Cryodrakon

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Cryodrakon
Temporal range: Campanian, 76.7–74.3  Ma
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Cryodrakon holotype left humerus.jpg
Holotype left humerus
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Order:† Pterosauria
Suborder:† Pterodactyloidea
Family:† Azhdarchidae
Subfamily:† Quetzalcoatlinae
Genus:† Cryodrakon
Hone et al., 2019
Species:
†C. boreas
Binomial name
†Cryodrakon boreas
Hone et al., 2019

Cryodrakon is a genus of azhdarchid pterosaur that lived during the late Campanian age of the Late Cretaceous period in what is now Canada, around 76.7 and 74.3 million years ago. Starting in 1972, fossil remains of large azhdarchid pterosaurs have been reported from Alberta. Paleontologists assigned them to the genus Quetzalcoatlus , given that it was the only known azhdarchid from North America back then and because they had limited information about its actual remains, so they simply could not deduce anything different. In 1992, a partial pterosaur skeleton was uncovered in the Dinosaur Park Formation of Alberta. It was partially described in 1995 by paleontologist Philip J. Currie and colleagues, with a subsequent and more complete description in 2005. But it was not until 2019 that it received a new genus and type species, Cryodrakon boreas, named and described by paleontologists David Hone, Michael Habib, and François Therrien. The partial skeleton was made the holotype specimen of this new pterosaur. Its generic name means "cold dragon" in Ancient Greek, in reference to its Canadian origin, while its specific name refers to the Greek god of the north winds, Boreas. All azhdarchid remains from the Dinosaur Park Formation were subsequently referred to Cryodrakon.

Contents

Most of the fossil remains of Cryodrakon belong to either young or subadult individuals, from which a wingspan of about 5 m (16 ft) could be estimated. However, much larger fossils have been unearthed, including an incomplete fifth cervical (neck) vertebra that measures 40 cm (1 ft 4 in). If complete, it could have measured at least 50 cm (1 ft 8 in). Adult individuals of Cryodrakon would have had a similar size to Quetzalcoatlus northropi based on the size of its fossils. A wingspan of about 10 m (33 ft) is the most commonly estimated for Cryodrakon, which makes it one of the largest flying animals to ever exist.

In its description, Cryodrakon was assigned to family Azhdarchidae, which includes the already mentioned Quetzalcoatlus. A phylogenetic analysis was not performed due to the fragmentary nature of its remains, but the describers pointed out certain features that would exclude Cryodrakon from being in a basal (primitive) postion within Azhdarchidae. Subsequent phylogenetic analyses would place it within the subfamily Quetzalcoatlinae, though in varying position depending on the study. Cryodrakon would have coexisted with many different types of dinosaurs in the Dinosaur Park Formation, as demonstrated by the abundance of dinosaur fossils found there.

Discovery and history

Location of the discovery of Cryodrakon (top; Canada) Significant pterosaur discoveries in western North America.png
Location of the discovery of Cryodrakon (top; Canada)

Since 1972, bones of large pterosaurs (flying reptiles) belonging to the family Azhdarchidae have been reported from Alberta. These were the first pterosaur fossils discovered in Canada. [1] [2] [3] [4] They were sometimes referred to as Quetzalcoatlus sp., indicating an uncertain species of Quetzalcoatlus, or were just assigned to Quetzalcoatlus altogether, without specifying anything. [5] [2] [4] This was a common theme back then. Paleontologists who uncovered large Cretaceous azhdarchid remains in North America would provisionally assign them to Quetzalcoatlus due to it being the only known azhdarchid from North America until then, coupled with the fact that information about said pterosaur's remains were limited, so they could not conclude anything different. Later research by American paleontologist Michael Habib would indicate that these fossil remains represented a taxon that was new to science. [3]

The holotype specimen consists of a wingbone (upper left), a fourth neck vertebra (middle top), a left humerus (upper right), and a tibia (bottom) Cryodrakon holotype cervical IV, humerus, pteroid, and tibia..jpg
The holotype specimen consists of a wingbone (upper left), a fourth neck vertebra (middle top), a left humerus (upper right), and a tibia (bottom)

