Pterosaur - Knowledge

1149: 3230: 903: 4324: 1022: 766: 1169:. The actinofibrils themselves consisted of three distinct layers in the wing, forming a crisscross pattern when superimposed on one another. The function of the actinofibrils is unknown, as is the exact material from which they were made. Depending on their exact composition (keratin, muscle, elastic structures, etc.), they may have been stiffening or strengthening agents in the outer part of the wing. The wing membranes also contained a thin layer of muscle, fibrous tissue, and a unique, complex circulatory system of looping blood vessels. The combination of actinofibrils and muscle layers may have allowed the animal to adjust the wing slackness and 1598:

the time, different from other contemporary feathers that did not carry this formation. The feather fossils obtained from this specimen also suggested the presence of Stage IIIa feathers, a new discovery which may also suggest that more complex feather structures were present at this time. Previously, no Stage III feather forms had been discovered in this time. This study contains multiple indications about the development of feather forms. These include a more precise estimate for the development of avian feather forms, as well as a more ancient ancestor that contained the origins of feather-specific melanosome signaling found in extant birds.

2287: 17180: 17327: 1685: 1618: 2995: 1327:, metacarpals I-III are small and do not connect to the carpus, instead hanging in contact with the fourth metacarpal. With these derived species, the fourth metacarpal has been enormously elongated, typically equalling or exceeding the length of the long bones of the lower arm. The fifth metacarpal had been lost. In all species, the first to third fingers are much smaller than the fourth, the "wingfinger", and contain two, three and four phalanges respectively. The smaller fingers are clawed, with the ungual size varying among species. In 3289:. Earlier suggestions were that pterosaurs were largely cold-blooded gliding animals, deriving warmth from the environment like modern lizards, rather than burning calories. In this case, it was unclear how the larger ones of enormous size, with an inefficient cold-blooded metabolism, could manage a bird-like takeoff strategy, using only the hind limbs to generate thrust for getting airborne. Later research shows them instead as being warm-blooded and having powerful flight muscles, and using the flight muscles for walking as quadrupeds. 4014: 382: 12521: 12510: 15947: 2703: 4551: 1411:

upside-down like bats, hanging from branches and using the fifth toes as hooks. Another hypothesis held that they stretched the brachiopatagia, but in articulated fossils the fifth digits are always flexed towards the tail. Later it became popular to assume that these toes extended an uropatagium or cruropatagium between them. As the fifth toes were on the outside of the feet, such a configuration would only have been possible if these rotated their fronts outwards in flight. Such a rotation could be caused by an

2547: 2182:. A 2007 reply by Dave Hone and Michael Benton could not reproduce this result, finding pterosaurs to be closely related to dinosaurs even without hindlimb characters. They also criticized David Peters for drawing conclusions without access to the primary evidence, that is, the pterosaur fossils themselves. Hone and Benton concluded that, although more basal pterosauromorphs are needed to clarify their relationships, current evidence indicates that pterosaurs are avemetatarsalians, as either the sister group of 1129: 14749: 3222: 986: 1578:. The various forms of filament structure present on the anurognathids in the 2018 study would also require a form of decomposition that would cause the different 'filament' forms seen. They therefore conclude that the most parsimonious interpretation of the structures is that they are filamentous protofeathers. But Liliana D'Alba points out that the description of the preserved integumentary structures on the two anurognathid specimens is still based upon gross morphology. She also points out that 3404: 1056: 135: 188: 2912: 1358: 1316:

in mature specimens, while three of the distal carpals fuse to form a distal syncarpal. The remaining distal carpal, referred to here as the medial carpal, but which has also been termed the distal lateral, or pre-axial carpal, articulates on a vertically elongate biconvex facet on the anterior surface of the distal syncarpal. The medial carpal bears a deep concave fovea that opens anteriorly, ventrally and somewhat medially, within which the pteroid articulates, according to Wilkinson.

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vertically below the body but were somewhat sprawling. The shinbone was often fused with the upper ankle bones into a tibiotarsus that was longer than the thighbone. It could attain a vertical position when walking. The calf bone tended to be slender, especially at its lower end that in advanced forms did not reach the ankle, sometimes reducing total length to a third. Typically, it was fused to the shinbone. The ankle was a simple, "mesotarsal", hinge. The, rather long and slender,

1936: 3015: 4305: 1882: 2807: 14755: 1557:, found that pterosaurs had a wide array of pycnofiber shapes and structures, as opposed to the homogeneous structures that had generally been assumed to cover them. Some of these had frayed ends, very similar in structure to four different feather types known from birds or other dinosaurs but almost never known from pterosaurs prior to the study, suggesting homology. A response to this study was published in 2020, where it was suggested that the structures seen on the 840: 12556: 14132: 4120:

Bennett and David Unwin, to conclude that the young were dependent on their parents for a relatively short period of time, during a period of rapid growth while the wings grew long enough to fly, and then left the nest to fend for themselves, possibly within days of hatching. Alternatively, they may have used stored yolk products for nourishment during their first few days of life, as in modern reptiles, rather than depend on parents for food. Fossilised

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together made for a rigid closed loop, able to withstand considerable forces. A peculiarity was that the breastbone connections of the coracoids often were asymmetrical, with one coracoid attached in front of the other. In advanced species the shoulder joint had moved from the shoulder blade to the coracoid. The joint was saddle-shaped and allowed considerable movement to the wing. It faced sideways and somewhat upwards.

13606: 1449: 3479: 2151: 15955: 2086:). As this would make them also rather close relatives of the dinosaurs, these results were seen by Kevin Padian as confirming his interpretation of pterosaurs as bipedal warm-blooded animals. Because these early analyses were based on a limited number of taxa and characters, their results were inherently uncertain. Several influential researchers who rejected Padian's conclusions offered alternative hypotheses. 4523: 2735: 4509: 1513: 2574: 3273: 1811:, then the largest known pterosaur, the first toothless one and the first from America. These layers too rendered thousands of fossils, also including relatively complete skeletons that were three-dimensionally preserved instead of being strongly compressed as with the Solnhofen specimens. This led to a much better understanding of many anatomical details, such as the hollow nature of the bones. 1189: 977:. The symphysis was often very thin transversely and long, accounting for a considerable part of the jaw length, up to 60%. If a crest was present on the snout, the symphysis could feature a matching mandible crest, jutting out to below. Toothed species also bore teeth in their dentaries. The mandible opened and closed in a simple vertical or "orthal" up-and-down movement. 1665:

important role in the progress of modern paleontology and geology. Scientific opinion at the time was that if such creatures were still alive, only the sea was a credible habitat; Collini suggested it might be a swimming animal that used its long front limbs as paddles. A few scientists continued to support the aquatic interpretation even until 1830, when German zoologist

466:') were smaller animals with fully toothed jaws and, typically, long tails. Their wide wing membranes probably included and connected the hind legs. On the ground, they would have had an awkward sprawling posture, but the anatomy of their joints and strong claws would have made them effective climbers, and some may have even lived in trees. Basal pterosaurs were 1526:(warm-blooded). They aided thermoregulation, as is common in warm-blooded animals who need insulation to prevent excessive heat-loss. Pycnofibers were flexible, short filaments, about five to seven millimetres long and rather simple in structure with a hollow central canal. Pterosaur pelts might have been comparable in density to many Mesozoic mammals. 3381:

to process. The low relative mass of the flocculi in birds is also a result of birds having a much larger brain overall; though this has been considered an indication that pterosaurs lived in a structurally simpler environment or had less complex behaviour compared to birds, recent studies of crocodilians and other reptiles show that it is common for

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number to nine. Instead, the vertebrae themselves became more elongated, up to eight times longer than wide. Nevertheless, the cervicals were wider than high, implying a better vertical than horizontal neck mobility. Pterodactyloids have lost all neck ribs. Pterosaur necks were probably rather thick and well-muscled, especially vertically.

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extensive than the simple pteroid-to-shoulder connection traditionally depicted in life restorations. The position of the pteroid bone itself has been controversial. Some scientists, notably Matthew Wilkinson, have argued that the pteroid pointed forward, extending the forward membrane and allowing it to function as an adjustable

4069:. Egg-burying would have been beneficial to the early evolution of pterosaurs, as it allows for more weight-reducing adaptations, but this method of reproduction would also have put limits on the variety of environments pterosaurs could live in and may have disadvantaged them when they began to face ecological competition from 781:. Traditionally, it was assumed that pterosaurs were extremely light relative to their size. Later, it was understood that this would imply unrealistically low densities of their soft tissues. Some modern estimates therefore extrapolate a weight of up to 250 kilograms (550 pounds) for the largest species. 1436:, natural casts and transformations of the original material. They may include horn crests, beaks or claw sheaths as well as the various flight membranes. Exceptionally, muscles were preserved. Skin patches show small round non-overlapping scales on the soles of the feet, the ankles and the ends of the 4319:

Pterosaurs have been a staple of popular culture for as long as their cousins the dinosaurs, though they are usually not featured as prominently in films, literature or other art. While the depiction of dinosaurs in popular media has changed radically in response to advances in paleontology, a mainly

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While very little is known about pterosaur reproduction, it is believed that, similar to all dinosaurs, all pterosaurs reproduced by laying eggs, though such findings are very rare. The first known pterosaur eggs were found in the quarries of Liaoning, the same place that yielded feathered dinosaurs,

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and several unidentified species show that pterosaurs walked with an erect posture with their four limbs held almost vertically beneath the body, an energy-efficient stance used by most modern birds and mammals, rather than the sprawled limbs of modern reptiles. Indeed, erect-limbs may be omnipresent

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A 2021 study showcases that niches previously occupied by small pterosaurs were increasingly occupied by the juvenile stages of larger species in the Late Cretaceous. Rather than being outcompeted by birds, pterosaurs essentially specialized a trend already occurring in previous eras of the Mesozoic.

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had. This discovery is one of many that leads us away from many previous theories of feathers evolving directly from scales in reptiles, given the significant distinction of melanosome organization and content between the two. This indicates a distinct form of melanosomes within feather structures at

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provides the main propulsive force for the hindlimb, was relatively unimportant. The tail vertebrae were amphicoelous, the vertebral bodies on both ends being concave. Early species had long tails, containing up to fifty caudal vertebrae, the middle ones stiffened by elongated articulation processes,

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Von Soemmerring, S. T., 1812, "Über einen Ornithocephalus oder über das unbekannten Thier der Vorwelt, dessen Fossiles Gerippe Collini im 5. Bande der Actorum Academiae Theodoro-Palatinae nebst einer Abbildung in natürlicher Grösse im Jahre 1784 beschrieb, und welches Gerippe sich gegenwärtig in der

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The flocculus sends out neural signals that produce small, automatic movements in the eye muscles. These keep the image on an animal's retina steady. Pterosaurs may have had such a large flocculus because of their large wing size, which would mean that there was a great deal more sensory information

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is a brain region that integrates signals from joints, muscles, skin and balance organs. The pterosaurs' flocculi occupied 7.5% of the animals' total brain mass, more than in any other vertebrate. Birds have unusually large flocculi compared with other animals, but these only occupy between 1 and 2%

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predecessors have not so far been described, the ancestry of pterosaurs is not fully understood. The oldest known pterosaurs were already fully adapted to a flying lifestyle. Since Seeley, it was recognised that pterosaurs were likely to have had their origin in the "archosaurs", what today would be

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was found to have melanosomes in forms that signal an earlier than anticipated development of the patterns found in extant feathers than previously thought. In these fossils, it appears as though the feather melanosomes took on a more complex form than the melanosome organization in scales that near

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of the thighbone, meaning that the legs would be spread. This would also turn the feet into a vertical position. They then could act as rudders to control yaw. Some specimens show membranes between the toes, allowing them to function as flight control surfaces. The uropatagium or cruropatagium would

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the forelimb digits besides the wingfinger have been lost altogether. The wingfinger accounts for about half or more of the total wing length. It normally consists of four phalanges. Their relative lengths tend to vary among species, which has often been used to distinguish related forms. The fourth

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birds, which fly after hatching without the need of parental care. A further study compares evidence for superprecociality and "late term flight" and overwhelmingly suggests that most if not all pterosaurs were capable of flight soon after hatching. A later study suggested that while smaller-bodied

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For the majority of pterosaur species, it is not known whether they practiced any form of parental care, but their ability to fly as soon as they emerged from the egg and the numerous flaplings found in environments far from nests and alongside adults has led most researchers, including Christopher

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The relative size of the hands and feet in pterosaurs (by comparison with modern animals such as birds) may indicate the type of lifestyle pterosaurs led on the ground. Azhdarchid pterosaurs had relatively small feet compared to their body size and leg length, with foot length only about 25–30% the

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and all their descendants." However, these types of definition would inevitably leave any related species that are slightly more primitive out of the Pterosauria. To remedy this, a new definition was proposed that would anchor the name not to any particular species but to an anatomical feature, the

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David Peters in 2000, published analyses finding pterosaurs to be protorosaurs or closely related to them. However, Peters gathered novel anatomical data using an unverified technique called "Digital Graphic Segregation" (DGS), which involves digitally tracing over images of pterosaur fossils using

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was the first scientist to describe a pterosaur fossil. At that time the concepts of evolution and extinction were imperfectly developed. The bizarre build of the pterosaur was shocking, as it could not clearly be assigned to any existing animal group. The discovery of pterosaurs would thus play an

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The front of the pubic bones articulated with a unique structure, the paired prepubic bones. Together these formed a cusp covering the rear belly, between the pelvis and the belly ribs. The vertical mobility of this element suggests a function in breathing, compensating the relative rigidity of the

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The pterosaur wrist consists of two inner (proximal, at the side of the long bones of the arm) and four outer (distal, at the side of the hand) carpals (wrist bones), excluding the pteroid bone, which may itself be a modified distal carpal. The proximal carpals are fused together into a "syncarpal"

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on the hind feet and folding the wing finger upward to walk on the three-fingered "hand". They could take off from the ground, and fossil trackways show at least some species were able to run and wade or swim. Their jaws had horny beaks, and some groups lacked teeth. Some groups developed elaborate

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in 1983 proposed a hypothetical "propterosaurus": a lizard-like arboreal animal developing a membrane between its limbs, first to safely parachute and then, gradually elongating the fourth finger, to glide. However, subsequent cladistic results did not fit this model well. Neither protorosaurs nor

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in Brazil began to produce chalk nodules that, though often limited in size and the completeness of the fossils they contained, perfectly preserved three-dimensional pterosaur skeletal parts. German and Dutch institutes bought such nodules from fossil poachers and prepared them in Europe, allowing

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In contrast, English and American paleontologists by the middle of the twentieth century largely lost interest in pterosaurs. They saw them as failed evolutionary experiments, cold-blooded and scaly, that hardly could fly, the larger species only able to glide, being forced to climb trees or throw

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A bone unique to pterosaurs, known as the pteroid, connected to the wrist and helped to support the forward membrane (the propatagium) between the wrist and shoulder. Evidence of webbing between the three free fingers of the pterosaur forelimb suggests that this forward membrane may have been more

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eggs suggests that the young pterosaurs had well-developed thigh bones for walking, but weak chests for flight. It is unknown if this holds true for other pterosaurs. Fossils of pterosaurs only a few days to a week old (called "flaplings") have been found, representing several pterosaur families,

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Tracks made by ctenochasmatoids indicate that these pterosaurs swam using their hindlimbs. In general, these have large hindfeet and long torsos, indicating that they were probably more adapted for swimming than other pterosaurs. Pteranodontians conversely have several speciations in their humeri

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Though traditionally depicted as ungainly and awkward when on the ground, the anatomy of some pterosaurs (particularly pterodactyloids) suggests that they were competent walkers and runners. Early pterosaurs have long been considered particularly cumbersome locomotors due to the presence of large

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were found in an Early Cretaceous deposit in northwest China. Examination of the shells by scanning electron microscopy showed the presence of a thin calcareous eggshell layer with a membrane underneath. A study of pterosaur eggshell structure and chemistry published in 2007 indicated that it is

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was robust and not very shortened. It was connected to the ankle in a higher position than the other metatarsals. It bore a long, and often curved, mobile clawless fifth toe consisting of two phalanges. The function of this element has been enigmatic. It used to be thought that the animals slept

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While historically thought of as simple leathery structures composed of skin, research has since shown that the wing membranes of pterosaurs were highly complex dynamic structures suited to an active style of flight. The outer wings (from the tip to the elbow) were strengthened by closely spaced

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Ezcurra, Martín D.; Nesbitt, Sterling J.; Bronzati, Mario; Dalla Vecchia, Fabio Marco; Agnolin, Federico L.; Benson, Roger B. J.; Brissón Egli, Federico; Cabreira, Sergio F.; Evers, Serjoscha W.; Gentil, Adriel R.; Irmis, Randall B.; Martinelli, Agustín G.; Novas, Fernando E.; Roberto da Silva,

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The necks of pterosaurs were relatively long and straight. In pterodactyloids, the neck is typically longer than the torso. This length is not caused by an increase of the number of vertebrae, which is invariably seven. Some researchers include two transitional "cervicodorsals" which brings the

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discovered pterosaur species had risen to 130. Over ninety percent of known taxa has been named during the "renaissance". Many of these were from groups the existence of which had been unknown. Advances in computing power enabled researchers to determine their complex relationships through the

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that is relatively long in pterosaurs. In advanced species, their combined whole, the scapulocoracoid, was almost vertically oriented. The shoulder blade in that case fitted into a recess in the side of the notarium, while the coracoid likewise connected to the breastbone. This way, both sides

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show considerable variation in the extent of their wing membranes and it is possible that, like these groups, different species of pterosaur had different wing designs. Indeed, analysis of pterosaur limb proportions shows that there was considerable variation, possibly reflecting a variety of

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of many of the pterosaurs. It was thought that by the end of the Cretaceous, only large species of pterosaurs were present (no longer true; see below). The smaller species were thought to have become extinct, their niche filled by birds. However, pterosaur decline (if actually present) seems

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after a comparable structure in birds. This was an adaptation to withstand the forces caused by flapping the wings. The notarium included three to seven vertebrae, depending on the species involved but also on individual age. These vertebrae could be connected by tendons or a fusion of their

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Since the 1990s, new discoveries and a more thorough study of old specimens have shown that crests are far more widespread among pterosaurs than previously assumed. That they were extended by or composed completely of keratin, which does not fossilize easily, had misled earlier research. For

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have again named many new taxa. As discoveries also increased in other parts of the world, a sudden surge in the total of named genera took place. By 2009, when they had increased to about ninety, this growth showed no sign of levelling-off. In 2013, M.P. Witton indicated that the number of

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The hindlimbs of pterosaurs were strongly built, yet relative to their wingspans smaller than those of birds. They were long in comparison to the torso length. The thighbone was rather straight, with the head making only a small angle with the shaft. This implies that the legs were not held

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When standing, pterosaurs probably rested on their metacarpals, with the outer wing folded to behind. In this position, the "anterior" sides of the metacarpals were rotated to the rear. This would point the smaller fingers obliquely to behind. According to Bennett, this would imply that the

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or upper arm bone is short but powerfully built. It sports a large deltopectoral crest, to which the major flight muscles are attached. Despite the considerable forces exerted on it, the humerus is hollow or pneumatised inside, reinforced by bone struts. The long bones of the lower arm, the

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control pitch. When walking the toes could flex upwards to lift the membrane from the ground. In Pterodactyloidea, the fifth metatarsal was much reduced and the fifth toe, if present, little more than a stub. This suggests that their membranes were split, increasing flight maneuverability.

