus of Pterosaur. It lived in the late Triassic period in the late Norian age (about 221 to 210 million years ago), and at a wingspan of around 60 cm (24 in), was one of the smallest and earliest Pterosaurs.[2]

Peteinosaurus
Temporal range: Late Triassic, 221–210 Ma
Peteinosaurus zambellii
Paratype MCSNB 3359
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Order: Pterosauria
Clade: Eopterosauria
Genus: Peteinosaurus
Wild, 1978
Species:
P. zambelli
Binomial name
Peteinosaurus zambelli
Wild, 1978

Discovery

Peteinosaurus fossil
Fossil in Museo civico di scienze naturali di Bergamo

Three fossils have been found near Cene, Italy.[3] The first fossil, the holotype MCSNB 2886, is fragmentary and disarticulated. The second, the articulated paratype MCSNB 3359, lacks any diagnostic features of Peteinosaurus and thus might be a different species. This paratype has a long tail (20 cm) made more stiff by long extensions of the vertebrae; this feature is common among pterosaurs of the Triassic. The third example is MCSNB 3496, another fragmentary skeleton. All specimens are those of subadults and of none has the skull been preserved.

Like most pterosaurs, Peteinosaurus had bones that were strong but very light. Peteinosaurus is trimorphodontic, with three types of conical teeth. An insectivorous lifestyle has been attributed to Peteinosaurus.[3] The fifth toe of Peteinosaurus was long and clawless. Its joint allowed it to flex in a different plane than the other phalanges in order to control the cruropatagium, as seen preserved in the specimen of Sordes pilosus PIN 2585.3.

The genus has been described by the German paleontologist Rupert Wild in 1978.[4] The type species is Peteinosaurus zambellii. The genus name is derived from Greek peteinos, "winged" and sauros, "lizard", the latter being used to indicate any saurian. The specific name, zambellii, honours Rocco Zambelli, the curator of the Bergamo natural history museum.

Phylogeny and classification

Peteinosaurus is one of the oldest-known pterosaurs, and at a mere sixty centimetres, had a tiny wingspan when compared to some later genera, such as Pteranodon whose wingspan exceeded twenty feet. Its wings were also proportionally smaller than those of later pterosaurs, as its wing length was only twice the length of the hindlimb.[3] All other known pterosaurs have wingspans at least three times the length of their hindlimbs.[3] It also had single cusped teeth that lacked the specialized heterodonty present in the other Italian Triassic pterosaur genus, Eudimorphodon.[3]

All these factors converge to hint that Peteinosaurus belongs to a group that possibly represents the most basal known pterosaurs: the Dimorphodontidae, to which it was assigned in 1988 by Robert L. Carroll.[5] The only other known member of that group is the later genus Dimorphodon, which lent its name to the family including both genera.[3] Later cladistic analyses however, have not shown a close connection between the two forms. Nevertheless, the possible basal position of Peteinosaurus has been affirmed by Fabio Marco Dalla Vecchia who suggested that Preondactylus, according to David Unwin the most basal pterosaur, might be a subjective junior synonym of Peteinosaurus. A 2010 cladistic analysis by Brian Andres and colleagues placed Peteinosaurus in Lonchognatha which includes Eudimorphodon and Austriadactylus as more basal.[6] The following phylogenetic analysis follows the topology of Upchurch et al. (2015).[7]

Coelophysis and Peteinosaurus
Restoration of Coelophysis and Peteinosaurus in their environment
Eopterosauria

Preondactylus buffarinii

Austriadactylus cristatus

Peteinosaurus zambellii

Eudimorphodontidae
Raeticodactylinae

Raeticodactylus filisurensis

Caviramus schesaplanensis

Eudimorphodontinae

Arcticodactylus cromptonellus

Carniadactylus rosenfeldi

Eudimorphodon ranzii

Mark Witton, however, considers Peteinosaurus to be a true dimorphodontid.[8]

