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[1]). The genus contains a single species, Carniadactylus rosenfeldi.

Carniadactylus
Temporal range: Late Triassic, 228 Ma
Carniadactylus rosenfeldi
Holotype
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Order: Pterosauria
Family: Eudimorphodontidae
Subfamily: Eudimorphodontinae
Genus: Carniadactylus
Dalla Vecchia, 2009
Species:
C. rosenfeldi
Binomial name
Carniadactylus rosenfeldi
(Dalla Vecchia, 1995)
Synonyms

Eudimorphodon rosenfeldi Dalla Vecchia, 1995

Description

Carniadactylus was similar in appearance and anatomy to its close relative Eudimorphodon, though it was significantly smaller. Like Eudimorphodon, it is notable for its complex multi-cusped teeth. Despite their similarities, the size difference between these two pterosaurs likely meant that they occupied different niches and relied on different food sources. This is supported by studies of their teeth. While similar in construction, the teeth of Carniadactylus show little to no wear, unlike the larger, fish-eating Eudimorphodon, which may have been able to chew its food. The smaller Carniadactylus probably fed on smaller, soft-bodied prey like worms and insect larvae.[2]

Classification

In 1995 the Italian paleontologist Fabio Marco Dalla Vecchia named a new species of the genus Eudimorphodon: E. rosenfeldi. The specific name honours the finder Corrado Rosenfeld.[3] The holotype was MFSN 1797, a partial fossil skeleton with parts of the skull and lower jaws, but lacking the tail, found near Udine.

It soon became clear however, that in cladistic analyses E. rosenfeldi was not the sister taxon of the type species of Eudimorphodon: E. ranzii. This made, dependent on the precise analysis, the genus paraphyletic or polyphyletic.

To avoid this Dalla Vecchia in 2009 created the new genus Carniadactylus. The type species is Carniadactylus rosenfeldi. The genus name is derived from Carnia, the name of the region the fossil was found, and Greek daktylos, "finger", a reference to the wing finger typical of pterosaurs. A second specimen, MPUM 6009, is the paratype, consisting of an almost complete skeleton that however has been largely preserved as an impression only. It is a third shorter than the holotype, that itself indicated a wingspan of about seventy centimetres. The disparity was by Dalla Vecchia explained as intraspecific variability.[4] In 2015 however, Alexander Kellner named a separate genus for MPUM 6009: Bergamodactylus.[5]

According to earlier analyses by Alexander Kellner, Carniadactylus was thought to be related to Peteinosaurus within the Dimorphodontidae. David Unwin later placed it into the Campylognathoididae. This was supported by an analysis by Dalla Vecchia that showed Carniadactylus as the sister taxon of Caviramus. However, a more thorough phylogenetic analysis by Andres & Myers in 2013 supported the original interpretation of Carniadactylus as the sister taxon to the type species of Eudimorphodon, and they reclassified it within that genus.[6] The following phylogenetic analysis follows the topology of Upchurch et al. (2015).[7]

Eopterosauria

Preondactylus buffarinii

Austriadactylus cristatus

Peteinosaurus zambellii

Eudimorphodontidae
Raeticodactylinae

Raeticodactylus filisurensis

Caviramus schesaplanensis

Eudimorphodontinae

Arcticodactylus cromptonellus

Carniadactylus rosenfeldi

Eudimorphodon ranzii

See also

References

  1. ^ Barrett, P. M., Butler, R. J., Edwards, N. P., & Milner, A. R. (2008). Pterosaur distribution in time and space: an atlas. Zitteliana, 61-107. [1]
  2. ^ Osi, A. (2010). "Feeding-related characters in basal pterosaurs: implications for jaw mechanism, dental function and diet." Lethaia, doi:10.1111/j.1502-3931.2010.00230.x
  3. ^ Dalla Vecchia F.M., 1995, "A new pterosaur (Reptilia, Pterosauria) from the Norian (Late Triassic) of Friuli (Northeastern Italy), Preliminary note". Gortania, 16 (1994): 59-66
  4. ^ Dalla Vecchia, Fabio M. (2009). "Anatomy and systematics of the pterosaur Carniadactylus (gen. n.) rosenfeldi (Dalla Vecchia, 1995)". Rivista Italiana de Paleontologia e Stratigrafia 115 (2): 159-188.
  5. ^ Kellner, Alexander W.A., 2015, "Comments on Triassic pterosaurs with discussion about ontogeny and description of new taxa", Anais da Academia Brasileira de Ciências 87(2): 669-689
  6. ^ Andres, B.; Myers, T. S. (2013). "Lone Star Pterosaurs". Earth and Environmental Science Transactions of the Royal Society of Edinburgh: 1. doi:10.1017/S1755691013000303.
  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.
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.

