Bergamodactylus
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''Bergamodactylus'' is a putative genus of basal
pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 ...
which lived during the
Late Triassic The Late Triassic is the third and final epoch of the Triassic Period in the geologic time scale, spanning the time between Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch and followed by the Early Jurassic Epoch ...
(early
Norian The Norian is a division of the Triassic Period. It has the rank of an age ( geochronology) or stage (chronostratigraphy). It lasted from ~227 to million years ago. It was preceded by the Carnian and succeeded by the Rhaetian. Stratigraphic ...
) in the area of present-day
Bergamo province The Province of Bergamo ( it, provincia di Bergamo; lmo, proìnsa de Bèrghem) is a province in the Lombardy region of Italy. It has a population of 1,112,187 (2017), an area of , and contains 243 ''comuni''. Its capital is the city of Bergamo. ...
in
Italy Italy ( it, Italia ), officially the Italian Republic, ) or the Republic of Italy, is a country in Southern Europe. It is located in the middle of the Mediterranean Sea, and its territory largely coincides with the homonymous geographical ...
. 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 ''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 (m ...
'' or as identical to ''
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 agoBarrett, P. M., Butler, R. J., Edwards, N. P., & Milner, A. R. (2008). Pterosaur distribu ...
''. 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.


History

In 1978,
Rupert Wild Rupert may refer to: People * Rupert (name), various people known by the given name or surname "Rupert" Places Canada *Rupert, Quebec, a village * Rupert Bay, a large bay located on the south-east shore of James Bay *Rupert River, Quebec * Ruper ...
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 In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to anchor or centralizes th ...
but these were explained as reflecting the young age of the animal.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 In 2009,
Fabio Marco Dalla Vecchia Fabio is a given name descended from Latin '' Fabius'' and very popular in Italy and Latin America (due to Italian migration). Its English equivalent is Fabian. The name is written without an accent in Italian and Spanish, but is usually accented ...
confirmed an earlier conclusion by
Alexander Kellner Alexander Wilhelm Armin Kellner (born September 26, 1961) is a Brazilian geologist and paleontologist who is a leading expert in the field of studying pterosaurs. His research has focused mainly on fossil reptiles from the Cretaceous Period, i ...
that the specimen must have been at least subadult in view of the fusion of the scapula and the
coracoid A coracoid (from Greek κόραξ, ''koraks'', raven) is a paired bone which is part of the shoulder assembly in all vertebrates except therian mammals (marsupials and placentals). In therian mammals (including humans), a coracoid process is prese ...
, 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 ''Carniadactylus'' is a genus of pterosaur which existed in Europe during the Late Triassic period (late Carnian or early Norian, about 228 million years agoBarrett, P. M., Butler, R. J., Edwards, N. P., & Milner, A. R. (2008). Pterosaur distri ...
''. 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 Bergamo (; lmo, Bèrghem ; from the proto- Germanic elements *''berg +*heim'', the "mountain home") is a city in the alpine Lombardy region of northern Italy, approximately northeast of Milan, and about from Switzerland, the alpine lakes Com ...
with a Greek δάκτυλος, ''daktylos'', "finger", a usual suffix in pterosaur names since ''
Pterodactylus ''Pterodactylus'' (from Greek () meaning 'winged finger') is an extinct genus of pterosaurs. It is thought to contain only a single species, ''Pterodactylus antiquus'', which was the first pterosaur to be named and identified as a flying rept ...
''. The specific name honours Wild.Kellner placed ''Bergamodactylus'', within the
Novialoidea Novialoidea (meaning "new wings") is an extinct clade of macronychopteran pterosaurs that lived from the latest Early Jurassic to the latest Late Cretaceous (early Toarcian to late Maastrichtian ageBarrett, P. M., Butler, R. J., Edwards, N. P., ...
, in the
Campylognathoidea Novialoidea (meaning "new wings") is an extinct clade of macronychopteran pterosaurs that lived from the latest Early Jurassic to the latest Late Cretaceous (early Toarcian to late Maastrichtian ageBarrett, P. M., Butler, R. J., Edwards, N. P., & ...
. 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''.


Description

The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of seve ...
, MPUM 6009, was found in a layer of the
Calcari di Zorzino Formation The Calcare di Zorzino, Italian for Zorzino Limestone is a Late Triassic (Norian) geological formation in Italy ( Cene and ).Norian The Norian is a division of the Triassic Period. It has the rank of an age ( geochronology) or stage (chronostratigraphy). It lasted from ~227 to million years ago. It was preceded by the Carnian and succeeded by the Rhaetian. Stratigraphic ...
(upper
Alaunian The Norian is a division of the Triassic Period. It has the rank of an age (geochronology) or stage (chronostratigraphy). It lasted from ~227 to million years ago. It was preceded by the Carnian and succeeded by the Rhaetian. Stratigraphic d ...
). 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 The ''postorbital'' is one of the bones in vertebrate skulls which forms a portion of the dermal skull roof and, sometimes, a ring about the orbit. Generally, it is located behind the postfrontal and posteriorly to the orbital fenestra. In some v ...
is slender with a thin branch towards the
frontal bone The frontal bone is a bone in the human skull. The bone consists of two portions.'' Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bony part of the forehead, ...
. The
praemaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has ...
does not reach the lower rim of the external nostril. The fourth
metacarpal In human anatomy, the metacarpal bones or metacarpus form the intermediate part of the skeletal hand located between the phalanges of the fingers and the carpal bones of the wrist, which forms the connection to the forearm. The metacarpal bones ar ...
is short, with only 40% of the length of the
humerus The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a r ...
and 30% of the length of the
ulna The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of t ...
. 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 Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 to 6 ...
. ''Bergamodactylus'' has a short second phalanx of the wing finger in common with ''Carniadactylus''.


See also

*
List of pterosaur genera This list of pterosaurs is a comprehensive listing of all genera that have ever been included in the order Pterosauria, excluding purely vernacular terms. The list includes all commonly accepted genera, but also genera that are now considered inval ...


References

{{Taxonbar, from=Q20668509 Late Triassic pterosaurs of Europe Fossil taxa described in 2015 Taxa named by Alexander Kellner Novialoids Fossils of Italy