''Carniadactylus'' is a genus of

pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the Order (biology), order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cre ...

which existed in

Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a subcontinent of Eurasia and it is located enti ...

during the

Late Triassic The Late Triassic is the third and final epoch (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch ...

period (late

Carnian The Carnian (less commonly, Karnian) is the lowermost stage of the Upper Triassic Series (or earliest age of the Late Triassic Epoch). It lasted from 237 to 227 million years ago (Ma). The Carnian is preceded by the Ladinian and is followe ...

or 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 ...

, about 228 million years agoBarrett, P. M., Butler, R. J., Edwards, N. P., & Milner, A. R. (2008). Pterosaur distribution in time and space: an atlas. ''Zitteliana'', 61-107

/ref>). The genus contains a single species, ''Carniadactylus rosenfeldi''.

Description

''Carniadactylus'' was similar in appearance and anatomy to its close relative ''

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 ...

'', 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.Osi, A. (2010). "Feeding-related characters in basal pterosaurs: implications for jaw mechanism, dental function and diet." ''Lethaia'',

Classification

In 1995 the Italian

paleontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of foss ...

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 ...

named a new species of the genus ''

'': ''E. rosenfeldi''. The

specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...

honors the finder

Corrado Rosenfeld Corrado may refer to: Places * Anticoli Corrado, comune in the City of Rome * Monte Vidon Corrado, comune in the Province of Fermo People Given name *See Corrado (given name) Surname * Andrea Corrado (1873-1963), Italian ship owner * Andrea Di Co ...

. 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 ...

was MFSN 1797, a partial fossil skeleton with parts of the skull and lower jaws, but lacking the tail, found near

Udine Udine ( , ; fur, Udin; la, Utinum) is a city and ''comune'' in north-eastern Italy, in the middle of the Friuli Venezia Giulia region, between the Adriatic Sea and the Alps (''Alpi Carniche''). Its population was 100,514 in 2012, 176,000 with ...

. It soon became clear however, that in

cladistic Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived ch ...

analyses ''E. rosenfeldi'' was not the sister taxon of the

type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen( ...

of ''Eudimorphodon'': ''E. ranzii''. This made, dependent on the precise analysis, the genus

paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...

polyphyletic A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of conver ...

. 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 Carnia ( fur, Cjargne or ''Cjargna''/''Cjargno'' in local variants, vec, Ciargna, german: Karnien, sl, Karnija) is a historical-geographic region in the northeastern Italian area of Friuli. Its 27 municipalities all belong to the Province of U ...

, 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 In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype nor a syntype). O ...

, 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. In 2015 however,

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 ...

named a separate genus for MPUM 6009: ''

Bergamodactylus ''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 p ...

''. According to earlier analyses by

, ''Carniadactylus'' was thought to be related to '' Peteinosaurus'' within the

Dimorphodontidae Dimorphodontidae (or dimorphodontids) is a group of early " rhamphorhynchoid" pterosaurs named after ''Dimorphodon'', that lived in the Late Triassic to Early Jurassic. While fossils that can be definitively referred to the group are rare, dim ...

. David Unwin later placed it into the

Campylognathoididae 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., ...

. 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. The following phylogenetic analysis follows the topology of Upchurch ''et al.'' (2015). In 2020 however, a study upheld by Matthew G. Baron about early pterosaur interrelationships found ''Carniadactylus'' to group with '' Caviramus'', ''

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 ...

'', and the

Austriadraconidae Caviramidae is a group of basal pterosaurs. It was erected by paleontologist Matthew G. Baron in 2020. It was defined as the least inclusive clade that includes ''Arcticodactylus cromptonellus'' and '' Caviramus schesaplanensis''. The members ...

, which in turn were within a clade called Caviramidae.Matthew G. Baron (2020). "Testing pterosaur ingroup relationships through broader sampling of avemetatarsalian taxa and characters and a range of phylogenetic analysis techniques". PeerJ. 8: e9604. doi:10.7717/peerj.9604. PMC 7512134. .

References

{{Taxonbar, from=Q2078471 Pterosaurs Late Triassic pterosaurs of Europe Fossil taxa described in 2009

Description

Classification

See also

References