Wellnhopterus

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Wellnhopterus
Temporal range: Late Cretaceous (Maastrichtian), 68–66  Ma
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TMM 42489-2.png
Illustration of specimen TMM 42489-2, the holotype
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Order:† Pterosauria
Suborder:† Pterodactyloidea
Family:† Azhdarchidae
Genus:† Wellnhopterus
Andres & Langston Jr., 2021
Species:
†W. brevirostris
Binomial name
†Wellnhopterus brevirostris
Andres & Langston Jr., 2021
Synonyms
  • Javelinadactylus sagebieli
    Campos, 2021

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.

Contents

Back in July 2021, the jaw remains of Wellnhopterus had actually been given a separate genus and type species, "Javelinadactylus sagebieli", named and described by paleontologist Hebert Campos. He assigned it to the group Thalassodrominae. The article describing it, however, has now been retracted due to Campos lacking ownership of the reported data and because Andres had submitted his paper describing Wellnhopterus earlier than he did. As of the present day, Wellnhopterus brevirostris is the formal name of this pterosaur.

In its description, Andres and Langston Jr. estimated a wingspan of about 3 m (9 ft 10 in), making it a mid-sized pterosaur. It is comparable to the wingspan estimate of the contemporary species Quetzalcoatlus lawsoni, which sits at about 4.5 m (15 ft). The blunt structure of the snout, coupled with the more robustly-built and shorter neck of Wellnhopterus is quite different from the contemporary azhdarchid Quetzalcoatlus, which had more slender jaws and a longer neck. This anatomy of Wellnhopterus has been compared to other "blunt-jawed" azhdarchids, such as Hatzegopteryx , which are more adapted to hunt proportionally large prey in a raptorial manner. Wellnhopterus has been assigned to the family Azhdarchidae, but its position within the group has been somewhat disputed. Its describers placed it within the subfamily Quetzalcoatlinae, but other studies have recovered it in a more basal position outside Quetzalcoatlinae.

Discovery and history

Size chart of specimen TMM 42489-2 reconstructed as an azhdarchid, including the jaws and neck vertebrae TMM 42489-2.1 & TMM 42489-2.3 to 2.7 Size Chart Ver. 1.png
Size chart of specimen TMM 42489-2 reconstructed as an azhdarchid, including the jaws and neck vertebrae

In March 1986, the remains of jaws and cervical (neck) vertebrae of a large pterosaur were discovered in the Javelina Formation at Big Bend National Park in Brewster County, western Texas, by preparator Robert Rainey. In 1991, German paleontologist Peter Wellnhofer illustrated the find by a picture in his book The Illustrated Encyclopedia of Pterosaurs. In the photo caption, Wellnhofer assigned it to Quetzalcoatlus sp., indicating an indeterminate species of said pterosaur. [1] [2] Accordingly, his illustrator John Sibbick reconstructed Quetzalcoatlus itself with a snout that was too blunt, a mistake repeated in many popular science works. In 1996, paleontologists Alexander Kellner and Wann Langston Jr. pointed out that the remains dated from older layers than those which had produced Quetzalcoatlus sp. and that it represented a pterosaur with a shorter snout and neck. [3] In 2004, Kellner, limiting himself to the skull material which he reconstructed, stated that it was a member of the family Tapejaridae. [4] In 2006, British paleontologists David Martill and Darren Naish concluded that the snout resembled that of Tupuxuara more than that of Quetzalcoatlus and referred to the taxon as the "Javelina Tupuxuara". [5] However, the fossils were studied in 2013 by American paleontologist Brian Andres, whose cladistic analysis recovered the taxon as a member of the family Azhdarchidae, both when the cervical vertebrae were included and when only the skull material was entered in the analysis. [6] In the same year, British paleontologist Mark Witton similarly favored an azhdarchid interpretation over a thalassodromid one such as Tupuxuara, comparing it to other azhdarchids that had blunt jaws. [7]

