Eurazhdarcho
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''Eurazhdarcho'' is a
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 ...
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
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 ...
age 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 Romania, about 69 million years ago. Starting in 2009, pterosaur fossil remains were unearthed in a layer of the
Sebeș Formation The Sebeș Formation is a geological formation in Romania. It is of Maastrichtian age. It is laterally equivalent to the Sard Formation. The base of the formation consists of claystones interbedded with sandstones and conglomerates. It is well know ...
in Lancrăm, southwestern
Transylvania Transylvania ( or ; ; or ; Transylvanian Saxon dialect, Transylvanian Saxon: ''Siweberjen'') is a List of historical regions of Central Europe, historical and cultural region in Central Europe, encompassing central Romania. To the east and ...
by paleontologist Mátyás Vremir. In 2013, he, along with paleontologists
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 ...
,
Darren Naish Darren William Naish (born 26 September 1975) is a British vertebrate palaeontologist, author and science communicator. As a researcher, he is best known for his work describing and reevaluating dinosaurs and other Mesozoic reptiles, including ...
, and Gareth Dyke would name 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 ...
''Eurazhdarcho langendorfensis''. Its generic name is a combination of
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
and the 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 ...
'', while its
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 ...
is in reference to its origin from Langendorf (the name of Lancrăm in the language of the German ethnic minority in Romania). 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 ''Eurazhdarcho'' consists of a partial skeleton that includes cervical (neck) vertebrae,
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 ...
, and
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 ...
fragments. The related and much larger azhdarchid ''
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 ...
'' has also been found in the same location, indicating a possibility that ''Eurazhdarcho'' may have a juvenile of this pterosaur. However, the describers deemed this unlikely as its fossils seem to represent an adult specimen. With extrapolations from the comparisons of cervical vertebrae and wingbones of ''Eurazhdarcho'' with those of the related azhdarchid '' Zhejiangopterus'' from China, Vremir and colleagues would estimate a
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 ...
not surpassing . This makes ''Eurazhdarcho'' a medium size member of its family, Azhdarchidae. The distinctive traits that the describers set for ''Eurazhdarcho'' are all found in its cervical vertebrae. Within Azhdarchidae, the position of ''Eurazhdarcho'' has been somewhat disputed, with some studies recovering it within the subfamily Quetzalcoatlinae, while others have found it in a basal (primitive) position within the family. Due to the abundance of terrestrial fauna in the Sebeș Formation, it has been suggested that ''Eurazhdarcho'' most likely fed by snatching small terrestrial prey while walking on all fours instead of being a fish-eater on the coast.


