Mistralazhdarcho
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''Mistralazhdarcho'' is a genus of
azhdarchid 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 Cre ...
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 ...
that lived during the
Campanian The Campanian is the fifth of six ages of the Late Cretaceous epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campa ...
and
Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
ages of the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent 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 ''cre ...
period in what is now France. A rich fossil site was discovered in 1992 by paleontologist Xavier Valentin at Velaux–La Bastide Neuve, in the south of France. Pterosaur fossil remains would be subsequently uncovered in the site. They were found in a layer of the Aix-en-Provence Basin at Velaux–La Bastide Neuve and consist of a partial skeleton that includes the skull. These remains would later be made 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 ...
specimen of the new
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
and
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 ...
''Mistralazhdarcho maggii'', named and described in 2018 by Valentin, along with paleontologists Romain Vullo, Géraldine Garcia, Pascal Godefroit, and Aude Cincotta. The generic name combines the mistral, a typical northern wind from the area, and the related genus ''
Azhdarcho ''Azhdarcho'' is a genus of azhdarchid pterosaur from the late Cretaceous Period (geology), Period of the Bissekty Formation (middle Turonian stage, about 92 million years ago) of Uzbekistan, as well as the Zhirkindek Formation of Kazakhstan a ...
''. 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 Jean-Pierre Maggi, the mayor of
Velaux Velaux (; ) is a commune in the Bouches-du-Rhône department in southern France. It is near Coudoux and Ventabren. Velaux is an old Provençal village set on a hill overlooking the high plains of the inland west side of Aix. It is locate ...
, who supported the La Bastide Neuve paleontological project. Based on extrapolations of the size of its
humerus The humerus (; : humeri) 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 (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...
and the
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 the related pterosaur '' Zhejiangopterus'', the estimated wingspan for the holotype specimen of ''Mistralazhdarcho'' sits between . However, the holotype is not fully grown. Wingspans of adult individuals are estimated to have measured between . By comparing its fossil remains to those of its relatives, ''Mistralazhdarcho'' seems to have shared the most features with the genus ''
Aerotitan ''Aerotitan'' is a genus of azhdarchid pterosaur that lived during the Maastrichtian age of the Late Cretaceous period in what is now Argentina. Its only remains, which consist of a partial snout, were found in the Allen Formation of the Neuqué ...
'' from Argentina. ''Mistralazhdarcho'' was assigned to the family Azhdarchidae by Vullo and colleagues, though a
cladistic Cladistics ( ; from Ancient Greek 'branch') 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 ...
analysis was not made in its description. Later studies would include ''Mistralazhdarcho'' in their
phylogenetic analyses In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as Computational phylogenetics, phylogenetic inference. It infers the relationship among organ ...
, but its placement varies depending on the study. Within Azhdarchidae, some have recovered ''Mistralazhdarcho'' in the subfamily Azhdarchinae, while others in Quetzalcoatlinae. However, its position as the
sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
of ''Aerotitan'' has always been consistent. Because of its size, ''Mistralazhdarcho'' most likely represents a mid-sized class of European azhdarchid, its contemporary relatives in Europe were either small or gigantic.