The known fossil remains of Wellnhopterus consist of specimen TMM 42489-2, which is a set of upper and lower jaws, specimens TMM 42489-2.3 to 42489-2.8, which are several cervical vertebrae, and specimen TMM 42489-2.9, which is a fragmentary long bone. The layer of the Javelina Formation where they were found dated back to the middle Maastrichtian of the Late Cretaceous. The complete specimen, including the jaws and the cervical vertebrae, were given the new genus and type species Wellnhopterus brevirostris in December 2021. It was named and described by Andres and, posthumously, Langston Jr. The generic name Wellnhopterus honors Wellnhofer by combining his name with the Greek word πτερόν (pteron, meaning "wing"), a typical suffix for pterosaur genera, and would translate as "Wellnhofer's wing". The specific name brevirostris is a combination of the Latin words brevis, meaning "short", and rostrum, meaning "beak", which references its description in the literature as a “short-faced animal”. Specimen TMM 42489-2 was made the holotype of Wellnhopterus and it includes the front of the paired (both left-side and right-side) premaxillae, the paired maxillae, the jugals, the front of the mandibular symphysis and the dentary branch. The remains had been partially articulated. [2]

"Javelinadactylus"

Size chart of specimen TMM 42489-2 reconstructed as a thalassodromine Javelinadactylus sagebieli (TMM 42489-2).png
Size chart of specimen TMM 42489-2 reconstructed as a thalassodromine

Back in July 2021, the genus and type species "Javelinadactylus sagebieli" had been named and described by Brazilian paleontologist Hebert Campos based only on specimen TMM 42489-2. The neck vertebrae (specimens TMM 42489-2.3 to 2.7) were not considered to be part of the holotype in this paper. The generic name Javelinadactylus is a combination of the Javelina Formation, in reference to its origin there, with the Ancient Greek word δάκτυλος (daktylos, meaning "finger"), a usual suffix in the names of pterosaurs. The specific name sagebieli honors James Sagebiel, head of the vertebrate fossils collection of the University of Texas. It was the second pterosaur species named from the latest Cretaceous of Texas, after Quetzalcoatlus . Campos had assigned it to the subfamily Thalassodrominae within Tapejaridae, which would make "Javelinadactylus" the first known tapejarid from North America and would extend the range of the group to the Maastrichtian. However, in July 2022, Campos' paper was retracted due to the following reasons: [8]

"The editors have retracted this article, because the author did not have the ownership of the data reported. This was confirmed by the University of Texas at Austin. In addition, this article described a new species that was based on photographs of specimen TMM 42489-2. This specimen has been described and named, as Wellnhopterus brevirostris, gen. et sp. nov., in a different article that was submitted for publication prior to the submission of this article, and itself was part of a larger study. Hebert Bruno Nascimento Campos agrees to this retraction. The online version of this article contains the full text of the retracted article as Supplementary Information." [8]

Description

Size of Wellnhopterus (middle bottom) compared to contemporary pterosaurs, birds, and a human. Quetzalcoatlus northropi (lower left) and Q. lawsoni (above Wellnhopterus) are also depicted Size disparity of late Maastrichtian pterosaurs and birds.svg
Size of Wellnhopterus (middle bottom) compared to contemporary pterosaurs, birds, and a human. Quetzalcoatlus northropi (lower left) and Q. lawsoni (above Wellnhopterus) are also depicted

In its description by Andres and Langston Jr., Wellnhopterus was given an estimated wingspan of about 3 m (9 ft 10 in), [2] much smaller than that of the contemporary species Quetzalcoatlus northropi, which had an estimated wingspan of around 10 to 11 m (33 to 36 ft). [9] [10] However, the wingspan of Wellnhopterus is comparable to another contemporary species of Quetzalcoatlus, Q. lawsoni, which had estimated wingspan of around 4.5 m (15 ft). [2] With regards to the structure of its snout, Wellnhopterus has a well-defined bluntness in its jaws and a stockier and shorter neck, which is very different from the slender-jawed and longer-necked structure of its coeval Quetzalcoatlus. Due to these features, Wellnhopterus has been compared to other "blunt-jawed" members of the Azhdarchidae (the family to which Wellnhopterus belongs), which have a more robustly-built but shorter neck and thicker jaws. [7] [2] One such example is Hatzegopteryx from Romania. [11]