In 1992, a partial pterosaur skeleton was found and excavated in Upper Cretaceous strata of Dinosaur Provincial Park in Alberta, more precisely in a layer of the Dinosaur Park Formation that dated back to the late Campanian (between 76.7 and 74.3 million years ago). The excavation site was located near the lower or older boundary of this range. It was subsequently reported and partially described by Canadian paleontologist Philip J. Currie and colleagues in 1995. They noticed that the bones had tooth marks and even a broken tip of a tooth imbedded in one of them, most likely coming from Saurornitholestes langstoni of the same formation. [6] [3] [2] The bones of the skeleton were not articulated, but associated, representing an immature animal. This is the only pterosaur found in Canada with fossil remains that consist of more than just a single bone so far. Due to the skeleton's initial description only being partial, a more detailed description of was made in 2005 by Currie and Eva Koppelhus. [7] [2] This partial skeleton, specimen TMP 1992.83, lacks the skull and consists of a fourth cervical (neck) vertebra, a rib, a humerus, a pteroid (wing) bone, a fourth metacarpal, a tibia and a metatarsal. [3] [2]

The specific name of Cryodrakon refers to the Greek god of the north wind, Boreas WindGod2.JPG
The specific name of Cryodrakon refers to the Greek god of the north wind, Boreas

In 2019, specimen TMP 1992.83 received a new separate genus and type species, Cryodrakon boreas, and became the holotype of this new pterosaur. Cryodrakon boreas was named and described by paleontologists David Hone and François Therrien, alongside Habib. The generic name Cryodrakon is derived from the Ancient Greek words κρύος (kryos, meaning "cold") and δράκων (drakon, meaning "dragon"), while the specific name boreas refers to the Greek god the north wind, Boreas. Hence, Cryodrakon boreas could be translated as "cold dragon of the north winds". [2] Habib had previously considered the name Cryodrakon viserion , as a reference to the ice dragon in Game of Thrones . [3]

Through comparisons with other azhdarchid taxa, Hone and colleagues concluded that all azhdarchid material from Alberta could be referred to a single distinct taxon. Therefore, they assigned all the known azhdarchid remains from the Dinosaur Park Formation to Cryodrakon boreas. The specimens consisted of numerous cervical vertebrae, a scapulocoracoid, an ulna, several fourth metacarpals, wing finger phalanges and a femur. These bones represent individuals of various biological ages, among them juveniles and a large mature exemplar. However, most of the bones are from medium-sized individuals. In their 2019 study, only the cervical vertebrae were described in detail, since bones from other parts of the body had already been treated in 2005. [2] [3]

Description

Cryodrakon holotype cervical IV.jpg
Cryodrakon holotype pteroid.jpg
Close-up view of the holotype fourth neck vertebra (left) and wingbone (right)

The majority of the fossil remains of Cryodrakon, among them its holotype bones, most likely represented young and subadult individuals. Their dimensions are similar to those of Quetzalcoatlus sp. (now known as the species Q. lawsoni [8] ). A wingspan of at least 5 m (16 ft) has been set for Cryodrakon based on these smaller remains. [2] [4] [3] Q. lawsoni is estimated to have had a similar wingspan of around 4.5 to 5 m (15 to 16 ft). [8] [9] However, the remaining azhdarchid material from the Dinosaur Park Formation that was assigned to Cryodrakon by Hone and colleagues indicates many individuals of varying size, from specimens less than half the size of Q. lawsoni to others than double it. For example, specimen TMP 1996.12.369, a fifth cervical vertebra with a length of only 10.6 mm (0.42 in), is from a juvenile Cryodrakon, which was given an estimated wingspan of about 2 m (6 ft 7 in). At the same time, specimen TMP 1980.16.1367 is an incomplete fifth neck vertebra with a preserved length of 40 cm (1 ft 4 in), which is estimated to have been at least 50 cm (1 ft 8 in) if complete. It presumably belonged to an adult individual. [2] This specimen was originally described in 1982 by Currie and Dale Russell. They assigned it to Quetzalcoatlus northropi and mistakenly identified it as a femur. A wingspan of 13 m (43 ft) was estimated. [5] [2] However, nowadays, wingspan estimates for Q. northropi have been more moderate, at around 10 to 11 m (33 to 36 ft), which would still make it one of the largest flying animals to ever exist. [10] [11]

