Quetzalcoatlus

''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 from the ''Q. northropi'' locality. In 2021, these remains were assigned to the name ''Quetzalcoatlus lawsoni'' by Brian Andres and (posthumously) Wann Langston Jr, as part of a series of publications on the genus.

''Quetzalcoatlus northropi'' has gained fame as a candidate for the largest flying animal ever discovered, though estimating its size has been difficult due to the fragmentary nature of the only known specimen. While wingspan estimates over the years have ranged from , more recent estimates hover around . The smaller and more complete ''Q. lawsoni'' had a wingspan of around . Unlike most azhdarchids, ''Q. lawsoni'' had a small head crest, an extension of the premaxilla. Two different forms have been identified: one had a rectangular head crest and a taller nasoantorbital fenestra (a structure combining the naris and antorbital fenestra in many pterosaurs), and the other had a more rounded head crest and a shorter nasoantorbital fenestra. The proportions of ''Quetzalcoatlus'' behind the skull were typical of azhdarchids, with a very long neck and beak, shortened non-wing digits that were well adapted for walking, and a very short tail.

Historical interpretations of the diet of ''Quetzalcoatlus'' have ranged from scavenging to skim-feeding like the modern skimmer bird. However, more recent research has found that it most likely hunted small prey on the ground, in a similar way to storks and ground hornbills. This has been dubbed the terrestrial stalking hypothesis and is thought to be a common feeding behavior among large azhdarchids. On the other hand, the second species, ''Q. lawsoni'', appears to have been associated with alkaline lakes, and a diet of small aquatic invertebrates has been suggested. Similarly, while ''Q. northropi'' is speculated to have been fairly solitary, ''Q. lawsoni'' appears to have been highly gregarious (social). Azhdarchids like ''Quetzalcoatlus'' were highly terrestrial by pterosaur standards, though even the largest were nonetheless capable of flight. Based on the work of Mark P. Witton and Michael Habib in 2010, it now seems likely that pterosaurs, especially larger taxa such as ''Quetzalcoatlus'', launched quadrupedally (from a four-legged posture), using the powerful muscles of their forelimbs to propel themselves off the ground and into the air.

Research history and taxonomy

Discovery and naming

The genus ''Quetzalcoatlus'' is based on fossils discovered in rocks pertaining to the Late Cretaceous Javelina Formation in Big Bend National Park, Texas. Remains of dinosaurs and other prehistoric life had been found in the area since the beginning of the 20th century. The first ''Quetzalcoatlus'' fossils were discovered in 1971 by the graduate student Douglas A. Lawson while conducting field work for his Master's degree project on the paleoecology of the Javelina Formation. This field work was supervised by Wann Langston Jr., an experienced paleontologist who had been doing field work in the region since 1938 and since 1963 led expeditions through his position as curator at the Texas Science and Natural History Museum. The two had first visited the park together in March 1970, with Lawson discovering the first ''Tyrannosaurus rex'' fossil from Texas. Returning in 1971, Lawson discovered a bone while investigating an arroyo on the western edge of the park, and returned to Austin with a section of it. He and Langston then identified it as a pterosaur fossil based on its hollow internal structure with thin walls. Returning in November 1971 for further excavations, they were struck by the unprecedented size of the remains compared to known pterosaurs. The initial material consisted of a giant radius and ulna, two fused wristbones known as syncarpals, and the end of the wing finger. Altogether, the material comprised a partial left wing from an individual (specimen number TMM 41450-3) later estimated at over in wingspan. Lawson described the remains in his 1972 thesis as "Pteranodon gigas", and diagnosed it as being "nearly twice as large as any previously described species of ''Pteranodon''". As a thesis is not recognized as a published worked by the International Code for Zoological Nomenclature (ICZN), "Pteranodon gigas" is not a valid name. Further field work at the site was conducted in March 1973, when fragments were found alongside a long and delicate bone connected to an apparently larger element. This fossil was left in the ground until April 1974, when they fully excavated the larger element, a humerus. Due to the close association of discovered remains, Langston felt confident there were nothing more to be found at the site. Several later excavations of the site have indeed been unsuccessful.

Lawson announced his discovery in the journal ''Science'' in March 1975, with a depiction of the animal's size compared to a large aircraft and a ''Pteranodon'' gracing the cover of the issue. Lawson wrote that it was "without doubt the largest flying animal presently known". He illustrated and briefly described the remains known at the time, but did not offer a name and indicated that a more extensive description was in preparation that would diagnose the species. In May, he submitted a short response to his original paper to the journal, considering how such an enormous animal could have flown. Within the paper, he briefly established the name ''Quetzalcoatlus northropi'', but did still not provide a diagnosis or a more detailed description, which would later cause nomenclatural problems. Though not specified in the original publication, Lawson's named the genus after the Aztec feathered serpent god Quetzalcōātl, while the specific name honors John Knudsen Northrop, the founder of Northrop Corporation, who drove the development of large tailless Northrop YB-49 aircraft designs resembling ''Quetzalcoatlus''. The discovery of the giant pterosaur left a strong impression on both the scientific community and the general public, and was reported on throughout the world. It was featured in Time Magazine and appeared on the cover of Scientific American in 1981 alongside an article on pterosaurs by Langston. The species would become referenced by over 500 scientific publications, with ''Quetzalcoatlus northropi'' becoming the single most cited pterosaur species and ''Quetzalcoatlus'' the fourth most cited pterosaur genus after ''Pteranodon'', '' Rhamphorhynchus'', and '' Pterodactylus'', much older genera with many more species than ''Quetzalcoatlus''.