Classification

In its description in 2021 by Andres and Langston Jr., Wellnhopterus was recovered as a member of the family Azhdarchidae. [2] In the same year, Andres performed a phylogenetic analysis in which he recovered Wellnhopterus in the subfamily Quetzalcoatlinae within Azhdarchidae. Wellnhopterus was more specifically found in a trichotomy with Cryodrakon and a clade consisting of Hatzegopteryx, Arambourgiania , and Quetzalcoatlus. The first cladogram below shows the results of his study. [12] In 2023, paleontologist Rodrigo Pêgas and colleagues performed their own phylogenetic analysis and recovered Wellnhopterus in a much more different position than Andres in 2021. Within Azhdarchidae, it formed a trichotomy with both Phosphatodraco and Aralazhdarcho . This clade, which was joined by Eurazhdarcho , was recovered outside Quetzalcoatlinae in a more basal (primitive) position. The second cladogram below shows the results of their analysis. [13]

Map showing global distribution of faunas containing small-medium and giant-sized azhdarchids, with Wellnhopterus (TMM 42489-2; lower left) alongside Quetzalcoatlus Azhdarchid map.png
Map showing global distribution of faunas containing small-medium and giant-sized azhdarchids, with Wellnhopterus (TMM 42489-2; lower left) alongside Quetzalcoatlus

Paleobiology

Wellnhopterus was likely raptorial, as "blunt-jawed" azhdarchids are thought to have been specialized to hunt proportionally large prey. [7] [14] [15] Its co-existence with the much larger Quetzalcoatlus and a possible additional pterosaur taxon in the Javelina Formation have been used as examples of pterosaur diversity being higher in the Maastrichtian than traditionally thought. [16]

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>Tupuxuara</i> Genus of thalassodromid pterosaur from the Early Cretaceous

Tupuxuara is a genus of thalassodromid pterosaur that lived during the Albian age of the Early Cretaceous, about 112 million years ago. Its remains were found in what is now the Romualdo Formation of the Santana Group in Brazil. Tupuxuara was named in 1988 by paleontologists Alexander Kellner and Diógenes de Almeida Campos. The name Tupuxuara means "familiar spirit" referring to a familiar spirit in the mythology of the Tupi people in Brazil. Two species have been named, T. longicristatus, the type species, and T. leonardii. An additional species has been named in 2013, T. deliradamus. However, the validity of this species has been put into question and it may not even belong to Tupuxuara.

<i>Thalassodromeus</i> Genus of thalassodromid pterosaur from the Early Cretaceous

Thalassodromeus is a genus of pterosaur that lived in what is now Brazil during the Early Cretaceous period, about a hundred million years ago. The original skull, discovered in 1983 in the Araripe Basin of northeastern Brazil, was collected in several pieces. In 2002, the skull was made the holotype specimen of Thalassodromeus sethi by palaeontologists Alexander Kellner and Diogenes de Almeida Campos. The generic name means "sea runner", and the specific name refers to the Egyptian god Seth due to its crest being supposedly reminiscent of Seth's crown. Other scholars have pointed out that the crest was instead similar to the crown of Amon. A jaw tip was assigned to T. sethi in 2005, became the basis of the new genus Banguela in 2015, and assigned back to Thalassodromeus as the species T. oberlii in 2018, though other researchers consider it a valid genus. Another species was described in 2015 based on a supposed crest fragment, but this was later shown to be part of a turtle shell.

<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.

<span class="mw-page-title-main">Tapejaridae</span> Family of azhdarchoid pterosaurs from the Cretaceous period

Tapejaridae is a family of azhdarchoid pterosaurs from the Cretaceous period. Members are currently known from Brazil, England, Hungary, Morocco, Spain, the United States, and China. The most primitive genera were found in China, indicating that the family has an Asian origin.

<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 paleontologists Eric Buffetaut, Dan Grigorescu, and Zoltan Csiki 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>Phosphatodraco</i> Genus of azhdarchid pterosaur from the Late Cretaceous

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 pterodactyloid pterosaurs

Azhdarchoidea 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, Tendaguripterus, that lived in the Late Jurassic period. Remains of this group have been found in the Americas, Africa, and Eurasia, suggesting that they probably had a global distribution.

<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.

<span class="mw-page-title-main">Thalassodromidae</span> Family of azhdarchoid pterosaurs from the Cretaceous period

Thalassodromidae is a group of azhdarchoid pterosaurs from the Cretaceous period. Its traditional members come from Brazil, which include the type genus Thalassodromeus, as well as Tupuxuara. Additionally, the recently named genus Kariridraco, also from Brazil, is another undisputed thalassodromid. Proposals of additional thalassodromid genera have proved more controversial.

