Arambourgiania

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Arambourgiania
Temporal range: Maastrichtian
Arambourgiania philadelphiae.JPG
Holotype fossil cast at Museum Histoire Naturelle, Paris
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Order:† Pterosauria
Suborder:† Pterodactyloidea
Family:† Azhdarchidae
Subfamily:† Quetzalcoatlinae
Genus:† Arambourgiania
Nessov vide Nessov & Yarkov, 1989
Species:
†A. philadelphiae
Binomial name
†Arambourgiania philadelphiae
(Arambourg, 1959)
Synonyms
Genus synonymy
Species synonymy
  • Titanopteryx philadelphiae
    Arambourg, 1959

Arambourgiania is a genus of azhdarchid pterosaur that lived during the Maastrichtian age of the Late Cretaceous, in what is now Jordan. Additional fossil remains from the United States and Morocco have also been found, but their assignment to Arambourgiania is only tentative. The original specimen was discovered in the 1940s by a railway worker near Russeifa, Jordan. After examination by paleontologist Camille Arambourg, a new species was named in 1959, Titanopteryx philadelphiae. The generic name means "titan wing", as the fossil was initially misidentified as a huge wing metacarpal (it would be later identified as a cervical (neck) vertebra), while the specific name refers to the ancient name of Amman (the capital of Jordan), Philadelphia. The genus "Titanopteryx" would later be problematic, as it had already been taken by a fly. Because of this, paleontologist Lev Nessov in 1989 named a new genus, Arambourgiania, in honor of Arambourg. The new species was now known as Arambourgiania philadelphiae.

Contents

Arambourgiania is one of the largest flying animals ever discovered. Initial wingspan estimates ranged from 11 to 13 m (36 to 43 ft), which would have made it the largest known pterosaur. However, due to Arambourgiania only presenting fragmentary specimens, recent research has suggested more moderate wingspan estimates. Some of the latest studies put the wingspan anywhere between 8 to 10 m (26 to 33 ft), which would still be among the largest known flying creatures. Arambourgiania has often been compared to other gigantic pterosaurs such as Quetzalcoatlus and Hatzegopteryx in terms of size.

Arambourgiania is a member of the family Azhdarchidae, which incudes some of the largest known pterosaurs ever. One of the closest relatives of Arambourgiania is Quetzalcoatlus, as multiple studes have found both pterosaurs to be grouped together within Azhdarchidae.

History of discovery

Holotype cervical vertebra of Arambourgiania (F) compared to those of other azhdarchids Characteristics of azhdarchid cervical vertebrae.png
Holotype cervical vertebra of Arambourgiania (F) compared to those of other azhdarchids

In the early 1940s, a railway worker during repairs on the Amman-Damascus railroad, near the city of Russeifa in Jordan, found a fossil bone measuring 60.96 cm (2 ft). This specimen was acquired in 1943 by the director of a nearby phosphate mine, Amin Kawar, who brought it to the attention of British archeologist Fielding after the Second World War. This generated some publicity — the bone was even shown to Abdullah I of Jordan — but more importantly, it made the scientific community aware of the find. [1]

In 1953, the fossil was sent to the National Museum of Natural History in Paris, France, where it was examined by French paleontologist Camille Arambourg. In 1954, he concluded the bone was the wing metacarpal of a giant pterosaur. Afterwards, in 1959, he named a new genus and species: Titanopteryx philadelphiae. The generic name meaning "titan wing" in Greek, referring to the enormous size of the fossil, while the specific name refers to the ancient name of Amman that was used by the Greeks: Philadelphia. Arambourg let a plaster cast be made and then sent the fossil back to the phosphate mine; this last aspect was later forgotten and the bone was assumed lost. [1]

While studying and describing the closely related pterosaur Quetzalcoatlus from Texas in 1975, American paleontologist Douglas A. Lawson concluded that the bone was not a metacarpal but a cervical (neck) vertebra. [2]

Arambourgiania was named after paleontologist Camille Arambourg Camille Arambourg Harcourt.jpg
Arambourgiania was named after paleontologist Camille Arambourg