1222:("arm membrane") was the primary component of the wing, stretching from the highly elongated fourth finger of the hand to the hindlimbs. Finally, at least some pterosaur groups had a membrane that stretched between the legs, possibly connecting to or incorporating the tail, called the 1144:

and extended along the sides of the body. Where they ended has been very controversial but since the 1990s a dozen specimens with preserved soft tissue have been found that seem to show they attached to the ankles. The exact curvature of the trailing edge, however, is still equivocal.

4100:) for their age, and wing proportions similar to adults. In fact, many pterosaur flaplings have been considered adults and placed in separate species in the past. Additionally, flaplings are normally found in the same sediments as adults and juveniles of the same species, such as the 1051:

into a "supraneural plate". Their ribs also would be tightly fused into the notarium. In general, the ribs are double headed. The sacrum consisted of three to ten sacral vertebrae. They too, could be connected via a supraneural plate that, however, would not contact the notarium.

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was long and low, its front and rear blades projecting horizontally beyond the edges of the lower pelvic bones. Despite this length, the rod-like form of these processes indicates that the hindlimb muscles attached to them were limited in strength. The, in side view narrow,

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teeth, varying in build, and some still had teeth in the palate. In later groups the teeth mostly became conical. Front teeth were often longer, forming a "prey grab" in transversely expanded jaw tips, but size and position were very variable among species. With the derived

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suggested that pterosaurs used a vaulting mechanism to obtain flight. The tremendous power of their winged forelimbs would enable them to take off with ease. Once aloft, pterosaurs could reach speeds of up to 120 km/h (75 mph) and travel thousands of kilometres.

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Pterosaurs had a wide range of sizes, though they were generally large. The smallest species had a wingspan no less than 25 centimetres (10 inches). The most sizeable forms represent the largest known animals ever to fly, with wingspans of up to 10–11 metres (33–36 feet).

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mammals. Their hind limbs, on the other hand, were not built for speed, but they were long compared with most pterosaurs, and allowed for a long stride length. While azhdarchid pterosaurs probably could not run, they would have been relatively fast and energy efficient.

510:, suggesting that early feathers evolved in the common ancestor of pterosaurs and dinosaurs, possibly as insulation. In life, pterosaurs would have had smooth or fluffy coats that did not resemble bird feathers. They were warm-blooded (endothermic), active animals. The 2363:-sized adult azhdarchid further indicate that small pterosaurs from the Late Cretaceous might actually have simply been rarely preserved in the fossil record, helped by the fact that there is a strong bias against terrestrial small sized vertebrates such as juvenile 2478:

The precise relationships between pterosaurs is still unsettled. Many studies of pterosaur relationships in the past have included limited data and were highly contradictory. However, newer studies using larger data sets are beginning to make things clearer. The

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The first to fourth toes were long. They had two, three, four and five phalanges respectively. Often the third toe was longest; sometimes the fourth. Flat joints indicate a limited mobility. These toes were clawed but the claws were smaller than the hand claws.

1299:. This view was contradicted in a 2007 paper by Chris Bennett, who showed that the pteroid did not articulate as previously thought and could not have pointed forward, but rather was directed inward toward the body as traditionally interpreted. Specimens of 10268: 4152:, due to the fast rate the limb bones closest to the body grew compared to any other element of their skeleton after hatching. Other factors mentioned were the limits of soft shelled eggs and the size of the pelvic opening of large female pterosaurs. 1561:

were actually a result of the decomposition of aktinofibrils: a type of fibre used to strengthen and stiffen the wing. However, in a response to this, the authors of the 2018 paper point to the fact that the presence of the structures extend past the

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was described to have feathers to support the claim that feathers had a common origin with Ornithodirans but was argued against by several authors. The only method to assure if it was homologous to feathers is to use a scanning electron microscope.

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Traditionally, almost all pterosaurs were seen as surface-feeding piscivores or fish-eaters, a view that still dominates popular science. Today, many pterosaurs groups are thought to have been terrestrial carnivores, omnivores or insectivores.

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Like the upper jaws, the paired lower jaws of pterosaurs were very elongated. In advanced forms, they tended to be shorter than the upper cranium because the jaw joint was in a more forward position. The front lower jaw bones, the dentaries or

557:, which excludes the pterosaurs. Pterosaurs are nonetheless more closely related to birds and other dinosaurs than to crocodiles or any other living reptile, though they are not bird ancestors. Pterosaurs are also colloquially referred to as 478:) evolved many sizes, shapes, and lifestyles. Pterodactyloids had narrower wings with free hind limbs, highly reduced tails, and long necks with large heads. On the ground, they walked well on all four limbs with an upright posture, standing 2352:. Small azhdarchoid pterosaurs were also present in the Campanian. This suggests that late Cretaceous pterosaur faunas were far more diverse than previously thought, possibly not even having declined significantly from the early Cretaceous. 802:

Compared to the other vertebrate flying groups, the birds and bats, pterosaur skulls were typically quite large. Most pterosaur skulls had elongated jaws. Their skull bones tend to be fused in adult individuals. Early pterosaurs often had

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A 2009 study showed that pterosaurs had a lung-and-air-sac system and a precisely controlled skeletal breathing pump, which supports a flow-through pulmonary ventilation model in pterosaurs, analogous to that of birds. The presence of a

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were unusually long compared to other pterosaurs, and, in azhdarchids, the bones of the arm and hand (metacarpals) were particularly elongated. Furthermore, as a whole, azhdarchid front limbs were proportioned similarly to fast-running

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Pterosauria has usually been defined as node-based and anchored to several extensively studied taxa as well as those thought to be primitive. One 2003 study defined Pterosauria as "The most recent common ancestor of the Anurognathidae,

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A related problem is the origin of pterosaur flight. Like with birds, hypotheses can be ordered into two main varieties: "ground up" or "tree down". Climbing a tree would cause height and gravity to provide both the energy and a strong

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Pterosaurs had a variety of lifestyles. Traditionally seen as fish-eaters, the group is now understood to have also included hunters of land animals, insectivores, fruit eaters and even predators of other pterosaurs. They reproduced by

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ornithodirans are biologically equivalent to lizards. Furthermore, the transition between gliding and flapping flight is not well-understood. More recent studies on basal pterosaur hindlimb morphology seem to vindicate a connection to

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Butler, Richard J.; Barrett, Paul M.; Nowbath, Stephen & Upchurch, Paul (2009). "Estimating the effects of sampling biases on pterosaur diversity patterns: implications for hypotheses of bird/pterosaur competitive replacement".

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are known to possess prehensile feet and hands respectively; all other known pterosaurs have flat, plantigrade feet with no opposable toes, and the feet are generally proportionally small, at least in the case of the Pteranodontia.

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itself was named after a fishing method known as "skim-feeding", later understood to be biomechanically impossible. Perhaps it pursued relatively large prey, in view of its reinforced jaw joints and relatively high bite force.

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show the pteroid in articulation with the proximal syncarpal, suggesting that the pteroid articulated with the 'saddle' of the radiale (proximal syncarpal) and that both the pteroid and preaxial carpal were migrated centralia.

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to be seen as a generalist or a terrestrial predator of small vertebrates, but the highly robust humerus and high-aspect wing morphology, suggest it may have been capable of grabbing prey on the wing. The small insectivorous

1080:. Such tails acted as rudders, sometimes ending at the rear in a vertical diamond-shaped or oval vane. In pterodactyloids, the tails were much reduced and never stiffened, with some species counting as few as ten vertebrae. 7469:

Cuvier, G., 1809, "Mémoire sur le squelette fossile d'un Reptil volant des environs d'Aichstedt, que quelques naturalistes ont pris pour un oiseau, et donc nous formons un genre de Sauriens, sous le nom de Ptero-Dactyle",

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produced thousands of pterosaur fossils, that however, were of poor quality, consisting mostly of strongly eroded fragments. Nevertheless, based on these, numerous genera and species would be named. Many were described by

3548:), adapted to walking in soft muddy soil, similar to modern wading birds. Though clearly forelimb-based launchers, basal pterosaurs have hindlimbs well adapted for hopping, suggesting a connection with archosaurs such as 11292:

Frey, E., Martill, D., and Buchy, M. (2003). "A new crested ornithocheirid from the Lower Cretaceous of northeastern Brazil and the unusual death of an unusual pterosaur" in: Buffetaut, E., and Mazin, J.-M. (eds.).

1997:. Insights from other fields of biology were applied to the data obtained. All this resulted in a substantial progress in pterosaur research, rendering older accounts in popular science books completely outdated. 1037:

The torso was relatively short and egg-shaped. The vertebrae in the back of pterosaurs originally might have numbered eighteen. With advanced species a growing number of these tended to be incorporated into the

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Yang, Zixiao; Jiang, Baoyu; McNamara, Maria E.; Kearns, Stuart L.; Pittman, Michael; Kaye, Thomas G.; Orr, Patrick J.; Xu, Xing; Benton, Michael J. (December 2020). "Reply to: No protofeathers on pterosaurs".

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photo editing software. Bennett only recovered pterosaurs as close relatives of the protorosaurs after removing characteristics of the hindlimb from his analysis, to test the possibility of locomotion-based

1725:

redescribed the original specimen and an additional one. He saw them as affiliated to birds and bats. Although he was mistaken in this, his "bat model" would be influential during the 19th century. In 1843,

3426:(thigh bone) is only moderately inward facing, suggesting that pterosaurs had an erect stance. It would have been possible to lift the thigh into a horizontal position during flight, as gliding lizards do. 832:. This feature likely evolved to lighten the skull for flight. In contrast, the bones behind the eye socket contracted and rotated, strongly inclining the rear skull and bringing the jaw joint forward. The 1919:

especially propagated the new views, publishing a series of studies depicting pterosaurs as warm-blooded, active and running animals. This coincided with a revival of the German school through the work of

4054:, described in 2011, which also had a leathery shell and, also like modern reptiles but unlike birds, was fairly small compared to the size of the mother. In 2014 five unflattened eggs from the species 1710:. At first most species were assigned to this genus and ultimately "pterodactyl" was popularly and incorrectly applied to all members of Pterosauria. Today, paleontologists limit the term to the genus 899:

covered their teeth with jawbone tissue for a crushing function. If teeth were present, they were placed in separate tooth sockets. Replacement teeth were generated behind, not below, the older teeth.

1832:. This material gave birth to a German school of pterosaur research, which saw flying reptiles as the warm-blooded, furry and active Mesozoic counterparts of modern bats and birds. In 1882, Marsh and 925:

in which case it is called a "supraoccipital crest". Front and rear crests can be present simultaneously and might be fused into a single larger structure, the most expansive of which is shown by the

4415:

had to add inaccurate bat-like wing fingers to his stop motion models in order to keep the membranes from falling apart, though this particular error was common in art even before the film was made.

2204:

included various proposed pterosaur relatives, yet also found pterosaurs to be closer to dinosaurs and unrelated to more basal taxa. Working from his 1996 analysis, Bennett published a 2020 study on

6210:

Wang X, Zhou Z, Zhang F, Xu X (2002). "A nearly completely articulated rhamphorhynchoid pterosaur with exceptionally well-preserved wing membranes and "hairs" from Inner Mongolia, northeast China".

4461:

was intended to be depicted, nesting behavior that was known to be inaccurate by 2001, and leathery wings, rather than the taut membranes of muscle fiber required for pterosaur flight. Petrie from

2315:. It seems that the K-Pg extinction event at the end of the Cretaceous, which wiped out all non-avian dinosaurs and many other animals, was the direct cause of the extinction of the pterosaurs. 6658:"Pterosaurs evolved a muscular wing–body junction providing multifaceted flight performance benefits: Advanced aerodynamic smoothing, sophisticated wing root control, and wing force generation" 1734:. Ironically, as the "bat model" depicted pterosaurs as warm-blooded and furred, it would turn out to be more correct in certain aspects than Cuvier's "reptile model" in the long run. In 1834, 1340:

wingfinger, able to describe the largest arc of any wing element, up to 175°, was not folded by flexion but by an extreme extension. The wing was automatically folded when the elbow was bowed.

873:

beak tissue has been preserved, though in toothed forms, the beak is small and restricted to the jaw tips and does not involve the teeth. Some advanced beaked forms were toothless, such as the

9497: 10395: 1243:

There has been considerable argument among paleontologists about whether the main wing membranes (brachiopatagia) attached to the hindlimbs, and if so, where. Fossils of the rhamphorhynchoid

9273: 6479:

Zhou, Chang-Fu; Schoch, Rainer R. (2011). "New material of the non-pterodactyloid pterosaur Changchengopterus pani Lü, 2009 from the Late Jurassic Tiaojishan Formation of western Liaoning".

1349:

also identified a membranous "fairing" (area conjunctioning the wing with the body at the neck), as opposed to the feathered or fur-composed "fairing" seen in birds and bats respectively.

3608:

were nocturnal, aerial insectivores. With highly flexible joints on the wing finger, a broad, triangular wing shape, large eyes and short tail, these pterosaurs were likely analogous to

1534:

Pterosaur filaments could share a common origin with feathers, as speculated in 2002 by Czerkas and Ji. In 2009, Kellner concluded that pycnofibers were structured similarly to theropod

2455:. Starting from the 21st century, new discoveries are now filling in these gaps and giving a better picture of the evolution of pterosaurs. Traditionally, they were organized into two 6253:

Frey, E.; Tischlinger, H.; Buchy, M.-C.; Martill, D. M. (2003). "New specimens of Pterosauria (Reptilia) with soft parts with implications for pterosaurian anatomy and locomotion".

10311:

Pêgas, Rodrigo V; Costa, Fabiana R; Kellner, Alexander W A (24 September 2021). "Reconstruction of the adductor chamber and predicted bite force in pterodactyloids (Pterosauria)".

1814:

Meanwhile, finds from the Solnhofen had continued, accounting for the majority of complete high-quality specimens discovered. They allowed to identify most new basal taxa, such as

935:

sported a bizarre antler-like crest. The crests were only a few millimetres thin transversely. The bony crest base would typically be extended by keratinous or other soft tissue.

10758:

Xiaolin Wang, Kellner Alexander W.A.; Cheng, Xin; Jiang, Shunxing; Wang, Qiang; Sayão Juliana, M.; Rordrigues Taissa, Costa Fabiana R.; Li, Ning; Meng, Xi; Zhou, Zhonghe (2015).

1639:

are very rare, due to their light bone construction. Complete skeletons can generally only be found in geological layers with exceptional preservation conditions, the so-called

10009:"Evidence for tactile foraging in pterosaurs: a sensitive tip to the beak of Lonchodraco giganteus (Pterosauria, Lonchodectidae) from the Upper Cretaceous of southern England" 2311:

unrelated to bird diversity, as ecological overlap between the two groups appears to be minimal. In fact, at least some avian niches were reclaimed by pterosaurs prior to the

4457:. Paleontologist Dave Hone noted that the pterosaurs in this film had not been significantly updated to reflect modern research. Errors persisting were teeth while toothless 812:, the skulls became even more elongated, sometimes surpassing the combined neck and torso in length. This was caused by a stretching and fusion of the front snout bone, the 9964:

Lü J.; Xu L.; Chang H.; Zhang X. (2011). "A new darwinopterid pterosaur from the Middle Jurassic of western Liaoning, northeastern China and its ecological implications".

1240:

pterosaurs had a broader uro/cruropatagium stretched between their long fifth toes, with pterodactyloids, lacking such toes, only having membranes running along the legs.

2108:

was also suggested. Some basal archosauromorphs seem at first glance to be good candidates for close pterosaur relatives due to their long-limbed anatomy; one example is

4144:

pterosaurs were most likely superprecocial or precocial, owing to the consistent or decreasing wing aspect ratio during growth, certain large-bodied pterosaurs, such as

3341:

air sac system in at least some pterodactyloids would have further reduced the density of the living animal. Like modern crocodilians, pterosaurs appeared to have had a

9218:

Lü J.; Unwin D.M.; Xu L.; Zhang X. (2008). "A new azhdarchoid pterosaur from the Lower Cretaceous of China and its implications for pterosaur phylogeny and evolution".

1332:

phalanx is usually the shortest. It lacks a claw and has been lost completely by nyctosaurids. It is curved to behind, resulting in a rounded wing tip, which reduces

6960:

Yang, Zixiao; Jiang, Baoyu; McNamara, Maria E.; Kearns, Stuart L.; Pittman, Michael; Kaye, Thomas G.; Orr, Patrick J.; Xu, Xing; Benton, Michael J. (January 2019).

