Field crews from The University of Texas at Austin first identified pterosaur remains from the Upper Cretaceous Javelina Formation of Big Bend National Park in 1971 and continued excavation of these animals for decades. The announcement of the giant Quetzalcoatlus northropi in 1975 by graduate student Douglas Lawson drew worldwide attention, and fossil preparators William Amaral and Robert Rainey discovered several key localities in a region informally called Pterodactyl Ridge that have been thoroughly collected and documented. The Pterodactyl Ridge sites produced hundreds of bones from surface collection and quarries through 1986, but later surface collection yielded poorer results. The majority of these elements represent an animal substantially smaller than Q. northropi, Quetzalcoatlus lawsoni Andres and Langston, 2021—historically referred to as Quetzalcoatlus sp. These and subsequent field expeditions from several institutions have reported occurrences of pterosaurs from both the Aguja and Javelina formations, but this study limits only the Javelina Formation material to pterosaurs. Quetzalcoatlus northropi is known within Big Bend National Park only from stream channel facies, and the smaller Q. lawsoni from the upper abandoned channel-lake facies at Pterodactyl Ridge. The lower abandoned channel-lake facies strata of Pterodactyl Ridge produce a third genus and species, Wellnhopterus brevirostris Andres and Langston, 2021. In addition, a smaller azhdarchid is found in the overbank floodplain facies.

The Maastrichtian Javelina Formation of southwestern Texas comprises a thick sequence of stream channel and floodplain deposits accumulated in a broad southeast-trending valley, several hundred kilometers inland from the Late Cretaceous shoreline. Three pterosaur species are found here. Remains of Quetzalcoatlus lawsoni, sp. nov., are concentrated in deposits of shallow alkaline lakes that developed in abandoned reaches of stream channels. Areas surrounding the lakes were vegetated with fan palms, and the higher floodplain supported a subtropical forest dominated by the dicot tree Javelinoxylon and araucariacean conifers. The shallow lakes were inhabited by a diverse invertebrate fauna of arthropods, gastropods, and bivalves, a likely food source for the slender-beaked Quetzalcoatlus lawsoni, sp. nov., which may have had a lifestyle similar to modern large gregarious wading birds. In contrast, remains of the giant Q. northropi are rare and found instead only in stream channel facies. It may have had a more solitary lifestyle and preferred riparian habitats. The warm, dry, subtropical but nonseasonal conditions of the region may represent a preferred climatic regime for azhdarchid pterosaurs generally.

Quetzalcoatlus is the largest flying organism ever known and one of the most familiar pterosaurs to the public. Despite a half century of interest, it remains very incompletely described. This shortfall is addressed here through a full morphological description of Quetzalcoatlus and the other pterosaur material of Big Bend National Park, Texas. The first reported material was described and named Quetzalcoatlus northropi by Douglas Lawson in 1975, but in two separate publications. A ruling by the International Commission of Zoological Nomenclature was required for the name to be made available. Review of the pterosaur fauna of the Park recovers three valid species of azhdarchid pterosaurs in the latest Cretaceous Period Javelina and Black Peaks formations. The size and occurrence of these species are correlated with depositional environment. The holotype of the giant Quetzalcoatlus northropi and six other giant specimens referred to it occur in stream-channel deposits, including the youngest reported pterosaur. The vast majority of specimens (200+) are from large pterosaurs found in the abandoned channel-lake deposits at Pterodactyl Ridge; they form a diagnosable natural group erected as the new species Quetzalcoatlus lawsoni. A moderate-sized partial skull and cervical series also found in the abandoned channel-lake deposits at Pterodactyl Ridge, but lower in the section, is distinct from both species and is erected as Wellnhopterus brevirostris, gen. et sp. nov. Overbank flood-plain facies preserve another eleven specimens of extreme size variation, including small azhdarchids. The Big Bend pterosaur fauna provides the greatest known sample of azhdarchid pterosaurs and three-dimensional pterosaur morphology.

The Azhdarchidae have come to be known as the most diverse clade of Late Cretaceous pterosaurs and the largest flying creatures in existence. Since the erection of the taxon nearly four decades ago, many partial specimens have been referred to it from the Early Cretaceous and Late Jurassic, but none of these identifications can be confirmed. The most comprehensive phylogenetic analysis and taxonomy of Pterosauria is presented, and the evolutionary history of the Azhdarchidae is reviewed. As currently known, azhdarchids are restricted to the Late Cretaceous (Turonian–Maastrichtian). Fourteen species are currently included in the Azhdarchidae: Quetzalcoatlus northropi and Q. lawsoni are recovered as sister taxa in a monophyletic Quetzalcoatlus, with Arambourgiania philadelphiae, Hatzegopteryx thambema, a trichotomy with Cryodrakon boreas and Wellnhopterus brevirostris, Zhejiangopterus linhaiensis, Eurazhdarcho langendorfensis, a Phosphatodraco mauritanicus + Aralazhdarcho bostobensis sister group, as well as an Azhdarcho lancicollis + Albadraco tharmisensis + Aerotitan sudamericanus + Mistralazhdarcho maggii clade are recovered as successive outgroups to Quetzalcoatlus in the Azhdarchidae. The previous azhdarchid species Montanazhdarcho minor and Radiodactylus langstoni are recovered as non-azhdarchid azhdarchiforms; Alanqa saharica and Argentinadraco barrealensis are thalassodromines; Cretornis hlavaci and Volgadraco bogolubovi are pteranodontians; and Bakonydraco galaczi is a tapejarine. Up to a dozen pterosaur lineages persist into the latest Cretaceous (Maastrichtian Age) including azhdarchids, pteranodontids, and nyctosauromorphs. In the Late Cretaceous, an ornithocheirid, cimoliopterids, a lonchodrachonid, a lonchodectid, pteranodontians, tapejarines, thalassodromines, a chaoyangopterine, and azhdarchiforms are present. The pterosaurs did not have a terminal decline in diversity and were increasing in species number at the end of the Cretaceous Period.

We reconstruct the proportions and possible motions of the skeleton of the giant azhdarchid pterosaur Quetzalcoatlus. The neck had substantial dorsoventral mobility, and the head and the neck could swing left and right through an arc of ca. 180°. In flight, it is most plausible that the hind limbs were drawn up bird-like, with the knee anterior to the acetabulum. In this position, an attachment of the wing membrane to the hind limb would have been useless. A straight-legged posterior extension of the hind limb, such as rotation of the hind limb into a fully ‘bat-like’ pose, was likely prevented by soft tissues of the hip joint. Given these difficulties, the traditional ‘broad-winged’ bat-like restoration is unrealistic. On the ground, Quetzalcoatlus, like other ornithodirans, had an erect stance and a parasagittal gait. Terrestrial locomotion was powered almost entirely by the hind limbs. The pace length would have been limited to the length of the glenoacetabular distance, except that Quetzalcoatlus (like other pterodactyloids) had a unique gait in which the forelimb was elevated out of the way of the hind limb from step to step. If the humerus were retracted 80° and adducted nearly to the body wall, the elbow and wrist may have been able to extend to effect a quadrupedal launch with assistance from the hind limbs, assuming sufficient long bone strength and sufficient extensor musculature at these forelimb joints. A bipedal launch using the hind limbs alone also appears plausible: despite the animal’s great size, the hind limb to torso length ratio is the greatest for all known pterosaurs.