In the 1980s, Russian paleontologist Lev Nesov was informed by an entomologist that the name Titanopteryx had already been given by Günther Enderlein to a fly from the Simulidae family in 1935. Therefore, in 1989, he renamed the genus into Arambourgiania, honoring Arambourg. [3] However, the name "Titanopteryx" was informally kept in use in the West, partially because the new name Arambourgiania was assumed by many to be a nomen dubium (dubious name). [4]

Skeletal reconstructions of Arambourgiania (C) and the related Hatzegopteryx (A), and Quetzalcoatlus lawsoni (D), with known parts in gray Hatzegopteryx-Witton-and-Naish-2017.png
Skeletal reconstructions of Arambourgiania (C) and the related Hatzegopteryx (A), and Quetzalcoatlus lawsoni (D), with known parts in gray

In early 1995, paleontologists David Martill and Eberhard Frey traveled to Jordan in an attempt to clarify matters. In a cupboard of the office of the Jordan Phosphate Mines Company they discovered some other pterosaur bones: a smaller vertebra and the proximal and distal extremities of a wing phalanx — but not the original material of Arambourgiania. However, after their departure to Europe, engineer of the mine Rashdie Sadaqah investigated further and in 1996 he established that the original fossil had been bought from the company in 1969 by geologist Hani N. Khoury, who had then donated it in 1973 to the University of Jordan. It was still present in the collection of this institute and now it could be restudied by Martill and Frey. [5] [6] [4]

Later, Frey and Martill rejected the suggestion that Arambourgiania was a nomen dubium or an identical pterosaur to Quetzalcoatlus, affirming its validity to replace the preoccupied name "Titanopteryx". [5] [4]

In 2016, an azhdarchid cervical vertebra (MPPM 2000.23.1) was described from the Coon Creek Formation of McNairy County in Tennessee, United States. It was referred to A. philadelphiae, which may potentially extend the geographic range of Arambourgiania to North America. [7] However, this specimen was later referred to as cf. Arambourgiania, indicating a species of uncertain affinities. [8] In 2022, American paleontologist Gregory S. Paul stated that it may not even belong to Arambourgiania itself. [9]

Ulna bone from the Ouled Abdoun Basin of Morocco that may belong to Arambourgiania Possible Arambourgiania ulna.png
Ulna bone from the Ouled Abdoun Basin of Morocco that may belong to Arambourgiania

In 2018, topotype specimens were located in Bavarian State Collection for Palaeontology and Geology in Munich, Germany that were placed there in 1966 from Jordan. These probably represent additional elements of the holotype individual. They are identified as cf. Arambourgiania and include the fragments of two cervical vertebrae, a neural arch, a left femur, a possible radius, and a metacarpal IV, as well as other indeterminate fragments. [10] The incomplete left ulna of the "Sidi Chennane azhdarchid" from Morocco has also been identified as cf. Arambourgiania. [11]

In 2024, the describers of the genus Inabtanin discovered a large partial right humerus of a large pterosaur in the Ruseifa Phosphate Mines, near the Jordanian capital of Amman, which was where the holotype of Arambourgiania was recovered. They concluded that the specimen belonged to A. philadelphiae and that it is comparable in size and shape to the humerus of the type species of Quetzalcoatlus, Q. northropi. [12]

Description

Length of the holotype cervical vertebra of Arambourgiania (first above) compared to other azhdarchid cervical vertebrae Azhdarchid pterosaur neck lengths.png
Length of the holotype cervical vertebra of Arambourgiania (first above) compared to other azhdarchid cervical vertebrae

The holotype of Arambourgiania, UJA VF1, consists of a very elongated cervical vertebra, probably the fifth. Today the middle section is missing; the original material was about 62 cm (2 ft 0.41 in) long, but had been sawed into three parts. Most of the fossil now consists of an internal infilling or mold; the thin bone walls are missing on most of the surface. The holotype does not present the whole vertebra; a piece is absent from its posterior end as well. [4]