Cast of the holotype of Arambourgiania. Specimen TMP 1980.16.1367 of Cryodrakon would have been of comparable size Arambourgiania philadelphiae.JPG
Cast of the holotype of Arambourgiania . Specimen TMP 1980.16.1367 of Cryodrakon would have been of comparable size

An fully-grown Cryodrakon would have most likely been of similar size to Q. northropi, based on the size of its remains. A wingspan of about 10 m (33 ft) has been estimated for adult individuals. [4] [3] In his 2022 book about pterosaurs, American paleontologist Gregory S. Paul had also set the same wingspan measurement for Cryodrakon. This would make it one of the largest known flying animals as well. [9] The previously mentioned specimen TMP 1980.16.1367 of Cryodrakon is also comparable in size to the holotype of Arambourgiania , which measured around 60 cm (1 ft 11.62 in), indicating that Cryodrakon would have also been of similar size to this pterosaur. [2] Initial wingspan estimates of Arambourgiania ranged from 11 to 13 m (36 to 43 ft), [12] [13] but more recent estimates have been more moderate, ranging from 8 to 10 m (26 to 33 ft), mostly due to the fragmentary nature of its remains. This would Arambourgiania equal to or even slightly smaller in size than Cryodrakon. [9] [14]

Life reconstruction of Cryodrakon in terrestrial pose Cryodrakon.jpg
Life reconstruction of Cryodrakon in terrestrial pose

In terms of body structure, Cryodrakon was proportionally similar to Quetzalcoatlus and other long-necked advanced members of the Azhdarchidae, though its somewhat more robust bones may indicate that it was slightly heavier. [2]

Cryodrakon is distinguished from all other known azhdarchids by two features of its neck vertebrae. The lateral pneumatic fossae or pneumatophores, a pair of small openings leading to air pockets on either side of the neural canal, were positioned near the lower edge of the neural canal, while those of other azhdarchids (with the purported exception of Eurazhdarcho ) were positioned higher up. The second distinguishing feature related to its postexapophyses, large bony knobs adjacent to the protruding rear connection surface of each vertebra, the cotyle. Cryodrakon's postexapophyses were prominent in width but short in length, clearly separated from the cotyle, and their facets were directed downwards. [2]

Classification

In its description, Hone and colleagues placed Cryodrakon in the family Azhdarchidae. No exact cladistic analysis had been given to clarify its precise relationship with other azhdarchids. However, the describers were confident that it did not belong in the basalmost (most primitive) position within Azhdarchidae due to the lack of distinct cervical zygapophyses in its middle cervicals. Additionally, based on the geological age of its fossil remains, Cryodrakon would have been one of the oldest known North American azhdarchids. [2] In 2021, American paleontologist Brian Andres performed a phylogenetic analysis on Quetzalcoatlus, in which he included Cryodrakon. He recovered Cryodrakon in a derived (advanced) position in the subfamily Quetzalcoatlinae within Azhdarchidae. Cryodrakon formed a trichotomy with Wellnhopterus and a clade containing the quetzalcoatlines Hatzegopteryx , Arambourgiania , and Quetzalcoatlus. His analysis is shown in the first cladogram below. [15] In 2023, a study by paleontologist Rodrigo Pêgas and collagues also recovered Cryodrakon within Quetzalcoatlinae, but differed from Andres in its specific position within the subfamily. In their analysis, they found Cryodrakon as the basalmost member of Quetzalcoatlinae. Their study is shown in the second cladogram below. [16]

The related Quetzalcoatlus was the pterosaur to which all the azhdarchid remains from the Dinosaur Park Formation were referred back then Gfp-quetzalcaotlus.jpg
The related Quetzalcoatlus was the pterosaur to which all the azhdarchid remains from the Dinosaur Park Formation were referred back then

Paleobiology

Cryodrakon is thought to have been capable of flight, [3] which would make it one of the largest flying animals known to have existed, just like Quetzalcoatlus. [4] Azhdarchid pterosaurs similar to Cryodrakon are currently thought to have fed by hunting for smaller animals while on the ground, similar to modern day marabou storks. [17]

Some specimens of Cryodrakon show signs of being eaten by other archosaurs, either due to predation or scavenging. The holotype partial skeleton of Cryodrakon was found to have been scavenged by a dromaeosaurid, possibly Saurornitholestes langstoni. Aside from tooth marks, a broken tooth was found in one of the bones, which, according to the authors, meant that the thin-walled bone of Cryodrakon "must have been very tough." [6] [3] In 2025, a fifth cervical vertebra of a juvenile Cryodrakon (TMP 2023.012.0237) showed probable crocodilian bite marks. [18]

