Argentinadraco
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''Argentinadraco'' (meaning "Argentina dragon") is an extinct genus of
azhdarchoid Azhdarchoidea (or azhdarchoids) is a group of pterosaurs within the suborder Pterodactyloidea, more specifically within the group Ornithocheiroidea. Pterosaurs belonging to this group lived throughout the Early and Late Cretaceous periods, with ...
pterosaur from the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...
Portezuelo Formation The Portezuelo Formation is a geologic formation of Late Cretaceous ( Late Turonian to Early Coniacian) age, outcropping in the Mendoza, Río Negro and Neuquén provinces of Argentina.Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
. It contains a single species, ''A. barrealensis'', named in 2017 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 ...
and Jorge Calvo. ''Argentinadraco'' is unusual for bearing a bottom jaw with a concave bottom edge, as well as a pair of ridges and depressions on the top surface. These features distinguish it from all other azhdarchoid groups, complicating its assignment, but it may belong to the
Chaoyangopteridae Chaoyangopteridae (or chaoyangopterids) is a family of pterosaurs within the larger group Azhdarchoidea. Chaoyangopterids lived mostly during the Early Cretaceous period, though possible members, ''Microtuban'', ''Xericeps'' and ''Argentinadraco' ...
. The ridges on the lower jaw may have been used to feed on small invertebrates in loose sediment within the system of lakes and rivers that it resided in.


Discovery and naming

''Argentinadraco'' is known from a single partial lower jaw, missing the rear end. The specimen is also compressed, especially near the tip of the jaw. Catalogued as MUCPv-1137 in the Centro Paleontológico Lago Barreales (CePaLB) of the
National University of Comahue The National University of Comahue ( es, Universidad Nacional del Comahue, UNCNavarro, Fernando A. ''. Tremédica, Asociación Internacional de Traductores y Redactores de Medicina y Ciencias Afines/UNCoPortal de Prensa de la Uncoma, Universidad ...
, it was found in layers of yellow
sandstone Sandstone is a clastic sedimentary rock composed mainly of sand-sized (0.0625 to 2 mm) silicate grains. Sandstones comprise about 20–25% of all sedimentary rocks. Most sandstone is composed of quartz or feldspar (both silicates) ...
and red/green
claystone Mudrocks are a class of fine-grained siliciclastic sedimentary rocks. The varying types of mudrocks include siltstone, claystone, mudstone, slate, and shale. Most of the particles of which the stone is composed are less than and are too sm ...
within the Futalognko quarry, which is located on the northern shore of Lake Barreales. The quarry is northwest of
Neuquén Neuquén (; arn, Nehuenken) is the capital city of the Argentine province of Neuquén and of the Confluencia Department, located in the east of the province. It occupies a strip of land west of the confluence of the Limay and Neuquén rivers w ...
, in
Neuquén Province Neuquén () is a province of Argentina, located in the west of the country, at the northern end of Patagonia. It borders Mendoza Province to the north, Rio Negro Province to the southeast, and Chile to the west. It also meets La Pampa Province a ...
,
Patagonia Patagonia () refers to a geographical region that encompasses the southern end of South America, governed by Argentina and Chile. The region comprises the southern section of the Andes Mountains with lakes, fjords, temperate rainforests, and gl ...
,
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
. Exposed deposits at the site belong to the
Portezuelo Formation The Portezuelo Formation is a geologic formation of Late Cretaceous ( Late Turonian to Early Coniacian) age, outcropping in the Mendoza, Río Negro and Neuquén provinces of Argentina.Neuquén Group The Neuquén Group is a group of geologic formations found in Argentina. Rocks in the Neuquén Group fall within the Cenomanian to early Campanian stages of the Late Cretaceous Period. It overlies the older Lohan Cura Formation and is itself over ...
of the
Neuquén Basin Neuquén Basin ( es, Cuenca Neuquina) is a sedimentary basin covering most of Neuquén Province in Argentina. The basin originated in the Jurassic and developed through alternating continental and marine conditions well into the Tertiary. The bas ...
, that dates to the
Turonian The Turonian is, in the ICS' geologic timescale, the second age in the Late Cretaceous Epoch, or a stage in the Upper Cretaceous Series. It spans the time between 93.9 ± 0.8 Ma and 89.8 ± 1 Ma (million years ago). The Turonian is preceded by t ...
or
Coniacian The Coniacian is an age or stage in the geologic timescale. It is a subdivision of the Late Cretaceous Epoch or Upper Cretaceous Series and spans the time between 89.8 ± 1 Ma and 86.3 ± 0.7 Ma (million years ago). The Coniacian is preceded by t ...
epochs of the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
period.
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 ...
, Jorge Calvo, Juan Porfiri, and Domenica dos Santos briefly described the specimen in an abstract at the 2011 Fourth Latin American Congress of Vertebrate Paleontology. Kellner and Calvo would go on to describe and name the specimen formally in 2017. The genus name ''Argentinadraco'' is derived from Argentina plus the suffix ''-draco'', from the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
word for "dragon", while the species name ''barrealensis'' is a reference to the locality of Lake Barreales.