Arambourgiania scale.png
Size of Arambourgiania (far right) compared to a human, Tyrannosaurus , and Balaur , the latter two lived roughly at the same time as Arambourgiania
Arambourgiania, Nyctosaurus and Quetzalcoatlus scale.jpg
Size of Arambourgiania (center left) compared to a human, Nyctosaurus (beside the human), and Quetzalcoatlus northropi (right). Note the very similar size of Arambourgiania and Q. northropi

Frey and Martill estimated the total length of the holotype to have been 78 cm (2 ft 7 in), using for comparison the relative position of the smallest diameter of the shaft of the fifth cervical vertebra of Quetzalcoatlus. The total neck length was extrapolated at about 3 m (9 ft 10 in) using the same method. From the relatively slender vertebra, the length dimension was then selected to be compared to that of Quetzalcoatlus as well, estimated at 66 cm (2 ft 2 in) long, which results in a ratio of 1.18. Applying that ratio to the overall size, Frey and Martill in the late 1990s concluded that the wingspan of Arambourgiania was 11 to 13 m (36 to 43 ft), larger than the estimated wingspan of Quetzalcoatlus, which measured 10 to 11 m (33 to 36 ft). This would have made Arambourgiania the largest pterosaur ever known. [5] [4] In 1997, paleontologist Lorna Steel and colleagues reconstructed a life-sized skeleton of Arambourgiania based on better-known related pterosaurs. They set its wingspan at 11.5 m (38 ft), within the range of Frey and Martill's estimate. [6]

Estimated size of Arambourgiania (dark turquoise; upper left) compared to contemporary pterosaurs, birds, and a human. Hatzegopteryx and Quetzalcoatlus northropi are also depicted (dark brown; lower left) Size disparity of late Maastrichtian pterosaurs and birds.svg
Estimated size of Arambourgiania (dark turquoise; upper left) compared to contemporary pterosaurs, birds, and a human. Hatzegopteryx and Quetzalcoatlus northropi are also depicted (dark brown; lower left)

Subsequently, the estimates proposed by Frey and Martill in the late 1990s were taken into question, with later estimates of the wingspan of Arambourgiania being more moderate. This was due to the remains being too fragmentary to estimate a gigantic size. In 2003, the researchers who described the related pterosaur Phosphatodraco stated that the wingspan of Arambourgiania was more likely at 7 m (23 ft), though this measurement was not given a rationale. [13] In 2010, paleontologists Mark Witton and Michael Habib argued that a 7 m (23 ft) wingspan is an underestimate for Arambourgiania, while a 11 to 13 m (36 to 43 ft) wingspan would be too much. [14]

In his 2022 pterosaur book, Paul proposed that Arambourgiania had a wingspan of 8 to 9 m (26 to 30 ft), which would be smaller than that of Quetzalcoatlus northropi, the which he kept at 10 to 11 m (33 to 36 ft). Arambourgiania would have also had a smaller wingspan than that of the related Hatzegopteryx from Romania, which Paul situated at 10 to 12 m (33 to 39 ft). Just like both Arambourgiania and Quetzalcoatlus, Hatzegopteryx is also another pterosaur that consistently ranks among the largest known flying animals. [9] In a 2024 study, the wingspan of Arambourgiania was estimated to be around 10 m (33 ft) based on a large humerus comparable in size to that of Q. northropi. This new estimate for the wingspan of Arambourgiania is slightly larger than Paul's 2022 estimate, but does not surpass the wingspan of Quetzalcoatlus. [12]

Classification

Quetzalcoatlus is among the closest relatives of Arambourgiania, with numerous studies recovering them as sister taxa Gfp-quetzalcaotlus.jpg
Quetzalcoatlus is among the closest relatives of Arambourgiania, with numerous studies recovering them as sister taxa

Arambourgiania was initially assigned to a newly named subfamily called Azhdarchinae by Nesov in 1984. It was still known as "Titanopteryx" at the time of the assignment. Azhdarchinae also included the pterosaurs Azhdarcho and Quetzalcoatlus. Nesov assigned the subfamily as part of the family Pteranodontidae, based on its members featuring toothless beaks just like the pteranodontids. [15] Unaware of the creation of Azhdarchinae, American paleontologist Kevin Padian created the family Titanopterygiidae, which included both "Titanopteryx" and Quetzalcoatlus. [16] Two years later, in 1986, Padian would synonymize the two terms and elevate Azhdarchinae to family level, creating the Azhdarchidae as it is known today. [17]