Bergamodactylus

Bergamodactylus is a genus of basal pterosaur living during the Late Triassic (early Norian) in the area of present-day Bergamo province in Italy. Its only species is Bergamodactylus wildi. It was previously regarded as a juvenile Eudimorphodon or as identical to Carniadactylus.In 1978, Rupert Wild described a small pterosaur specimen in the collection of the Museo di Paleontologia dell´Università di Milano, found near Cene in Lombardy. He referred to it as the "Milan Exemplar" and identified it as a juvenile of Eudimorphodon ranzii. Wild noted considerable differences with the latter's type specimen but these were explained as reflecting the young age of the animal.In 2009, Fabio Marco Dalla Vecchia confirmed an earlier conclusion by Alexander Kellner that the specimen must have been at least subadult in view of the fusion of the scapula and the coracoid, the upper wristbones being fused into a syncarpal, and the fusion of the extensor process on the first wing phalanx. Dalla Vecchia referred the specimen to Carniadactylus.In 2015, Kellner concluded that the Milan Exemplar represented a different species from Carniadactylus. It showed differences in build that could not be explained by individual variation, it was much smaller though of similar age, and it was of a younger geological age. He named a separate genus and species Bergamodactylus wildi. The generic name combines a reference to Bergamo with a Greek δάκτυλος, daktylos, "finger", a usual suffix in pterosaur names since Pterodactylus. The specific name honours Wild.The holotype, MPUM 6009, was found in a layer of the Calcari di Zorzino Formation dating from the early Norian (upper Alaunian). It consists of a partial skeleton including the skull, compressed on a single plate. It is largely articulated and includes the lower jaws, most of the wings, much of the vertebral column except the tail, and hindlimb elements. Some bones have only been preserved as impressions.Bergamodactylus is one of the smallest known pterosaurs: Kellner in 2015 estimated the wingspan at just 465 millimetres. He also established some distinguishing traits. The postorbital bone is slender with a thin branch towards the frontal bone. The praemaxilla does not reach the lower rim of the external nostril. The fourth metacarpal is short, with only 40% of the length of the humerus and 30% of the length of the ulna. The thighbone is short, attaining just half of the length of either the ulna or the first wing finger phalanx.Bergamodactylus has multi-cusped teeth like Eudimorphodon but their number strongly differs: fourteen in both the upper jaw and the lower jaw as against respectively twenty-nine and twenty-eight in the latter species. Additional differences with Carniadactylus include a tooth row that extends further to the rear, a lower mandibula, a higher placed deltopectoral crest on the humerus and a shorter upper part of the kinked pteroid. Bergamodactylus has a short second phalanx of the wing finger in common with Carniadactylus.Kellner placed Bergamodactylus, within the Novialoidea, in the Campylognathoidea.

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.

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.

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).

Eudimorphodon

Eudimorphodon was a pterosaur that was discovered in 1973 by Mario Pandolfi in the town of Cene, Italy and described the same year by Rocco Zambelli. The nearly complete skeleton was retrieved from shale deposited during the Late Triassic (mid to late Norian stage), making Eudimorphodon one of the oldest pterosaurs known. It had a wingspan of about 100 centimetres (3.3 ft) and at the end of its long bony tail may have been a diamond-shaped flap like in the later Rhamphorhynchus. If so, the flap may have helped it steer while maneuvering in the air. Eudimorphodon is known from several skeletons, including juvenile specimens.

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.

Orionides

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

Peteinosaurus

Peteinosaurus ( peh-TY-nə-SOR-əs; 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.

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.

Seazzadactylus

Seazzadactylus is a basal pterosaur genus that during the late Triassic lived in the area of present Italy.

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