Description

The size of ''Argentinadraco'' is difficult to estimate, but the preserved segment of the lower jaw measures long. The
symphysis A symphysis (, pl. symphyses) is a fibrocartilaginous fusion between two bones. It is a type of cartilaginous joint, specifically a secondary cartilaginous joint. # A symphysis is an amphiarthrosis, a slightly movable joint. # A growing together ...
of the lower jaw was likely long, taking up some 50% of the jaw's total length in life. The external
cortical bone A bone is a rigid organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red and white blood cells, store minerals, provide structure and support for the body, and ...
is thin, and the bones of the jaw are thoroughly fused, which characterizes ''Argentinadraco'' as a relatively
derived Derive may refer to: * Derive (computer algebra system), a commercial system made by Texas Instruments * ''Dérive'' (magazine), an Austrian science magazine on urbanism *Dérive, a psychogeographical concept See also * *Derivation (disambiguatio ...
(specialized)
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 ...
. While the top margin of the lower jaw is straight, like members of the
Azhdarchidae Azhdarchidae (from the Persian word , , a dragon-like creature in Persian mythology) is a family of pterosaurs known primarily from the Late Cretaceous Period, though an isolated vertebra apparently from an azhdarchid is known from the Early Cret ...
, the bottom margin of the symphysis is deep ( tall) at the rear but becomes shallower in front, making it markedly concave. This sets ''Argentinadraco'' apart from other members of the
Azhdarchoidea Azhdarchoidea (or azhdarchoids) is a group of pterosaurs within the suborder Pterodactyloidea, more specifically within the group Ornithocheiroidea. Pterosaurs belonging to this group lived throughout the Early and Late Cretaceous periods, with ...
. The entire bottom margin is blunt and thickened. There is also, unusually, a small crest along the concave portion of the bottom margin, but it is not as developed as the crests of the
Tapejaridae Tapejaridae (from a Tupi word meaning "the old being") are a family of pterodactyloid pterosaurs from the Cretaceous period. Members are currently known from Brazil, England, Hungary, Morocco, Spain, the United States, and China. The most primi ...
. The top surface of the symphysis is bordered by blunt outer margins (unlike the sharp margins of ''
Thalassodromeus ''Thalassodromeus'' is a genus of pterosaur that lived in what is now Brazil during the Early Cretaceous period, about a hundred million years ago. The original skull, discovered in 1983 in the Araripe Basin of northeastern Brazil, was collect ...
''), separated by a narrow depression. This depression becomes a concave shelf at the back of the jaw, and it is unusually flanked by a pair of well-developed ridges inset from the actual outer margins. The ridges are separated from the outer margins by narrow depressions. Most other toothless pterosaurs have a simple flattened or concave surface. There is a shallow pit, or fossa, at the back of the symphysis, caused by it ending in two segments as in ''
Caupedactylus ''Caupedactylus'' is an extinct genus of tapejarid azhdarchoid pterosaur known from the Early Cretaceous period (Albian stage) of what is now the Romualdo Formation of the Araripe Basin in northeastern Brazil. The type and only species of ''Caup ...
'' and ''
Quetzalcoatlus ''Quetzalcoatlus'' is a genus of pterosaur known from the Late Cretaceous period of North America (Maastrichtian stage); its members were among the largest known flying animals of all time. ''Quetzalcoatlus'' is a member of the Azhdarchidae, ...
''.


