''Eurazhdarcho'' is a
genus
Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial n ...
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 (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the Order (biology), order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cre ...
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'', ...
period (
Maastrichtian
The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interv ...
stage) of what is now the Transylvanian Basin of
Romania
Romania ( ; ro, România ) is a country located at the crossroads of Central, Eastern, and Southeastern Europe. It borders Bulgaria to the south, Ukraine to the north, Hungary to the west, Serbia to the southwest, Moldova to the east, a ...
. Its fossil remains dated back 69 million years ago.
Discovery and naming

In 2009,
Mátyás Vremir
Mátyás Vremir (13 November 1970 – 24 July 2020) was a Romanian geologist and palaeontologist.
Education and career
Vremir was born on 13 November 1970 in Cluj, the son of artists Mircea Vremir and Ildikó Kováts. He studied geology at Bab ...
at
Lancrăm near
Sebeş-Glod in
Transylvania
Transylvania ( ro, Ardeal or ; hu, Erdély; german: Siebenbürgen) is a historical and cultural region in Central Europe, encompassing central Romania. To the east and south its natural border is the Carpathian Mountains, and to the west the ...
at the SbG-B site uncovered the remains of a pterosaur. 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 ( ro, Universitatea Babeș-Bolyai , hu, Babeș-Bolyai Tudományegyetem, commonly known as UBB) is a public research university located in Cluj-Napoca, Romania. UBB has a long academic tradition, started by Universitas ...
.
In 2013, Vremir,
Alexander Wilhelm Armin 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 Perio ...
,
Darren Naish
Darren William Naish 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 '' Eotyrannus'', ''Xeno ...
, and
Gareth Dyke
Gareth John Dyke is a paleontologist whose work is concerned with the evolutionary history of birds and their dinosaurian relatives. His specific research interests include the phylogenetics of birds, the functional morphology of aves and non-avia ...
named and described 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( ...
''Eurazhdarcho langendorfensis''. The generic name combines the name of
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 subcontinent of Eurasia and it is located enti ...
with that of the related form ''
Azhdarcho
''Azhdarcho'' is a genus of azhdarchid pterosaur from the late Cretaceous Period of the Bissekty Formation (middle Turonian stage, about 92 million years ago) of Uzbekistan, as well as the Zhirkindek Formation of Kazakhstan. It is known from ...
''. The
specific name refers to
Langendorf, the name of Lancrǎm in the language of the
German ethnic minority in Romania. The article appeared in the
electronic journal
An academic journal or scholarly journal is a periodical publication in which scholarship relating to a particular academic discipline is published. Academic journals serve as permanent and transparent forums for the presentation, scrutiny, and ...
''
PLoS ONE'' without an accompanying printed version; it nevertheless validly names the
taxon
In biology, a taxon ( back-formation from '' taxonomy''; plural 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 n ...
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 publisher, the Int ...
.
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 seve ...
, EME VP 312, was found in a layer of the
Sebeş Formation dating from the upper Early
Maastrichtian
The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interv ...
, about 69 million years old. It consists of a partial skeleton lacking the skull. It includes three 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 form the intermediate part of the skeletal hand located between the phalanges of the fingers and the carpal bones of the wrist, which forms the connection to the forearm. The metacarpal bones ...
; the upper part of the first
phalanx
The phalanx ( grc, φάλαγξ; plural phalanxes or phalanges, , ) was a rectangular mass military formation, usually composed entirely of heavy infantry armed with spears, pikes, sarissas, or similar pole weapons. The term is particular ...
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 ps ...
. Later covered by a thin layer of dirt, it was damaged by
beetle
Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
s and
termite
Termites are small insects that live in colonies and have distinct castes ( eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blat ...
s.
The authors noted that from the same Romanian layers the related 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, ''Hatzegopter ...
'' is known; the known fossil material from both genera does not overlap. The authors considered an identity to be unlikely because the much smaller EME VP 312 seems to represent an adult individual.
Description