Prior to the announcement of the discovery, Langston had returned to Big Bend with a group of fossil preparators in February 1973, primarily aiming to excavate bones of the dinosaur '' Alamosaurus''. One of the preparators, a young man named Bill Amaral who went on to be a respected field worker, had been skipping his lunches to conduct additional explorations of the area. He came across some additional fragments of pterosaur bone on a different portion of the ridge, around 50 kilometers away from the original site. Two more new sites quickly followed nearby, producing many fragments which the crew figured could be fit back together, in addition to a complete carpal and intact wing bone. Langston noted in his field notes that none of these bones suggested animals as large as Lawson's original specimen. Further remains came from Amaral's first site in April 1974, after Lawson's site had been exhausted; a long neck vertebra and a pair of jawbones appeared. Associated structures were initially hoped to represent filamentous pycnofibres, but were later confirmed to be conifer needles. Near the end of the 1974 season, Langston stumbled over a much more complete pterosaur skeleton; it consisted of a wing, multiple vertebrae, a femur and multiple other long bones. They lacked time to fully excavate it, leaving it in the ground until the next field season. This area, where many smaller specimens began to emerge, came to be known as Pterodactyl Ridge. Two of the smaller individuals were reported in the first 1975 paper by Lawson, presumed to belong to the same species, though Langston would begin to question the idea they belonged to ''Q. northropi'' by the early 1980s. Excavations continued in 1976, and eight new specimens emerged in 1977; in 1979, despite complications due to losing the field notes form 1977, Langston discovered another new site that would produce an additional ten specimens. Most importantly, a humerus of the smaller animal was finally found, which Langston considered of great importance to understanding ''Quetzalcoatlus''. Several further new localities followed in 1980, but 1981 proved less successful and Langston began to suspect the ridge may have been mostly depleted of pterosaur fossils. There was similarly little success in 1982, and visits during 1983 and 1985 proved to provide the last substantive discoveries of ''Quetzalcoatlus'' fossils. Langston returned in 1989, 1991, 1992, and 1996, but only found isolated bones and fragments. Eventually a handful of additional specimens were discovered by former student Thomas Lehman. A visit to Lawson's initial site during 1991 showed that all traces of excavation had by now eroded away. Langston would visit Big Bend for the last time in 1999, having concluded the pterosaur expeditions to focus on the excavation of two skulls of ''Deinosuchus'', another famous fossil of the area.

Later research

The expected further description implicated by Lawson never came. For the next 50 years, the material would remain under incomplete study, and few concrete anatomical details were documented within the literature. Much confusion surrounded the smaller individuals from Pterodactyl Ridge. In a 1981 article on pterosaurs, Langston expressed reservations whether they were truly the same species as the immense ''Q. northropi''. In the meantime, Langston focused on the animal's publicity. He worked on a life-sized gliding replica of ''Quetzalcoatlus northropi'' with aeronautical engineer Paul MacCready between 1981 and 1985, promoting it in a dedicated IMAX film. The model was created to understand the flight of the animal — prior to Lawson's discovery such a large flier wasn't thought possible, and the subject remained controversial at the time. Furthermore, the model was intended to allow people to experience the animal in a more dynamic manner than a mere static display or film. Around this time he also created a skeletal mount of the genus that was exhibited at the Texas Memorial Museum.

The next scientific effort of note was a 1996 paper by Langston and pterosaur specialist Alexander Kellner. By this time, Langston was confident the smaller animals were a separate species. A full publication establishing such a species was still in preparation at the time, but due to the importance of the skull material for the understanding of azhdarchid anatomy, the skull anatomy was published first. In this publication, he animal was referred to as ''Quetzalcoatlus'' sp., a placeholder designation for material not assigned to any particularly valid species. Once again, the planned further publication failed to materialize for decades, and ''Quetzalcoatlus'' sp. remained in limbo. A publication on the bioaeromechanics of the genus was also planned by Langston and James Cunningham, but this failed to materialize and the partially completed manuscript later became lost. Ultimately, a comprehensive publication on ''Quetzalcoatlus'' sp. would not appear before Langston's death in 2013. By this point he had produced many notes and individual descriptions, but had not begun writing any formal manuscript that could be published.

In 2021, a comprehensive description of the genus was finally published, the 19th entry in the Memoir series of special publications by the Society of Vertebrate Paleontology in the '' Journal of Vertebrate Paleontology''. It consisted of five studies published together. Kevin Padian was the primary organizer of the project. A paper on the history of discoveries in Big Bend National Park was authored by Matthew J. Brown, Chris Sagebiel, and Brian Andres. It focused on curating a comprehensive list of specimens belonging to each species to ''Quetzalcoatlus'' and the locality information of each within Big Bend. Thomas Lehman contributed a study on the paleonvironment that ''Quetzalcoatlus'' would have resided within, based upon work he had begun with Langston as early as 1993. Brian Andres published a study on the morphology and taxonomy of the genus, established the species ''Quetzalcoatlus lawsoni'' for the smaller animal that had gone for decades without a name. The specific name honoured Lawson, who discovered ''Quetzalcoatlus''. Despite not contributing directly to the written manuscript, the authors of the memoir and Langston's family agreed that he posthumously be considered a co-author due the basis of the work in the decades of research he dedicates to the subject. Also authored by Andres was a phylogenetic study of ''Quetzacoatlus'' and its relationships within Pterosauria, with a focus on the persistence of many lineages into the Late Cretaceous contra classical interpretations of ''Quetzalcoatlus'' as the last of a dying lineage. Finally, a study on the functional morphology of the genus was authored by Padian, James Cunningham, and John Conway (who contributed scientific illustrations and cover art to the Memoir), with Langston once again considered a posthumous co-author due to his foundational work on the subject. Brown and Padian prefaced the Memoir, who once again emphasized their gratitude to Langston for his decades of work on the animal leading up to the publication.

Taxonomy and material

The genus ''Quetzalcoatlus'' consists of two valid species: the type species ''Q. northropi'' and the second species ''Q. lawsoni''. Though the name was introduced in 1975, the lack of a formal description complicated its validity for several decades. The oldest name for the species is "''Pteranodon'' gigas", from Lawson's 1973 thesis. However, the International Code of Zoological Nomenclature (ICZN) does not consider a thesis to be a formal publication capable of establishing of a taxon, and the name has not been used since. Regarding the genus ''Quetzalcoatlus'' and species ''Q. northropi'', the name being established in a separate publication than the anatomical diagnosis also failed ICZN standards. As such, it was argued that the name was a ''nomen nudum'', an intended but invalid scientific name, though some authors argued the second publication referencing the initial description was sufficient. The species received a diagnosis in a 1991 paper by Lev Nessov, but no action was taken to formalize the name. Furthermore, a study by Mark Witton and colleagues in 2010 doubted whether ''Quetzalcoatlus'' could be validly diagnosed at all. They noted that the bones preserved in the holotype of ''Q. northropi'' were not typically considered to be taxonomically informative between close relatives, and that they appeared extremely similar to those of other giant azhdarchids such as the Romanian azhdarchid '' Hatzegopteryx''. Both of these issues were settled in the 2021 paper, whose rediagnosis affirmed ''Quetzalcoatlus'' as distinct. The authors agreed that the original paper did not constitute a valid establishment of the name. The authors noted their publication could serve as the a basis for the name, but did not wish to change the previously presumed authorship of the name. Thus, they submitted an application for the ICZN in 2017 to make an exception to the requirements, and had Lawson's second 1975 paper to be declared as the valid authority of the genus and species. The approval of this ICZN petition on August 30, 2019, conserved and formalized the binomen ''Quetzalcoatlus northropi'' as the type species.