Classification

Although being toothless clearly places ''Argentinadraco'' in the
Dsungaripteroidea Dsungaripteroidea is a group of pterosaurs within the suborder Pterodactyloidea. Evolutionary history The earliest known fossils attributed to this group are from the Kimmeridgian-age Upper Jurassic Argiles d'Octeville Formation of France, date ...
''sensu'' Kellner, more specifically within the Azhdarchoidea, its precise classification within this group is more elusive. Lower jaws of the
Thalassodrominae Thalassodrominae or Thalassodromidae (meaning "sea runners", due to previous misconceptions of skimming behavior; they are now thought to be terrestrial predators) is a group of azhdarchoid pterosaurs from the Cretaceous period. Its traditional ...
,
Chaoyangopteridae Chaoyangopteridae (or chaoyangopterids) is a family of pterosaurs within the larger group Azhdarchoidea. Chaoyangopterids lived mostly during the Early Cretaceous period, though possible members, ''Microtuban'', ''Xericeps'' and ''Argentinadraco' ...
, and
Azhdarchidae Azhdarchidae (from the Persian word , , a dragon-like creature in Persian mythology) is a family of pterosaurs known primarily from the Late Cretaceous Period, though an isolated vertebra apparently from an azhdarchid is known from the Early Cret ...
tend to be either very similar or radically divergent within the same group. The proportions of at least the front portion of the lower jaw of ''Argentinadraco'' resembles those of the azhdarchid '' Zhejiangopterus'', and the chaoyangopterids '' Chaoyangopterus'' and '' Shenzhoupterus''. ''Argentinadraco'' bears differences, however, from all three groups. Its jaw is more robust and shorter than those of ''Quetzalcoatlus'' and some other azhdarchids. The deep rear end and the blunt ridges on the top surface also separate it from thalassodromines and chaoyangopterids. While the thalassodromine ''
Tupuxuara ''Tupuxuara'' is a genus of large, crested, and toothless Pterodactyloidea, pterodactyloid pterosaur from the Early Cretaceous period (Albian stage) of what is now the Romualdo Formation of the Santana Group, Brazil, about 125 to 112 million ye ...
'' has a crest on the bottom margin of its jaw, it does not have the shallow depression on the top surface. Kellner and Calvo tentatively assigned ''Argentinadraco'' to the Azhdarchidae based on provenance. It may instead represent an entirely new group, but this is difficult to test. Nevertheless, ''Argentinadraco'' can be confidently excluded from the
Pteranodontidae The Pteranodontidae are a family of large pterosaurs of the Cretaceous Period of North America and Africa. The family was named in 1876 by Othniel Charles Marsh. Pteranodontids had a distinctive, elongated crest jutting from the rear of the head ( ...
,
Nyctosauridae Nyctosauridae (meaning "night lizards" or "bat lizards") is a family of specialized soaring pterosaurs of the late Cretaceous Period of North America, Africa, and possibly Europe. It was named in 1889 by Henry Alleyne Nicholson and Richard Lydek ...
, and
Tapejarinae Tapejaridae (from a Tupi word meaning "the old being") are a family of pterodactyloid pterosaurs from the Cretaceous period. Members are currently known from Brazil, England, Hungary, Morocco, Spain, the United States, and China. The most primit ...
, the other groups of toothless dsungaripteroid pterosaurs. The back of the symphysis in ''Argentinadraco'' is deep like the pteranodontids, but the deepest point is further forward; the shelf of the symphysis is also inclined downwards instead of being arched as in pteranodontids, and the two-segment ending of the symphysis is not known in pteranodontids either. Nyctosaurids have an additional shelf on the symphysis, and the top margin of the lower jaw curves upwards. Finally, ''Argentinadraco'' does not have a downturned lower jaw, a step-like upper margin, or a deep crest, which all characterize tapejarines, and its jaw is much less robust. A study describing ''Javelinadactylus'' (now ''
Wellnhopterus ''Wellnhopterus'' () is an azhdarchid pterosaur recovered from the Late Cretaceous Javelina Formation in Texas that was previously identified as a thalassodromine. It consists of a set of upper and lower jaws (TMM 42489-2), as well as some cerv ...
'') recovered ''Argentinadraco'' as a closely related thalassodromine. The study has since been retracted. Conversely, a study focused on ''
Aerotitan ''Aerotitan'' is a genus of large azhdarchid pterosaur known from the Late Cretaceous period (Maastrichtian stage) of what is now the Allen Formation of the Neuquén Basin in northern Patagonia, Argentina. Discovery and naming The type speci ...
'' recovers it as a chaoyangopterid closely related to '' Xericeps''.Pêgas, R.V.; Holgado, B.; Ortiz David, L.D.; Baiano, M.A.; Costa, F.R. (August 21, 2021). "On the pterosaur Aerotitan sudamericanus (Neuquén Basin, Upper Cretaceous of Argentina), with comments on azhdarchoid phylogeny and jaw anatomy". Cretaceous Research. in press: Article 104998. doi:10.1016/j.cretres.2021.104998.


Paleobiology

The unusually elaborate combination of ridges and depressions on the top surface of the lower jaw in ''Argentinadraco'' suggests that the lower jaw interlocked with the upper jaw in some way. Meanwhile, as has been inferred for ''
Pteranodon ''Pteranodon'' (); from Ancient Greek (''pteron'', "wing") and (''anodon'', "toothless") is a genus of pterosaur that included some of the largest known flying reptiles, with ''P. longiceps'' having a wingspan of . They lived during the late Cr ...
'', the depth of the rear of the symphysis suggests that it had strong bite forces. Kellner and Calvo speculatively suggested that ''Argentinadraco'' used its peculiar jaw to slice through loose sediment in the rivers or lakes of its environment, potentially feeding on small invertebrates such as
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s. They interpreted such a lifestyle as being consistent with the terrestrial feeding habits of azhdarchids.


