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''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, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 to ...
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 Continent#Subcontinents, subcontinent of Eurasia ...
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 followed by t ...
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 defi ...
, 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 (mi ...
'', 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 Italian(s) may refer to: * Anything of, from, or related to the people of Italy over the centuries ** Italians, an ethnic group or simply a citizen of the Italian Republic or Italian Kingdom ** Italian language, a Romance language *** Regional Ita ...
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 fossi ...
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 ''
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 (mi ...
'': ''E. rosenfeldi''. The specific name honors the finder Corrado Rosenfeld. 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 several ...
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 t ...
. 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 char ...
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 ...
or
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 converg ...
. 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 Ud ...
, 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). Of ...
, 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
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 ...
, ''Carniadactylus'' was thought to be related to ''
Peteinosaurus ''Peteinosaurus'' ( ; 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 , was one of the smallest and earli ...
'' 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, dimorp ...
.
David Unwin David (; , "beloved one") (traditional spelling), , ''Dāwūd''; grc-koi, Δαυΐδ, Dauíd; la, Davidus, David; gez , ዳዊት, ''Dawit''; xcl, Դաւիթ, ''Dawitʿ''; cu, Давíдъ, ''Davidŭ''; possibly meaning "beloved one". w ...
later placed it into the Campylognathoididae. This was supported by an analysis by Dalla Vecchia that showed ''Carniadactylus'' as the sister taxon of ''
Caviramus ''Caviramus'' is a genus of caviramid pterosaur from the Late Triassic (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 ...
''. 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 ''Caviramus'' is a genus of caviramid pterosaur from the Late Triassic (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 ...
'', ''
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 dis ...
'', 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 of ...
, which in turn were within a clade called
Caviramidae 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 o ...
.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. .


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 ...
*
Timeline of pterosaur research This timeline of pterosaur research is a chronologically ordered list of important fossil discoveries, controversies of interpretation, and taxonomic revisions of pterosaurs, the famed flying reptiles of the Mesozoic era. Although pterosaurs w ...


References

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