4378:, whose name means "toy finger" for its resemblance to old, inaccurate children's toys. Pterosaurs have sometimes been incorrectly identified as (the ancestors of) 3723:

were first considered aerial predators. Lacking a robust jaw structure or powerful flying muscles, they are now seen as arboreal or semiterrestrial insectivores.

2249:

similarities with pterosaurs. The results of the latter study were subsequently supported by an independent analysis of early pterosauromorph interrelationships.

1176:

As shown by cavities in the wing bones of larger species and soft tissue preserved in at least one specimen, some pterosaurs extended their system of respiratory

1980:

that since the 1990s has brought forth hundreds of exquisitely preserved two-dimensional fossils, often showing soft tissue remains. Chinese researchers such as

8941:"Does morphology reflect osteohistology-based ontogeny? A case study of Late Cretaceous pterosaur jaw symphyses from Hungary reveals hidden taxonomic diversity" 4135:

may have practiced some form of parental care. However, this study has since been criticised. Most evidence currently leans towards pterosaur hatchlings being

1432:

or ultraviolet light photography has revealed many traces not visible to the naked eye. These are often imprecisely called "impressions" but mostly consist of

9370: 9047: 7797: 5493:

Witton, Mark P.; Martill, David M.; Loveridge, Robert F. (2010). "Clipping the Wings of Giant Pterosaurs: Comments on Wingspan Estimations and Diversity".

3991: 915:

The public image of pterosaurs is defined by their elaborate head crests. This was influenced by the distinctive backward-pointing crest of the well-known

4801:"Testing pterosaur ingroup relationships through broader sampling of avemetatarsalian taxa and characters and a range of phylogenetic analysis techniques" 3245:

Katsufumi Sato, a Japanese scientist, did calculations using modern birds and concluded that it was impossible for a pterosaur to stay aloft. In the book

3455:

were later found with a distinctive four-toed hind foot and three-toed front foot; these are the unmistakable prints of pterosaurs walking on all fours.

3257:

based their research on the now-outdated theories of pterosaurs being seabird-like, and the size limit does not apply to terrestrial pterosaurs, such as

2214:

and pterosaurs were non-archosaur archosauromorphs, albeit not particularly closely related to each other. By contrast, a later 2020 study proposed that

1403:

was always splayed to some degree. The foot was plantigrade, meaning that during the walking cycle the sole of the metatarsus was pressed onto the soil.

1386:

was not perforated and allowed considerable mobility to the leg. It was directed obliquely upwards, preventing a perfectly vertical position of the leg.

726:

The two groups overlapped in time, but the earliest pterosaurs in the fossil record are basal pterosaurs, and the latest pterosaurs are pterodactyloids.

4342:

The vague generic term "pterodactyl" is often used for these creatures. The animals depicted in fiction and pop culture frequently represent either the

12613: 3625:

analogue in the past, is indicated by its jaw structure, gait, and poor flight capabilities, as a terrestrial/semiarboreal predator of small mammals,

8880: 1216:("fore membrane"), was the forward-most part of the wing and attached between the wrist and shoulder, creating the "leading edge" during flight. The 1120:, jutted obliquely upwards. The rear edge of the breastbone was the deepest point of the thorax. Clavicles or interclavicles were completely absent. 10403: 10247: 9281: 7789: 1796:

had suggested pterosaurs were the direct ancestors of birds. Owen opposed the views of both men, seeing pterosaurs as cold-blooded "true" reptiles.

921:. The main positions of such crests are the front of the snout, as an outgrowth of the premaxillae, or the rear of the skull as an extension of the 9304: 545:

Pterosaurs are often referred to by popular media or the general public as "flying dinosaurs", but dinosaurs are defined as the descendants of the

11431:

Zhou, X.; Pêgas, R. V.; Ma, W.; Han, G.; Jin, X.; Leal, M. E. C.; Bonde, N.; Kobayashi, Y.; Lautenschlager, S.; Wei, X.; Shen, C.; Ji, S. (2021).

17694: 17228: 5072: 4096:

including pterodactylids, rhamphorhinchids, ctenochasmatids and azhdarchids. All preserve bones that show a relatively high degree of hardening (

1382:

into an ischiopubic blade. Sometimes, the blades of both sides were also fused, closing the pelvis from below and forming the pelvic canal. The

1112:, was wide. It had only a shallow keel. Via sternal ribs, it was at its sides attached to the dorsal ribs. At its rear, a row of belly ribs or 9070:"Small, immature pterosaurs from the Cretaceous of Africa: implications for taphonomic bias and palaeocommunity structure in flying reptiles" 4574: 4188:

A 2021 study indicates that pterosaur juveniles of larger species increasingly took the roles previously occupied by adult small pterosaurs.

2258: 2087: 3979:, embedded in it. The vertebrae are known not to have been eaten and exposed to digestion, as the joints are still articulated. Fossils of 1463:-like filaments known as pycnofibers on the head and torso. The term "pycnofiber", meaning "dense filament", was coined by palaeontologist 4087:

Wing membranes preserved in pterosaur embryos are well developed, suggesting that pterosaurs were ready to fly soon after birth. However,

4084:, as opposed to the single functional ovary in birds, dismissing the reduction of functional ovaries as a requirement for powered flight. 3315:. The replica was launched with a ground-based winch. It flew several times in 1986 and was filmed as part of the Smithsonian's IMAX film 1136:

Pterosaur wings were formed by bones and membranes of skin and other tissues. The primary membranes attached to the extremely long fourth

7451:

Cuvier G (1801). ". In: Extrait d'un ouvrage sur les espèces de quadrupèdes dont on a trouvé les ossemens dans l'intérieur de la terre".

5717:

Czerkas, S.A., and Ji, Q. (2002). A new rhamphorhynchoid with a headcrest and complex integumentary structures. In: Czerkas, S.J. (Ed.).

3668:

likely had a strong bite force, indicating an adaptation towards hard food items that might have been chewed in view of the tooth wear.

1406:

There was a clear difference between early pterosaurs and advanced species regarding the form of the fifth digit. Originally, the fifth

9427: 4444: 2471:(unnatural) group, since the pterodactyloids evolved directly from them and not from a common ancestor, so, with the increasing use of 751:

habits, mouth bristles, and feet adapted for clinging. Parallel adaptations are seen in birds and bats that prey on insects in flight.

8263:

Hone D.W.E.; Benton M.J. (2007). "An evaluation of the phylogenetic relationships of the pterosaurs to the archosauromorph reptiles".

4048:, a pterosaur known by abundant material. This was supported by the description of an additional pterosaur egg belonging to the genus 743:) is debated. Anurognathids were highly specialized. Small flyers with shortened jaws and a wide gape, some had large eyes suggesting 17584: 11589: 9479: 7424:

Collini, C.A. (1784). "Sur quelques Zoolithes du Cabinet d'Histoire naturelle de S. A. S. E. Palatine & de Bavière, à Mannheim."

2312: 9904:"A new rhamphorhynchid pterosaur from the Upper Jurassic of Xinjiang, China, and the phylogenetic relationships of basal pterosaurs" 5445:"A new rhamphorhynchid pterosaur from the Upper Jurassic of Xinjiang, China, and the phylogenetic relationships of basal pterosaurs" 17740: 11386: 10612:

Lü J.; Unwin D.M.; Deeming D.C.; Jin X.; Liu Y.; Ji Q. (2011). "An egg-adult association, gender, and reproduction in pterosaurs".

7097:"The Soft Tissue of Jeholopterus (Pterosauria, Anurognathidae, Batrachognathinae) and the Structure of the Pterosaur Wing Membrane" 1148: 2420:

in 2020 as "he clade characterized by the apomorphy fourth manual digit hypertrophied to support a wing membrane, as inherited by

1905:, whose spectacular traits refuted what had become entrenched orthodoxy. In 1970, likewise the description of the furry pterosaur 1861:

themselves from cliffs to achieve a take-off. In 1914, for the first-time pterosaur aerodynamics were quantitatively analysed, by

494:, which covered their bodies and parts of their wings. Pycnofibers grew in several forms, from simple filaments to branching down 11337: 8349:"The phylogenetic relationships of basal archosauromorphs, with an emphasis on the systematics of proterosuchian archosauriforms" 2467:, "advanced" pterosaurs with short tails. However, this traditional division has been largely abandoned. Rhamphorhynchoidea is a 4200:

of pterosaurs and modern birds and reptiles have been used to infer daily activity patterns of pterosaurs. The pterosaur genera

4124:

nests were shown preserving many male and female pterosaurs together with their eggs in a manner to a similar to that of modern

4038:

were squashed flat with no signs of cracking, so evidently the eggs had leathery shells, as in modern lizards. The egg from the

3931:

was corroborated as a specialised consumer of hard plant material with a relatively high BFQ and high mechanical advantage, and

3229: 8885:

Varricchio, 2002 from the Two Medicine Formation (Upper Cretaceous) of Western USA (Montana) as a pterosaur rather than a bird"

6961: 1899:

took place, a quick increase in the number of studies and critical ideas, influenced by the discovery of additional fossils of

3269:

concluded that atmospheric differences between the present and the Mesozoic were not needed for the giant size of pterosaurs.

1792:. Seeley thought that pterosaurs were warm-blooded and dynamic creatures, closely related to birds. Earlier, the evolutionist 17589: 12592: 11533: 11514: 11495: 9340: 5563: 5338: 5280: 4917: 4867: 4271: 973:, were at the tip tightly fused into a central symphysis. This made the lower jaws function as a single connected whole, the 7144:

Unwin, David M.; Bakhurina, Natasha N. (September 1994). "Sordes pilosus and the nature of the pterosaur flight apparatus".

2359:, indicating a higher diversity of Late Cretaceous pterosaurs than previously accounted for. The recent findings of a small 902: 10269:"An unusual modification of the jaws in cf. Alanqa, a mid-Cretaceous azhdarchid pterosaur from the Kem Kem beds of Morocco" 3345:, seeing as their shoulder-pectoral girdles were too inflexible to move the sternum as in birds, and they possessed strong 2416:

presence of an enlarged fourth finger that supports a wing membrane. This "apomorophy-based" definition was adopted by the

1993:. New and old fossils yielded much more information when subjected to modern ultraviolet light or roentgen photography, or 1428:

The rare conditions that allowed for the fossilisation of pterosaur remains, sometimes also preserved soft tissues. Modern

9397: 1497:

and ultraviolet examination of pterosaur specimens have provided incontrovertible proof: pterosaurs had pycnofiber coats.

615:

between elements disappeared. In some later pterosaurs, the backbone over the shoulders fused into a structure known as a

17221: 8787:"Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary" 4155:

Growth rates of pterosaurs once they hatched varied across different groups. In more primitive, long-tailed pterosaurs ("

1924:, who in 1970s laid the foundations of modern pterosaur science. In 1978, he published the first pterosaur textbook, the 3522:

length of the lower leg. This suggests that azhdarchids were better adapted to walking on dry, relatively solid ground.

17745: 10487:(4 (supplement, abstracts from XX Jornadas Argentinas de Paleontología de Vertebrados, La Plata, 26-28 May 2004)): 40R. 9068:

Smith, Roy E.; Chinsamy, Anusuya; Unwin, David M.; Ibrahim, Nizar; Zouhri, Samir; Martill, David M. (16 October 2021).

6553: 6392: 1965:

their scientists to describe many new species and revealing a whole new fauna. Soon, Brazilian researchers, among them

3877:, estimating the bite force and potential dietary habits of nine selected species. The study corroborated the view of 1014:. Advanced pterosaurs are unique in possessing special processes projecting adjacent to their condyle and cotyle, the 11507:

The Illustrated Encyclopedia of Pterosaurs: An Illustrated Natural History of the Flying Reptiles of the Mesozoic Era

11358: 11309: 9181: 5726: 4670: 2391: 2295: 17179: 9048:"Pterosaur remains (Archosauria, Ornithodira) from the early Late Cretaceous of "La Buitrera", Río Negro, Argentina" 8515: 5263:(2004). "Origin and relationships of Dinosauria". In Weishampel, David B.; Dodson, Peter; Osmólska, Halszka (eds.). 4323: 591:

The anatomy of pterosaurs was highly modified from their reptilian ancestors by the adaptation to flight. Pterosaur

9602: 9360: 4569: 611:

able to coordinate complex flying behaviour. Pterosaur skeletons often show considerable fusion. In the skull, the

8082: 4163:, the average growth rate during the first year of life was 130% to 173%, slightly faster than the growth rate of 3652:

has been found with fish remains in its stomach, but its dentition suggests an opportunistic diet. Slender-winged

1262:

seem to demonstrate that the wing membrane did attach to the hindlimbs, at least in some species. However, modern

1021: 603:

attachment surface for a given skeletal weight. The bone walls were often paper-thin. They had a large and keeled

3563:

interpreted to have been suggestive of a water-based version of the typical quadrupedal launch, and several like

1369:

of pterosaurs was of moderate size compared to the body as a whole. Often the three pelvic bones were fused. The

8540: 2266:. Like this archosaur, basal pterosaur lineages have plantigrade hindlimbs that show adaptations for saltation. 17214: 9205: 9158: 8940: 4599: 4329: 1607: 14754: 11139:

Yang, Zixiao; Jiang, Baoyu; Benton, Michael J.; Xu, Xing; McNamara, Maria E.; Hone, David W. E. (2023-07-26).

6457: 5151:(March 1975). "Pterosaur from the Latest Cretaceous of West Texas: Discovery of the Largest Flying Creature". 3943:

were proposed to be ground-feeding generalists with intermediate bite force values and less specialised jaws.

3782:

at the tip of its beak; birds with similarly numerous foramina have sensitive beaks used to feel for food, so

11582: 11221:

Schmitz, L.; Motani, R. (2011). "Nocturnality in Dinosaurs Inferred from Scleral Ring and Orbit Morphology".

10092:(Pterodactyloidea: Azhdarchoidea) from the Early Cretaceous Jehol Biota and its paleoecological implications" 1722: 1042:. Such species also often show a fusion of the front dorsal vertebrae into a rigid whole which is called the 12520: 10197:"Neck biomechanics indicate that giant Transylvanian azhdarchid pterosaurs were short-necked arch predators" 6506:

Wang, Xiao-Lin; Kellner, Alexander W. A.; Jiang, Shun-Xing; Cheng, Xin; Meng, Xi; Rodrigues, Taissa (2010).

765: 462:

There were two major types of pterosaurs. Basal pterosaurs (also called 'non-pterodactyloid pterosaurs' or '

4391: 1866: 1727: 35: 15946: 14748: 10859: 8915: 4167:. Growth in these species slowed after sexual maturity, and it would have taken more than three years for 3835:

were specialist molluscivores, using their powerful jaws to crush the shells of molluscs and crustaceans.

1857:

determined that the brains of pterosaurs more resembled those of birds than modern cold-blooded reptiles.

17348: 17326: 10476: 10253: 7822: 4542: 4442:

released during the 1960s, 1970s, 1990s, and 2000s, and also appeared in the 2019 American-produced film

2015: 1485: 881:, and had larger, more extensive, and more bird-like beaks. Some groups had specialised tooth forms. The 17: 12509: 5091:"Discovery of a rare arboreal forest-dwelling flying reptile (Pterosauria, Pterodactyloidea) from China" 4474:, not reflecting their full dietary variation. They are also often shown as aerial predators similar to 187: 17440: 17276: 12559: 10760:"Eggshell and Histology Provide Insight on the Life History of a Pterosaur with Two Functional Ovaries" 4905: 4564: 4451:

After the 1960s, pterosaurs remained mostly absent from notable American film appearances until 2001's

4396: 3810:

were suspension feeders, using their numerous fine teeth to filter small organisms from shallow water.

1538:. Others were unconvinced, considering the difference with the "quills" found on many of the bird-like 1440:. They covered pads cushioning the impact of walking. Scales are unknown from other parts of the body. 452: 12527: 10008: 4354:, or a fictionalized hybrid of the two. Many children's toys and cartoons feature "pterodactyls" with 17750: 14131: 12585: 11695: 9723:"New pterosaur tracks (Pteraichnidae) from the Late Cretaceous Uhangri Formation, southwestern Korea" 6986: 3895: 2448: 2196:

benefited from far more data and found strong support for pterosaurs being avemetatarsalians, though

1793: 1700:. Cuvier agreed in 1801, understanding it was an extinct flying reptile. In 1809, he coined the name 1661: 1628: 10252:. The Annual Symposium of Vertebrate Paleontology and Comparative Anatomy. Edinburgh. Archived from 9107:

Kellner, A. W. (2003). "Pterosaur phylogeny and comments on the evolutionary history of the group".

8202:

Irmis, R. B.; Nesbitt, S. J.; Padian, K.; Smith, N. D.; Turner, A. H.; Woody, D.; Downs, A. (2007).

4969: 12689: 12438: 11610: 11575: 11197: 8457:

Lúcio; Smith, Nathan D.; Stocker, Michelle R.; Turner, Alan H.; Langer, Max C. (17 December 2020).

5752: 4579: 4364:-like tails and teeth, a combination that never existed in nature. However, at least one pterosaur 4360: 4335: 3311: 3298: 3294: 1535: 1274:

The bony elements of the arm formed a mechanism to support and extend the wing. Near the body, the

13590: 9620:

Codorniú, Laura; Paulina Carabajal, Ariana; Pol, Diego; Unwin, David; Rauhut, Oliver W.M. (2016).

6021:(Pterosauria, Anurognathidae, Batrachognathinae) and the structure of the pterosaur wing membrane" 14167: 13598: 5048: 4584: 3791: 3349:. Thus, their respiratory system had characteristics comparable to both modern archosaur clades. 3317: 2757: 861: 17712: 14136: 11035:"Prenatal development in pterosaurs and its implications for their postnatal locomotory ability" 8735: 4131:. Due to how underdeveloped the chests of the hatchlings were for flying, it was suggested that 3709:), and shorter, broader wings. These were either terrestrial/aerial predators of vertebrates or 2114:, a "protorosaur" with skin membranes on its hindlimbs likely used for gliding. A 1999 study by 17611: 17579: 17523: 17518: 17503: 17087: 15245: 12132: 11009: 10148:

Pêgas, R. V., & Kellner, A. W. (2015). Preliminary mandibular myological reconstruction of

9171: 8989:

Martin-Silverstone, Elizabeth; Witton, Mark P.; Arbour, Victoria M.; Currie, Philip J. (2016).