''Eurazhdarcho'' is a medium-sized
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 ...
. The authors estimated its
wingspan
The wingspan (or just span) of a bird or an airplane is the distance from one wingtip to the other wingtip. For example, the Boeing 777–200 has a wingspan of , and a wandering albatross (''Diomedea exulans'') caught in 1965 had a wingspan ...
at , extrapolating from an estimated length for the fourth metacarpal of about .
The authors established some distinctive traits, 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 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
The spinal column, a defining synapomorphy shared by nearly all vertebrates, Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...
are small and placed in a low position.
Phylogeny
''Eurazhdarcho'' was by the authors assigned to the 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 ...
; a
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 ch ...
analysis was not performed.
In 2018 however, Nicholas Longrich and colleagues included ''Eurazhdarcho'' in a large
phylogenetic analysis
In biology
Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that pro ...
of the Pterosauria, where it was recovered within the family Azhdarchidae.
Their
cladogram
A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
is presented below:
Paleobiology

The area where ''Eurazhdarcho'' was found, in the Upper Cretaceous was localized on the
Hațeg Island
Hațeg Island was a large offshore 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 modern-day Hațeg, ...
, part of the
European Archipelago
European, or Europeans, or Europeneans, may refer to:
In general
* ''European'', an adjective referring to something of, from, or related to Europe
** Ethnic groups in Europe
** Demographics of Europe
** European cuisine, the cuisines of Europe a ...
. The SbG-B site, though encompassing a surface of just 200 m³, has yielded several distinct animal species among which the turtle ''
Kallokibotion bajazidi
''Kallokibotion'' is an extinct genus of stem-turtle from the Upper Cretaceous (Maastrichtian, 86–66 million years ago), known from fossils found in Romania. One species is known, ''Kallokibotion bajazidi'', which was named by Franz Nopcsa afte ...
'', the hadrosaur ''
Telmatosaurus
''Telmatosaurus'' (meaning "marsh lizard") is a genus of 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 been explained as an ...
'' and a form referred to 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 th ...
''
Magyarosaurus
''Magyarosaurus'' (" Magyar 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 only in length and in body mass. The ...
''. This terrestrial fauna suggests that ''Eurazdarcho'' was not a coastal
piscivore
A piscivore () is a carnivorous animal that eats primarily fish. The name ''piscivore'' is derived . Piscivore is equivalent to the Greek-derived word ichthyophage, both of which mean "fish eater". Fish were the diet of early tetrapod evoluti ...
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, niche differentiation (also known as niche segregation, niche separation and niche partitioning) refers to the process by which competing species use the environment differently in a way that helps them to coexist. The competitive exclu ...
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. 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 abo ...
from the
Maastrichtian
The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interv ...
of
Texas
Texas (, ; Spanish language, Spanish: ''Texas'', ''Tejas'') is a state in the South Central United States, South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2 ...
has brought forth the giant ''
Quetzalcoatlus northropi
''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, a ...
'' but also a smaller ''Quetzalcoatlus sp.'' and the
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 Cretaceous, Early and Late Cretaceo ...
represented by specimen TMM 42489-2. The
Two Medicine Formation
The Two Medicine Formation is a geological formation, or rock body, in northwestern Montana and southern Alberta that was deposited between and (million years ago), during Campanian (Late Cretaceous) time. It crops out to the east of the Rocky M ...
fauna from 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. Campani ...
of
Montana
Montana () is a U.S. state, state in the Mountain states, Mountain West List of regions of the United States#Census Bureau-designated regions and divisions, division of the Western United States. It is bordered by Idaho to the west, North ...
includes the smaller ''
Montanazhdarcho
''Montanazhdarcho'' is a genus of azhdarchoid pterosaur from the Late Cretaceous period (Campanian stage) of what is now the state of Montana, United States. ''Montanazhdarcho'' is known from only one species, ''M. minor''.
Discovery
The holo ...
minor'', with a wingspan of , but also fragments of larger forms with a span of . In the coeval
Dinosaur Park Formation
The Dinosaur Park Formation is the uppermost member of the Belly River Group (also known as the Judith River Group), a major geologic unit in southern Alberta. It was deposited during the Campanian stage of the Late Cretaceous, between about 7 ...
of
Canada
Canada is a country in North America. Its ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world's second-largest country by tota ...
the smaller specimen RTMP 92.83 was discovered with a wingspan of but also the large specimen PMA P.80.16.1367 indicating a wingspan of about wide.
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 Biological taxonomy, taxonomic revisions of pterosaurs, the famed flying reptiles of the Mesozoic Era (ge ...
References
{{Portal bar, Paleontology, Cretaceous, Romania
Azhdarchids
Maastrichtian life
Late Cretaceous pterosaurs of Europe
Cretaceous Romania
Fossils of Romania
Hațeg fauna
Fossil taxa described in 2013