Life reconstruction of two Arambourgiania fighting over a small theropod Arambourgiania philadelphiae.png
Life reconstruction of two Arambourgiania fighting over a small theropod

The placement of Arambourgiania within the family Azhdarchidae has been consistent in various studies, which is in a derived (advanced) position in the subfamily Quetzalcoatlinae. However, with respect to other quetzalcoatlines in the group, it's specific location has been somewhat disputed. One of its closest relatives is Quetzalcoatlus, as both pterosaurs have consistently been found together in multiple phylogenetic analyses, either as sister taxa or close to each other. [18] [11] [19] [20] However, there have also been several studies opposing this placement and have instead favored a closer relationship between Arambourgiania and the azhdarchids Mistralazhdarcho and Aerotitan . [21] [22] [23]

Below are two cladograms showing different studies regarding the position of Arambourgiania within Azhdarchidae. The first one is based on the phylogenetic analysis by American paleontologist Brian Andres in 2021, which places Arambourgiania within Quetzalcoatlinae as the sister taxon to both species of Quetzalcoatlus, Q. northropi and Q. lawsoni. [19] The second cladogram is based on the 2023 study by paleontologist Rodrigo Pêgas and colleagues, in which they placed Arambourgiania in a trichotomy with Mistralazhdarcho and Aerotitan within Quetzalcoatlinae, contrasting its placement as the sister taxon of Quetzalcoatlus. [23]

Mistralazhdarcho.png
Aerotitan.jpg
In some studies, the azhdarchids Mistralazhdarcho (above) and Aerotitan (below) have been recovered as the closest relatives of Arambourgiania

Paleobiology

A 2024 study by paleontologist Kierstin Rosenbach included the description of the humerus of Arambourgiania. They compared it to that of soaring birds and suggested that Arambourgiania itsef was also a soarer. This is in contrast to the contemporary pterosaur Inabtanin, which had a style of flight closer to those of continuously flapping birds. [12]

See also

Related Research Articles

<i>Quetzalcoatlus</i> Genus of azhdarchid pterosaurs from the Late Cretaceous

Quetzalcoatlus is a genus of azhdarchid pterosaur that lived during the Maastrichtian age of the Late Cretaceous in what is now North America. The first specimen, recovered in 1971 from the Javelina Formation of Texas, United States, consists of several wing fragments. It was made the holotype of Quetzalcoatlus northropi in 1975 by Douglas Lawson. The generic name refers to the Aztec serpent god of the sky, Quetzalcōātl, while the specific name honors Jack Northrop, designer of tailless fixed-wing aircraft. The remains of a second species were found between 1972 and 1974, also by Lawson, around 40 km (25 mi) from the type of Q. northropi. In 2021, these remains were assigned the name Quetzalcoatlus lawsoni by Brian Andres and (posthumously) Wann Langston Jr.

<span class="mw-page-title-main">Azhdarchidae</span> Family of large azhdarchoid pterosaurs

Azhdarchidae 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 Cretaceous as well. Azhdarchids are mainly known for including some of the largest flying animals discovered, but smaller cat-size members have also been found. Originally considered a sub-family of Pteranodontidae, Nesov (1984) named the Azhdarchinae to include the pterosaurs Azhdarcho, Quetzalcoatlus, and Titanopteryx. They were among the last known surviving members of the pterosaurs, and were a rather successful group with a worldwide distribution. Previously it was thought that by the end of the Cretaceous, most pterosaur families except for the Azhdarchidae disappeared from the fossil record, but recent studies indicate a wealth of pterosaurian fauna, including pteranodontids, nyctosaurids, tapejarids and several indeterminate forms.