4953: 4407: 3901: 3886: 3373: 3367: 1941: 1784:, at the time the main English expert on the subject, who also wrote the first pterosaur book, 1623: 1232:

seem to suggest that it simply connected the legs but did not involve the tail (rendering it a

1010:) was concave and into it fitted a convex extension at the rear of the preceding vertebra, the 17186: 12503: 10249:

Pterosaur overlords of Transylvania: short-necked giant azhdarchids in Late Cretaceous Romania

9498:"The wingtips of the pterosaurs: Anatomy, aeronautical function and 3 ecological implications" 9069: 5328: 5272: 3612:

or extant insectivorous bats, being capable of high manoeuvrability at relatively low speeds.

3575:, which presumably still needed to launch from water in case they found themselves in it. The 3458:

Fossil footprints show that pterosaurs stood with the entire foot in contact with the ground (

2356: 955:

photography. While fossil crests used to be restricted to the more advanced Pterodactyloidea,

17707: 17649: 17594: 17192: 12222: 9330: 4401: 4254:, being active throughout the day for short intervals. As a result, the possibly fish-eating 2397: 1862: 1800: 1490: 1170: 17699: 11387:"'Godzilla: King of the Monsters' Trailer Turns Mothra, Rodan, and More Into Epic Spectacle" 10715:

Grellet-Tinner G, Wroe S, Thompson MB, Ji Q (2007). "A note on pterosaur nesting behavior".

10498:

Chiappe, Luis M.; Codorniú, Laura; Grellet-Tinner, Gerald; Rivarola, David (December 2004).

9453: 5264: 4399:. They appeared in a number of films and television programs since, including the 1933 film 4262:

may have had similar activity patterns to modern nocturnal seabirds, and the filter-feeding

3831:

were arboreal omnivores, supplementing seeds and fruits with small insects and vertebrates.

2286: 17668: 17533: 17460: 17445: 15954: 15340: 13779: 13689: 13605: 12578: 12514: 11960: 11446: 11230: 11095: 10963: 10901: 10812: 10724: 10679: 10621: 10568: 10433: 10351: 10280: 10103: 10020: 9973: 9918: 9832: 9734: 9687: 9227: 9116: 9002: 8955: 8845: 8746: 8573: 8473: 8218: 7916: 7644: 7311: 7197: 7153: 7053: 6814: 6669: 6262: 6219: 6176: 6084: 5976: 5459: 5377: 5218: 5160: 5102: 4855: 4763: 4463: 4436: 4246: 3927: 3249:

it is theorized that they were able to fly due to the oxygen-rich, dense atmosphere of the

2175: 1896: 1476: 849: 707: 546: 15962: 3864:

was a robustly built predator of relatively large prey, including medium-sized dinosaurs.

2487:

analysis presented by Longrich, Martill and Andres in 2018, with clade names after Andres

518:, which hollowed out their bones to an extreme extent. Pterosaurs spanned a wide range of 8: 17735: 17528: 17455: 16844: 16533: 16220: 15519: 15446: 13214: 12934: 11668: 8516:"Paleontologists find pterosaur precursors that fill a gap in early evolutionary history" 4556: 4109: 4039: 4027: 3939: 3933: 3922: 3580: 3242:

The mechanics of pterosaur flight are not completely understood or modeled at this time.

3010: 2569: 2367:, and that their diversity might actually have been much larger than previously thought. 2066: 1986: 1781: 1776: 1684: 1666: 1653: 1412: 825: 410: 11450: 11234: 11173: 11140: 11099: 10967: 10905: 10816: 10728: 10683: 10625: 10572: 10437: 10355: 10284: 10107: 10024: 9977: 9922: 9836: 9738: 9691: 9587: 9231: 9120: 9006: 8959: 8849: 8750: 8577: 8477: 8222: 7920: 7648: 7552:"Versuch einer Eintheilung der Säugethiere in 6 Stämme und der Amphibien in 6 Ordnungen" 7315: 7201: 7157: 7057: 7018: 6818: 6673: 6266: 6223: 6180: 6088: 5980: 5463: 5381: 5222: 5164: 5106: 5049:"It's Official: Those Flying Reptiles Called Pterosaurs Were Covered in Fluffy Feathers" 4859: 4767: 3750:

caused ornithocheirans and the later nyctosaurids to be aerial dip-feeders like today's

3687:, were fish-eaters with long, slender wings, needle-like dentition and long, thin jaws. 2090:

proposed an ancestry among the basal Archosauromorpha, specifically long-necked forms ("

619:, which served to stiffen the torso during flight, and provide a stable support for the 17543: 17508: 17261: 16373: 14847: 14690: 13833: 13636: 13623: 13537: 13379: 13010: 12985: 12461: 11254: 11116: 11083: 11059: 11034: 10986: 10947: 10925: 10917: 10838: 10740: 10645: 10477:"Primer reporte de un embrión de pterosaurio (Cretácico inferior, San Luis, Argentina)" 10457: 10377: 10223: 10196: 10126: 10087: 10044: 9989: 9934: 9848: 9788: 9750: 9703: 9648: 9622:"A Jurassic pterosaur from Patagonia and the origin of the pterodactyloid neurocranium" 9621: 9556: 9251: 9132: 9089: 9023: 8990: 8971: 8907: 8861: 8813: 8786: 8762: 8659: 8632: 8497: 8430: 8399: 8375: 8348: 8329: 8280: 8242: 8187: 8170: 8154: 8125: 8035: 7940: 7932: 7814: 7660: 7332: 7299: 7280: 7231: 7169: 7121: 7096: 7077: 6999: 6837: 6802: 6713: 6700: 6657: 6575: 6548: 6461: 6414: 6387: 6327: 6278: 6235: 6192: 6107: 6073:"Respiratory evolution facilitated the origin of pterosaur flight and aerial gigantism" 6072: 6045: 6016: 6015:

Kellner, A.W.A.; Wang, X.; Tischlinger, H.; Campos, D.; Hone, D.W.E.; Meng, X. (2009).

5992: 5696:

Naish D, Martill DM (2003). "Pterosaurs – a successful invasion of prehistoric skies".

5475: 5401: 5242: 5184: 5125: 5090: 4932: 4827: 4800: 4291: 3747: 3325: 3146: 3108: 2994: 2857: 2460: 2254: 2200:

was not included due to its poor preservation. A 2016 archosauromorph-focused study by

1833: 1468: 1296: 1237: 748: 542:, a good oxygen supply and strong muscles made pterosaurs powerful and capable flyers. 511: 499: 398: 182: 10418: 9769: 9672: 8203: 7501:

Naturalien-Sammlung der königlichen Akademie der Wissenschaften zu München befindet",

7044:

Unwin, David M.; Martill, David M. (December 2020). "No protofeathers on pterosaurs".

5362: 5203: 4181:, grew to adult size within the first year of life. Additionally, pterodactyloids had 1617: 1018:, and the cotyle also may possess a small prong on its midline called a hypapophysis. 17559: 17538: 17498: 17378: 17296: 16799: 15614: 15543: 14785: 14772: 14715: 14478: 14376: 14278: 14196: 13974: 13951: 13917: 13847: 13388: 13304: 13256: 13230: 12943: 12848: 11529: 11510: 11491: 11464: 11305: 11246: 11178: 11160: 11121: 11064: 10991: 10830: 10781: 10697: 10649: 10637: 10594: 10537: 10529: 10449: 10369: 10228: 10131: 10048: 10036: 9993: 9985: 9903: 9852: 9707: 9653: 9548: 9361:"Pterosaurs breathed in bird-like fashion and had inflatable air sacs in their wings" 9336: 9243: 9201: 9177: 9154: 9136: 9093: 9028: 8818: 8718: 8664: 8501: 8489: 8458: 8435: 8380: 8234: 8204:"A Late Triassic Dinosauromorph Assemblage from New Mexico and the Rise of Dinosaurs" 8027: 8022: 8005: 7892:

Studies of the structure, evolution, and flight of pterosaurs (reptilia: Pterosauria)

7337: 7272: 7235: 7223: 7126: 7081: 7069: 6991: 6842: 6717: 6705: 6687: 6580: 6529: 6438: 6419: 6319: 6314: 6297: 6282: 6112: 6050: 5722: 5663: 5559: 5444: 5393: 5334: 5276: 5265: 5260: 5234: 5176: 5148: 5130: 5066: 4913: 4863: 4832: 4781: 4453: 3672: 3631: 3571:-like aerial hawking. These adaptations are also seen in terrestrial pterosaurs like 3533: 2884: 2822: 2770: 2675: 2596: 2002: 1966: 1961: 1946: 1735: 1464: 1361:

An anhanguerid pelvis seen from above, with the right side rotated towards the viewer

1301: 1259: 1212:

The pterosaur wing membrane is divided into three basic units. The first, called the

1077: 644: 484: 266: 11258: 10929: 10744: 10668:"Sexually Dimorphic Tridimensionally Preserved Pterosaurs and Their Eggs from China" 9938: 9754: 9560: 9255: 8991:"A small azhdarchoid pterosaur from the latest Cretaceous, the age of flying giants" 8975: 8911: 8865: 8766: 8585: 8333: 8284: 8039: 7944: 7818: 7664: 7551: 7284: 7003: 6524: 6507: 6492: 6465: 6331: 6239: 5996: 5479: 5363:"Neuroanatomy of flying reptiles and implications for flight, posture and behaviour" 5246: 5188: 4116:

flaplings from Argentina. All are found in deep aquatic environment far from shore.

4013: 3342: 3338: 3225:

Diagrams showing breathing motion (top two) and internal air sac system (bottom two)

1873:

as a pure glider. Little research was done on the group during the 1940s and 1950s.

381: 17569: 17343: 17028: 16911: 16903: 16695: 16624: 16549: 16091: 15994: 15981: 15913: 15793: 15782: 15032: 14889: 14826: 14601: 14583: 14421: 14245: 14061: 14047: 14005: 13511: 13497: 13429: 13108: 12658: 12116: 11876: 11680: 11454: 11238: 11168: 11152: 11111: 11103: 11084:"Powered flight in hatchling pterosaurs: Evidence from wing form and bone strength" 11054: 11046: 10981: 10971: 10909: 10842: 10820: 10771: 10732: 10687: 10629: 10584: 10576: 10519: 10511: 10461: 10441: 10381: 10359: 10320: 10288: 10218: 10208: 10175: 10121: 10111: 10028: 9981: 9926: 9840: 9784: 9742: 9695: 9643: 9633: 9583: 9538: 9235: 9124: 9081: 9018: 9010: 8963: 8899: 8853: 8808: 8798: 8754: 8708: 8654: 8644: 8581: 8481: 8425: 8415: 8370: 8360: 8321: 8311: 8272: 8246: 8226: 8182: 8149: 8141: 8017: 7924: 7806: 7652: 7327: 7319: 7262: 7213: 7205: 7173: 7161: 7116: 7108: 7061: 6981: 6973: 6832: 6822: 6695: 6677: 6570: 6562: 6519: 6488: 6453: 6409: 6401: 6309: 6270: 6227: 6196: 6184: 6102: 6092: 6040: 6032: 5984: 5967:

Bennett SC (2000). "Pterosaur flight: the role of actinofibrils in wing function".

5659: 5467: 5405: 5385: 5226: 5168: 5120: 5110: 5023: 5003: 4822: 4812: 4771: 4724: 4629: 4528: 4185:, meaning that the animals reached a fixed maximum adult size and stopped growing. 4156: 4018: 3957: 3874: 3836: 3832: 3807: 3509: 2977: 2787: 2715: 2464: 2451:

of pterosaurs has historically been difficult, because there were many gaps in the

2193: 2099: 2071: 2022:, and it has been described as the world's best-preserved skeleton of a pterosaur. 2000:

In 2017 a fossil from a 170-million-year-old pterosaur, later named as the species

1921: 1847: 1773: 1715: 1437: 999: 896: 886: 809: 612: 572: 463: 456: 248: 10892:

from the Solnhofen Limestone of Germany: Year-classes of a single large species".

10776: 10759: 10292: 10032: 9128: 9085: 7756:

Zittel, K.A. (1882). "Über Flugsaurier aus dem lithografischen Schiefer Bayerns".

6941:

Goldfuss, A (1831). "Beiträge zur Erkentniss verschiedner Reptilien der Vorwelt".

6274: 3422:

Pterosaurs' hip sockets are oriented facing slightly upwards, and the head of the

3385:

to achieve high intelligence levels with small brains. Studies on the endocast of

2702: 2546: 2165:" which some controversial studies have posited as a close relative of pterosaurs. 17513: 17417: 17400: 17271: 17117: 16960: 16809: 16594: 16498: 16473: 16466: 16453: 16386: 16272: 16170: 16136: 16032: 16012: 15866: 15724: 15665: 15652: 15623: 15499: 15410: 15269: 15142: 15097: 14969: 14898: 14877: 14703: 14364: 14344: 14316: 14292: 14228: 14220: 14213: 14205: 13960: 13940: 13770: 13746: 13696: 13504: 13487: 13477: 13454: 13410: 13402: 13370: 13359: 13338: 13292: 13284: 13223: 13139: 13079: 12998: 12967: 12857: 12836: 12789: 12674: 12649: 12549: 12427: 12370: 11981: 11759: 11737: 11707: 11598: 11437: 10976: 10952:

inferred from bone histology and the diversity of pterosaurian growth strategies"

10580: 10324: 10164:"Azhdarchid pterosaurs: water-trawling pelican mimics or "terrestrial stalkers"?" 10116: 10007:

Martill, David M.; Smith, Roy E.; Longrich, Nicholas; Brown, James (2021-01-01).

9574:

Hopson J.A. (1977). "Relative Brain Size and Behavior in Archosaurian Reptiles".

9196:

Padian, K. (1997). "Pterosauromorpha", pp. 617–18 in Currie, P.J. and Padian, K.

8803: 8300:"The early evolution of archosaurs: relationships and the origin of major clades" 7531:

Newman, E (1843). "Note on the Pterodactyle Tribe considered as Marsupial Bats".

6827: 6097: 5204:"A new giant pterosaur with a robust skull from the latest cretaceous of Romania" 5172: 4937: 4589: 4514: 4483: 4412: 4350: 4236: 3917: 3841: 3796: 3787: 3756: 3743: 3714: 3701: 3677: 3654: 3452: 3392: 3250: 2529: 2371: 2339: 2274: 2143: 2075: 2019: 1887: 1816: 1267: 1154: 1089: 1006:. The vertebrae in front of the tail were "procoelous": the cotyle (front of the 961: 908: 882: 874: 739: 695: 650: 429: 320: 232: 54: 11561: 6379: 1641: 1128: 393:

locations. Colored species or genera names correspond to their taxonomic group.

17574: 17465: 17388: 17335: 17306: 17266: 17241: 17094: 17067: 17045: 16869: 16783: 16768: 16720: 16659: 16644: 16610: 16435: 16419: 16407: 16393: 16346: 16303: 16263: 16177: 16123: 15851: 15844: 15837: 15809: 15716: 15637: 15564: 15475: 15231: 15123: 15058: 14999: 14939: 14905: 14625: 14552: 14524: 14443: 14393: 14332: 14301: 13901: 13877: 13855: 13801: 13759: 13703: 13671: 13352: 13269: 13246: 13125: 13051: 13019: 12872: 12821: 12769: 12544: 12400: 11901: 11814: 11780: 11277: 11107: 10556: 8758: 7323: 7209: 7065: 4691: 4594: 4492: 4197: 4172: 4001: 3962: 3907: 3878: 3786:

may have used its beak to feel for fish or invertebrates in shallow water. The

3735: 3638: 3605: 3306: 3221: 3165: 3127: 2907: 2698: 2542: 2348: 2334: 2238: 2221:

were the sister clade to pterosauria. This was based on newly described fossil

2201: 2115: 1851: 1840:. German studies continued well into the 1930s, describing new species such as 1808: 1772:, also the first non-pterodactyloid pterosaur known. Later in the century, the 1697: 1693: 1558: 1550: 1467:

and colleagues in 2009. Pycnofibers were unique structures similar to, but not

1097: 1064: 1007: 991: 985: 795: 760: 719: 701: 671: 528: 519: 475: 361: 162: 30:"Pterodactyl" redirects here. For the genus commonly called "pterodactyl", see 11459: 11432: 10946:

Prondvai, E.; Stein, K.; Ősi, A.; Sander, M. P. (2012). Soares, Daphne (ed.).

10913: 10736: 10692: 10667: 10246:

Witton, M.; Brusatte, S.; Dyke, G.; Naish, D.; Norell, M.; Vremir, M. (2013).

10152:(Pterodactyloidea: Tapejaridae). Flugsaurier 2015 Portsmouth, abstracts, 47–48 9930: 9746: 9699: 9239: 8857: 8838:

Earth and Environmental Science Transactions of the Royal Society of Edinburgh

8697:"Early penguin fossils, plus mitochondrial genomes, calibrate avian evolution" 8485: 8276: 7928: 7907:

Padian K (1983). "A Functional Analysis of Flying and Walking in Pterosaurs".

7267: 7250: 6977: 6803:"A reappraisal of azhdarchid pterosaur functional morphology and paleoecology" 6656:

Pittman, Michael; Barlow, Luke A.; Kaye, Thomas G.; Habib, Michael B. (2021).

