''Bennettazhia'' is a genus of

tapejaromorph Azhdarchoidea ( , meaning " azhdarchid-like forms") is a group of pterosaurs within the suborder Pterodactyloidea. Pterosaurs belonging to this group lived throughout the Early and Late Cretaceous periods, with one tentative member, '' Tendagur ...

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 ...

which lived during the

Albian The Albian is both an age (geology), age of the geologic timescale and a stage (stratigraphy), stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early Cretaceous, Early/Lower Cretaceous epoch (geology), Epoch/s ...

stage of the

Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...

from what is now the

Hudspeth Formation The Hudspeth Formation is a Cretaceous sedimentary Geological formation, rock formation, found in Oregon of the United States of America. The formation dates to the Albian age of the Early Cretaceous period. During the Albian much of this formatio ...

of the state of

Oregon Oregon ( , ) is a U.S. state, state in the Pacific Northwest region of the United States. It is a part of the Western U.S., with the Columbia River delineating much of Oregon's northern boundary with Washington (state), Washington, while t ...

in the

United States The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...

. Although originally identified as a species of the

pteranodontoid Pteranodontoidea (or pteranodontoids, from Greek meaning "toothless wings") is an extinct clade of ornithocheiroid pterosaurs from the Early to Late Cretaceous (early Valanginian to late Maastrichtian stages) of Asia, Africa, Europe, North Ameri ...

pterosaur ''

Pteranodon ''Pteranodon'' (; from and ) is a genus of pterosaur that included some of the largest known flying reptiles, with ''P. longiceps'' having a wingspan of over . They lived during the late Cretaceous geological period of North America in presen ...

'', ''Bennettazhia'' is now thought to have been a different animal. The type and only species is ''B. oregonensis''.

Discovery and history

In 1928, Charles Gilmore named a new species of ''

'': ''P. oregonensis''. A

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 ...

(

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 ...

MPUC V.126713), two fused dorsal vertebrae and the broken-off end of some joint bone had been unearthed from the

Lower Cretaceous Lower may refer to: * ''Lower'' (album), 2025 album by Benjamin Booker * Lower (surname) * Lower Township, New Jersey *Lower Receiver (firearms) * Lower Wick Gloucestershire, England See also * Nizhny {{Disambiguation ...

(

stage) beds of the

Wheeler County Wheeler County is the name of several counties in the United States: * Wheeler County, Georgia * Wheeler County, Nebraska * Wheeler County, Oregon Wheeler County is a county in the U.S. state of Oregon. As of the 2020 census, the populatio ...

, United States, to which the

specific epithet In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...

refers. Gilmore noted similarities to ''

Nyctosaurus ''Nyctosaurus'' (meaning "night lizard") is a genus of nyctosaurid pterosaur from the Late Cretaceous period of what is now the Niobrara Formation of the mid-western United States, which, during the time ''Nyctosaurus'' was alive, was covered in ...

'' though the specimens were larger. In 1989, S. Christopher Bennett concluded that the remains might be those of a member 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 Cre ...

instead of a

pteranodontid The Pteranodontidae are a family of large pterosaurs from the Late Cretaceous of North America and possibly other continents including Europe and Africa. The family was named in 1876 by Othniel Charles Marsh. Pteranodontids had a distinctive, elon ...

. In 1991, Russian paleontologist

Lev Nesov Lev or LEV may refer to: People and fictional characters *Lev (given name) *Lev (surname) Places *Lev, Azerbaijan, a village * Lev (crater), a tiny lunar crater Religion *an abbreviation for Leviticus, the third book of the Hebrew Bible and th ...

therefore named a new azhdarchid genus: ''Bennettazhia''. The genus name honors Bennett and combines his name with

Persian Persian may refer to: * People and things from Iran, historically called ''Persia'' in the English language ** Persians, the majority ethnic group in Iran, not to be conflated with the Iranic peoples ** Persian language, an Iranian language of the ...

''azhdarha'', "dragon", a reference to ''

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 the Azhdarchidae. Bennett himself in 1994 changed his opinion and stated that it belonged to the

Dsungaripteridae Dsungaripteridae is a group of pterosaurs within the suborder Pterodactyloidea. They were robust pterosaurs with good terrestrial abilities and flight honed for inland settings, and were commonly interpreted as durophagous and possibly piscivoro ...