Though Lawson originally considered all ''Quetzalcoatlus'' remains to belong to one species, today two species are recognized: the large ''Quetzalcoatlus northropi'' and the smaller ''Quetzalcoatlus lawsoni''. The exact nature of what material belongs to each species remains unclear, due in part to the distribution of specimens across various localities and stratigraphic levels found at Big Bend, as well as the limited scope of ''Q. northropi'' material to compare to. The name-bearing type specimen of ''Q. northropi'' is TMM 41450-3, a partial wing. It includes a humerus, radius, ulna, wrist bones, finger bones, and many elements of the elongate wing finger, in addition to thousands of unidentifiable fragments. It is from the uppermost rocks of the Javelina Formation, making it one of the youngest pterosaur specimens known. Only a single other specimen can confidently be referred to the same species, a left ulna designated TMM 44036-1 known from the Black Peaks Formation, around three quarters the size of the type specimen and sharing distinctive anatomy. Four other specimens share a similarly giant size, but cannot be definitively assigned to ''Q. northropi'' in lack of overlapping material or distinguishing anatomical regions. TMM 41047-1 and TMM 41398-3, are both partial femurs, the former twice the size of that seen in ''Q. lawsoni''. Their anatomy indicates they belonged to the same species, and is distinct from that of ''Q. lawsoni''. Part of a wing finger, TMM 41398-4, is also of the correct size to belong to ''Q. northropi'' but does not preserve the essential anatomy to confirm its identity. This specimen and the smaller femur were the first two specimens Lawson discovered, prior to uncovering the type specimen. Finally, one of the oldest pterosaur specimens in Big Bend is a giant cervical vertebra not matching that of smaller species from the formation. Whilst conventional pterosaur research would assign all of these to ''Q. northropi'', the 2021 redescription preferred to be cautious and merely assigned them to ''Q. cf. northropi'', indicating uncertainty.

The assignment of remains to ''Q. lawsoni'' has proved more simplistic; a large quantity of similar remains were found together in nearby sites, 50 km from the ''Q. northropi'' locality. In total, 305 different fossil elements from 214 specimens are known, all of which are considered consistent with assignment to the same species. This is the most amount of remains referred to any singular species of pterosaur. The vast majority of the dozens of specimens are disarticulated individual bones. A few individual animals are, however, represented by associated remains; identification of these individuals was complicated by each bone being catalogued under a separate number, which was revised as part of the 2021 study. The most complete specimen is TMM 41961-1, which possesses the most complete skull as well as several neck vertebrae, much of both wings, femurs, tibiotarsi, two metatarsals, and one of the toe bones. It was one of the original specimens described by Langston in 1975, and in accordance with Langston's wishes and its completeness was designated as the type specimen. Two less complete specimens, TMM 42180-14 and TMM 42161-1, were also preserved in partial articulation, the former mostly composed of limb and neck bones whilst the latter consists of neck and skull bones. Beyond this, identification of individual specimens is difficult. Two other specimens are more loosely associated, and others may have belonged to a single individual but are too weathered to identify with confidence. In some cases two or three neck vertebrae were found in presumed association, and similar loose associations of one or two limbs bones are seen in several cases. Taken together, nearly the entirely skeleton is represented, with the exception of some of the back of the skull. Eight different specimens preserve various portions of the skull, together allowing for a rather complete picture (excepting a portion of the back of the skull), and similarly the entire mandible is represented when cross-referencing four specimens. All nine neck vertebrae are known, and most torso vertebrae are known through the preservation of the notarium and synsacrum, structures consisting of several fused vertebrae in ornithocheiroid pterosaurs; it is unknown how many unfused vertebrae may have existed between these structures. Every single bone in the arm is known from at least one complete specimen, and the hindlimbs and pelvis are also more or less all present, though the femur and pelvis suffer from poor preservation.

Other referred and reclassified material

Though most pterosaur remains from Big Bend have been referred to ''Quetzalcoatlus'', some other material exists. Most prominent amongst these is a specimen discovered around north of the Pterodactyl Ridge localities, designated as TMM 42489-2 and compromising a partial skull and jaws as well as five articulated neck vertebrae. It was immediately noted for its distinct shorted-jawed anatomy compared to what had come to be expected from ''Q. lawsoni'' specimens. Despite this distinctiveness, it was referred to as a separate ''Quetzalcoatlus sp.'' in a 1991 book by Langston. As early as 1996, however, this was revised with recognition it was certainly a new genus informally known as the "short-faced pterosaur". It was formally named in the 2021 paper alongside ''Q. lawsoni'', as the genus '' Wellnhopterus''. In addition to this specimen, several indeterminate azhdarchid remains and some remains too fragmentary to assign beyond Pterosauria are known from Big Bend. Some of these represent smaller animals than the uniformly sized ''Q. lawsoni'' remains. Whether any of these remains represented separate animals from ''Quetzalcoatlus'' cannot be determined.

Several specimens from across Late Cretaceous North America were historically referred to ''Quetzalcoatlus''. In 1982 a femur from the Campanian aged Dinosaur Park Formation of Alberta, Canada was referred to the genus by Philip J. Currie and Dale Russell. Currie later described further remains from Dinosaur Park in a 2005 book, noting their morphological similarity to ''Quetzalcoatlus'' but expressing caution against referral to the genus. In 2019, however, all azhdarchid remains from the formation were revised as distinct from ''Quetzalcoatlus'' and named as the new genus '' Cryodrakon''. A humerus from the Two Medicine Formation in Montana was also provisionally referred to the genus; it was considered an indeterminate azhdarchid or a specimen of '' Montanazhdarcho'' by subsequent studies. A neck vertebra from the Hell Creek Formation, also from Montana but dating to the Maastrichtian, was discovered in 2002 and initially referred in 2006 to ''Quetzalcoatlus''. The 2021 paper merely considered it an azhdarchiform specimen of uncertain affinities, but the 2025 study named it as the holotype of a distinct genus '' Infernodrakon''.