5988: 5471: 5230: 4776: 4751: 3403: 2911: 1981: 1055: 134: 17729: 17634: 17395: 17311: 17301: 17131: 17102: 17078: 17059: 17017: 17002: 16951: 16940: 16833: 16825: 16775: 16712: 16683: 16667: 16651: 16637: 16578: 16556: 16507: 16400: 16161: 16104: 16023: 15858: 15830: 15802: 15673: 15556: 15347: 15277: 15149: 15132: 14988: 14925: 14869: 14634: 14590: 14471: 14464: 14457: 14450: 14435: 14385: 14309: 14266: 14259: 14185: 14032: 14025: 14018: 13867: 13809: 13794: 13787: 13735: 13518: 13187: 13147: 13097: 13066: 13027: 12922: 12914: 12892: 12796: 12780: 12393: 12386: 12342: 12321: 12276: 12139: 11930: 11567: 11164: 10598: 10533: 10040: 6691: 4488: 4208: 4202: 4050: 3860: 3824: 3802: 3739: 3731: 3719: 3695: 3644: 3550: 3538: 3488: 3089: 2990: 2927: 2894: 2801: 2512: 2452: 2407: 2375: 2343: 2323: 2318:

In the early 2010s, several new pterosaur taxa were discovered dating to the

2206: 2162: 2156: 2137: 2120: 2110: 2095: 2091: 1912: 1854: 1822: 1706: 1674: 1650: 1588: 1433: 1357: 1345: 1320: 1307: 1284: 1165: 1092:

was a strong structure that transferred the forces of flapping flight to the

947: 941: 922: 855: 689: 665: 656: 638: 567: 534: 351: 340: 310: 293: 174: 150: 92: 50: 31: 11242: 10633: 8713: 8696: 8230: 6682: 5115: 4478:, grasping human victims with talons on their feet. However, only the small 3014: 2131: 1935: 828:

of pterodactyloid pterosaurs merged into a single large opening, called the

769:

Size disparity of late Maastrichtian Pterosaurs compared to birds and humans

17124: 17052: 16923: 16818: 16752: 16704: 16514: 16442: 16320: 16288: 16056: 15823: 15751: 15692: 15587: 15224: 15209: 15193: 15184: 15173: 15013: 14976: 14862: 14812: 14655: 14568: 14561: 14541: 14513: 14506: 14492: 14485: 14400: 14357: 14324: 14040: 13982: 13925: 13910: 13890: 13715: 13446: 13438: 13418: 13345: 13324: 13317: 13172: 12960: 12906: 12865: 12415: 12334: 12266: 12211: 12146: 12104: 12097: 12089: 12047: 11938: 11885: 11829: 11805: 11468: 11250: 11182: 11156: 11125: 11068: 11050: 10995: 10860:"Pterosaur hatchlings needed their parents, trove of eggs reveals (Update)" 10834: 10785: 10701: 10641: 10541: 10453: 10373: 10232: 10180: 10163: 10135: 9657: 9552: 9365: 9356: 9247: 9032: 8822: 8722: 8668: 8493: 8459:"Enigmatic dinosaur precursors bridge the gap to the origin of Pterosauria" 8439: 8384: 8238: 8145: 8031: 7341: 7276: 7227: 7130: 7112: 7073: 6995: 6846: 6709: 6584: 6566: 6533: 6508:"New long-tailed pterosaurs (Wukongopteridae) from western Liaoning, China" 6423: 6405: 6323: 6116: 6054: 6036: 5761:

Occasional Papers of the Natural History Museum of the University of Kansas

5397: 5238: 5180: 5134: 4836: 4785: 4479: 4475: 4427: 4374: 4304: 4270:

birds that feed at night. The differences between activity patterns of the

4267: 4230: 4136: 4044: 3882: 3847: 3812: 3689: 3544: 3505: 3471: 3438: 3408: 3266: 3258: 2806: 2729: 2652: 2484: 2468: 2246: 2054: 2046: 2007: 1916: 1881: 1842: 1769: 1678: 1660:, became much sought after by rich collectors. In 1784, Italian naturalist 1568: 1539: 1505: 1370: 1333: 1254: 1250: 1073: 1048: 891: 878: 735: 731: 554: 523: 156: 8171:"The phylogenetic position of the Pterosauria within the Archosauromorpha" 5008: 4991: 4618:

Adapted from Witton (2013). Taxonomic groups based on Unwin et al. (2010).

4080:

specimen showcases that at least some pterosaurs had a pair of functional

3648:

were highly aerial animals and fast, agile flyers with long robust wings.

17681: 17660: 17643: 17472: 17422: 17405: 17363: 17358: 17353: 17010: 16970: 16889: 16853: 16587: 16362: 16295: 16238: 16184: 16041: 15709: 15580: 15572: 15532: 15491: 15483: 15358: 15325: 15293: 15285: 15256: 15217: 15201: 15160: 15089: 15081: 15074: 15020: 14961: 14953: 14614: 14576: 14499: 14154: 14087: 14054: 13990: 13331: 13200: 13155: 13132: 12807: 12379: 12358: 12350: 12307: 12160: 12153: 12125: 12068: 12061: 12022: 11916: 11909: 11859: 11836: 11822: 11791: 9527:"Breathing in a box: Constraints on lung ventilation in giant pterosaurs" 4320:

outdated picture of pterosaurs has persisted since the mid-20th century.

4224: 4128: 3996: 3985: 3828: 3774: 3751: 3683: 3617: 3576: 3568: 3564: 3498: 3459: 3443: 3290: 3262: 3184: 2747: 2437: 2270: 2170: 1973: 1901: 1828: 1764: 1759: 1429: 1328: 1324: 1068: 1015: 952: 931: 926: 844: 839: 683: 632: 479: 467: 369: 285: 144: 67: 17206: 11433:"A new darwinopteran pterosaur reveals arborealism and an opposed thumb" 10524: 9014: 6962:"Pterosaur integumentary structures with complex feather-like branching" 5389: 3705:

had more robust jaws and teeth (which were ziphodont, dagger-shaped, in

3615:

Interpretations of the habits of basal groups have changed profoundly.

3305:

In 1985, the Smithsonian Institution commissioned aeronautical engineer

1116:

was present, covering the entire belly. To the front, a long point, the

1063:

The tails of pterosaurs were always rather slender. This means that the

17373: 17316: 17286: 17281: 16978: 16933: 16861: 16736: 16728: 16563: 16542: 16487: 16355: 16330: 16280: 16245: 16191: 16079: 16071: 16064: 15816: 15739: 15731: 15630: 15594: 15432: 15333: 15312: 15301: 15238: 14855: 14648: 14641: 14076: 13967: 12974: 12631: 12286: 12256: 12239: 12174: 12167: 12082: 12012: 12002: 11923: 11894: 11843: 11653: 11628: 10921: 10213: 9844: 9722: 9638: 8903: 8649: 8420: 8365: 7936: 7810: 7300:"Pterosaur melanosomes support signalling functions for early feathers" 7218: 4817: 4389:

Pterosaurs were used in fiction in Sir Arthur Conan Doyle's 1912 novel

4344: 4312: 4251: 4219: 4177: 4088: 4056: 3890: 3761: 3589: 3572: 3524: 3447:, might have walked or even run bipedally, in addition to flying, like 3382: 3278: 2472: 2230: 2215: 2188: 2104: 2083: 2050: 2041: 1990: 1746: 1731: 1480: 1407: 1400: 1375: 917: 813: 804: 789: 677: 604: 550: 539: 459:

stretching from the ankles to a dramatically lengthened fourth finger.

448: 444: 112: 77: 17686: 11198:"July: Pterosaurs parents | News and features | University of Bristol" 11082:

Naish, Darren; Witton, Mark P.; Martin-Silverstone, Elizabeth (2021).

10589: 9151:

Anatomy, Phylogeny and Palaeobiology of Early Archosaurs and Their Kin

8134:

Philosophical Transactions of the Royal Society B: Biological Sciences

6546: 5646:

Frey E, Martill DM (1998). "Soft tissue preservation in a specimen of

3893:

based on them being relatively weak but fast biters, and suggest that

3501:, but they too appear to have been generally efficient on the ground. 2035: 1895:

The situation for dinosaurs was comparable. From the 1960s onwards, a

1096:. It was probably covered by thick muscle layers. The upper bone, the 538:, which reached wingspans of at least nine metres. The combination of 17450: 17038: 16480: 16428: 16150: 16143: 16111: 16049: 15775: 15768: 15760: 15680: 15458: 15424: 15374: 15367: 15104: 15042: 14946: 14534: 13933: 13650: 13277: 13117: 12952: 12829: 12640: 12450: 12075: 11637: 11418:

From Abba to Zoom A Pop Culture Encyclopedia of the Late 20th Century

11394: 11363: 10497: 9543: 9526: 8878: 8325: 7774:

Broili, F., 1927, "Ein Ramphorhynchus mit Spuren von Haarbedeckung",

7656: 7503:

Denkschriften der königlichen bayerischen Akademie der Wissenschaften

7165: 6231: 6188: 5302: 4471: 4383: 4241: 4214: 4164: 4149: 4062: 4031: 3975: 3970: 3952: 3912: 3851: 3660: 3626: 3609: 3448: 3430: 3387: 3346: 2871: 2774: 2600: 2480: 2456: 2417: 2364: 2319: 2218: 2125:

and named the group Ornithodira to encompass pterosaurs and dinosaurs

1762:

found in England the first pterosaur genus outside Germany, named as

1696:

first suggested that it represented a flying creature in a letter to

1523: 1494: 1383: 1288: 1113: 1003: 833: 821: 744: 713: 624: 507: 471: 199: 117: 61: 17605: 10825: 10800: 10515: 10445: 10364: 10339: 8988: 5419: 1522:

The presence of pycnofibers strongly indicates that pterosaurs were

1493:, but had been widely doubted. Since the 1990s, pterosaur finds and 1287:, are much longer than the humerus. They were probably incapable of 1197:, as depicted here, evidences the possibility that pterosaurs had a 17628: 17482: 17383: 17291: 17110: 16994: 16986: 16760: 16744: 16675: 16252: 15602: 15507: 15439: 15111: 15066: 14839: 14068: 13728: 12625: 12296: 12194: 12033: 11622: 8455: 8006:"The shape of pterosaur evolution: evidence from the fossil record" 7489:

Analyse de la Nature ou tableau de l'univers et des corps organisés

7023: 6549:"High lift function of the pteroid bone and forewing of pterosaurs" 6388:"High lift function of the pteroid bone and forewing of pterosaurs" 5271:(2nd ed.). Berkeley: University of California Press. pp. 4140: 4035: 3966: 3873:

A 2021 study reconstructed the adductor musculature of skulls from

3817: 3514: 2931: 2826: 2629: 2516: 2441: 2226: 2179: 2079: 2018:

claims that it is the largest of its kind ever discovered from the

2011: 1994: 1977: 1563: 1546: 1218: 1177: 1101: 1043: 974: 616: 592: 515: 455:. Their wings were formed by a membrane of skin, muscle, and other 440: 436: 219: 107: 102: 87: 82: 72: 11141:"Allometric wing growth links parental care to pterosaur giantism" 10499: 8967: 8316: 8299: 7845:

Hankin E.H. & Watson D.S.M.; "On the Flight of Pterodactyls",

4315:, created by Mark Witton for the Royal Society's 350th anniversary 1928:, and in 1991 the second ever popular science pterosaur book, the 17410: 11549: 10611: 9619: 9525:

Geist, N.; Hillenius, W.; Frey, E.; Jones, T.; Elgin, R. (2014).

8451: 8449: 7610:

Ornithosauria – an elementary study of the bones of Pterodactyles

4952:

Colbert, Edwin H. (Edwin Harris); Knight, Charles Robert (1951).

4125: 4081: 3779: 3769: 3765: 3492:

were quadrupeds, and some rather efficient terrestrial predators.

3441:

suggested that smaller pterosaurs with longer hindlimbs, such as

3286: 2307: 2242: 2062: 1956:

This development accelerated through the exploitation of two new

1657: 1448: 1379: 1275: 1011: 870: 817: 778: 620: 495: 122: 97: 17673: 10555:

Codorniú, Laura; Chiappe, Luis M.; Cid, Fabricio D. (May 2013).

9328: 5326: 3760:), while boreopterids were freshwater diving animals similar to 3478: 3285:

Another issue that has been difficult to understand is how they

2734: 2150: 2118:

found that pterosaurs were avemetatarsalians closely related to

951:, the true extent of these crests has only been uncovered using 561:, particularly in fiction and journalism. However, technically, 447:(228 to 66 million years ago). Pterosaurs are the earliest 17368: 15701: 15468: 15050: 13658: 13525: 12619: 11616: 11552:, multi-authored website about all aspects of pterosaur science 10396:"Prehistoric sharks feasted on flying reptiles, fossil reveals" 9673:"Pterosaur Stance and Gait and the Interpretation of Trackways" 8938: 5330:

Nature's Flyers: Birds, Insects, and the Biomechanics of Flight

4536: 4066: 3899:

was specialised in consuming relatively large prey compared to

3866: 3855: 3730:

Among pterodactyloids, a greater variation in diet is present.

3710: 3622: 3585: 3483: 3412: 2624: 2573: 2233:

similarities with pterosaurs and reconstructions of lagerpetid

1950: 1907: 1636: 1574: 1512: 1499: 1472: 1366: 1245: 1228: 1193: 1137: 1093: 1039: 600: 390: 209: 168: 12570: 11555: 9170:

de Queiroz, K.; Cantino, P. D.; Gauthier, J. A., eds. (2020).

8564:

Baron, Matthew G. (October 2021). "The origin of Pterosaurs".

8446: 4659: 4638: 4171:

to attain maximum size. In contrast, the more advanced, large

3850:

are now understood to be terrestrial predators akin to ground

3584:

may display adaptations for wing-propelled diving like modern

3429:

There was considerable debate whether pterosaurs ambulated as

3272: 2370:

At least some non-pterodactyloid pterosaurs survived into the

2074:. In the 1980s, early cladistic analyses found that they were 14799: 13395: 11338:"The One Born of Fire: a pterosaurological analysis of Rodan" 11081: 8083:"'Superbly preserved' pterosaur fossil unearthed in Scotland" 6439:"Articulation and Function of the Pteroid Bone of Pterosaurs" 4421: 4416: 3921:

proposed to share this feeding habit based on high estimated

3529: 3463: 3434: 3423: 3358: 2964: 2401: 2355:

Small-sized pterosaur species apparently were present in the

2328: 2234: 2222: 1554: 889:

used combs of numerous needle-like teeth for filter feeding;

608: 425: 11359:"A Monster-Sized Breakdown of Every Insane 'Godzilla' Movie" 10714: 9770:"Pterosaur tracks and the terrestrial ability of pterosaurs" 9335:. Boulder, Colo: Geological Society of America. p. 60. 9217: 7985: 7776:

Sitzungsberichte der Bayerischen Akademie der Wissenschaften

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Claessens LP, O'Connor PM, Unwin DM (2009). Sereno P (ed.).

6014: 5443:

Andres, Brian; Clark, James M.; Xing, Xu (29 January 2010).

5360: 4647: 3567:

must have foraged while swimming, as they seem incapable of

1226:; the extent of this membrane is not certain, as studies on 1100:, was a straight bar. It was connected to a lower bone, the 514:

had efficient unidirectional "flow-through" breathing using

17564: 17432: 17245: 12467: 10801:"Palaeontology: pterosaur embryo from the Early Cretaceous" 9819:

Bennett, S. C. (2007). "A second specimen of the pterosaur

6760: 6758: 6756: 6754: 6752: 6750: 6737: 6735: 6733: 6731: 6729: 6727: 6637: 6627: 6625: 6623: 6621: 6481:

Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen

6345: 6343: 6341: 6252: 5894: 5892: 5890: 5888: 5886: 5873: 5871: 5793: 5791: 5789: 5753:"Taxonomy and systematics of the Late Cretaceous pterosaur 5652:

Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen

5592: 5590: 5046: 4662: 4641: 4379: 4070: 3467: 2303: 1836:

published studies about the wing membranes of specimens of

1460: 1280: 1201:

patagium – a membrane connecting the legs that, unlike the

1188: 1132:

Various configurations proposed for the wings of pterosaurs

596: 503: 10757: 9963: 6070: 5089:

Wang, X.; Kellner, A.W.A.; Zhou, Z.; Campos, D.A. (2008).

4955:

The dinosaur book: the ruling reptiles and their relatives

4908:(2003) , Peter Roach; James Hartmann; Jane Setter (eds.), 3995:

has been interpreted as being attacked or scavenged by an

2346:, several tapejarids and the indeterminate non-azhdarchid 1566:, and the presence of both aktinofibrils and filaments on 17253: 17237: 10337: 10055: 10006: 9169: 9067: 9052:

XXIII Jornadas Argentinas de Paleontología de Vertebrados

8836:

Andres, B.; Myers, T. S. (2013). "Lone Star Pterosaurs".

8105: 7872:

Padian, K (1979). "The wings of pterosaurs: A new look".

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Dragons of the Air: An account of extinct flying reptiles

7590: 7578: 7566: 7512: 6458:

10.1671/0272-4634(2007)27[881:AAFOTP]2.0.CO;2

4752:"The Earliest Pterodactyloid and the Origin of the Group" 3727:, in particular, seems to have been a beetle specialist. 2463:, a "primitive" group of long-tailed pterosaurs, and the 2360: 1263: 1202: 1141: 777:

Standing, such giants could reach the height of a modern

580: 490:

Pterosaurs sported coats of hair-like filaments known as

27:

Flying reptiles of the extinct clade or order Pterosauria

11272: 11270: 11268: 9721:

Hwang K, Huh M, Lockley MG, Unwin DM, Wright JL (2002).

9045: 8683:

Walking with dinosaurs (episode 4 ) – Giant Of The Skies

8610:

Rupert Wild, 1983, "Über die Ursprung der Flugsaurier",

8065: 8063: 7363: 6747: 6724: 6618: 6338: 5883: 5868: 5786: 5732: 5691: 5689: 5687: 5685: 5670: 5587: 5201: 11145:

Proceedings of the Royal Society B: Biological Sciences

11039:

Proceedings of the Royal Society B: Biological Sciences

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Nesbitt, S.J., Desojo, J.B., & Irmis, R.B. (2013).