<i>Azhdarcho</i> Genus of azhdarchid pterosaur from the Late Cretaceous

Azhdarcho is a genus of azhdarchid pterosaur from the late Cretaceous Period of the Bissekty Formation of Uzbekistan, as well as the Zhirkindek Formation of Kazakhstan and possibly also the Ialovachsk Formation of Tajikistan. It is known from fragmentary remains including the distinctive, elongated neck vertebrae that characterizes members of the family Azhdarchidae, a family that includes many giant pterosaurs such as Quetzalcoatlus. The name Azhdarcho comes from the Persian word azhdar (اژدر), a dragon-like creature in Persian mythology. The type species is Azhdarcho lancicollis. The specific epithet lancicollis is derived from the Latin words lancea and collum ("neck").

<i>Hatzegopteryx</i> Genus of large azhdarchid pterosaur from the Late Cretaceous

Hatzegopteryx 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, Hatzegopteryx thambema, named by Buffetaut et al. in 2002 based on parts of the skull and humerus. Additional specimens, including a neck vertebra, were later placed in the genus, representing a range of sizes. The largest of these remains indicate it was among the biggest pterosaurs, with an estimated wingspan of 10 to 12 m.

<i>Zhejiangopterus</i> Genus of azhdarchid pterosaur from the Late Cretaceous

Zhejiangopterus is a genus of azhdarchid pterosaur known from one species, which lived in China during the late Cretaceous Period. The genus was named in 1994 by Chinese paleontologists Cai Zhengquan and Wei Feng. The type species is Zhejiangopterus linhaiensis. The genus name refers to Zhejiang Province and a Latinized Greek pteron, "wing". The specific name refers to the city of Linhai.

<i>Bennettazhia</i> Genus of tapejaromorph pterosaur from the Early Cretaceous

Bennettazhia is a genus of tapejaromorph pterosaur which lived during the Albian stage of the Early Cretaceous from what is now the Hudspeth Formation of the state of Oregon in the United States. Although originally identified as a species of the pteranodontoid pterosaur Pteranodon, Bennettazhia is now thought to have been a different animal. The type and only species is B. oregonensis.

Bogolubovia is a genus of pterosaur from the Upper Cretaceous Rybushka Formation of Petrovsk, Saratov Oblast, Russia. It is named for Nikolai Nikolaevich Bogolubov, the paleontologist who discovered the remains in 1914.

<i>Phosphatodraco</i> Late Cretaceous genus of pterosaur

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 (neck) vertebrae was discovered in the Ouled Abdoun Phosphatic Basin. The specimen was made the holotype of the new genus and species Phosphatodraco mauritanicus in 2003; the genus name means "dragon from the phosphates", and the specific name refers to the region of Mauretania. Phosphatodraco was the first Late Cretaceous pterosaur known from North Africa, and the second pterosaur genus described from Morocco. It is one of the only known azhdarchids preserving a relatively complete neck, and was one of the last known pterosaurs. Additional cervical vertebrae have since been assigned to the genus, and it has been suggested that fossils of the pterosaur Tethydraco represent wing elements of Phosphatodraco.

<i>Aralazhdarcho</i> Genus of azhdarchid pterosaur from the Late Cretaceous

Aralazhdarcho is a genus of azhdarchid pterosaur that lived during the Santonian to the early Campanian ages of the Late Cretaceous period in what is now Kazakhstan. The type and only known species is Aralazhdarcho bostobensis, named and described by Russian paleontologist Alexander Averianov. Its remains were found in the Bostobe Formation of Kazakhstan. The generic name combines the Aral Sea and the related genus Azhdarcho, while the specific name refers to its origin from the Bostobe Formation.

<span class="mw-page-title-main">Azhdarchoidea</span> Superfamily of ornithocheiroid pterosaurs

Azhdarchoidea 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 one tentative member, Tendaguripterus, that lived in the Late Jurassic period. The largest azhdarchoids include members of the family Azhdarchidae, examples of these are Quetzalcoatlus, Hatzegopteryx, and Arambourgiania. Other notable pterosaur families belonging to the Azhdarchoidea are the Tapejaridae, the Thalassodromidae, and the Chaoyangopteridae. This group contains some of the most advanced members of the pterosaur order.