Another neck vertebra, discovered in the similarly aged Lance Formation in Wyoming and first described in 1964, was later referred to ''Quetzalcoatlus'' by Brent H. Breithaupt in 1982; later studies referred it to '' Azhdarcho'' or as an indeterminate azhdarchid or azhdarchiform. The 2025 ''Infernodrakon'' study found it to be distinguishable from taxon, but anatomically compatible with ''Q. lawsoni''; when tested in a phylogenetic analysis, they found it to form a polytomy with ''Q. lawsoni'', ''Q. northropi'', and a Moroccan specimen. Therefore, they considered it plausible it belonged to a small ''Q. lawsoni'' individually, but decided to merely make a tentative referral to ''Quetzalcoatlus'' at the genus level due to the incomplete nature of the specimen. The Moroccan specimen, from the Ouled Abdoun Basin (Maastrichtian), is designated as FSAC-OB 14 and was referred to aff. ''Quetzalcoatlus'' by a 2018 study. This indicates it is considered unlikely to belong to the genus but bears extreme anatomical similarity to it.

Description

''Quetzalcoatlus northropi'' was among the largest azhdarchids, though was rivalled in size by ''Arambourgiania'' and '' Hatzegopteryx'' (and possibly '' Cryodrakon''). Azhdarchids were split into two primary categories: short-necked taxa with short, robust beaks (i.e. ''Hatzegopteryx'' and ''Wellnhopterus''), and long-necked taxa with longer, slenderer beaks (i.e. '' Zhejiangopterus''). Of these, ''Quetzalcoatlus'' falls squarely into the latter. Based on the limb morphology of ''Q. lawsoni'', related azhdarchids such as ''Zhejiangopterus'', and pterosaurs at large, in addition to azhdarchid tracks from South Korea, ''Quetzalcoatlus'' was likely quadrupedal. As a pterosaur, ''Quetzalcoatlus'' would have been covered in hair-like pycnofibres, and had extensive wing-membranes, which would have been distended by a long wing-finger. There have been various models of the morphology of pterodactyloid wings, though based on multiple well-preserved pterosaur specimens, it is likely that azhdarchids had broad wings, with a brachiopatagium extending down to the ankle. The aspect ratio of azhdarchid wings is 8.1, similar to that of storks and birds of prey that engage in static soaring (relying on air currents to gain altitude and remain aloft).

Size

''Quetzalcoatlus'' is regarded as one of the largest pterosaurs, though its exact size has been difficult to determine. In 1975, Douglas Lawson compared the wing bones of ''Q. northropi'' to equivalent elements in '' Dsungaripterus'' and ''Pteranodon'' and suggested that it represented an individual with a wingspan of around , or, alternatively, or . Estimates put forward in subsequent years varied dramatically, ranging from , owing to differences in methodology. Among the supporters of the initial size estimates was Robert T. Bakker, who, in his 1988 book '' The Dinosaur Heresies'', contended that ''Quetzalcoatlus'' may indeed have reached the upper estimates, and that it may have remained flighted by altering its method of flapping. Other estimates contemporary to Bakker's, however, consistently supported a smaller size of . More recent estimates based on greater knowledge of azhdarchid proportions place its wingspan at . This would approach the maximum size possible for azhdarchids, estimated at ; while higher wingspans may technically be possible, they would require significant morphological changes, and the animal would struggle to become airborne due to increased strain on its joints and long bones. In one paper from the 2021 Memoir which redescribed ''Quetzalcoatlus'', ''Q. lawsoni'' was estimated to have a wingspan of around . In 2022, Gregory S. Paul suggested that it had a somewhat larger wingspan of around and a body length, from beak to tail, of . Large azhdarchids such as ''Q. northropi'' have been estimated to have a shoulder height of about , and the head may have been held at a similar height to that of an extant giraffe.

Body mass estimates for ''Quetzalcoatlus'' have, similarly, been historically variable. Mass estimates for giant azhdarchids are problematic because no existing species shares a similar size or body plan, and in consequence, published results vary widely. Crawford Greenewalt gave mass estimates of between for ''Q. northropi'', with the former figure assuming a small wingspan of , and the latter assuming a far larger wingspan of . In 2010, Donald M. Henderson recovered the body mass of ''Quetzalcoatlus'' at , twice that of other contemporary estimates, citing it as evidence for secondary flightlessness in the genus. However, the vast majority of estimates published since the 2000s have hovered around , due largely to a greater understanding of how aberrant the anatomy of azhdarchids was in comparison to other pterosaur clades. In 2021, Kevin Padian and colleagues estimated that ''Q. northropi'' would have weighed around , and that ''Q. lawsoni'' would have weighed , while a year later, Gregory S. Paul estimated a body mass of for the latter species.

Skull

Complete skulls are not known from either ''Quetzalcoatlus'' species. Reconstructions of its skull anatomy must therefore draw from eight separate ''Q. lawsoni'' specimens which preserve skull elements. Based on the length of the mandible (lower jaw), the skull of ''Q. lawsoni'' likely measured about in length. The distance between the condyloid (articular) processes of the mandible is around . The ratio between the length of the skull and the length of the average dorsal (back) vertebra is very high in ''Q. lawsoni'', and is surpassed only by ''Pteranodon'' and '' Tupuxuara''. The nasoantorbital fenestra, a massive opening found in many pterosaurs which combined the nasal cavity (which housed the external nostril) and antorbital fenestra, was very large in ''Q. lawsoni'', occupying about a third of the total length of the skull. In the largest specimen, TMM 41961-1.1, it measured in length and in height. ''Q. lawsoni'''s choanae (internal nostrils) were confluent and anteriorly expanded, meaning that they were essentially fused and expanded towards the front. The orbit was small and obovate, resembling an inverted egg in shape. It was fairly low on the skull, similar to other azhdarchoids, like tapejarids. Like in other pterosaurs, the orbit was positioned in a way that would have allowed ''Quetzalcoatlus'' to see both forwards and directly to the sides, giving it a degree of stereoscopic vision.

Like other azhdarchids, ''Quetzalcoatlus'' had a long, slender, toothless beak, with a gape of around 52 degrees. It was constructed from the premaxilla and maxilla on the upper jaw, and the dentary on the lower jaw. Beak tips are not preserved in any specimen, so it is not clear whether its tip was sharp or had some other morphology, such as a keel. The dentary had a slight sinusoidal curve, which is also observed in '' Hatzegopteryx''. The mandibular symphyses would have widened slightly as the jaw opened, slightly separating the mandibles, which has led to suggestions that some sort of gular pouch was present. At the base of the beak, formed from the premaxilla, was a crest, referred to by some authors as a sagittal crest. A premaxillary crest is also observed in ''Wellnhopterus'', though is smaller and more anterior. The exact form of the crest in ''Q. lawsoni'' has yet to be determined, due to the poor preservation of the rear half of ''Q. lawsoni''s skull. From what is known, two distinct morphotypes have been suggested: one with a square sagittal crest and a tall nasoantorbital fenestra, and one with a more semicircular sagittal crest and a shorter nasoantorbital fenestra. Additionally, one morphotype is larger than the other, and has a proportionally shorter beak. Despite their differences, both share the diagnostic traits of ''Q. lawsoni'', and are considered the same species. The reason for there being two morphotypes is unclear, though it may correlate to individual variation, ontogeny, or sexual dimorphism.