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Proceedings of the Royal Society B: Biological Sciences

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Geggel 2018-12-17T19:23:17Z, Laura (17 December 2018).

3965:

discusses an Early Cretaceous fossil of three cervical

2065:

has been so heavily modified for flight, and immediate

1891:

by John Conway exemplifies the "new look" of pterosaurs

1002:

of pterosaurs numbered between thirty-four and seventy

14165: 10945: 10260: 10067: 9945: 9883: 9871: 9859: 9274:"Pterodactyls were too heavy to fly, scientist claims" 8262: 7635:

Mivart, G (1881). "A popular account of chamaeleons".

7453:

Journal de Physique, de Chimie et d'Histoire Naturelle

7408: 7406: 7404: 7402: 7353: 7351: 6655: 5750: 5492: 4962: 4470:

In most media appearances, pterosaurs are depicted as

1972:

Even more productive was the Early Cretaceous Chinese

663:

Pterodactyloids include the clades Ornithocheiroidea (

630:

Basal pterosaurs include the clades Dimorphodontidae (

11315: 11265: 11220: 10548: 10468: 9480:"With Wings Flapping, Model Pterodactyl Takes to Air" 9428:"Did giant pterosaurs vault aloft like vampire bats?" 8879:

Agnolin, Federico L. & Varricchio, David (2012).

8784: 8695:

Slack KE, Jones CM, Ando T, et al. (June 2006).

8633:"Were early pterosaurs inept terrestrial locomotors?" 8592: 8093: 8060: 8051: 8049: 7894:, Ph.D. diss., Department of Biology, Yale University 7387: 6505: 5682: 5088: 4671: 4644: 3983:

have been found with tooth marks from sharks such as

3276:

Skeletal reconstruction of a quadrupedally launching

2278:

is anatomically similar to that of early pterosaurs.

2169:

Two researchers, S. Christopher Bennett in 1996, and

1969:, intercepted the trade and named even more species. 965:

show that even some early pterosaurs possessed them.

623:. Likewise, the sacral vertebrae could form a single 12424: 12318: 12303: 12292: 12282: 12272: 12262: 12252: 12208: 12044: 12029: 12018: 12008: 11998: 11873: 11802: 11787: 11776: 10491: 10419:"Palaeontology: pterosaur egg with a leathery shell" 10417:

Ji Q, Ji SA, Cheng YN, et al. (December 2004).

9720: 9063: 9061: 8780: 8778: 8776: 8253: 7951: 6913: 6901: 6889: 6877: 6865: 6853: 6787: 6770: 6591: 6367: 6355: 6153: 6138: 6123: 6066: 6064: 5945: 5933: 5916: 5904: 5851: 5834: 5815: 5803: 5771: 5636: 5619: 5602: 5572: 5517: 5505: 5361:

Witmer LM, Chatterjee S, Franzosa J, Rowe T (2003).

5356: 5354: 5352: 5350: 4970:"Pterosaur distribution in time and space: an atlas" 4656: 4635: 4504: 4467:(1988), is a notable example from an animated film. 4431:, is portrayed as an enormous irradiated species of 2475:, it has fallen out of favor among most scientists. 2192:. A 2011 archosaur-focused phylogenetic analysis by 366: 358: 347: 336: 328: 316: 306: 298: 290: 282: 11562:"Comments on the phylogeny of the pterodactyloidea" 10086:Wu, Wen-Hao; Zhou, Chang-Fu; Andres, Brian (2017). 9421: 9419: 9332:

Posture, locomotion, and paleoecology of pterosaurs

8829: 8004:Dyke, G.J. McGowan; Nudds, R.L.; Smith, D. (2009). 7798:

Zeitschrift für Anatomie und Entwicklungsgeschichte

7399: 7375: 7348: 6547:Wilkinson M.T.; Unwin D.M.; Ellington C.P. (2006). 6167:and the nature of the pterosaur flight apparatus". 4653: 4632: 3255:

Posture, Locomotion, and Paleoecology of Pterosaurs

3247:

Posture, Locomotion, and Paleoecology of Pterosaurs

1926:

Handbuch der Paläoherptologie, Teil 19: Pterosauria

1673:used its wings as flippers and was affiliated with 13213: 11033:Unwin, David Michael; Deeming, D. Charles (2019). 11010:"First 3D pterosaur eggs found with their parents" 10557:"First occurrence of stomach stones in pterosaurs" 10554: 9392: 9390: 9388: 8785:Longrich, N.R.; Martill, D.M.; Andres, B. (2018). 8304:Bulletin of the American Museum of Natural History 8046: 7737:Marsh, O.C. (1882). "The wings of Pterodactyles". 6010: 6008: 6006: 3664:were likely terrestrial/semiarboreal generalists. 1336:. The wingfinger is also bent somewhat downwards. 14689: 13634: 11138: 10338:Buffetaut E, Martill D, Escuillié F (July 2004). 10310: 9058: 8773: 6540: 6061: 5420:"Pterosaur.net :: Origins and Relationships" 5347: 5202:Buffetaut E, Grigorescu D, Csiki Z (April 2002). 5195: 4266:may have had similar activity patterns to modern 4061:likely pterosaurs buried their eggs, like modern 3858:, eating any prey item they could swallow whole. 3794:obtained food in coastal or freshwater habitats. 3764:, and pteranodonts pelagic plunge-divers akin to 3754:(with the exception of the plunge-diving adapted 3629:, and large insects. Its robust dentition caused 2269:At least one study found that the early Triassic 1850:discovered hair follicles in pterosaur skin, and 526:to the largest known flying creatures, including 17727: 14783: 11872: 10475:Codorniú, L.; Chiappe, L.; Rivarola, D. (2014). 10267:Martill, David M.; Ibrahim, Nizar (March 2015). 10155: 9416: 9143: 9109:Geological Society, London, Special Publications 6255:Geological Society, London, Special Publications 5327:Alexander, David E. & Vogel, Steven (2004). 4008: 3528:had slightly larger feet (47% the length of the 2378:situation for late Cretaceous pterosaur faunas. 2302:It was once thought that competition with early 15992: 15912: 14825: 11526:Pterosaurs: Natural History, Evolution, Anatomy 10941: 10939: 10883: 10881: 10879: 9901: 9385: 7980:Handbuch der Paläoherpetologie XIX. Pterosauria 7426:Acta Theodoro-Palatinae Mannheim 5 Pars Physica 6662:Proceedings of the National Academy of Sciences 6295: 6209: 6003: 5442: 5095:Proceedings of the National Academy of Sciences 5084: 5082: 4886:and its implications for pterosaur phylogeny", 4852:Pterosaurs: Natural History, Evolution, Anatomy 2385: 1704:, "wing-finger". This was in 1815 Latinised to 895:could have over a thousand bristle-like teeth. 12673: 11757: 11597: 11430: 11332: 11330: 10888:Bennett S. C. (1995). "A statistical study of 10410: 10266: 9802: 9800: 9798: 9445: 8694: 8626: 8624: 8622: 8620: 8612:Weltenberger Akademie, Erwin Rutte-Festschrift 8003: 7902: 7900: 7143: 6162: 5721:. The Dinosaur Museum: Blanding, Utah, 15–41. 5650:(Wagner) from the Upper Jurassic of Germany". 5024:"Pterosaur.net :: Terrestrial Locomotion" 4881: 4849: 4745: 4743: 4725:expansion of ecological niches in the Mesozoic 4148:showed possible evidence of their young being 3253:period. However, both Sato and the authors of 583:, some fossils of which have been discovered. 571:, and more broadly to members of the suborder 17222: 12586: 11583: 11424: 10887: 9355: 8400:"Reassessment of the Triassic archosauriform 8117: 7505:, München: mathematisch-physikalische Classe 6801:Witton MP, Naish D (2008). McClain CR (ed.). 6386:Wilkinson MT, Unwin DM, Ellington CP (2006). 6298:"Limb disparity and wing shape in pterosaurs" 4992:"The Extent of the Pterosaur Flight Membrane" 4989: 4951: 4749: 4575:List of pterosaur-bearing stratigraphic units 4386:dinosaurs and not descendants of pterosaurs. 3734:contained many piscivorous taxa, such as the 3536:had very large feet (69% of tibial length in 1059:The shoulder girdle connected to the notarium 16577: 12207: 11032: 10936: 10876: 10085: 9957: 9100: 8939:Prondvai, E.; Bodor, E. R.; Ösi, A. (2014). 7043: 6499: 5719:Feathered Dinosaurs and the Origin of Flight 5695: 5322: 5320: 5079: 3532:), while filter-feeding pterosaurs like the 3462:), in a manner similar to many mammals like 142:Six pterosaurs (top left to bottom right): 15691: 11327: 10605: 10194: 10188: 10161: 9902:Andres, B.; Clark, J. M.; Xing, X. (2010). 9795: 9573: 9398:"Why pterosaurs weren't so scary after all" 9329:Templin, R. J.; Chatterjee, Sankar (2004). 9211: 9190: 8835: 8617: 8130:and the origin of dinosaurs and pterosaurs" 7897: 7418: 6800: 5645: 5071:: CS1 maint: numeric names: authors list ( 4740: 4700: 4694: 3951:Pterosaurs are known to have been eaten by 3331: 2440:more closely related to pterosaurs than to 2025: 1939:The three-dimensionally preserved skull of 784: 17229: 17215: 12749: 12593: 12579: 11590: 11576: 11504: 11420:. Andrews MacMeel Publishing. p. 272. 9761: 9176:. CRC Press Boca Raton, FL. p. 2072. 7991: 7724: 7712: 7697: 7680: 7596: 7584: 7572: 7518: 7444: 7431: 7369: 6953: 6764: 6741: 6643: 6631: 6478: 6472: 6436: 6349: 6296:Dyke GJ, Nudds RL, Rayner JM (July 2006). 5966: 5898: 5877: 5797: 5738: 5676: 5596: 5486: 5141: 4435:. Rodan has appeared in multiple Japanese 4191: 3969:of a pterosaur with the broken tooth of a 3604:Early-on it was recognised that the small 3595: 729:The position of the clade Anurognathidae ( 595:were hollow and air-filled, like those of 380: 133: 17585:Tradeoffs for locomotion in air and water 17236: 11652: 11458: 11295:Evolution and Palaeobiology of Pterosaurs 11172: 11115: 11058: 10985: 10975: 10824: 10775: 10691: 10588: 10523: 10416: 10363: 10313:Zoological Journal of the Linnean Society 10306: 10304: 10302: 10222: 10212: 10179: 10125: 10115: 9647: 9637: 9542: 9451: 9022: 8812: 8802: 8712: 8658: 8648: 8429: 8419: 8374: 8364: 8315: 8186: 8175:Zoological Journal of the Linnean Society 8153: 8021: 7331: 7266: 7217: 7120: 6987:1983/1f7893a1-924d-4cb3-a4bf-c4b1592356e9 6985: 6836: 6826: 6699: 6681: 6574: 6523: 6413: 6313: 6106: 6096: 6044: 5744: 5317: 5300: 5253: 5147: 5124: 5114: 5007: 4983: 4912:, Cambridge: Cambridge University Press, 4826: 4816: 4775: 3989:, and a fossil with tooth marks from the 3870:may have been a specialist molluscivore. 3324:Large-headed species are thought to have 1960:. During the 1970s, the Early Cretaceous 1545:A 2018 study of the remains of two small 1529: 11564:, by Alexander W. A. Kellner (technical) 11356: 10798: 10764:Anais da Academia Brasileira de Ciências 10500:"Argentinian unhatched pterosaur fossil" 10340:"Pterosaurs as part of a spinosaur diet" 9767: 9670: 9576:Annual Review of Ecology and Systematics 9477: 9267: 9265: 7906: 7450: 7297: 6940: 6512:Anais da Academia Brasileira de Ciências 5549: 5547: 5545: 5543: 5541: 5539: 5537: 5296: 5294: 5292: 4425:) which first appeared in the 1956 film 4322: 4303: 4297: 4012: 3477: 3402: 3271: 3228: 3220: 2285: 2149: 2130: 2034: 1934: 1880: 1876: 1745: 1683: 1616: 1511: 1503:(which translates as "hairy demon") and 1447: 1356: 1187: 1147: 1127: 1054: 1020: 984: 901: 843:Reconstruction of crests: three crested 838: 788: 764: 474:of small vertebrates. Later pterosaurs ( 11415: 11286: 9966:Acta Geologica Sinica - English Edition 9818: 9271: 9173:Phylonyms: A Companion to the PhyloCode 9106: 8404:: neither runner nor biped, but hopper" 8397: 8346: 8297: 8168: 7787: 7094: 5713: 5711: 3451:. However, a large number of pterosaur 3309:to build a half-scale working model of 1601: 1509:show pycnofibers on the head and body. 1343:A laser-simulated fluorescence scan on 1108:The breastbone, formed by fused paired 14: 17728: 13622: 11523: 11299:Geological Society Special Publication 10854: 10852: 10661: 10659: 10299: 10073: 10061: 9951: 9895: 9889: 9877: 9865: 9806: 9600: 9373:from the original on February 21, 2009 9302: 8630: 8598: 8291: 8123: 8111: 8099: 8069: 8055: 7966: 7871: 7755: 7634: 7530: 7412: 7393: 7381: 7357: 7248: 7019:"Fur flies over new pterosaur fossils" 7016: 6928: 6907: 6895: 6883: 6871: 6859: 6781: 6612: 6373: 6361: 6147: 6132: 5954: 5939: 5927: 5910: 5862: 5845: 5828: 5809: 5780: 5630: 5613: 5581: 5528: 5511: 5303:"Pterosaurs: Myths and Misconceptions" 5259: 4750:Andres, B.; Clark, J.; Xu, X. (2014). 3925:(BFQ) and absolute bite force values. 2146:theorized to be related to pterosaurs. 1788:, and in 1901 the first popular book, 1483:was first reported from a specimen of 17610: 17609: 17210: 15979: 14771: 14770: 14152: 13621: 12736: 12612: 12574: 11609: 11571: 11485: 11384: 11321: 9425: 9349: 9262: 8563: 7862:, Londen: Penguin Books, 1988, p. 283 7736: 7472:Annales du Musée d'Histoire Naturelle 5553: 5534: 5289: 4958:. New York: McGraw-Hill. p. 153. 4904: 4798: 4021:juvenile from the Solnhofen Limestone 2432:(Sömmerring 1812)". A broader clade, 2313:Cretaceous–Paleogene extinction event 1741: 1067:retractor muscle which in most basal 15980: 10665: 8347:Ezcurra, Martín D. (28 April 2016). 7549: 5708: 4605: 3946: 3839:were likely terrestrial carnivores. 3827:mostly were terrestrial pterosaurs. 3486:trackways show that pterosaurs like 3361:cavities revealed that the animals ( 1183: 885:had recurved teeth for eating meat. 11357:Gonzales, Dave (October 12, 2016). 11195: 10849: 10656: 9601:Anthes, Emily (November 18, 2013). 9588:10.1146/annurev.es.08.110177.002241 9478:Molotsky, Irvin (28 January 1986). 9272:Alleyne, Richard (1 October 2008). 9046:Haluza, A.; Apesteguía, S. (2007). 8541:"Pterosaur Origins Flap into Focus" 8265:Journal of Systematic Palaeontology 5558:. New York: Pi Press. p. 246. 2306:species might have resulted in the 2186:or a branch between the latter and 1915:named a renaissance of pterosaurs. 1236:). A common interpretation is that 980: 836:was relatively large for reptiles. 627:while the pelvic bones fused also. 607:for flight muscles and an enlarged 565:may refer to members of the genus 502:to the down feathers found on both 24: 10561:Journal of Vertebrate Paleontology 9911:Journal of Vertebrate Paleontology 9789:10.1111/j.1502-3931.1996.tb01673.x 9603:"Coldblooded Does Not Mean Stupid" 8188:10.1111/j.1096-3642.1996.tb01267.x 6554:Proceedings of the Royal Society B 6446:Journal of Vertebrate Paleontology 6393:Proceedings of the Royal Society B 6025:Proceedings of the Royal Society B 5452:Journal of Vertebrate Paleontology 4990:Elgin RA, Hone DW, Frey E (2011). 3790:were likely primarily scavengers. 3398: 1158:, preserve the membrane structure 1083: 435:. They existed during most of the 25: 17762: 11543: 11385:Sharf, Zack (December 10, 2018). 10402:. October 3, 2018. Archived from 9305:"Were pterosaurs too big to fly?" 8538: 3352: 2392:List of pterosaur classifications 2296:Arizona Museum of Natural History 17325: 17191: 17185: 17178: 15961: 15953: 15945: 14753: 14747: 14135: 14130: 13604: 13597: 13589: 12555: 12554: 12526: 12519: 12513: 12508: 12502: 11409: 11378: 11350: 11278:"Pterosaurs In Popular Culture." 11214: 11189: 11132: 11075: 11026: 11002: 10792: 10751: 10388: 10331: 10239: 10195:Naish, D.; Witton, M.P. (2017). 10162:Witton, M.P.; Naish, D. (2015). 10142: 10079: 10000: 9986:10.1111/j.1755-6724.2011.00444.x 9812: 9714: 9664: 9613: 9594: 9567: 9518: 9490: 9471: 9426:Hecht, Jeff (16 November 2010). 9322: 9303:Powell, Devin (2 October 2008). 9296: 9163: 9153:. Geological Society of London. 9039: 8982: 8932: 8881:"Systematic reinterpretation of 8872: 8729: 8688: 8675: 8604: 8557: 8532: 8508: 8391: 8340: 8195: 8169:Bennett, S. Christopher (1996). 8162: 8075: 8023:10.1111/j.1420-9101.2008.01682.x 7997: 7972: 7884: 7865: 7852: 7839: 7781: 7768: 7749: 7730: 7628: 7615: 7602: 7543: 7524: 7494: 7481: 7439:Natürliches System der Amphibien 7249:D’Alba, Liliana (January 2019). 6315:10.1111/j.1420-9101.2006.01096.x 6163:Unwin DM, Bakhurina NN (1994). " 5757:(Pterosauria, Pterodactyloidea)" 4628: 4570:Graphical timeline of pterosaurs 4549: 4535: 4521: 4507: 4419:, a fictional giant monster (or 3955:. In the 1 July 2004 edition of 3013: 2993: 2930: 2910: 2825: 2805: 2773: 2733: 2701: 2628: 2599: 2572: 2545: 2515: 1471:(sharing a common origin) with, 1319:In derived pterodactyloids like 1258:, and a pterodactyloid from the 491: 186: 65: 17741:Late Triassic first appearances 14153: 12737: 12600: 9454:"The Great Pterodactyl Project" 8701:Molecular Biology and Evolution 8586:10.1016/j.earscirev.2021.103777 8010:Journal of Evolutionary Biology 7463: 7291: 7242: 7180: 7137: 7088: 7037: 7010: 6943:Nova Acta Academiae Leopoldinae 6934: 6649: 6525:10.1590/s0001-37652010000400024 6430: 6289: 6246: 6203: 5960: 5751:S. Christopher Bennett (1994). 5436: 5412: 5040: 5016: 4717: 4684: 3437:. In the 1980s, paleontologist 3216: 2006:in 2022, was discovered on the 1730:thought pterosaurs were flying 1423: 11528:. Princeton University Press. 7982:, Urban & Fischer, München 7298:Cincotta; et al. (2022). 7255:Nature Ecology & Evolution 7190:Nature Ecology & Evolution 7046:Nature Ecology & Evolution 6966:Nature Ecology & Evolution 5556:The Pterosaurs: From Deep Time 4945: 4938:Merriam-Webster.com Dictionary 4925: 4910:English Pronouncing Dictionary 4898: 4875: 4854:, Princeton University Press, 4843: 4792: 4621: 4612: 4600:Timeline of pterosaur research 4445:Godzilla: King of the Monsters 4330:When Dinosaurs Ruled the Earth 4026:and in Loma del Pterodaustro ( 2483:(family tree) below follows a 2326:, such as the ornithocheirids 1612: 1608:Timeline of pterosaur research 1443: 1208:patagium, leaves the tail free 816:, with the upper jawbone, the 586: 13: 1: 10777:10.1590/0001-3765201520150364 10293:10.1016/j.cretres.2014.11.001 10168:Acta Palaeontologica Polonica 10033:10.1016/j.cretres.2020.104637 9198:The Encyclopedia of Dinosaurs 9129:10.1144/gsl.sp.2003.217.01.10 9086:10.1016/j.cretres.2021.105061 7790:"Das Gehirn der Pterosaurier" 7095:Kellner; et al. (2009). 6275:10.1144/GSL.SP.2003.217.01.14 4996:Acta Palaeontologica Polonica 4733: 4009:Reproduction and life history 3391:show that brain evolution in 3233:Pterosaur flight adaptations. 2281: 2057:similarities with pterosaurs. 1738:coined the term Pterosauria. 1723:Samuel Thomas von Soemmerring 1688:Newman's marsupial pterosaurs 1308:Darwinopterus linglongtaensis 1152:Some specimens, such as this 793:Conical tooth, possibly from 10977:10.1371/journal.pone.0031392 10799:Wang X, Zhou Z (June 2004). 10581:10.1080/02724634.2013.731335 10117:10.1371/journal.pone.0185486 9825:Paläontologische Zeitschrift 8804:10.1371/journal.pbio.2001663 7612:, Cambridge University Press 7017:Briggs, Helen (2018-12-17). 6828:10.1371/journal.pone.0002271 6098:10.1371/journal.pone.0004497 5173:10.1126/science.187.4180.947 3372:) had massive flocculi. The 3357:An X-ray study of pterosaur 3328:in order to better balance. 2386:Classification and phylogeny 1867:David Meredith Seares Watson 1393: 36:Pterodactyl (disambiguation) 7: 11490:. McFarland & Company. 7739:American Journal of Science 6493:10.1127/0077-7749/2011/0131 4543:Evolutionary biology portal 4500: 3806:were piscivores, while the 3557: 3326:forwardly swept their wings 2016:National Museum of Scotland 1645:. The pieces from one such 1589:Tupandactylus cf. imperator 1569:Jeholopterus ningchengensis 1542:specimens too fundamental. 1486:Scaphognathus crassirostris 1459:Most or all pterosaurs had 10: 17767: 17477: 17441:Flying and gliding animals 17277:Fin and flipper locomotion 11599:Avemetatarsalia / Pan-Aves 11505:Wellnhofer, Peter (1991). 11479: 11283:, Accessed 27 August 2010. 11108:10.1038/s41598-021-92499-z 10325:10.1093/zoolinnean/zlaa163 8995:Royal Society Open Science 8759:10.1666/0094-8373-35.3.432 7849:, October 1914, pp. 324–35 7324:10.1038/s41586-022-04622-3 7210:10.1038/s41559-020-01309-8 7066:10.1038/s41559-020-01308-9 5664:10.1127/njgpa/210/1998/421 5333:. JHU Press. p. 191. 4799:Baron, Matthew G. (2020). 4565:Flying and gliding animals 2436:, has been defined as all 2389: 2290:Reconstructed skeleton of 2030: 1930:Encyclopedia of Pterosaurs 1621:Engraving of the original 1605: 1506:Jeholopterus ninchengensis 869:In some cases, fossilized 758: 648:), and Rhamphorhynchidae ( 428:of flying reptiles in the 389:Distribution of pterosaur 29: 17746:Maastrichtian extinctions 17618: 17552: 17491: 17431: 17334: 17323: 17252: 17175: 17077: 17027: 16959: 16950: 16922: 16898: 16887: 16843: 16808: 16798: 16694: 16632: 16623: 16619: 16608: 16573: 16532: 16497: 16461: 16452: 16418: 16381: 16372: 16345: 16341: 16318: 16262: 16233: 16229: 16218: 16160: 16131: 16122: 16099: 16090: 16031: 16022: 16007: 16003: 15988: 15975: 15942: 15908: 15792: 15750: 15660: 15651: 15613: 15551: 15542: 15527: 15518: 15457: 15419: 15408: 15357: 15320: 15311: 15264: 15255: 15183: 15168: 15159: 15131: 15122: 15031: 15008: 14997: 14986: 14934: 14923: 14888: 14834: 14821: 14810: 14794: 14779: 14766: 14744: 14714: 14698: 14685: 14624: 14609: 14600: 14551: 14523: 14430: 14419: 14375: 14352: 14343: 14291: 14287: 14276: 14254: 14243: 14195: 14180: 14176: 14161: 14148: 14128: 14086: 14013: 14004: 13950: 13900: 13885: 13876: 13865: 13842: 13831: 13769: 13754: 13745: 13723: 13714: 13684: 13680: 13669: 13645: 13630: 13617: 13586: 13536: 13486: 13476: 13428: 13378: 13369: 13312: 13303: 13264: 13255: 13244: 13209: 13198: 13182: 13170: 13107: 13092: 13088: 13077: 13061: 13049: 13009: 12993: 12984: 12942: 12933: 12901: 12890: 12847: 12816: 12806: 12779: 12764: 12760: 12745: 12732: 12700: 12682: 12669: 12608: 12539: 12500: 12449: 12423: 12414: 12369: 12329: 12317: 12248: 12237: 12203: 12192: 12115: 12056: 12043: 11994: 11990: 11979: 11948: 11884: 11868: 11857: 11801: 11772: 11768: 11753: 11734: 11706: 11688: 11679: 11661: 11648: 11605: 11460:10.1016/j.cub.2021.03.030 10914:10.1017/S0022336000034946 10737:10.1080/08912960701189800 10693:10.1016/j.cub.2014.04.054 10088:"The toothless pterosaur 9931:10.1080/02724630903409220 9747:10.1017/S0016756802006647 9700:10.1080/10420940390255501 9461:Engineering & Science 9240:10.1007/s00114-008-0397-5 8858:10.1017/S1755691013000303 8486:10.1038/s41586-020-3011-4 8277:10.1017/S1477201907002064 7929:10.1017/S009483730000765X 7268:10.1038/s41559-018-0767-0 6978:10.1038/s41559-018-0728-7 5989:10.1080/10292380009380572 5472:10.1080/02724630903409220 5231:10.1007/s00114-002-0307-1 4777:10.1016/j.cub.2014.03.030 4240:have been inferred to be 4218:have been inferred to be 3896:Tropeognathus mesembrinus 3725:Darwinopterus robustidens 3416:Haenamichnus uhangriensis 3363:Rhamphorhynchus muensteri 3237: 3008: 2988: 2981: 2961: 2954: 2925: 2905: 2898: 2888: 2868: 2861: 2820: 2798: 2791: 2768: 2761: 2751: 2726: 2719: 2696: 2689: 2672: 2665: 2650: 2643: 2621: 2614: 2594: 2587: 2567: 2560: 2540: 2533: 2510: 2503: 1794:St. George Jackson Mivart 1662:Cosimo Alessandro Collini 1586:In 2022, a new fossil of 1352: 1029:was longer than the torso 599:. This provided a higher 404: 397: 388: 379: 279: 274: 183:Scientific classification 181: 141: 132: 45: 11488:The Dinosaur Filmography 11196:Bristol, University of. 10400:Science & Innovation 8685:at 22', Tim Haines, 1999 8631:Witton, Mark P. (2015). 7847:The Aeronautical Journal 7487:Rafinesque, C.S., 1815, 6212:Chinese Science Bulletin 5554:Unwin, David M. (2006). 