Discovery and naming

In 1992, a rich fossil site was discovered by French paleontologist Xavier Valentin at Velaux–La Bastide Neuve, in the south of France. Between 2009 and 2012, pterosaur remains were excavated in the site. They were subsequently reported in the scientific literature in 2015 by paleontologist Aude Cincotta and colleagues, along with numerous other non-pterosaur fossil remains. Among the pterosaur remains is specimen MMS/VBN.09.C.001, which was discovered in a
sandstone Sandstone is a Clastic rock#Sedimentary clastic rocks, clastic sedimentary rock composed mainly of grain size, sand-sized (0.0625 to 2 mm) silicate mineral, silicate grains, Cementation (geology), cemented together by another mineral. Sand ...
layer of the Aix-en-Provence Basin at Velaux–La Bastide Neuve. The layer dates back to the late
Campanian The Campanian is the fifth of six ages of the Late Cretaceous epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campa ...
, about 72 million years ago. Specimen MMS/VBN.09.C.001 consists of a partial skeleton with the skull. It contains the symphysis of the lower jaws (the part where they join), the atlas-axis complex of the front neck, a middle neck vertebra, the left
humerus The humerus (; : humeri) 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 (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...
, a piece of the right humerus, the left radius, the right pteroid, the shaft 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 ...
, the proximal part of the first
phalanx The phalanx (: phalanxes or phalanges) was a rectangular mass military formation, usually composed entirely of heavy infantry armed with spears, pikes, sarissas, or similar polearms tightly packed together. The term is particularly used t ...
of the wing finger, the distal part of the same phalanx, and four bone fragments that could not be identified, which include some articular surface and two shafts. The skeleton was not found articulated but the bones were discovered on a limited surface of within the total surface of formed by the fossil layer. Therefore, it was concluded they represent a single individual, probably a subadult. In 2018, specimen MMS/VBN.09.C.001 was made 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 ...
of the newly named
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
and
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 ...
''Mistralazhdarcho maggii'', named and described by Valentin, along with paleontologists Romain Vullo, Géraldine Garcia, Pascal Godefroit, and Aude Cincotta. The generic name ''Mistralazhdarcho'' is a combination of the mistral, a northern wind typical of the area of discovery, and the related pterosaur ''
Azhdarcho ''Azhdarcho'' is a genus of azhdarchid pterosaur from the late Cretaceous Period (geology), Period of the Bissekty Formation (middle Turonian stage, about 92 million years ago) of Uzbekistan, as well as the Zhirkindek Formation of Kazakhstan a ...
'', the
type genus In biological taxonomy, the type genus (''genus typica'') is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearin ...
of
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 Cre ...
, the family to which ''Mistralazhdarcho'' belongs. 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 ...
''maggii'' honors Jean-Pierre Maggi, the mayor of
Velaux Velaux (; ) is a commune in the Bouches-du-Rhône department in southern France. It is near Coudoux and Ventabren. Velaux is an old Provençal village set on a hill overlooking the high plains of the inland west side of Aix. It is locate ...
, for his support of the La Bastide Neuve paleontological project.


Description

''Mistralazhdarcho'' is a large pterosaur. The humerus was estimated to have had an original length of . Knowing this measurement, there are a few methods that can be used in order to estimate its
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 ...
. One such method involves using a formula devised by Russian paleontologist Alexander Averianov. This would indicate a wingspan of about for ''Mistralazhdarcho''. Another possible method is to extrapolate the wingspan from the known skeleton of the azhdarchid '' Zhejiangopterus'', which is more complete. This would result in a wingspan estimate of . Both estimates seem to corroborate the hypothesis that the holotype individual of ''Mistralazhdarcho'' had a wingspan of about . However, said holotype was not yet fully grown. An adult exemplar could have had an estimated wingspan measuring around . In its description, Vullo and colleagues indicated some distinguishing traits of ''Mistralazhdarcho''. One of these was an
autapomorphy In phylogenetics, an autapomorphy is a distinctive feature, known as a Synapomorphy, derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or Outgroup (cladistics), outgroup taxa, not ...
, a unique derived character. The upper surface of the symphysis of the lower jaws shows a well-developed elevation on the midline in a relatively forward position of behind the mandible tip, compared to in the related pterosaur ''
Alanqa ''Alanqa'' is a genus of Azhdarchoidea, azhdarchoid pterodactyloid pterosaur from the Late Cretaceous period (Cenomanian stage) of what is now the Kem Kem Group of southeastern Morocco. The name ''Alanqa'' comes from the Arabic language, Arabic w ...
''. A second trait is also a possible autapomorphy, which is that the tip of the lower jaws is somewhat curved downwards. However, this trait would only be unique to ''Mistralazhdarcho'' if the related azhdarchid ''
Aerotitan ''Aerotitan'' is a genus of azhdarchid pterosaur that lived during the Maastrichtian age of the Late Cretaceous period in what is now Argentina. Its only remains, which consist of a partial snout, were found in the Allen Formation of the Neuqué ...
'' from Argentina does not have a curved tip itself. This latter pterosaur was originally described as not having one, but Vullo and colleagues, while studying ''Mistralazhdarcho'', concluded that the original description of ''Aerotitan'' was likely mistaken and that its curved holotype did not represent the middle of the jaws but their front end. Additionally, Vullo and colleagues gave a unique combination of traits that in themselves are not unique. The upper surface of the symphysis or fusion of the lower jaws shows elevated but blunt ridges on its edges. This symphysis has a V-shaped cross-section, thus lacking a lower crest. The underside of the atlas-axis complex of the neck is flat. In the middle neck vertebrae the front articulation processes, the
prezygapophyses The articular process or zygapophysis ( + apophysis) of a vertebra is a projection of the vertebra that serves the purpose of fitting with an adjacent vertebra. The actual region of contact is called the ''articular facet''.Moore, Keith L. et al. ...
, lightly diverge. The humerus is relatively short compared to the radius, with 66% of its length. In 2021, in a more thorough analysis of the already mentioned genus ''Aerotitan'', paleontologist Rodrigo Pêgas and colleagues compared its holotype to those of other azhdarchids, where they concluded that, among them, ''Mistralazhdarcho'' shared that most common features, especially its mandibular tip.


