Arcticodactylus
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''Arcticodactylus'' is a genus of basal
pterosaur Pterosaurs are 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 million to 66 million years ago). Pterosaurs are the earli ...
living 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 a ...
in the area of present-day
Greenland Greenland is an autonomous territory in the Danish Realm, Kingdom of Denmark. It is by far the largest geographically of three constituent parts of the kingdom; the other two are metropolitan Denmark and the Faroe Islands. Citizens of Greenlan ...
. Its only species was previously attributed to ''Eudimorphodon'', and its closest relatives may have been ''
Eudimorphodon ''Eudimorphodon'' is an extinct genus of pterosaur that was discovered in 1973 by Mario Pandolfi in the town of Cene, Lombardy, Cene, Italy and described the same year by Rocco Zambelli. The nearly complete skeleton was retrieved from shale depos ...
'' or '' Austriadraco''.


History of discovery

In 1989, William Amaral on the McKnight Bjerg in the east of Greenland discovered a rich fossil site. It was excavated in 1991 and 1992. Part of the material was a small skeleton of a pterosaur. In 2001, Farish Jenkins, Neil Shubin, Stephen Gatesy and Kevin Padian named and described it as a new species of ''Eudimorphodon'': ''Eudimorphodon cromptonellus''. The specific name honors Professor Alfred Walter Crompton. The suffix ''~ellus'', in
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
indicating a diminutive, alluded to the small size of the specimen. The
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
, MGUH VP 3393, was found in the Carlsberg Fjord Beds of the Ørsted Dal Member of the Fleming Fjord Formation dating from the
Norian The Norian is a division of the Triassic geological period, Period. It has the rank of an age (geology), age (geochronology) or stage (stratigraphy), stage (chronostratigraphy). It lasted from ~227.3 to Mya (unit), million years ago. It was prec ...
–
Rhaetian The Rhaetian is the latest age (geology), age of the Triassic period (geology), Period (in geochronology) or the uppermost stage (stratigraphy), stage of the Triassic system (stratigraphy), System (in chronostratigraphy). It was preceded by the N ...
. It consists of a partial skeleton with skull. It is largely disarticulated. The reference to ''Eudimorphodon'' had been essentially based on the similarity in tooth form, especially the distinctive multi-cuspid build with three, four or five points on the crown. In 2003,
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 ...
pointed out that other basal pterosaurs also possess such teeth. In 2014, Fabio Marco Dalla Vecchia noted that ''E. cromptonellus'' shared not a single trait with ''Eudimorphodon ranzii'' not present in other pterosaurs but lacked the distinguishing fang-like teeth, pterygoid teeth and striated tooth enamel. In 2015, Kellner named a separate genus ''Arcticodactylus''. The generic name is derived from the
Arctic The Arctic (; . ) is the polar regions of Earth, polar region of Earth that surrounds the North Pole, lying within the Arctic Circle. The Arctic region, from the IERS Reference Meridian travelling east, consists of parts of northern Norway ( ...
, and Greek δάκτυλος, ''daktylos'', "finger", a usual suffix in pterosaur names since ''
Pterodactylus ''Pterodactylus'' (from ) is a genus of extinct 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 reptile and one of the first prehis ...
''. The '' Life Science Identifier'' is 72AE012A-018A-4B4B-950F-3CCB4C1D2471. The
type species In International_Code_of_Zoological_Nomenclature, 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 spe ...
is ''Eudimorphodon cromptonellus'', the ''
combinatio nova In biological taxonomy, a combinatio nova (abbreviated comb. nov. or n. comb.) refers to the formal renaming of an organism's scientific name when it is transferred to a different genus, reclassified within a different species, or its taxonomic ...
'' is ''Arcticodactylus cromptonellus''.


Description

The holotype individual of ''Arcticodactylus'' is the smallest pterosaur known, with an estimated
wingspan The wingspan (or just span) of a bird or an airplane is the distance from one wingtip to the opposite wingtip. For example, the Boeing 777–200 has a wingspan of , and a wandering albatross (''Diomedea exulans'') caught in 1965 had a wingsp ...
of just . In 2001, on the basis of
histological Histology, also known as microscopic anatomy or microanatomy, is the branch of biology that studies the microscopic anatomy of biological tissue (biology), tissues. Histology is the microscopic counterpart to gross anatomy, which looks at large ...
research on its bone structure, it was considered not to have been fully grown yet, though not newly born. In 2015, Kellner established some distinguishing traits, correcting and adding to the 2001 diagnosis. The jaws have eleven or twelve multi-cusped teeth per side. The articulation surface of the fourth
metacarpal In human anatomy, the metacarpal bones or metacarpus, also known as the "palm bones", are the appendicular bones that form the intermediate part of the hand between the phalanges (fingers) and the carpal bones ( wrist bones), which articulate ...
with the fourth finger shows two true condyles. The thighbone is only a little shorter than the shinbone, with 96% of its length. The scapula is much longer, 93%, than the
coracoid A coracoid 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 present as part of the scapula, but this is n ...
. The humerus is only slightly shorter than the thighbone, with 92% of its length, or the ulna with 91% of ulnar length. The thighbone is somewhat longer than the first phalanx of the wing finger that has 91% of femoral length. The third
metatarsal The metatarsal bones or metatarsus (: metatarsi) are a group of five long bones in the midfoot, located between the tarsal bones (which form the heel and the ankle) and the phalanges ( toes). Lacking individual names, the metatarsal bones are ...
of the foot is elongated with 56% of shinbone length. These proportions imply that ''Arcticodactylus'' had relatively short wings and large feet. ''Arcticodactylus'' can furthermore be distinguished from ''Eudimorphodon'' in the lack of long fang-like teeth in the middle of the tooth row and from ''Eudimorphodon ranzii'', '' Carniadactylus'' and ''
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 ...
'' by a triangular instead of rectangular deltopectoral crest on the humerus. ''Articodactylus'' has fewer teeth than any other known Triassic pterosaur. Jenkins e.a. claimed that the unique articulation in ''Arcticodactylus'' between the main wing metacarpal and the wing finger, with two rounded condyles, was a transitional shape between the ancestral form that featured a single rounded articulation surface on the metacarpal allowing a considerable amount of lateral movement, and the condition in later pterosaurs that showed a gentle depression or trochlea. The two condyles, the upper one being the largest, would have forced the finger into the most optimal plane of movement during the upstroke of the wing.


Classification

In 2001, ''E. cromptonellus'' was placed in the Eudimorphodontidae. Kellner in 2015 indicated a basal position in the Pterosauria, the short coracoid suggesting a close affinity to '' Austriadraco'' within an Austriadraconidae. According to Kellner, the original describers had incorrectly identified a coracoid as a
quadrate bone The quadrate bone is a skull bone in most tetrapods, including amphibians, sauropsids ( reptiles, birds), and early synapsids. In most tetrapods, the quadrate bone connects to the quadratojugal and squamosal bones in the skull, and forms up ...
. 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 ''Arcticodactylus'' to group with ''Carniadactylus'', '' Raeticodactylus'', and the Austriadraconidae, which in turn were within a clade he 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. .


See also

*
List of pterosaur genera This list of pterosaurs is a comprehensive listing of all Genus, 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 considere ...


References

{{Taxonbar, from=Q20651334 Late Triassic pterosaurs of Europe Pterosaurs Fossil taxa described in 2015