Paleoenvironment

Map showing global distribution of faunas containing small-medium and giant-sized azhdarchids, with Cryodrakon in the upper left Azhdarchid map.png
Map showing global distribution of faunas containing small-medium and giant-sized azhdarchids, with Cryodrakon in the upper left

The Dinosaur Park Formation, where fossils of Cryodrakon have been excavated, preserves many fossils from alluvial and coastal plain environments. Both of these environments would have experienced high precipitation, warm temperatures, and high humidity, with conditions becoming more swamp-like as time progressed. [19] There was a great diversity of herbivorous dinosaurs including both lambeosaurine and saurolophine hadrosaurs, centrosaurine and chasmosaurine ceratopsians, and nodosaurid and ankylosaurid ankylosaurs, which could have been able to feed on different vegetation levels from one another. [7] [20] The predatory niches were occupied by small maniraptoran theropods (including dromaeosaurids and troodontids), medium-sized juvenile tyrannosaurids, and fully grown tyrannosaurids. [21]

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 North America. The type specimen, recovered in 1971 from the Javelina Formation of Texas, United States, consists of several wing fragments and was described as Quetzalcoatlus northropi in 1975 by Douglas Lawson. The first part of the name refers to the Aztec serpent god of the sky, Quetzalcōātl, while the second part honors Jack Northrop, designer of a 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 Q. northropi locality. In 2021, these remains were assigned the name Quetzalcoatlus lawsoni by Brian Andres and (posthumously) Wann Langston Jr, as part of a monograph on the genus.

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

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, while the specific name refers to the ancient name of Amman, 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.

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

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

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<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">Pterosaur size</span>

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

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

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

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

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 the Sebeș Formation in Lancrăm, southwestern Transylvania by paleontologist Mátyás Vremir. In 2013, he, along with paleontologists Alexander Kellner, Darren Naish, and Gareth Dyke would name the new genus and type species Eurazhdarcho langendorfensis. Its generic name is a combination of Europe and the genus Azhdarcho, while its specific name is in reference to its origin from Langendorf. The holotype specimen of Eurazhdarcho consists of a partial skeleton that includes cervical (neck) vertebrae, metacarpal, and phalanx fragments. The related and much larger azhdarchid Hatzegopteryx 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.

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

Mistralazhdarcho is a genus of azhdarchid pterosaur that lived during the Campanian and Maastrichtian ages of the Late Cretaceous 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 specimen of the new genus and type species 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. The specific name honors Jean-Pierre Maggi, the mayor of Velaux, who supported the La Bastide Neuve paleontological project.

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 Solomon and colleagues.

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

Wellnhopterus is a genus of azhdarchid pterosaur that lived during the Maastrichtian age of the Late Cretaceous period in what us now Texas, United States. Its fossil remains were discovered in the Javelina Formation at Big Bend National Park, located in Brewster County, Texas. The fossils consist of a set of upper and lower jaws, several cervical (neck) vertebrae, and a fragmentary long bone. Originally referred to an indeterminate species of Quetzalcoatlus, these remains would receive various interpretations ranging from a tapejarid to an azhdarchid identity. It was not until December 2021 that they were given a new genus and type species, Wellnhopterus brevirostris, named and described by paleontologists Brian Andres and, posthumously, Wann Langston Jr.. Its generic name means "Wellnhofer's wing", named in honor of paleontologist Peter Wellnhofer, while its specific name means "short-beaked", attributing to the blunt structure of its jaws.

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

Thanatosdrakon is a genus of azhdarchid pterosaur that lived during the Coniacian and Santonian ages of the Late Cretaceous period in what is now Argentina, around 89.6 and 86.3 million years ago. Its remains were found in the Plottier Formation of the Neuquén Basin in the Mendoza Province. The genus only consists of the type species, Thanatosdrakon amaru, named and described by paleontologists Leonardo Ortiz David, Bernardo González Riga, and Alexander Kellner. Its generic name means "dragon of death" in Greek, while its specific name is a Quechuan word meaning "flying serpent" and refers to the Incan deity Amaru. Thanatosdrakon is known from two specimens, the holotype, consisting of a partial postcranial skeleton, and the paratype, consisting of a left humerus. The holotype includes material that is previously undescribed in giant azhdarchid pterosaurs.

