Bergamodactylus Temporal range: Late Triassic, | |
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The "Milan Exemplar" | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Order: | † Pterosauria |
Family: | † Campylognathoididae |
Genus: | † Bergamodactylus Kellner, 2015 |
Type species | |
†Bergamodactylus wildi Kellner, 2015 |
Bergamodactylus is a putative genus of basal pterosaur which lived during the Late Triassic (early Norian) in the area of present-day Bergamo province in Italy. Its only species is Bergamodactylus wildi. It was named in 2015 based on a pterosaur specimen which had previously been regarded as a juvenile Eudimorphodon or as identical to Carniadactylus . [1] Some Triassic pterosaur specialists consider the distinguishing features of Bergamodactylus to be invalid or insufficient to distinguish it from Carniadactylus, and thus retain the specimen in that genus. [2]
In 1978, Rupert Wild described a small pterosaur specimen in the collection of the Museo di Paleontologia dell´Università di Milano , found near Cene, 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. [3]
In 2009, Fabio Marco Dalla Vecchia confirmed an earlier conclusion by Alexander Kellner [4] 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 rosenfeldi . [5]
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.Kellner placed Bergamodactylus, within the Novialoidea, in the Campylognathoidea. [1]
In 2018, Dalla Vecchia argued that Kellner's observations on development in pterosaurs were oversimplified, and that the Milan Exemplar's distinguishing features were ambiguous, invalid, or individual variation at best. As a result, Dalla Vecchia referred the specimen back to Carniadactylus, rendering Bergamodactylus wildi a junior synonym of Carniadactylus rosenfeldi. [2]
The holotype, MPUM 6009, was found in a layer of the Calcari di Zorzino Formation dating from the early Norian (upper Alaunian). [1] 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. [3]
Bergamodactylus is one of the smallest known pterosaurs: Kellner in 2015 estimated the wingspan at just 465 millimeters (18.3 inches). 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. [1]
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. [1]
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, making Eudimorphodon one of the oldest pterosaurs known. It had a wingspan of about 100 centimeters (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.
Preondactylus is a genus of long-tailed pterosaurs from the Late Triassic 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.
Peteinosaurus was a prehistoric genus of pterosaur. It lived in the late Triassic period in the late Norian age, and at a wingspan of around 60 cm (24 in), was one of the smallest and earliest pterosaurs, although other estimates suggest a wingspan of up to 1 m (3.3 ft).
Cene is a comune (municipality) in the Province of Bergamo in the Italian region of Lombardy, located about 60 kilometres (37 mi) northeast of Milan and about 15 kilometres (9 mi) northeast of Bergamo.
Caviramus is a genus of caviramid pterosaur from the Late Triassic lower Kössen Formation of the Northern Calcareous Alps of Switzerland.
Megalancosaurus is a genus of extinct reptile from the Late Triassic Dolomia di Forni Formation and Zorzino Limestone of northern Italy, and one of the best known drepanosaurids. The type species is M. preonensis; a translation of the animal's scientific name would be "long armed reptile from the Preone Valley."
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.
Vallesaurus is an extinct genus of Late Triassic elyurosaur drepanosauromorph. First found in Northern Italy in 1975, it is one of the most primitive drepanosaurs. V. cenenis is the type species, which was first mentioned in 1991 but only formally described in 2006. A second species, V. zorzinensis, was named in 2010.
Langobardisaurus is an extinct genus of tanystropheid archosauromorph reptile, with one valid species, L. pandolfii. Its fossils have been found in Italy and Austria, and it lived during the Late Triassic period, roughly 228 to 201 million years ago. Langobardisaurus was initially described in 1994, based on fossils from the Calcare di Zorzino Formation in Northern Italy. Fossils of the genus are also known from the Forni Dolostone of Northern Italy and the Seefeld Formation of Austria.
Bobosaurus is an extinct genus of sauropterygian reptile related to plesiosaurs. It is based on the holotype MFSN 27285, a partial skeleton found in Early Carnian-age rocks of the Rio del Lago Formation, northeastern Italy. Bobosaurus was named in 2006 by Fabio M. Dalla Vecchia and the type species is B. forojuliensis. It may be a pistosaurid, or closer to Plesiosauria. A recent cladistic analysis found it to be a pistosaur. It was relatively large animal, with more than 3 m (9.8 ft) in length.
The Forni Dolostone, also known as the Dolomia di Forni, is a Late Triassic dolomite geological formation in northeastern Italy. The formation was deposited in a lagoonal to shallow marine environment.
Eosaurichthys is an extinct genus of saurichthyid ray-finned fish that lived during the late Permian epoch in what is now China.
Carniadactylus is a genus of pterosaur which existed in Europe during the Late Triassic period. The genus contains a single species, Carniadactylus rosenfeldi.
Sinosaurichthys is an extinct genus of saurichthyid ray-finned fish, which existed in southwestern China during the Middle Triassic. Fossils have been found in the Upper Member of the Guanling Formation of two localities: Yangjuan of Panxian County, Guizhou Province, and Dawazi of Luoping, Yunnan Province, China.
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 is an extinct family of early pterosaurs from the Late Triassic 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, but they didn't defined the clade. Brian Andres define Eudimorphodontidae and found Peteinosaurus to be most closely related to it. Furthermore, he found monophyletic Eudimorphodon clade, 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.
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.
Austriadraco is a genus of pterosaur living during the Late Triassic in the area of present Austria. Its only species—Austriadraco dallavecchiai—was previously attributed to Eudimorphodon, and its closest relatives may have been Eudimorphodon or Arcticodactylus.
Seazzadactylus is a basal pterosaur genus that lived during the late Triassic in the area of present Italy.
Raibliania is an extinct genus of tanystropheid archosauromorph discovered in the Calcare del Predil Formation in Italy. It lived during the Carnian stage of the Late Triassic and it was related to Tanystropheus. Raibliania is distinct from Tanystropheus due to some distinct features of the cervical vertebrae and teeth. The type species is Raibliania calligarisi, named in 2020. The holotype consists of a partial post-cranial skeleton, with the known elements including vertebrae, a single tooth, several ribs, gastralia and parts of the pelvis.