Discovery and naming

In 2009, Romanian paleontologist Mátyás Vremir uncovered the remains of a pterosaur at the SbG-B site of the
Sebeș Formation The Sebeș Formation is a geological formation in Romania. It is of Maastrichtian age. It is laterally equivalent to the Sard Formation. The base of the formation consists of claystones interbedded with sandstones and conglomerates. It is well know ...
in Lancrăm, near Sebeș-Glod in southwestern
Transylvania Transylvania ( or ; ; or ; Transylvanian Saxon dialect, Transylvanian Saxon: ''Siweberjen'') is a List of historical regions of Central Europe, historical and cultural region in Central Europe, encompassing central Romania. To the east and ...
, Romania. He donated these to the ''Erdélyi Múzeum'' of the '' Societății Muzeului Ardelean'' (Transylvanian Museum Society). Subsequent excavations by Vremir discovered additional bones of the same individual animal and were added by him to the collection of the
Babeș-Bolyai University The Babeș-Bolyai University ( , , commonly known as UBB) is a public research university located in Cluj-Napoca, Romania. Established in 1581 as Academia Claudiopolitana, it underwent several reorganizations over the centuries, eventually taking ...
. In 2013, Vremir, along with paleontologists
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 ...
,
Darren Naish Darren William Naish (born 26 September 1975) is a British vertebrate palaeontologist, author and science communicator. As a researcher, he is best known for his work describing and reevaluating dinosaurs and other Mesozoic reptiles, including ...
, and Gareth Dyke named and described a 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 ...
, ''Eurazhdarcho langendorfensis''. The generic name ''Eurazhdarcho'' combines the name of
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
with that of 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
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 ...
''langendorfensis'' refers to its provenance from Langendorf, which is the name of Lancrăm in the language of the German ethnic minority in Romania. The article appeared in the
electronic journal An academic journal (or scholarly journal or scientific journal) is a periodical publication in which scholarship relating to a particular academic discipline is published. They serve as permanent and transparent forums for the dissemination, scr ...
'' PLoS ONE'' without an accompanying printed version; it nevertheless validly names the
taxon In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
under the new rules of the
ICZN The International Code of Zoological Nomenclature (ICZN) is a widely accepted convention in zoology that rules the formal scientific naming of organisms treated as animals. It is also informally known as the ICZN Code, for its formal author, t ...
. 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, EME VP 312, was found in a layer of the Sebeș Formation dating from the upper Early
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 ...
, about 69 million years old. It consists of a partial skeleton lacking the skull. It includes three cervical (neck) vertebrae among which the almost complete third and the fourth; the third and fourth right
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 upper 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 lower part of the second phalanx; a lower phalanx of one of the other fingers and a number of undetermined fragments. The Babeș-Bolyai University material is included within this enumeration and is not indicated by a separate inventory number. Generally, the quality of the bones is poor, with much of the outer cortex broken or eroded and internal structures present as impressions of natural molds. The fossils have not been completely flattened, preserving three-dimensionality, but compression has caused some distortion. The carcass had probably by flooding been deposed on its back in mud near a riverbank. Afterwards it was exposed to the air, weathering and being scavenged as proven by circular bite-marks inflicted by the conical teeth of some member of the
Crocodyliformes Crocodyliformes is a clade of crurotarsan archosaurs, the group often traditionally referred to as "crocodilians". They are the first members of Crocodylomorpha to possess many of the features that define later relatives. They are the only pseu ...
. Later covered by a thin layer of dirt, it was damaged by
beetle Beetles are insects that form the Taxonomic rank, order Coleoptera (), in the superorder Holometabola. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 40 ...
s and
termite Termites are a group of detritivore, detritophagous Eusociality, eusocial cockroaches which consume a variety of Detritus, decaying plant material, generally in the form of wood, Plant litter, leaf litter, and Humus, soil humus. They are dist ...
s. The describers of ''Eurazhdarcho'' noted that the related giant pterosaur ''
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 ...
'' is known from the same Romanian layers; the known fossil material from both genera does not overlap. Vremir and colleagues considered a juvenile identity for ''Eurazhdarcho'' with respect to ''Hatzegopteryx'' to be unlikely because the much smaller EME VP 312 seems to represent an adult individual.


Description

''Eurazhdarcho'' is a medium-sized member 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 it belongs. Comparing the cervical vertebrae and wingbones of the largest adult and most complete specimen of '' Zhejiangopterus'', Vremir and colleagues noted that the length of the fourth cervical is roughly the same as in specimen EME VP 312 of ''Eurazhdarcho'', while the estimated length of the fourth metacarpal is substantially less, in ''Eurazhdarcho'' versus in ''Zhejiangopterus''. Extrapolating from this data, Vremir and colleagues estimated 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 ''Eurazhdarcho'' at no more than . Vremir and colleagues established some distinctive traits for ''Eurazhdarcho'', all present in the cervical vertebrae. The third neck vertebra has three-quarters of the length of the fourth vertebra, whereas 60 percent would be normal with azhdarchids. The necks of 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. ...
, the front joint processes, are well-developed and elongated, obliquely pointing forwards and outwards under an angle of 30 degrees with the long axis of the vertebra. The preexapophysis, a secondary joint process on the side of the prezygapophysis, is well-developed with a forward pointing articulation facet and separated from a process, itself the remains of the diapophysis and possibly a neck rib, on the outer base of the prezygapophysis, by a deep trough on its underside. The pneumatic openings, the entrance holes for the air sacks, at the sides of the
neural arch Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spinal ...
are small and placed in a low position.


Classification

In its description, ''Eurazhdarcho'' was assigned to the family Azhdarchidae, based on the method of
comparative anatomy Comparative anatomy is the study of similarities and differences in the anatomy of different species. It is closely related to evolutionary biology and phylogeny (the evolution of species). The science began in the classical era, continuing in t ...
. However, 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 performed. In his 2021 study of ''
Quetzalcoatlus ''Quetzalcoatlus'' () is a genus of azhdarchid pterosaur that lived during the Maastrichtian age of the Late Cretaceous in North America. The Type (biology), type specimen, recovered in 1971 from the Javelina Formation of Texas, United States, ...
'', American paleontologist Brian Andres included ''Eurazhdarcho'' in his
phylogenetic analysis In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical data ...
. He placed ''Eurazhdarcho'' in the subfamily Quetzalcoatlinae within Azhdarchidae. His analysis is shown in the first
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 ...
below. In 2023, paleontologist Rodrigo Pêgas and colleagues recovered ''Eurazhdarcho'' in a different position within Azhdarchidae. They found it forming a
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
with azhdarchids ''
Phosphatodraco ''Phosphatodraco'' is a genus of azhdarchid pterosaur that lived during the Late Cretaceous of what is now Morocco. In 2000, a pterosaur specimen consisting of five Cervical vertebrae, cervical (neck) vertebrae was discovered in the Ouled Abdou ...
'', '' Wellnhopterus'', and '' Aralazhdarcho'' in a basal (primitive) position within the family. This classification model would be followed by a few subsequent studies. Their phylogenetic analysis is shown in the second cladogram below. Topology 1: Andres (2021). Topology 2: Pêgas and colleagues (2023).