Skeleton

Like other pterosaurs, ''Quetzalcoatlus'' had light, hollow bones, supported internally by struts called trabeculae. The neck of ''Q. lawsoni'', measured from the third cervical (neck) vertebra to the seventh, has been estimated at . It consisted of nine elongated vertebrae, which were procoelous, meaning that they were concave at the front. All of them were compressed dorsoventrally (top to bottom), and were better suited for dorsoventral motion than lateral (side-to-side) motion. However, the lateral range of motion was still extensive, and the neck and head could swing left and right in an arc of about 180 degrees. Like in other azhdarchoids, the cervical vertebrae were low, with neural arches that were essentially inside the centrum. In most azhdarchids, the neural spine of the seventh cervical vertebra was fairly long. This was not the case in ''Q. lawsoni'', where the neural spine was shorter. Internally, the cervical vertebrae were supported by trabeculae that increased their buckling load substantially (about 90%). This may have been an adaptation for counteracting shear forces exerted on the neck while in flight, though may have also enabled agonistic neck-bashing behaviors like those seen in giraffes. While airborne, the neck of ''Q. northropi'' would have likely assumed a slight S-shaped curve, similar to swans. Similar to other azhdarchids, the torso was proportionally small, about half as long again as the humerus. In ''Quetzalcoatlus'' specifically, the vertebrae at the base of the neck and the pectoral girdle (shoulder girdle) are poorly known. The first four dorsal (back) vertebrae are fused into a notarium, as in some other pterosaurs and birds, particularly ornithocheiroids. The vertebral count of the notarium is unlike '' Zhejiangopterus'', which had six notarial vertebrae, but like '' Azhdarcho''. Most other dorsal vertebrae are absent, except for three which had been integrated into the sacrum. Around seven dorsal vertebrae were free of the notarium and sacrum. Four true sacral vertebrae are preserved, though there were likely seven in all. No caudal (tail) vertebrae are preserved.
''Quetzalcoatlus''' scapulocoracoid, a fusion of the scapula and coracoid, bones, was U-shaped and broad. Throughout azhdarchid evolution, the scapula had been reducing in length, and this reached its pinnacle in ''Q. lawsoni'', where the scapula was only slightly longer than the coracoid. The sternum, like that of other pterosaurs, was broad. It had a transverse ridge on its posterior (rear) portion, also observed in ''Pteranodon'', albeit it is further forward in that genus. Like other pterosaurs, the bone walls of the sternum were very thin, and in the case of ''Q. lawsoni'', they were as little as thick. ''Quetzalcoatlus'' and other azhdarchids have forelimb and hind limb proportions more similar to modern running ungulate mammals than to members of other pterosaur clades, implying that they were uniquely suited to a terrestrial lifestyle. The humerus was short and robust, with considerable mobility. Its morphology differs somewhat between species, with ''Q. lawsoni''s humerus having a proportionally shorter deltopectoral crest, and ''Q. northropi''s being shaped more like a twisted hourglass. The ulna of ''Q. northropi'' was relatively shorter than that of ''Q. lawsoni'', measuring 1.36 times the length of the humerus, as opposed to 1.52 times the length of the humerus in ''Q. lawsoni'' and other azhdarchiforms. Like all pterosaurs, ''Quetzalcoatlus'' had a specialized carpal (wrist) bone, the pteroid, which attached to the sesamoid bone of the wrist. It faced inwards, pointing towards the deltopectoral crest, and supported the propatagium. Specialized muscles allowed it to move slightly. The first digit is the smallest, and the third is the biggest, with the exception of the wing finger. As with all pterosaurs, the wing finger was considerably longer than the others. It did not contact the ground, as evidenced by footprints. Whilst on land, it may have been held between the elbow and the torso. The wings were short and broad, and forelimb musclature was extensive. Flapping power came from several muscle groups on the torso, forearm and manus (hand).

Typically for a pterosaur, the pelvis of ''Q. lawsoni'' consists of an ilium, pubis and ischium that had fused into a single unit, articulating anteroventrally (towards the front, at the bottom) with the prepubis, a small bone that sat before the pubis and likely helped support the abdominal cavity during respiration. The pelvis of one ''Q. lawsoni'' specimen (TMM 41954-57) is large compared to that of other specimens, with deep posterior emargination and no preserved symphisis. This suggests that ''Q. lawsoni'' exhibited sexual dimorphism, similar to that suggested for other monofenestratans (i.e. '' Darwinopterus, Anhanguera'' and '' Nyctosaurus''). The femur was significantly more gracile than the humerus, though was still among the most robust bones in ''Quetzalcoatlus'' skeleton, judging by ''Q. lawsoni''. Based on ''Zhejiangopterus'', the humerus was around eighty percent of the length of the tibiotarsus. Azhdarchids overall had fairly narrow feet, no longer than 30% of the length of the tibia, which may have borne fleshy pads similar to those of tapejarids. They were plantigrade, meaning that they walked with the sole of the foot flat on the ground. ''Q. lawsoni'' possessed well-developed pedal (foot) unguals, which supported moderately curved claws, shorter and slightly straighter than those of tapejarids.

Classification

When describing ''Quetzalcoatlus'' in 1975, Douglas Lawson and Crawford Greenewalt opted not to assign it to a clade more specific than Pterodactyloidea, though comparisons with '' Arambourgiania'' (then ''Titanopteryx'') from Jordan had been drawn earlier that year. In 1984, Lev Alexandrovich Nessov erected the subfamily Azhdarchinae within Pteranodontidae to contain ''Azhdarcho'', ''Quetzalcoatlus'', and ''Titanopteryx''. Unaware of that subfamily, in the same year, Kevin Padian erected the family Titanopterygiidae to accommodate ''Quetzalcoatlus'' and ''Titanopteryx'', defining it based on the length and general morphology of the cervical vertebrae. Two years later, in 1986, noting commonalities not only in contained genera but in diagnostic features, he rendered Titanopterygiidae a junior synonym of Azhdarchinae, elevating the latter to family level and forming the family Azhdarchidae. In 2003, the clade Azhdarchoidea was defined by David Unwin. Azhdarchids were determined to form a clade, Neoazhdarchia, with Tapejaridae. '' Montanazhdarcho'' from North America and '' Zhejiangopterus'' from China were incorporated into Azhdarchidae.Unwin, D. M., (2003). "On the phylogeny and evolutionary history of pterosaurs." Pp. 139-190. in Buffetaut, E. & Mazin, J.-M., (eds.) (2003). ''Evolution and Palaeobiology of Pterosaurs''. Geological Society of London, Special Publications 217, London, 1-347. In the supplementary material for their 2014 paper describing ''Kryptodrakon progenitor'', Andres, James Clark and Xing Xu named a new subfamily, Quetzalcoatlinae, of which ''Quetzalcoatlus'' is the type genus.