. Since then, some authors including Bennett himself have classified this genus as a sister group to the

Tapejaridae Tapejaridae (from a Tupi language, Tupi word meaning 'the lord of the ways') is a Family (biology), family of azhdarchoid pterosaurs from the Cretaceous Period (geology), period. Members are currently known from Brazil, England, Hungary, Morocco, ...

or possibly a member of the

Thalassodrominae 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'', ...

. In 2023, the discovery of

guano Guano (Spanish from ) is the accumulated excrement of seabirds or bats. Guano is a highly effective fertiliser due to the high content of nitrogen, phosphate, and potassium, all key nutrients essential for plant growth. Guano was also, to a le ...

and unusual fragmentation of

ammonite Ammonoids are extinct, (typically) coiled-shelled cephalopods comprising the subclass Ammonoidea. They are more closely related to living octopuses, squid, and cuttlefish (which comprise the clade Coleoidea) than they are to nautiluses (family N ...

s from the Hudspeth Formation have been interpreted as evidence that this pterosaur ate

mollusc Mollusca is a phylum of protostome, protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant taxon, extant species of molluscs are recognized, making it the second-largest animal phylum ...

s and formed large colonies on nearby cliffs similar to modern

seagull Gulls, or colloquially seagulls, are seabirds of the subfamily Larinae. They are most closely related to terns and skimmers, distantly related to auks, and even more distantly related to waders. Until the 21st century, most gulls were placed ...

rookeries. New remains of this pterosaur are also reported, including two isolated teeth (F127985A and F127910B) and an edentulous (teeth-lacking) section of the lower

mandible In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...

(F127960).

Description

''Bennettazhia'' was a medium-sized pterosaur with an estimated wingspan of . In 2007, American biologist Michael Habib revealed the result of a study by

CAT-scan A computed tomography scan (CT scan), formerly called computed axial tomography scan (CAT scan), is a medical imaging technique used to obtain detailed internal images of the body. The personnel that perform CT scans are called radiographers or ...

of the type specimen of ''Bennettazhia''. The humerus, long, is uncrushed, which is uncommon for a pterosaur fossil and therefore offered a rare opportunity to investigate the bone structure. Apart from the thin bone wall, the humerus was filled with a spongy tissue consisting of

trabecula A trabecula (: trabeculae, from Latin for 'small beam') is a small, often microscopic, biological tissue, tissue element in the form of a small Beam (structure), beam, strut or rod that supports or anchors a framework of parts within a body or ...

e, very thin bone layers and struts, forming a light yet strong construction. Habib inferred that such strength would have allowed even very large pterosaurs to launch themselves from the ground using their forelimbs. The same investigation made a better classification possible. The humerus has an elongated deltopectoral crest that is unwarped. Both dsungaripterids and azhdarchoids show this feature, but only the latter group is typified by such a very thin outer bone wall. Habib concluded that ''Bennettazhia'' was a member of the

Azhdarchoidea Azhdarchoidea ( , meaning "azhdarchid-like forms") is a group of pterosaurs within the suborder Pterodactyloidea. Pterosaurs belonging to this group lived throughout the Early and Late Cretaceous periods, with one tentative member, '' Tendaguri ...

, a more encompassing group than the Azhdarchidae.Habib, M. (2007). "Structural characteristics of the humerus of ''Bennettazhia oregonensis'' and their implications for specimen diagnosis and azhdarchoid biomechanics", p. 16 in: ''Flugsaurier: The Wellnhofer pterosaur meeting'', Bavarian State Collection for Palaeontology, Munich, 1, 16.

Classification

The

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 follows a 2014

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 ...

by Brian Andres and colleagues. In the analysis, they placed ''Bennettazhia'' within the clade

Tapejaromorpha Azhdarchoidea ( , meaning "azhdarchid-like forms") is a group of pterosaurs within the suborder Pterodactyloidea. Pterosaurs belonging to this group lived throughout the Early and Late Cretaceous periods, with one tentative member, '' Tendaguri ...

, as the

basalmost In phylogenetics, basal is the direction of the ''base'' (or root) of a rooted phylogenetic tree or cladogram. The term may be more strictly applied only to nodes adjacent to the root, or more loosely applied to nodes regarded as being close to th ...

member.

References

External links

The Pterosaur Database (pdf)
{{Portal bar, Paleontology, United States Early Cretaceous pterosaurs of North America Tapejaromorphs Early Cretaceous reptiles of North America Fossil taxa described in 1991 Fossil taxa described in 1928