Paleobiology

The area where ''Eurazhdarcho'' was found, in the Upper Cretaceous was localized on the
Hațeg Island Hațeg Island was a large offshore Former island, island in the Tethys Sea which existed during the Late Cretaceous period, probably from the Cenomanian to the Maastrichtian ages. It was situated in an area corresponding to the region around moder ...
, part of the prehistoric European archipelago in the ancient
Tethys Sea The Tethys Ocean ( ; ), also called the Tethys Sea or the Neo-Tethys, was a prehistoric ocean during much of the Mesozoic Era and early-mid Cenozoic Era. It was the predecessor to the modern Indian Ocean, the Mediterranean Sea, and the Eurasia ...
. The SbG-B site, though encompassing a surface of just , has yielded several distinct animal species, among which are the turtle '' Kallokibotion bajazidi'', the hadrosaur ''
Telmatosaurus ''Telmatosaurus'' (meaning "marsh lizard") is a genus of basal (phylogeny), basal hadrosauromorph dinosaur from the Late Cretaceous of Romania. It was a relatively small hadrosaur, measuring approximately in length and in body mass, which has b ...
'' and the
titanosaur Titanosaurs (or titanosaurians; members of the group Titanosauria) were a diverse group of sauropod dinosaurs, including genera from all seven continents. The titanosaurs were the last surviving group of long-necked sauropods, with taxa still thr ...
''
Magyarosaurus ''Magyarosaurus'' (" Hungarian lizard") is a genus of dwarf sauropod dinosaur from late Cretaceous Period (early to late Maastrichtian) in Romania. It is one of the smallest-known adult sauropods, measuring less than long and weighing less than ...
''. This terrestrial fauna suggests that ''Eurazdarcho'' was not a coastal
piscivore A piscivore () is a carnivorous animal that primarily eats fish. Fish were the diet of early tetrapod evolution (via water-bound amphibians during the Devonian period); insectivory came next; then in time, the more terrestrially adapted repti ...
catching fish on the wing, affirming the "superstork" model for azhdarchids, in which they are terrestrial stalkers snatching small prey animals while walking on all fours. If ''Eurazhdarcho'' was indeed distinct from ''Hatzegopteryx'', its discovery implies the presence of two azhdarchid forms in the Hațeg fauna, the one gigantic, the other medium-sized. This suggests a
niche partitioning In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for e ...
between them, although it is as yet unclear how this correlates with differences in prey preference and hunting techniques. This reflects a pattern seen in other Late Cretaceous faunae which also show a combination of a large azhdarchid species with a smaller one. One such example is the
Javelina Formation The Javelina Formation is a geological formation in Texas. Dating has shown that the strata date to the Maastrichtian stage of the Late Cretaceous, approximately 70 to 66.5 million years old. The middle part of the formation has been dated to ab ...
from the
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 ...
of
Texas Texas ( , ; or ) is the most populous U.S. state, state in the South Central United States, South Central region of the United States. It borders Louisiana to the east, Arkansas to the northeast, Oklahoma to the north, New Mexico to the we ...
, which has brought forth the giant ''Quetzalcoatlus northropi'', but also smaller
taxa In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
such as ''Quetzalcoatlus lawsoni'' and '' Wellnhopterus brevirostris''.


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 ...
*
Timeline of pterosaur research This timeline of pterosaur research is a chronologically ordered list of important fossil discoveries, controversies of interpretation, and Biological taxonomy, taxonomic revisions of pterosaurs, the famed flying reptiles of the Mesozoic Era (ge ...


References

{{Portal bar, Paleontology, Romania Azhdarchidae Maastrichtian life Late Cretaceous pterosaurs of Europe Cretaceous Romania Fossils of Romania Hațeg fauna Fossil taxa described in 2013