See also

References

  1. ^ Creisler, B. (2003). "Pterosauria Translation and Pronunciation Guide" Archived 2007-11-09 at the Wayback Machine, Dinosauria On-Line.
  2. ^ Haines, Tim; Chambers, Paul (2005). The complete guide to prehistoric life.
  3. ^ a b c d e f "Peteinosaurus." In: Cranfield, I. (ed.) (2000). The Illustrated Directory of Dinosaurs and Other Prehistoric Creatures. London: Salamander Books, Ltd. Pp. 282-283. ISBN 1-84065-241-1.
  4. ^ Wild, R. (1978). "Die Flugsaurier (Reptilia, Pterosauria) aus der Oberen Trias von Cene bei Bergamo, Italien", Bolletino della Societa Paleontologica Italiana, 17 (2): 176-256.
  5. ^ Carroll, R. L. (1988). Vertebrate Paleontology and Evolution. New York: W. H. Freeman and Company. ISBN 0-7167-1822-7.
  6. ^ Andres, B., J. M. Clark, and X. Xu (2010). "A new rhamphorhynchid pterosaur from the Upper Jurassic of Xinjiang, China, and the phylogenetic relationships of basal pterosaurs", Journal of Vertebrate Paleontology 30 (1): 163-187.
  7. ^ Upchurch, P.; Andres, B.B.; Butler, R.J.; Barrett, P.M. (2015). "An analysis of pterosaurian biogeography: implications for the evolutionary history and fossil record quality of the first flying vertebrates". Historical Biology. 27 (6): 697–717. doi:10.1080/08912963.2014.939077. PMC 4536946.
  8. ^ Pterosaurs: Natural History, Evolution, Anatomy, 2013
Arcticodactylus

Arcticodactylus is a genus of basal pterosaur living during the Late Triassic in the area of present Greenland. Its only species was previously attributed to Eudimorphodon, and its closest relatives may have been Eudimorphodon or Austriadraco.

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.

Caelestiventus

Caelestiventus ( sə-LES-tih-VEN-təs, meaning "heavenly wind") is a pterosaur genus from the Late Triassic (Norian or Rhaetian) found in western North America. The type species, Caelestiventus hanseni, honors Robin Hansen, the Bureau of Land Management geologist (BLM), who facilitated access to the excavation site.

Caelestiventus is important because it is the sole example of a desert-dwelling non-pterodactyloid pterosaur and is 65 million years older than other known desert-dwelling pterosaurs. Additionally, it shows that even the earliest pterosaurs were morphologically and ecologically diverse and that the Dimorphodontidae originated in the Triassic Period.

Carniadactylus

Carniadactylus is a genus of pterosaur which existed in Europe during the Late Triassic period (late Carnian or early Norian, about 228 million years ago). The genus contains a single species, Carniadactylus rosenfeldi.

Caviramus

Caviramus is a genus of "rhamphorhynchoid" pterosaur from the Late Triassic (late Norian-early Rhaetian-age) lower Kössen Formation of the Northern Calcareous Alps of Switzerland.

The genus was in 2006 named by Nadia Fröbisch and Jörg Fröbisch. The type species is Caviramus schesaplanensis. The genus name is derived from Latin cavus, "hollow" and ramus, "branch". The specific name refers to Mount Schesaplana.

Cerapoda

Cerapoda ("ceratopsians and ornithopods") is a clade of the dinosaur order Ornithischia.

Dimorphodon

Dimorphodon was a genus of medium-sized pterosaur from the early Jurassic Period. It was named by paleontologist Richard Owen in 1859. Dimorphodon means "two-form tooth", derived from the Greek δι (di) meaning "two", μορφη (morphe) meaning "shape" and οδων (odon) meaning "tooth", referring to the fact that it had two distinct types of teeth in its jaws – which is comparatively rare among reptiles.

Dimorphodontia

Dimorphodontia is a group of early "rhamphorhynchoid" pterosaurs named after Dimorphodon, that lived in the Late Triassic to Early Jurassic.

A family, Dimorphodontidae, was named in 1870 by Harry Govier Seeley (as "Dimorphodontae") with Dimorphodon as the only known member. In 2003 David Unwin defined a clade Dimorphodontidae, as the group consisting of the last common ancestor of Dimorphodon macronyx and Peteinosaurus zambellii, and all its descendants. However, later studies found that Dimorphodon may not be closely related to Peteinosaurus, so this definition of Dimorphodontidae would therefore be superfluous. In 2014, Brian Andres and colleagues defined another clade, Dimorphodontia, as a replacement. Dimorphodontia would include all pterosaurs more closely related to Dimorphodon than to Pterodactylus. According to the analysis published by Andres et al., Dimorphodontia is also a small group, including only Dimorphodon and Parapsicephalus.In 2018, a close relative of Dimorphodon was described from the Late Triassic of North America by Britt and colleagues, and was named Caelestiventus. This discovery expanded the geographic, temporal and also the ecological range of dimorphodontians, as it was discovered in the Late Triassic Nugget Sandstone in Utah, which was a desert at the time. Britt and colleagues also redefined Dimorphodontidae as the least inclusive clade containing Dimorphodon macronyx and Caelestiventus hanseni.