The relationship between ''Quetzalcoatlus'' and other giant azhdarchids, like ''Arambourgiania'' and ''Hatzegopteryx'', is not certain. In 2021, Brian Andres recovered them as sister taxa, with ''Arambourgiania'' being the sister taxon of ''Quetzalcoatlus'' and ''Hatzegopteryx'' being slightly more basal. However, Rodrigo V. Pêgas et al., in 2022, instead recovered ''Quetzalcoatlus'' as part of one of two quetzalcoatline branches, alongside ''Cryodrakon''; the other giant azhdarchid genera were recovered on the other branch. A similar dichotomy was recovered by Leonardo Ortiz David et al. that same year, with the inclusion of '' Thanatosdrakon'' as ''Quetzalcoatlus'' sister genus.

The first of the below phylogenetic analyses shows the results of Andres (2021). The second shows the results of Ortiz David et al. (2022).

Topology 1: Andres (2021).

Topology 2: Ortiz David and colleagues (2022).

Paleobiology

Terrestrial locomotion

Azhdarchids are the only group of pterosaurs to which trackways have been confidently assigned. '' Haenamichnus'', from South Korea, matches azhdarchids in foot shape, age, and size. One long trackway of this kind shows that azhdarchids walked with their limbs held directly underneath their body, and along with the morphology of their feet indicates they were more proficient on the ground than other pterosaurs. Terrestrial locomotion in azhdarchids like ''Quetzalcoatlus'' likely involved a pacing gait, wherein the limbs on one side of the body would move at roughly the same time, followed by those of the opposite side. For example, the forelimb on one side of the body would lift off the ground and move forward first, to avoid colliding with the hind foot, and the hind limb would follow suit. The forefoot would be planted in the ground just before the hind foot. Once the stride completed, the same process would repeat on the opposite side of the body.

Flight

The nature of flight in ''Quetzalcoatlus'' and other giant azhdarchids was poorly understood until serious biomechanical studies were conducted in the 21st century. A 1984 experiment by Paul MacCready used practical aerodynamics to test the flight of ''Quetzalcoatlus''. MacCready constructed a model flying machine or, ornithopter, with a simple computer functioning as an autopilot. The model successfully flew with a combination of soaring and wing flapping. The model was based on a then-current weight estimate of around , far lower than more modern estimates of over . The method of flight in these pterosaurs depends largely on their weight, which has been controversial, and widely differing masses have been favored by different scientists. Some researchers have suggested that these animals employed slow, soaring flight, while others have concluded that their flight was fast and dynamic. In 2010, Donald Henderson argued that the mass of ''Q. northropi'' had been underestimated, even the highest estimates, and that it was too massive to have achieved powered flight. He estimated it in his 2010 paper as , and argued that it may have been flightless.

Other flight capability estimates have disagreed with Henderson's research, suggesting instead an animal superbly adapted to long-range, extended flight. In 2010, Mike Habib, a professor of biomechanics at Chatham University, and Mark Witton, a British paleontologist, undertook further investigation into the claims of flightlessness in large pterosaurs. After factoring wingspan, body weight, and aerodynamics, computer modeling led the two researchers to conclude that ''Q. northropi'' was capable of flight up to for 7 to 10 days at altitudes of . Habib further suggested a maximum flight range of for ''Q. northropi''. Henderson's work was also further criticized by Witton and Habib in another study, which pointed out that, although Henderson used excellent mass estimations, they were based on outdated pterosaur models, which caused Henderson's mass estimations to be more than double what Habib used in his estimations and that anatomical study of ''Q. northropi'' and other big pterosaur forelimbs showed a higher degree of robustness than would be expected if they were purely quadrupedal. This study proposed that large pterosaurs most likely utilized a short burst of powered flight to then transition to thermal soaring. However, a study from 2022 suggests that they would only have flown occasionally and for short distances, like the Kori bustard (the world's heaviest bird that actively flies) and that they were not able to soar at all. Studies of ''Q. northropi'' and ''Q. lawsoni'' published in 2021 by Kevin Padian et al. instead suggested that ''Quetzalcoatlus'' was a very powerful flier. While Padian himself also suggested that the legs and feet were likely tucked under the body during flight as in modern birds, co-authors John Conway and James Cunningham endorsed a system more in line with conventional models of pterosaur flight, wherein the hind limbs were splayed out while the animal was airborne.

Launching

Early interpretations of ''Quetzalcoatlus'' launching relied on bipedal models. In 2004, Sankar Chatterjee and R.J. Templin used a model and utilized a running launch cycle powered by the hind limbs, in which ''Q. northropi'' was only barely able to take off. In 2008, Michael Habib suggested that the only feasible takeoff method for a ''Quetzalcoatlus'' was one that was mainly powered by the forelimbs. In 2010, Mark Witton and Habib noted that the femur of ''Quetzalcoatlus'' was only a third as strong as what would be expected from a bird of equal size, whereas the humerus is considerably stronger, and affirmed that an azhdarchid the size of ''Quetzalcoatlus'' would have great difficulty taking off bipedally. Thus, they considered a quadrupedal launching method, with the forelimbs applying most of the necessary force, a likelier method of takeoff. In 2021, Kevin Padian et al. attempted to resurrect the bipedal launch model, using a comparatively light weight estimate of . They suggested that ''Quetzalcoatlus'' hind limbs were more powerful than previously suggested, and that they were strong enough to launch its body as high as off the ground without the aid of the forelimbs. A large breastbone would support the necessary muscles to create a flight stroke, allowing ''Quetzalcoatlus'' to gain enough clearance to begin the downstrokes needed for takeoff. In an blog post written in response to the Padian et al. study, Mark Witton wrote that little of ''Quetzalcoatlus''' morphology indicated that a bipedal launch mechanism was employed, and that quadrupedal launching, even in the smaller ''Q. lawsoni'', remained the likeliest model.