Paleoecology

A diverse assemblage of animals has been recovered from the Futalognko quarry, which represents a continental deposit. It was deposited in a humid environment, and represents a system of meandering rivers. ''Argentinadraco'' in particular was probably preserved after being trapped on a
point bar A point bar is a depositional feature made of alluvium that accumulates on the inside bend of streams and rivers below the slip-off slope. Point bars are found in abundance in mature or meandering streams. They are crescent-shaped and located on ...
by low-energy water flow. Pterosaurs are rare in the Futalognko quarry, but remains (including an
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 ...
) have previously been assigned to the Azhdarchidae. Dinosaurs include the
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
s ''
Megaraptor ''Megaraptor'' () is a genus of large theropod dinosaur that lived in the ages of the Late Cretaceous. Its fossils have been discovered in the Patagonian Portezuelo Formation of Argentina, South America. Initially thought to have been a giant dr ...
'' and ''
Unenlagia ''Unenlagia'' (meaning "half-bird" in Latinized Mapudungun) is a genus of dromaeosaurid theropod dinosaur that lived in South America during the Late Cretaceous period. The genus ''Unenlagia'' has been assigned two species: ''U. comahuensis'', ...
'' (alongside
dromaeosaurid Dromaeosauridae () is a family of feathered theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek ('), meaning ...
and
carcharodontosaurid Carcharodontosauridae (carcharodontosaurids; from the Greek καρχαροδοντόσαυρος, ''carcharodontósauros'': "shark-toothed lizards") is a group of carnivorous theropod dinosaurs. In 1931, Ernst Stromer named Carcharodontosauridae ...
teeth), the
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their bo ...
'' Futalognkosaurus'', and indeterminate
iguanodontia Iguanodontia (the iguanodonts) is a clade of herbivorous dinosaurs that lived from the Middle Jurassic to Late Cretaceous. Some members include ''Camptosaurus'', ''Dryosaurus'', ''Iguanodon'', ''Tenontosaurus'', and the hadrosaurids or "duck-bi ...
n
ornithopod Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (), that started out as small, bipedal running grazers and grew in size and numbers until they became one of the most successful groups of herbivores in the Cretaceous wo ...
s. Additionally, a
crocodylomorph Crocodylomorpha is a group of pseudosuchian archosaurs that includes the crocodilians and their extinct relatives. They were the only members of Pseudosuchia to survive the end-Triassic extinction. During Mesozoic and early Cenozoic times, cro ...
similar to ''
Comahuesuchus ''Comahuesuchus'' is an extinct genus of notosuchian crocodylomorphs from the Santonian Bajo de la Carpa Formation of Argentina. It was described by palaeontologist José Bonaparte in 1991. The type species is ''C. brachybuccalis''.
'' is also known from Futalognko, as are teeth of the Peirosauridae. Turtles of the group
Pelomedusoidea :''Alternatively, "Pelomedusidae" may refer to the Pelomedusoidea. See below for details.'' Pelomedusidae is a family of freshwater turtles endemic to sub-Saharan Africa, including Madagascar, São Tomé, and the Seychelles(Although this pop. ...
have been found as well. Fish include a small member of the
Euteleostei Euteleostei, whose members are known as euteleosts, is a clade of bony fishes within Teleostei that evolved some 240 million years ago. It is divided into Protacanthopterygii (including the salmon and dragonfish) and Neoteleostei (including t ...
, two members of the Clupeomorpha, and a member of the Semionotidae, known from scales. Freshwater
bivalve Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, bival ...
s have also been found at Futalognko. Finally, plant fossils are dominated by
angiosperm Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s, specifically
dicotyledon The dicotyledons, also known as dicots (or, more rarely, dicotyls), are one of the two groups into which all the flowering plants (angiosperms) were formerly divided. The name refers to one of the typical characteristics of the group: namely, t ...
s, but leaves and fruiting bodies from
gymnosperm The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers, cycads, ''Ginkgo'', and gnetophytes, forming the clade Gymnospermae. The term ''gymnosperm'' comes from the composite word in el, γυμνό ...
s are also known alongside
conifer Conifers are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single ...
s.


See also

*
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 ...
*
List of pterosaurs 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 inva ...
* 2018 in paleontology


References

{{Portal bar, Paleontology, Cretaceous, Argentina Azhdarchoids Coniacian life Turonian life Late Cretaceous pterosaurs of South America Cretaceous Argentina Fossils of Argentina Portezuelo Formation Fossil taxa described in 2017 Taxa named by Alexander Kellner