<span class="mw-page-title-main">Pterosaur size</span>

Pterosaurs included the largest flying animals ever to have lived. They are a clade of prehistoric archosaurian reptiles closely related to dinosaurs. Species among pterosaurs occupied several types of environments, which ranged from aquatic to forested. Below are the lists that comprise the smallest and the largest pterosaurs known as of 2022.

<i>Alanqa</i> Genus of azhdarchid pterosaur from the Late Cretaceous

Alanqa is a genus of pterodactyloid pterosaur from the Late Cretaceous period of what is now the Kem Kem Beds of southeastern Morocco. The name Alanqa comes from the Arabic word العنقاءal-‘Anqā’, for a mythical bird of Arabian culture.

<i>Aerotitan</i> Genus of azhdarchid pterosaur from the Late Cretaceous

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, consisting of a partial snout, were found in the Allen Formation of the Neuquén Basin in northern Patagonia. This specimen was made the holotype of Aerotitan sudamericanus by paleontologist Fernando Novas and colleagues. The generic name combines the Greek word ἀήρ, meaning "air" and Titan, alluding the pterosaur's large size. The specific name is a reference to its origin, South America.

<i>Eurazhdarcho</i> Genus of azhdarchid pterosaur from the Late Cretaceous

Eurazhdarcho is a genus of azhdarchid pterosaur from the Late Cretaceous period of what is now the Transylvanian Basin of Romania. Its fossil remains dated back 69 million years ago.

<i>Cryodrakon</i> Genus of large azhdarchid pterosaur from the Late Cretaceous

Cryodrakon is a genus of azhdarchid pterosaur that lived during the Campanian age of the Late Cretaceous period in what is now Canada. Its remains contains a single species, Cryodrakon boreas, recovered from the Dinosaur Park Formation.

Albadraco is a genus of azhdarchid pterosaur that lived during the Maastrichtian age of the Late Cretaceous period in what is now Romania. The type species is Albadraco tharmisensis. Its remains, consisting of parts of the snout and a neck vertebra, were found in the Șard Formation, near the city of Alba Iulia in Transylvania. The species was named and described in 2019 by paleontologist Alexandru Adrian Solomon and colleagues.

<i>Apatorhamphus</i> Genus of azhdarchoid pterosaur from the Late Cretaceous

Apatorhamphus is an extinct genus of azhdarchoid pterosaur from the Kem Kem Group of Morocco. It might have been part of the Chaoyangopteridae. It is only known from a few snout fragments and it likely had a wingspan of between 3–7 metres (9.8–23.0 ft)

<i>Wellnhopterus</i> Genus of azhdarchid pterosaur from the Late Cretaceous

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, as well as some cervical vertebrae and a fragmentary long bone. In July 2021, the jaws were given the genus name "Javelinadactylus", with the type and only species as "J. sagebieli"; however, this article has now been retracted. In a paper published in December 2021, the complete holotype was independently named Wellnhopterus, with the only species being W. brevirostris. As of 2022, this is the formal name of this pterosaur.

<i>Thanatosdrakon</i> Genus of azhdarchid pterosaur from the Late Cretaceous

Thanatosdrakon is a genus of quetzalcoatline azhdarchid pterosaur from the Late Cretaceous Plottier Formation of the Neuquén Basin in western Argentina. The genus name is derived from the Greek words thanatos (=death) and drakon (=dragon), while the specific name is a Quechuan word meaning "flying serpent" and refers to the Incan deity Amaru. The type and only species is Thanatosdrakon amaru, known from two specimens consisting of several well-preserved axial and appendicular bones including material previously undescribed in giant azhdarchids. Thanatosdrakon is one of the oldest known members of the Quetzalcoatlinae. T. amaru lived from about 90 to 86 million years ago.

<i>Nipponopterus</i> Genus of azhdarchid pterosaurs

Nipponopterus is an extinct genus of azhdarchid pterosaurs from the Late Cretaceous Mifune Group of Japan. The genus contains a single species, N. mifunensis, known from a partial neck vertebra. Nipponopterus represents the first pterosaur to be named from Japan.