<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, which consist 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 to 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 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>Argentinadraco</i> Genus of azhdarchoid pterosaur from the Late Cretaceous

Argentinadraco is an extinct genus of azhdarchoid pterosaur from the Late Cretaceous Portezuelo Formation of Argentina. It contains a single species, A. barrealensis, named in 2017 by Alexander Kellner and Jorge Calvo. Argentinadraco is unusual for bearing a bottom jaw with a concave bottom edge, as well as a pair of ridges and depressions on the top surface. These features distinguish it from all other azhdarchoid groups, complicating its assignment, but it may belong to the Chaoyangopteridae. The ridges on the lower jaw may have been used to feed on small invertebrates in loose sediment within the system of lakes and rivers that it resided in.

<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.

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

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.

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>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

  1. ↑ Wellnhofer, P. (1991). The Illustrated Encyclopedia of Pterosaurs. London: Salamander Books, Ltd. p. 144. ISBN   978-0517037010.
  2. 1 2 3 4 5 6 Andres, B.; Langston Jr., W. (December 14, 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.
  3. ↑ Kellner, Alexander W. A.; Langston, Wann Jr. (June 5, 1996). "Cranial remains of Quetzalcoatlus (Pterosauria, Azhdarchidae) from Late Cretaceous sediments of Big Bend National Park, Texas". Journal of Vertebrate Paleontology. 16 (2): 222–231. Bibcode:1996JVPal..16..222K. doi:10.1080/02724634.1996.10011310.
  4. ↑ Kellner, A. W. A. (2004). "New information on the Tapejaridae (Pterosauria, Pterodactyloidea) and discussion of the relationships of this clade". Ameghiniana. 41 (4): 521–534. ISSN   1851-8044.
  5. ↑ Martill, D. M.; Naish, D. (2006). "Cranial crest development in the Azhdarchoid pterosaur Tupuxuara, with a review of the genus and tapejarid monophyly". Palaeontology. 49 (4): 925–941. Bibcode:2006Palgy..49..925M. doi: 10.1111/j.1475-4983.2006.00575.x . S2CID   15609202.
  6. ↑ Andres, B.; Myers, T.S. (2013). "Lone Star Pterosaurs". Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 103 (3–4): 383–398. Bibcode:2012EESTR.103..383A. doi:10.1017/S1755691013000303. S2CID   84617119.
  7. 1 2 3 Witton, Mark P. (2013). Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press. ISBN   978-0-691-15061-1.
  8. 1 2 Campos, H. B. N. (July 31, 2021). "A new azhdarchoid pterosaur from the Late Cretaceous Javelina Formation of Texas". Biologia. 77 (8): 2255. Bibcode:2022Biolg..77.2255C. doi: 10.1007/s11756-021-00841-7 . S2CID   238764420. (Retracted, see doi:10.1007/s11756-021-00841-7)
  9. ↑ 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.
  10. ↑ 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.
  11. ↑ Witton, M.; Brusatte, S.; Dyke, G.; Naish, D.; Norell, M.; Vremir, M. (2013). Pterosaur overlords of Transylvania: short-necked giant azhdarchids in Late Cretaceous Romania. The Annual Symposium of Vertebrate Paleontology and Comparative Anatomy. Edinburgh. Archived from the original on April 6, 2016. Retrieved January 18, 2017.
  12. ↑ Andres, B. (December 14, 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.
  13. ↑ 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.
  14. ↑ Pêgas, R. V.; Costa, F. R.; Kellner, A. W. A. (2018). "New Information on the osteology and a taxonomic revision of the genus Thalassodromeus (Pterodactyloidea, Tapejaridae, Thalassodrominae)". Journal of Vertebrate Paleontology. 38 (2): e1443273. Bibcode:2018JVPal..38E3273P. doi:10.1080/02724634.2018.1443273. S2CID   90477315.
  15. ↑ Naish, D.; Witton, M.P. (2017). "Neck biomechanics indicate that giant Transylvanian azhdarchid pterosaurs were short-necked arch predators". PeerJ. 5: e2908. doi: 10.7717/peerj.2908 . PMC   5248582 . PMID   28133577.
  16. ↑ Longrich, N.R.; Martill, D.M.; Andres, B.; Penny, D. (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.
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