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.

Eopterosauria

Eopterosauria is a group of basal pterosaurs from the Triassic, which form their own clade. The term was first used in Andres et al. (2014) to include Preondactylus, Austriadactylus, Peteinosaurus and Eudimorphodontidae. Inside the group were two other new clades, Preondactylia, which included Preondactylus and Austriadactylus, and Eudimorphodontoidea, to include Eudimorphodontidae and Raeticodactylidae. Eopterosauria was defined as "the least inclusive clade containing Preondactylus buffarinii and Eudimorphodon ranzii". The specimen BSP 1994, previously assigned to Eudimorphodon, was named the separate taxon Austriadraco in 2015, and assigned to the new family Austriadraconidae, but further classification was not described. The following phylogenetic analysis follows the topology of Andres et al. (2014).

Eudimorphodontidae

Eudimorphodontidae is an extinct family of early pterosaurs from the Late Triassic (early Norian to Rhaetian age) of Europe. It was named by Peter Wellnhofer in 1978 to include Eudimorphodon ranzii. Some phylogenetic analyses suggested that Eudimorphodontidae is a junior synonym of Campylognathoididae, however more comprehensive analyses found Eudimorphodontidae to be basal to Macronychoptera that includes Campylognathoididae and more derived pterosaurs (Breviquartossa). Wang et al. (2009) found Eudimorphodontidae to include six species (the monospecific Peteinosaurus, Raeticodactylus and Caviramus, and three species of Eudimorphodon), but they didn't defined the clade. Brian Andres (2010, in press) define Eudimorphodontidae and found Peteinosaurus to be most closely related to it. Furthermore, he found monophyletic Eudimorphodon clade (unlike Wang et al., 2009 and Dalla Vecchia, 2009), and defined two subfamilies within Eudimorphodontidae. The Eudimorphodontinae includes all taxa more closely related to Eudimorphodon ranzii than to Raeticodactylus filisurensis while the Raeticodactylinae includes all taxa more closely related to Raeticodactylus filisurensis than to Eudimorphodon ranzii. More recently, Raeticodactylus and Caviramus were moved into their own family, Raeticodactylidae. The below cladogram follows that analysis.

Jingshanosaurus

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

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.

Norian

The Norian is a division of the Triassic geological period. It has the rank of an age (geochronology) or stage (chronostratigraphy). The Norian lasted from ~227 to 208.5 million years ago. It was preceded by the Carnian and succeeded by the Rhaetian.

Orionides

Orionides is a clade of tetanuran theropod dinosaurs from the Middle Jurassic to the Present. The clade includes most theropod dinosaurs, including birds.

Preondactylus

Preondactylus is a genus of long-tailed pterosaurs from the Late Triassic (late Carnian or early Norian age, about 228 million years ago) that inhabited what is now Italy. It contains a single known species, Preondactylus buffarinii, which was discovered by Nando Buffarini in 1982 at the Forni Dolostone near Udine in the Preone valley of the Italian Alps.

Raeticodactylidae

Raeticodactylidae is a family of eudimorphodontoid eopterosaurian pterosaurs that lived in Switzerland during the Late Triassic. The family includes Caviramus, and the type genus Raeticodactylus, which are both known from the Kössen Formation, around 205 mya. Raeticodactylidae was first used in 2014 by Andres et al., as a group of all pterosaurs closer to Raeticodactylus than Eudimorphodon. The following phylogenetic analysis follows the topology of Andres et al. (2014).

Raeticodactylus

Raeticodactylus is a genus of non-pterodactyloid pterosaur from the late Norian-early Rhaetian-age Upper Triassic lower Kössen Formation of the central Austroalpine of Grisons, Switzerland. It is known from holotype BNM 14524, a single disarticulated partial skeleton including an almost complete skull, found in August 2005. This genus was named and described in 2008 by its discoverer Rico Stecher; the type species is Raeticodactylus filisurensis. The specific name refers to Filisur.

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