References

  1. 1 2 Arambourg, C. (1959). "Titanopteryx philadelphiae nov. gen., nov. sp. Ptérosaurien géant". Notes Mém. Moyen-Orient. 7: 229–234.
  2. ↑ Lawson, Douglas A. (1975). "Pterosaur from the Latest Cretaceous of West Texas: Discovery of the largest flying creature". Reports. Science. 187 (4180): 947–948. Bibcode:1975Sci...187..947L. doi:10.1126/science.187.4180.947. PMID   17745279. S2CID   46396417.
  3. ↑ Nessov, L.A. (1989). "New Cretaceous-Paleogene birds of the USSR and some remarks on the origin and evolution of the class Aves". Trudy Zoologicheskogo Instituta AN SSSR (in Russian). 197: 78–97.
  4. 1 2 3 4 5 Martill, D.M.; Frey, E.; Sadaqah, R.M.; Khoury, H.N. (1998). "Discovery of the holotype of the giant pterosaur Titanopteryx philadelphiae Arambourg, 1959, and the status of Arambourgiania and Quetzalcoatlus". Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen. 207: 57–76. doi:10.1127/njgpa/207/1998/57.
  5. 1 2 3 Frey, E.; Martill, D.M. (1996). "A reappraisal of Arambourgiania (Pterosauria, pterodactyloidea): one of the world's largest flying animals". Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen. 199 (2): 221–247. doi:10.1127/njgpa/199/1996/221.
  6. 1 2 Steel, L.; Martill, D.M.; Kirk, J.; Anders, A.; Loveridge, R.F.; Frey, E.; Martin, J.G. (1997). "Arambourgiania philadelphiae: giant wings in small halls". The Geological Curator. 6 (8): 305–313. doi: 10.55468/GC539 .
  7. ↑ Harrell, T. Lynn Jr.; Gibson, Michael A.; Langston, Wann Jr. (2016). "A cervical vertebra of Arambourgiania philadelphiae (Pterosauria, Azhdarchidae) from the Late Campanian micaceous facies of the Coon Creek Formation in McNairy County, Tennessee, USA". Bullettin Alabama Museum of Natural History. 33 (2): 94–103. ISSN   0196-1039.
  8. ↑ Andres, B.; Langston, W. Jr. (2021). "Morphology and taxonomy of Quetzalcoatlus Lawson 1975 (Pterodactyloidea: Azhdarchoidea)". Journal of Vertebrate Paleontology. 41 (sup1): 142. Bibcode:2021JVPal..41S..46A. doi: 10.1080/02724634.2021.1907587 . ISSN   0272-4634. S2CID   245125409.
  9. 1 2 Paul, Gregory S. (2022). The Princeton Field Guide to Pterosaurs. Princeton University Press. p. 161. doi:10.1515/9780691232218. ISBN   9780691232218. S2CID   249332375.
  10. ↑ Martill, David M.; Moser, Markus (2018). "Topotype specimens probably attributable to the giant azhdarchid pterosaur Arambourgiania philadelphiae (Arambourg 1959)". Geological Society, London, Special Publications. 455 (1): 159–169. Bibcode:2018GSLSP.455..159M. doi:10.1144/SP455.6. ISSN   0305-8719. S2CID   132649124.
  11. 1 2 Longrich, Nicholas R.; Martill, David M.; Andres, Brian; Penny, David (2018). "Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary". PLOS Biology. 16 (3): e2001663. doi: 10.1371/journal.pbio.2001663 . PMC   5849296 . PMID   29534059.
  12. 1 2 3 Rosenbach, K. L.; Goodvin, D. M.; Albshysh, M. G.; Azzam, H. A.; Smadi, A. A.; Mustafa, H. A.; Zalmout, I. S. A.; Wilson Mantilla, J. A. (2024). "New pterosaur remains from the Late Cretaceous of Afro-Arabia provide insight into flight capacity of large pterosaurs". Journal of Vertebrate Paleontology. 44 (1). e2385068. Bibcode:2024JVPal..44E5068R. doi: 10.1080/02724634.2024.2385068 .