Paleoecology

Definitive fossils of ''Quetzalcoatlus'' have only been found from the Javelina Formation of Texas, though similar and potentially congeneric azhdarchids are known from isolated bones across North America. The formation consists of around of fluvial deposits from the middle to late Maastrichtian, and is gradually overlain by Black Peaks Formation which contains the Cretaceous–Paleogene boundary. At the time of sedimentation, the coastline of the seaway was around southeast of the formation. The fauna is dominated by the large sauropod '' Alamosaurus'', which has been considered characteristic of the fauna of the Lancian of southern Laramidia, with ''Quetzalcoatlus'' noted in association despite its relative inabundance. Specimens of ''Quetzalcoatlus'' are only known from the upper half of the formation where they are the most common vertebrate, with magnetostratigraphy and uranium–lead dating demonstrating that these deposits were from the latest Maastrichtian around 67 to 66 million years ago. Additional pterosaur specimens that may be ''Quetzalcoatlus'' are found in older parts of the formation, though the single oldest pterosaur fossil belongs to the azhdarchid '' Wellnhopterus''. As ''Q. lawsoni'' is known only from the upper part of the formation, and those of ''Q. northropi'' are from the top of the Javelina or the base of the Black Peaks Formations, it is possible that the three taxa succeeded each other, though uncertainty allows for some overlap of the species of ''Quetzalcoatlus''.

Even if ''Q. lawsoni'' and ''Q. northropi'' coexisted, their remains are found in different facies so they may have lived in different habitats. Specimens of ''Q. northropi'' are found in stream-channel deposits consisting of sandstone and conglomerate, where scattered skeletons, limb bones of ''Alamosaurus'', and local log jams are also found. In places, the conglomerates contain accumulations of small bones such as vertebrae, scales, teeth, and osteoderms of gar, fish, turtles, and crocodilians, and shells of gastropods. The flow in these streams fluctuated dramatically and was possibly ephemeral with short duration flooding events. While abandoned channel-lake deposits are the least common in the formation, most specimens of ''Q. lawsoni'' have been found in them. As the channels were abandoned or cut off from active flow, the lakes would develop where sediments would accumulate and occasionally be flooded by nearby streams. Burrows can be found throughout these deposits, bones are normally excrusted with calcite, and the carbonates present suggest photosynthetic algae and microbes as found in alkaline lakes. The lakes were several hundred meters wide but only a few meters deep at their maximum extent, though the main fossil layer is from a single prolonged lake development.

The overbanks surrounding the channels and lakes were covered in a mature forest represented by '' Javelinoxylon'' and araucariaceaen conifers showing a warm, dry, and nonseasonal environment of a tropical lowland. Much of the wood is infested with dry-wood termites. Slender aquatic vines and palm fronds and stalks can be found throughout the abandoned channel-lake deposits. Bioturbation in the lakes show it supported an abundant fauna of soft-bodied invertebrates including crustaceans, insects and annelids with intermittent freshwater bivalves and gastropods. Common aquatic vertebrates found elsewhere are absent, suggesting an inhospitable environment that was perhaps too alkaline. ''Q. lawsoni'' may have fed on the invertebrate fauna, and has a body plan similar to modern wading birds found in alkaline environments such as cranes and storks. The area surrounding lakes was vegetated with palmetto palms and trees in an evergreen or semideciduous forest with a closed canopy over tall. No nearby elevation or branches were present to assist with launching for ''Q. lawsoni'' so it must have been capable of takeoff from limited open spaces and of aerial maneuvering in dense forest. ''Q. northropi'' in contrast is found in channels suggesting a riparian habitat and more solitary lifestyle, but its absence from other similar riparian habitats of the time suggests something was preferable about the Javelina environment. The floodplain would have has a mean average temperature of and a dry and nonseasonal climate, so it ''Quetzalcoatlus'' did migrate in flocks, it would not have been in response to seasonality. The closest modern equivalent to the environment of ''Quetzalcoatlus'' would be the coastal plains of southern Mexico.

Feeding and ecological niche

In 1975, Douglas Lawson rejected the notion that ''Quetzalcoatlus'' might have had a fish-eating (piscivorous) lifestyle like pteranodontids. The Big Bend site where the holotype was discovered is roughly removed from the coastline, and since Lawson believed that the river systems of the locality were too small to support an animal the size of ''Q. northropi'', he instead suggested that it was a scavenger, similar to vultures. The holotype was found in close association with the skeletons of '' Alamosaurus'', a titanosaur sauropod, which Lawson cited as further evidence. However, David Martill dismissed this connection, noting that the association was nothing more than circumstantial evidence. Due to the light construction of ''Quetzalcoatlus''' beak, it would have had to wait for other predators to open up a carcass before it could feed. Another issue is that, while many scavenging birds have extremely flexible necks allowing them to efficiently probe for meat, ''Quetzalcoatlus'' had a stiffer neck, and would not have had the same advantages. Since ''Q. lawsoni'''s beak was relatively weak and unhooked, being compared by Wann Langston Jr. to chopsticks, it is unlikely that it could have torn flesh from a carcass. In the coming decades, further hypotheses regarding ''Quetzalcoatlus''' ecology were put forward. Lev Alexandrovich Nesov, in 1984, suggested that ''Q. northropi'' and other azhdarchids were piscivorous after all, and that they were ecological analogs to modern skimmers, which feed by skimming the water's surface. He did not, however, indicate any morphological indicators for this behavior, and indeed, azhdarchids lack many of the adaptations necessary for such a feeding method (such as robust, aerodynamic beaks with shock-absorbing structures, and the neck flexibility necessary for skimming). Thus, skimming is very unlikely. Another hypothesis, proposed by Langston, is that azhdarchids probed for burrowing invertebrates, using its beak to pluck them from the substrate. Among the evidence provided by Langston was extensive bioturbation of the strata around ''Quetzalcoatlus'' fossils, indicative of extensive invertebrate activity; this has since been suggested to be totally circumstantial. Such a lifestyle has subsequently been suggested for ''Q. lawsoni''. However, wading animals tend to have large, long and broad feet, with a high surface area. This contrasts the morphology seen in ''Q. lawsoni'' and other azhdarchids, which had small, narrow feet.