Classification

In its description in 2018, ''Mistralazhdarcho'' was placed in the family Azhdarchidae, without an exact
cladistic Cladistics ( ; from Ancient Greek 'branch') 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 ...
analysis. The elevation on the symphysis shared with ''Alanqa'' suggests a close relationship between the two genera. However, later
phylogenetic analyses In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as Computational phylogenetics, phylogenetic inference. It infers the relationship among organ ...
do not corroborate their close relationship. For example, a 2021 study by American paleontologist Brian Andres recovered ''Mistralazhdarcho'' as the
sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
of ''Aerotitan'' in the subfamily Azhdarchinae within Azhdarchidae, in the most derived (advanced) position. ''Alanqa'' in this analysis was not even recovered as an azhdarchid, but as a
thalassodromid Thalassodromidae (meaning "sea runners") is a group of azhdarchoid pterosaurs from the Early Cretaceous (Albian) of Brazil. All known definitive members come from the Romualdo Formation of Brazil, which include the type genus ''Thalassodromeus'', ...
instead. The close relationship between ''Mistralazhdarcho'' and ''Aerotitan'' would later be demonstrated again in a phylogenetic analysis by Pêgas and colleagues in 2023, where they found both azhdarchids in a trichotomy with the genus ''
Arambourgiania ''Arambourgiania'' (meaning "Camille Arambourg, Camille Arambourg's") is a genus of pterosaur, an extinct group of flying Reptile, reptiles, that inhabited Jordan during the Maastrichtian age of the Cretaceous, Cretaceous period, around 72 to ...
''. However, unlike Andres in 2021, Pêgas and colleagues found them nested within the subfamily Quetzalcoatlinae.
Cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
by Andres (2021). Cladogram by Pêgas and colleagues (2023).


Paleobiology

''Mistralazhdarcho'' is the first pterosaur named from the Campanian to the Maastrichtian of
Western Europe Western Europe is the western region of Europe. The region's extent varies depending on context. The concept of "the West" appeared in Europe in juxtaposition to "the East" and originally applied to the Western half of the ancient Mediterranean ...
. Due to its size, ''Mistralazhdarcho'' possibly represents an intermediate size class of European azhdarchids, in between the smaller genus ''
Eurazhdarcho ''Eurazhdarcho'' is a genus of azhdarchid pterosaur that lived during the Maastrichtian age of the Late Cretaceous period in what is now Romania, about 69 million years ago. Starting in 2009, pterosaur fossil remains were unearthed in a layer of ...
'' and the giant form ''
Hatzegopteryx ''Hatzegopteryx'' (" Hațeg basin wing") is a genus of azhdarchid pterosaur found in the late Maastrichtian deposits of the Densuş Ciula Formation, an outcropping in Transylvania, Romania. It is known only from the type species, ''Hatzegopte ...
''.


References

{{Portal bar, Paleontology, France Late Cretaceous pterosaurs of Europe Azhdarchidae Fossil taxa described in 2018