  13. ↑ Suberbiola, Xabier Pereda; Bardet, Nathalie; Jouve, Stéphane; Iarochène, Mohamed; Bouya, Baâdi; Amaghzaz, Mbarek (2003). "A new azhdarchid pterosaur from the Late Cretaceous phosphates of Morocco". Geological Society, London, Special Publications. 217 (1): 79–90. Bibcode:2003GSLSP.217...79S. doi:10.1144/GSL.SP.2003.217.01.08. S2CID   135043714.
  14. ↑ Witton, M.; Habib, M.B. (2010). Laudet, V. (ed.). "On the Size and Flight Diversity of Giant Pterosaurs, the Use of Birds as Pterosaur Analogues and Comments on Pterosaur Flightlessness". PLOS ONE. 5 (11): e13982. Bibcode:2010PLoSO...513982W. doi: 10.1371/journal.pone.0013982 . PMC   2981443 . PMID   21085624.
  15. ↑ Nesov, L. A. (1984). "Upper Cretaceous pterosaurs and birds from Central Asia". Paleontologicheskii Zhurnal. 1984 (1): 47–57. Archived from the original on January 5, 2009.
  16. ↑ Padian, Kevin (1984). "A large pterodactyloid pterosaur from the Two Medicine Formation (Campanian) of Montana". Journal of Vertebrate Paleontology. 4 (4): 516–524. Bibcode:1984JVPal...4..516P. doi:10.1080/02724634.1984.10012027. ISSN   0272-4634.
  17. ↑ Padian, Kevin (September 2, 1986). "A taxonomic note on two pterodactyloid families". Journal of Vertebrate Paleontology. 6 (3): 289. Bibcode:1986JVPal...6..289P. doi:10.1080/02724634.1986.10011624. ISSN   0272-4634.
  18. ↑ Andres, B.; Myers, T. S. (2013). "Lone Star Pterosaurs". Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 103 (3–4): 1. doi:10.1017/S1755691013000303. S2CID   84617119.
  19. 1 2 Andres, Brian (December 7, 2021). "Phylogenetic systematics of Quetzalcoatlus Lawson 1975 (Pterodactyloidea: Azhdarchoidea)". Journal of Vertebrate Paleontology. 41 (sup1): 203–217. Bibcode:2021JVPal..41S.203A. doi: 10.1080/02724634.2020.1801703 . ISSN   0272-4634. S2CID   245078533.
  20. ↑ Zhou, Xuanyu; Ikegami, Naoki; Pêgas, Rodrigo V.; Yoshinaga, Toru; Sato, Takahiro; Mukunoki, Toshifumi; Otani, Jun; Kobayashi, Yoshitsugu (November 16, 2024). "Reassessment of an azhdarchid pterosaur specimen from the Mifune Group, Upper Cretaceous of Japan". Cretaceous Research . 167: 106046. doi:10.1016/j.cretres.2024.106046. ISSN   0195-6671.
  21. ↑ 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. 129: Article 104998. doi:10.1016/j.cretres.2021.104998. ISSN   0195-6671. S2CID   238725853.
  22. ↑ Ortiz David, Leonardo D.; González Riga, Bernardo J.; Kellner, Alexander W. A. (April 12, 2022). "Thanatosdrakon amaru, gen. ET SP. NOV., a giant azhdarchid pterosaur from the upper Cretaceous of Argentina". Cretaceous Research. 135: 105228. Bibcode:2022CrRes.13705228O. doi:10.1016/j.cretres.2022.105228. S2CID   248140163 . Retrieved April 12, 2022.
  23. 1 2 Pêgas, R. V.; Zhoi, X.; Jin, X.; Wang, K.; Ma, W. (2023). "A taxonomic revision of the Sinopterus complex (Pterosauria, Tapejaridae) from the Early Cretaceous Jehol Biota, with the new genus Huaxiadraco". PeerJ. 11. e14829. doi: 10.7717/peerj.14829 . PMC   9922500 . PMID   36788812.
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