<i>Inabtanin</i> Genus of azhdarchoid pterosaurs

Inabtanin is an extinct genus of azhdarchoid pterosaurs from the Late Cretaceous Muwaqqar Formation of Jordan. The genus contains a single species, I. alarabia, known from a partial skeleton. Inabtanin represents one of the most complete pterosaur taxa known from the Afro-Arabia region.

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

Nipponopterus is a genus of azhdarchid pterosaur that lived during the Turonian and Coniacian ages of the Late Cretaceous period in what is now Japan. Its remains, which consist of a partial cervical (neck) vertebra, were found in the Mifune Group, located in the Kumamoto Prefecture in Kyūshū. It was made the holotype specimen of the genus and type species Nipponopterus mifunensis, named and described by paleontologist Xuanyu Zhou and colleagues in 2024. Nipponopterus represents the first pterosaur to be named from Japan.

References

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  3. 1 2 3 4 5 6 7 8 9 10 Greshk, Michael (10 September 2019). "New 'frozen dragon' pterosaur found hiding in plain sight - The flying reptile was mostly head and neck—and had at least a 16-foot wingspan, if not bigger". National Geographic Society . Archived from the original on 11 September 2019. Retrieved 10 September 2019.
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  8. 1 2 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 3 Paul, Gregory S. (2022). The Princeton Field Guide to Pterosaurs. Princeton University Press. p. 159. doi:10.1515/9780691232218. ISBN   9780691232218. S2CID   249332375.
  10. ↑ Witton, M.P.; Martill, D.M.; Loveridge, R.F. (2010). "Clipping the wings of giant pterosaurs: comments on wingspan estimations and diversity". Acta Geoscientica Sinica. 31: 79–81.
  11. ↑ Padian, K.; Cunningham, J.R.; Langston, W. Jr.; Conway, J. (2021). "Functional morphology of Quetzalcoatlus Lawson 1975 (Pterodactyloidea: Azhdarchoidea)". Journal of Vertebrate Paleontology. 41 (sup1): 218–251. Bibcode:2021JVPal..41S.218P. doi: 10.1080/02724634.2020.1780247 . S2CID   245125427.
  12. ↑ 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.
  13. ↑ 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.
  14. ↑ 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 .
  15. ↑ Andres, Brian (14 December 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.
  16. ↑ 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.
  17. ↑ Witton, Mark P.; Naish, Darren (28 May 2008). "A Reappraisal of Azhdarchid Pterosaur Functional Morphology and Paleoecology". PLOS ONE. 3 (5): e2271. Bibcode:2008PLoSO...3.2271W. doi: 10.1371/journal.pone.0002271 . PMC   2386974 . PMID   18509539.
  18. ↑ Brown, C. M.; Bell, P. R.; Owers, H.; Pickles, B. J. (2025). "A juvenile pterosaur vertebra with putative crocodilian bite from the Campanian of Alberta, Canada". Journal of Paleontology: 1–10. doi:10.1017/jpa.2024.12.
  19. ↑ Matson, Christopher Cody (2010). Paleoenvironments of the Upper Cretaceous Dinosaur Park Formation in southern Alberta, Canada (Unpublished master's thesis). University of Calgary. p. 125. doi:10.11575/PRISM/18677.
  20. ↑ Mallon, Jordan C.; Evans, David C.; Ryan, Michael J.; Anderson, Jason S. (4 April 2013). "Feeding height stratification among the herbivorous dinosaurs from the Dinosaur Park Formation (upper Campanian) of Alberta, Canada". BMC Ecology. 13 (1): 14. Bibcode:2013BMCE...13...14M. doi: 10.1186/1472-6785-13-14 . PMC   3637170 . PMID   23557203.
  21. ↑ Holtz, Thomas R. (1 September 2021). "Theropod guild structure and the tyrannosaurid niche assimilation hypothesis: implications for predatory dinosaur macroecology and ontogeny in later Late Cretaceous Asiamerica". Canadian Journal of Earth Sciences. 58 (9): 778–795. doi:10.1139/cjes-2020-0174. hdl: 1903/28566 .
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