The predominant model of azhdarchid feeding behavior is the "terrestrial stalking" hypothesis, which suggests that they fed upon small terrestrial prey. A predecessor to this hypothesis was proposed by Gregory S. Paul in a 1987 correspondence published in the journal ''Nature.'' Paul suggested that ''Q. northropi'' was analogous to modern storks, specializing in plucking small vertebrates, such as frogs and turtles, from around watercourses. This initial iteration was supported by Sankar Chatterjee and R. J. Templin, who believed that many pterodactyloids adopted such a lifestyle. Lifestyles analogous to modern herons were also proposed by Kevin Padian and Donna Braginetz in 1988, and by S. Christopher Bennett in 2001. In a 2008 paper on the paleoecology of azhdarchids, Mark Witton and Darren Naish reviewed previous models of their ecology and found the majority to be untenable. The anatomy possessed by azhdarchids was, to them, indicative of terrestrial prey capture. They argued that the family were ecologically closest to storks or ground hornbills, and coined the term "terrestrial stalker" to collectively describe them. Witton elaborated in a 2013 book that the proportions of azhdarchids would have been consistent with them striding through vegetated areas with their long limbs, and their downturned skull and jaws reaching the ground. Their long, stiffened necks would be an advantage as it would help lowering and raising the head and give it a vantage point when searching for prey, and enable them to grab small animals and fruit. The internal anatomy of ''Q. lawsoni''s cervical vertebrae suggests that it, and other mid-sized azhdarchids, may have been able to pick up prey animals weighing at maximum; prey size would have been further limited by the size of ''Q. lawsoni''s skull and gullet rather than body mass. ''Q. lawsoni''s skull does not exhibit cranial kinesis, meaning that the skull was essentially a rigid unit, and as such it was likely incapable of masticating hard objects.

''Q. northropi'' fossils have been found in plains deposits, and due to the paucity and location of its remains, was speculated by Thomas Lehman to have been a solitary hunter that favored riparian environments. ''Q. lawsoni'', on the other hand, is found in great numbers in facies that likely represent alkaline lakes. It may have lived like modern gregarious wading birds, feeding on small invertebrates such as annelids, crustaceans and insects that inhabit such environments, and using its beak to probe for burrowing invertebrates. The two species, if contemporaneous, were likely separated by such behavioral and ecological differences. In a 2021 study, Claudio Labita and David Martill noted that azhdarchids might have been less terrestrial than suggested by Witton and Naish, since azhdarchid fossils were known from marine strata, such as '' Phosphatodraco'' from Morocco and '' Arambourgiania'' from the phosphates of Jordan. They noted that no azhdarchids had been found in truly terrestrial strata, and proposed they could instead have been associated with aquatic environments, such as rivers, lakes, marine and off-shore settings.

Pterosaurs were generally thought to have gone gradually extinct by decreasing in diversity towards the end of the Cretaceous, but Longrich and colleagues suggested this impression could be a result of the poor fossil records for pterosaurs (the Signor-Lipps Effect). Pterosaurs during this time had increased niche-partitioning compared to earlier faunas from the Santonian and Campanian ages, and they were able to outcompete birds in large size based niches, and birds therefore remained small, not exceeding wingspans during the Late Cretaceous (most pterosaurs during this time had larger wingspans, and thereby avoided the small-size niche). To these researchers, this indicated that the extinction of pterosaurs was abrupt instead of gradual, caused by the catastrophic Chicxulub impact. Their extinction freed up more niches that were then filled by birds, which led to their evolutionary radiation in the Early Cenozoic.

Contemporary fauna

Beyond the dominant sauropod ''Alamosaurus'' the Javelina Formation is known from a diverse vertebrate fauna throughout its channel, overbank, and lake deposits. Both the pterosaurs ''Quetzalcoatlus'' and ''Wellnhopterus'' are known, and the dinosaur megafauna is represented by the ceratopsids '' Bravoceratops'' and '' Torosaurus utahensis'', hadrosaurids assigned to '' Gryposaurus'', '' Kritosaurus'' and Saurolophinae, ankylosaurs, and an indeterminate species of ''Tyrannosaurus''. Smaller theropods including the dromaeosaurid '' Saurornitholestes'' and an indeterminate troodontid have been found, as well as a variety of fishes, smaller reptiles, and mammals. Most fishes, smaller reptiles and mammals were found through the screening of microfossil localities, which also discovered scraps of dinosaurs. Fish include the stingray '' Dasyatis'', the bowfin, the gar ''Atractosteus'', while amphibians include an indeterminate frog, and reptiles are represented by the lizard '' Glyptosaurus'', the snake '' Dunnophis'', and the crocodile '' Brachychampsa''. Mammals are very common in microfossil sites of both the Black Peaks and Javelina Formations, represented in the latter by the multituberculates '' Ptilodus'', '' Mesodma'', '' Stygimys'', and '' Viridomys'', indeterminate therians, and the metatherians '' Peratherium'', '' Gelastops'', '' Bryanictis'', '' Mixodectes'', '' Palaechthon'', '' Eoconodon'', '' Carsioptychus'', '' Haploconus'', '' Ellipsodon'', and '' Promioclaenus''.

Cultural significance

In 1975, artist Giovanni Caselli depicted ''Quetzalcoatlus'' as a small-headed scavenger with an extremely long neck in the book ''The evolution and ecology of the Dinosaurs'' by British paleontologist Beverly Halstead. Over the next twenty-five years prior to future discoveries, it would launch similar depictions colloquially known as "paleomemes" in various books, as noted by Darren Naish.

In 1985, the US Defense Advanced Research Projects Agency (DARPA) and AeroVironment used ''Q''. ''northropi'' as the basis for an experimental ornithopter unmanned aerial vehicle (UAV). They produced a half-scale model weighing , with a wingspan of . Coincidentally, Douglas A. Lawson, who discovered ''Q. northropi'' in Texas in 1971, named it after John "Jack" Northrop, a developer of tailless flying wing aircraft in the 1940s. The replica of ''Q. northropi'' incorporates a "flight control system/autopilot which processes pilot commands and sensor inputs, implements several feedback loops, and delivers command signals to its various servo-actuators". It is on exhibit at the National Air and Space Museum.

In 2010, several life-sized models of ''Q. northropi'' were put on display on London's South Bank as the centerpiece exhibit for the Royal Society's 350th-anniversary exhibition. The models, which included both flying and standing individuals with wingspans of over , were intended to help build public interest in science. The models were created by scientists from the University of Portsmouth.

See also

* List of pterosaur genera
* Timeline of pterosaur research

Notes

References

Bibliography

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External links

''Quetzalcoatlus''
at EarthArchives.org
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{{Portal bar, Paleontology, United States

Azhdarchidae
Late Cretaceous pterosaurs of North America
Ojo Alamo Formation
Maastrichtian genus extinctions
Taxa named by Douglas A. Lawson
Fossil taxa described in 1975
Quetzalcoatl