5495:Acta Geoscientica Sinica 4888:Acta Geoscientica Sinica 4882:David M. Unwin (2010), " 4580:List of pterosaur genera 4411:In the latter, animator 4348:or (non-pterodactyloid) 4250:has been inferred to be 4196:Comparisons between the 4057:Hamipterus tianshanensis 3332:Air sacs and respiration 3312:Quetzalcoatlus northropi 3299:Johns Hopkins University 3295:University of Portsmouth 2026:Evolution and extinction 1180:into the wing membrane. 1123: 785:Skull, teeth, and crests 14168:Archaeopterodactyloidea 11486:Berry, Mark F. (2005). 11416:Mansour, David (2005). 11342:Journal of Geek Studies 11243:10.1126/science.1200043 10894:Journal of Paleontology 10634:10.1126/science.1197323 8231:10.1126/science.1143325 7428:, pp. 58–103 (1 plate). 6683:10.1073/pnas.2107631118 5116:10.1073/pnas.0707728105 4850:Mark P. Witton (2013), 4585:Phylogeny of pterosaurs 4372:-like crest and teeth: 4358:-like crests and long, 4192:Daily activity patterns 4108:flaplings found in the 3792:Archaeopterodactyloidea 3596:Diet and feeding habits 3395:was a modular process. 2758:Archaeopterodactyloidea 2210:which argued that both 2178:between pterosaurs and 1869:, but they interpreted 862:Tupandactylus imperator 830:nasoantorbital fenestra 754: 636:), Campylognathididae ( 17580:Terrestrial locomotion 17524:Evolution of cetaceans 17519:Origin of avian flight 17504:Evolution of tetrapods 11556:The Pterosaur Database 11157:10.1098/rspb.2023.1102 11051:10.1098/rspb.2019.0409 10666:Wang, Xiaolin (2014). 10181:10.4202/app.00005.2013 9452:MacCready, P. (1985). 9402:The Observer newspaper 8398:Bennett, S.C. (2020). 8298:Nesbitt, S.J. (2011). 8146:10.1098/rstb.1999.0489 7978:Wellnhofer, P., 1978, 7858:Bakker, Robert, 1986, 7113:10.1098/rspb.2009.0846 6567:10.1098/rspb.2005.3278 6406:10.1098/rspb.2005.3278 6037:10.1098/rspb.2009.0846 4701: 4695: 4408:One Million Years B.C. 4339: 4333:depicting an outsized 4316: 4294:between these genera. 4022: 3911:was corroborated as a 3820:-like filter-feeding. 3504:The forelimb bones of 3493: 3419: 3282: 3234: 3226: 2299: 2257:for incipient flight. 2166: 2147: 2078:(archosaurs closer to 2058: 1953: 1892: 1755: 1689: 1632: 1624:Pterodactylus antiquus 1530:Relation with feathers 1519: 1456: 1362: 1302:Changchengopterus pani 1209: 1159: 1133: 1060: 1030: 995: 912: 866: 856:Tupandactylus navigans 847:. From top to bottom: 799: 770: 705:), and Azhdarchoidea ( 693:), Dsungaripteroidea ( 681:), Ctenochasmatoidea ( 522:, from the very small 451:known to have evolved 34:. For other uses, see 17708:Paleobiology Database 17595:Undulatory locomotion 17544:Homologous structures 11524:Witton, Mark (2013). 11336:Thomas, H.N. (2020). 10150:Thalassodromeus sethi 9531:The Anatomical Record 9359:(February 18, 2009). 8714:10.1093/molbev/msj124 8566:Earth-Science Reviews 8402:Scleromochlus taylori 8128:Scleromochlus taylori 8124:Benton, M.J. (1999). 8087:Associated Press (AP) 7860:The Dinosaur Heresies 5009:10.4202/app.2009.0145 4326: 4307: 4298:Cultural significance 4139:, similar to that of 4016: 3481: 3406: 3377:of total brain mass. 3275: 3232: 3224: 2398:phylogenetic taxonomy 2390:Further information: 2289: 2153: 2134: 2100:basal archosauriforms 2038: 1938: 1884: 1877:Pterosaur renaissance 1863:Ernest Hanbury Hankin 1801:Othniel Charles Marsh 1749: 1687: 1620: 1549:-age pterosaurs from 1515: 1491:Georg August Goldfuss 1455:preserved pycnofibers 1451: 1378:fused with the broad 1360: 1191: 1151: 1131: 1058: 1024: 988: 905: 842: 792: 768: 17539:Analogous structures 17534:Convergent evolution 13780:Archaeoistiodactylus 13690:Archaeoistiodactylus 11445:(11): 2429–2436.e7. 7621:Seeley, H.G., 1901, 7608:Seeley, H.G., 1870, 7441:Munich, 1830: 1–354. 6017:"The soft tissue of 4464:The Land Before Time 4397:1925 film adaptation 4175:pterosaurs, such as 4091:scans of fossilised 3928:Tapejara wellnhoferi 3923:bite force quotients 3279:Pteranodon longiceps 2176:convergent evolution 2154:Life restoration of 2135:Life restoration of 2098:. A placement among 2039:Life restoration of 1897:dinosaur renaissance 1602:History of discovery 1477:convergent evolution 1475:hair, an example of 850:Tapejara wellnhoferi 547:last common ancestor 17590:Rotating locomotion 17529:Comparative anatomy 16845:Targaryendraconidae 16534:Ornithocheiriformes 16221:Ornithocheiromorpha 15520:Neopterodactyloidea 15447:Dsungaripteromorpha 13215:Campylognathoididae 12935:Eudimorphodontoidea 11558:, by Paul Pursglove 11451:2021CBio...31E2429Z 11235:2011Sci...332..705S 11100:2021NatSR..1113130N 10968:2012PLoSO...731392P 10906:1995JPal...69..569B 10817:2004Natur.429..621W 10729:2007HBio...19..273G 10684:2014CBio...24.1323W 10626:2011Sci...331..321L 10573:2013JVPal..33..647C 10438:2004Natur.432..572J 10406:on October 3, 2018. 10356:2004Natur.429...33B 10285:2015CrRes..53...59M 10273:Cretaceous Research 10108:2017PLoSO..1285486W 10025:2021CrRes.11704637M 10013:Cretaceous Research 9978:2011AcGlS..85..507L 9923:2010JVPal..30..163A 9837:2007PalZ...81..376B 9821:Anurognathus ammoni 9739:2002GeoM..139..421H 9727:Geological Magazine 9692:2003Ichno..10..115P 9232:2008NW.....95..891L 9220:Naturwissenschaften 9121:2003GSLSP.217..105K 9074:Cretaceous Research 9015:10.1098/rsos.160333 9007:2016RSOS....360333M 8960:2014Pbio...40..288P 8850:2012EESTR.103..383A 8751:2009Pbio...35..432B 8578:2021ESRv..22103777B 8545:Scientific American 8478:2020Natur.588..445E 8223:2007Sci...317..358I 7921:1983Pbio....9..218P 7788:Edinger, T (1927). 7649:1881Natur..24..335. 7437:Wagler, J. (1830). 7316:2022Natur.604..684C 7251:"Pterosaur plumage" 7202:2020NatEE...4.1592Y 7158:1994Natur.371...62U 7058:2020NatEE...4.1590U 6819:2008PLoSO...3.2271W 6674:2021PNAS..11807631P 6668:(44): e2107631118. 6437:Bennett SC (2007). 6267:2003GSLSP.217..233F 6224:2002ChSBu..47..226W 6181:1994Natur.371...62U 6089:2009PLoSO...4.4497C 5981:2000HBio...14..255B 5648:Pterodactylus kochi 5464:2010JVPal..30..163A 5390:10.1038/nature02048 5382:2003Natur.425..950W 5223:2002NW.....89..180B 5211:Naturwissenschaften 5165:1975Sci...187..947L 5107:2008PNAS..105.1983W 4860:2013pnhe.book.....W 4768:2014CBio...24.1011A 4557:Paleontology portal 4382:, though birds are 4110:Solnhofen limestone 4040:Lagarcito Formation 4028:Lagarcito Formation 3940:Tupuxuara leonardii 3934:Caupedactylus ybaka 3746:and Nyctosauridae. 3713:-like generalists. 3011:Ornithocheiromorpha 2570:Campylognathoididae 2357:Csehbánya Formation 2067:transitional fossil 1987:quantitative method 1942:Anhanguera santanae 1803:in 1870 discovered 1782:Harry Govier Seeley 1777:Cambridge Greensand 1750:Seeley's dynamical 1667:Johann Georg Wagler 1654:Solnhofen Limestone 989:A neck vertebra of 826:antorbital openings 575:of the pterosaurs. 506:and some non-avian 17509:Evolution of birds 17262:Aquatic locomotion 16374:Istiodactyliformes 14848:Lonchognathosaurus 14691:Eupterodactyloidea 13834:Pterodactyliformes 13637:Pterodactylomorpha 13624:Pterodactylomorpha 13538:Pterodactylomorpha 13380:Angustinaripterini 13011:Eudimorphodontinae 12986:Eudimorphodontidae 11509:. Crescent Books. 11088:Scientific Reports 10717:Historical Biology 10214:10.7717/peerj.2908 10090:Jidapterus edentus 10064:, pp. 150–51. 9845:10.1007/bf02990250 9639:10.7717/peerj.2311 9607:The New York Times 9284:on 31 October 2009 9200:. Academic Press. 8904:10.5252/g2012n4a10 8650:10.7717/peerj.1018 8421:10.7717/peerj.8418 8366:10.7717/peerj.1778 8114:, pp. 14, 17. 7890:Padian, K., 1980, 7811:10.1007/bf02117933 5969:Historical Biology 5261:Benton, Michael J. 4941:. Merriam-Webster. 4818:10.7717/peerj.9604 4340: 4317: 4292:niche partitioning 4290:may also indicate 4183:determinate growth 4023: 3992:Toolebuc formation 3778:found clusters of 3748:Niche partitioning 3621:, envisioned as a 3494: 3470:. Footprints from 3420: 3297:and Mike Habib of 3283: 3235: 3227: 3147:Pterodactylomorpha 3109:Pterodactyliformes 2858:Eupterodactyloidea 2461:Rhamphorhynchoidea 2300: 2255:selection pressure 2167: 2148: 2061:Because pterosaur 2059: 1954: 1893: 1834:Karl Alfred Zittel 1790:Dragons of the Air 1756: 1742:Expanding research 1721:In 1812 and 1817, 1714:or members of the 1690: 1633: 1520: 1457: 1363: 1238:non-pterodactyloid 1210: 1160: 1134: 1061: 1031: 996: 913: 867: 800: 771: 512:respiratory system 443:to the end of the 17723: 17722: 17612:Taxon identifiers 17603: 17602: 17560:Animal locomotion 17499:Evolution of fish 17379:facultative biped 17204: 17203: 17200: 17199: 17173: 17172: 17169: 17168: 17165: 17164: 17161: 17160: 17157: 17156: 17153: 17152: 17149: 17148: 17145: 17144: 17141: 17140: 16883: 16882: 16879: 16878: 16800:Targaryendraconia 16794: 16793: 16604: 16603: 16528: 16527: 16524: 16523: 16314: 16313: 16214: 16213: 16210: 16209: 16206: 16205: 16202: 16201: 15971: 15970: 15940: 15939: 15936: 15935: 15932: 15931: 15904: 15903: 15900: 15899: 15896: 15895: 15892: 15891: 15888: 15887: 15884: 15883: 15880: 15879: 15876: 15875: 15647: 15646: 15615:Chaoyangopterinae 15544:Chaoyangopteridae 15404: 15403: 15400: 15399: 15396: 15395: 15392: 15391: 15388: 15387: 15384: 15383: 14919: 14918: 14915: 14914: 14786:Ornithocheiroidea 14773:Ornithocheiroidea 14762: 14761: 14742: 14741: 14738: 14737: 14734: 14733: 14716:Ornithocheiroidea 14681: 14680: 14677: 14676: 14673: 14672: 14669: 14668: 14665: 14664: 14479:Gladocephaloideus 14415: 14414: 14411: 14410: 14377:Aurorazhdarchidae 14279:Ctenochasmatoidea 14239: 14238: 14197:Germanodactylidae 14144: 14143: 14126: 14125: 14122: 14121: 14118: 14117: 14114: 14113: 14110: 14109: 14106: 14105: 14000: 13999: 13975:Dendrorhynchoides 13952:Batrachognathinae 13918:Dendrorhynchoides 13848:Changchengopterus 13827: 13826: 13823: 13822: 13819: 13818: 13613: 13612: 13584: 13583: 13580: 13579: 13576: 13575: 13572: 13571: 13568: 13567: 13564: 13563: 13560: 13559: 13556: 13555: 13472: 13471: 13468: 13467: 13464: 13463: 13389:Angustinaripterus 13305:Rhamphorhynchinae 13257:Rhamphorhynchidae 13240: 13239: 13231:Campylognathoides 13166: 13165: 13045: 13044: 13041: 13040: 13037: 13036: 12944:Raeticodactylidae 12886: 12885: 12882: 12881: 12849:Austriadraconidae 12728: 12727: 12724: 12723: 12720: 12719: 12568: 12567: 12535: 12534: 12498: 12497: 12494: 12493: 12490: 12489: 12486: 12485: 12482: 12481: 12478: 12477: 12410: 12409: 12233: 12232: 12188: 12187: 12184: 12183: 11975: 11974: 11971: 11970: 11853: 11852: 11749: 11748: 11745: 11744: 11730: 11729: 11726: 11725: 11535:978-0-691-15061-1 11516:978-0-517-03701-0 11497:978-0-7864-2453-5 11276:Hone, D. (2010). 11202:www.bristol.ac.uk 10948:"Life history of 10510:(7017): 571–572. 9768:Unwin DM (1997). 9671:Padian K (2003). 9342:978-0-8137-2376-1 8681:BBC Documentary: 8472:(7838): 445–449. 8140:(1388): 1423–46. 7994:, pp. 1–192. 7758:Palaeontographica 7727:, pp. 37–38. 7625:, Londen: Methuen 7550:Kaup, J. (1834). 7310:(7907): 684–688. 7196:(12): 1592–1593. 7052:(12): 1590–1591. 6646:, pp. 53–54. 5565:978-0-13-146308-0 5340:978-0-8018-8059-9 5282:978-0-520-24209-8 5159:(4180): 947–948. 4919:978-3-12-539683-8 4869:978-0-691-15061-1 4606:Explanatory notes 4454:Jurassic Park III 4157:rhamphorhynchoids 4034:). The eggs from 3961:, paleontologist 3947:Natural predators 3772:. An analysis of 3673:Rhamphorhynchidae 3632:Campylognathoides 3212: 3211: 3203: 3202: 3194: 3193: 3175: 3174: 3156: 3155: 3137: 3136: 3118: 3117: 3099: 3098: 3080: 3079: 3071: 3070: 3062: 3061: 3053: 3052: 3044: 3043: 3035: 3034: 3026: 3025: 2943: 2942: 2885:Ornithocheiroidea 2847: 2846: 2838: 2837: 2823:Ctenochasmatoidea 2771:Germanodactylidae 2676:Changchengopterus 2597:Rhamphorhynchidae 2076:Avemetatarsalians 2003:Dearc sgiathanach 1967:Alexander Kellner 1962:Santana Formation 1947:Santana Formation 1736:Johann Jakob Kaup 1465:Alexander Kellner 1260:Santana Formation 1184:Parts of the wing 485:sexual dimorphism 483:head crests with 464:rhamphorhynchoids 419: 418: 414: 270: 16:(Redirected from 17758: 17751:Pterosauromorpha 17716: 17715: 17703: 17702: 17690: 17689: 17677: 17676: 17664: 17663: 17654: 17653: 17652: 17639: 17638: 17637: 17607: 17606: 17570:Robot locomotion 17344:Limb development 17329: 17302:Lobe-finned fish 17231: 17224: 17217: 17208: 17207: 17195: 17189: 17183: 17182: 17029:Coloborhynchinae 16957: 16956: 16912:Ornithocheiridae 16904:Brasileodactylus 16896: 16895: 16806: 16805: 16696:Ornithocheirinae 16630: 16629: 16625:Ornithocheiridae 16621: 16620: 16617: 16616: 16575: 16574: 16571: 16570: 16550:Brasileodactylus 16459: 16458: 16379: 16378: 16347:Lonchodraconidae 16343: 16342: 16339: 16338: 16231: 16230: 16227: 16226: 16129: 16128: 16097: 16096: 16092:Nyctosauromorpha 16029: 16028: 16005: 16004: 16001: 16000: 15995:Pteranodontoidea 15990: 15989: 15982:Pteranodontoidea 15977: 15976: 15966: 15965: 15958: 15957: 15950: 15949: 15914:Pteranodontoidea 15910: 15909: 15794:Quetzalcoatlinae 15783:Mistralazhdarcho 15748: 15747: 15689: 15688: 15658: 15657: 15549: 15548: 15525: 15524: 15417: 15416: 15318: 15317: 15262: 15261: 15181: 15180: 15166: 15165: 15129: 15128: 15033:Thalassodromidae 15006: 15005: 14995: 14994: 14932: 14931: 14890:Dsungaripterinae 14832: 14831: 14827:Dsungaripteridae 14823: 14822: 14819: 14818: 14792: 14791: 14781: 14780: 14768: 14767: 14757: 14752: 14751: 14696: 14695: 14687: 14686: 14607: 14606: 14602:Ctenochasmatinae 14584:Plataleorhynchus 14428: 14427: 14422:Ctenochasmatidae 14350: 14349: 14289: 14288: 14285: 14284: 14252: 14251: 14246:Euctenochasmatia 14178: 14177: 14174: 14173: 14163: 14162: 14150: 14149: 14139: 14134: 14062:Propterodactylus 14048:Ningchengopterus 14011: 14010: 14006:Pterodactyloidea 13883: 13882: 13874: 13873: 13840: 13839: 13752: 13751: 13721: 13720: 13682: 13681: 13678: 13677: 13643: 13642: 13632: 13631: 13619: 13618: 13608: 13602: 13601: 13594: 13593: 13512:Jianchangopterus 13498:Fenghuangopterus 13484: 13483: 13430:Rhamphorhynchini 13376: 13375: 13310: 13309: 13262: 13261: 13253: 13252: 13211: 13210: 13207: 13206: 13180: 13179: 13109:Dimorphodontidae 13090: 13089: 13086: 13085: 13059: 13058: 12991: 12990: 12940: 12939: 12899: 12898: 12814: 12813: 12762: 12761: 12758: 12757: 12747: 12746: 12734: 12733: 12680: 12679: 12671: 12670: 12664: 12663: 12659:Pterosauromorpha 12610: 12609: 12595: 12588: 12581: 12572: 12571: 12558: 12557: 12530: 12524: 12523: 12517: 12512: 12506: 12426: 12421: 12420: 12327: 12326: 12320: 12305: 12294: 12284: 12274: 12264: 12254: 12246: 12245: 12210: 12205: 12204: 12201: 12200: 12117:Sulcimentisauria 12054: 12053: 12046: 12031: 12020: 12010: 12000: 11992: 11991: 11988: 11987: 11882: 11881: 11877:Pterosauromorpha 11875: 11870: 11869: 11866: 11865: 11804: 11789: 11778: 11770: 11769: 11766: 11765: 11755: 11754: 11696:Archosauromorpha 11686: 11685: 11681:Archosauromorpha 11659: 11658: 11650: 11649: 11643: 11642: 11607: 11606: 11592: 11585: 11578: 11569: 11568: 11539: 11520: 11501: 11473: 11472: 11462: 11428: 11422: 11421: 11413: 11407: 11406: 11404: 11402: 11391: 11382: 11376: 11375: 11373: 11371: 11354: 11348: 11334: 11325: 11319: 11313: 11290: 11284: 11274: 11263: 11262: 11229:(6030): 705–08. 11218: 11212: 11211: 11209: 11208: 11193: 11187: 11186: 11176: 11136: 11130: 11129: 11119: 11079: 11073: 11072: 11062: 11030: 11024: 11023: 11021: 11020: 11006: 11000: 10999: 10989: 10979: 10943: 10934: 10933: 10885: 10874: 10873: 10871: 10870: 10856: 10847: 10846: 10828: 10796: 10790: 10789: 10779: 10770:(3): 1599–1609. 10755: 10749: 10748: 10712: 10706: 10705: 10695: 10663: 10654: 10653: 10620:(6015): 321–24. 10609: 10603: 10602: 10592: 10552: 10546: 10545: 10527: 10495: 10489: 10488: 10472: 10466: 10465: 10423: 10414: 10408: 10407: 10392: 10386: 10385: 10367: 10335: 10329: 10328: 10308: 10297: 10296: 10264: 10258: 10257: 10243: 10237: 10236: 10226: 10216: 10192: 10186: 10185: 10183: 10159: 10153: 10146: 10140: 10139: 10129: 10119: 10083: 10077: 10071: 10065: 10059: 10053: 10052: 10004: 9998: 9997: 9961: 9955: 9949: 9943: 9942: 9908: 9899: 9893: 9887: 9881: 9875: 9869: 9863: 9857: 9856: 9816: 9810: 9804: 9793: 9792: 9774: 9765: 9759: 9758: 9718: 9712: 9711: 9677: 9668: 9662: 9661: 9651: 9641: 9617: 9611: 9610: 9598: 9592: 9591: 9571: 9565: 9564: 9546: 9544:10.1002/ar.22839 9522: 9516: 9515: 9513: 9511: 9502: 9494: 9488: 9487: 9475: 9469: 9468: 9458: 9449: 9443: 9442: 9440: 9438: 9423: 9414: 9413: 9411: 9409: 9404:. 11 August 2013 9394: 9383: 9382: 9380: 9378: 9353: 9347: 9346: 9326: 9320: 9319: 9317: 9315: 9300: 9294: 9293: 9291: 9289: 9280:. Archived from 9269: 9260: 9259: 9215: 9209: 9194: 9188: 9187: 9167: 9161: 9147: 9141: 9140: 9104: 9098: 9097: 9065: 9056: 9055: 9043: 9037: 9036: 9026: 8986: 8980: 8979: 8945: 8936: 8930: 8929: 8927: 8926: 8920: 8914:. Archived from 8889: 8883:Piksi barbarulna 8876: 8870: 8869: 8833: 8827: 8826: 8816: 8806: 8782: 8771: 8770: 8733: 8727: 8726: 8716: 8692: 8686: 8679: 8673: 8672: 8662: 8652: 8628: 8615: 8608: 8602: 8596: 8590: 8589: 8561: 8555: 8554: 8552: 8551: 8536: 8530: 8529: 8527: 8526: 8512: 8506: 8505: 8463: 8453: 8444: 8443: 8433: 8423: 8395: 8389: 8388: 8378: 8368: 8344: 8338: 8337: 8319: 8295: 8289: 8288: 8260: 8251: 8250: 8217:(5836): 358–61. 8208: 8199: 8193: 8192: 8190: 8166: 8160: 8159: 8157: 8121: 8115: 8109: 8103: 8097: 8091: 8090: 8079: 8073: 8067: 8058: 8053: 8044: 8043: 8025: 8001: 7995: 7989: 7983: 7976: 7970: 7964: 7949: 7948: 7904: 7895: 7888: 7882: 7881: 7869: 7863: 7856: 7850: 7843: 7837: 7836: 7834: 7833: 7827: 7821:. Archived from 7794: 7785: 7779: 7772: 7766: 7765: 7753: 7747: 7746: 7734: 7728: 7722: 7716: 7710: 7701: 7695: 7684: 7678: 7669: 7668: 7657:10.1038/024335f0 7632: 7626: 7619: 7613: 7606: 7600: 7594: 7588: 7582: 7576: 7570: 7564: 7563: 7547: 7541: 7540: 7528: 7522: 7516: 7510: 7498: 7492: 7485: 7479: 7467: 7461: 7460: 7448: 7442: 7435: 7429: 7422: 7416: 7410: 7397: 7391: 7385: 7379: 7373: 7367: 7361: 7355: 7346: 7345: 7335: 7295: 7289: 7288: 7270: 7246: 7240: 7239: 7221: 7184: 7178: 7177: 7166:10.1038/371062a0 7141: 7135: 7134: 7124: 7107:(1679): 321–29. 7092: 7086: 7085: 7041: 7035: 7034: 7032: 7031: 7014: 7008: 7007: 6989: 6957: 6951: 6950: 6938: 6932: 6926: 6911: 6905: 6899: 6893: 6887: 6881: 6875: 6869: 6863: 6857: 6851: 6850: 6840: 6830: 6798: 6785: 6779: 6768: 6762: 6745: 6739: 6722: 6721: 6703: 6685: 6653: 6647: 6641: 6635: 6629: 6616: 6610: 6589: 6588: 6578: 6561:(1582): 119–26. 6544: 6538: 6537: 6527: 6503: 6497: 6496: 6476: 6470: 6469: 6443: 6434: 6428: 6427: 6417: 6400:(1582): 119–26. 6383: 6377: 6371: 6365: 6359: 6353: 6347: 6336: 6335: 6317: 6293: 6287: 6286: 6250: 6244: 6243: 6232:10.1360/02tb9054 6207: 6201: 6200: 6189:10.1038/371062a0 6160: 6151: 6145: 6136: 6130: 6121: 6120: 6110: 6100: 6068: 6059: 6058: 6048: 6031:(1679): 321–29. 6012: 6001: 6000: 5964: 5958: 5952: 5943: 5937: 5931: 5925: 5914: 5908: 5902: 5896: 5881: 5875: 5866: 5860: 5849: 5843: 5832: 5826: 5813: 5807: 5801: 5795: 5784: 5778: 5769: 5768: 5748: 5742: 5736: 5730: 5715: 5706: 5705: 5693: 5680: 5674: 5668: 5667: 5643: 5634: 5628: 5617: 5611: 5600: 5594: 5585: 5579: 5570: 5569: 5551: 5532: 5526: 5515: 5509: 5503: 5502: 5490: 5484: 5483: 5449: 5440: 5434: 5433: 5431: 5430: 5416: 5410: 5409: 5376:(6961): 950–53. 5367: 5358: 5345: 5344: 5324: 5315: 5314: 5312: 5310: 5298: 5287: 5286: 5270: 5257: 5251: 5250: 5208: 5199: 5193: 5192: 5145: 5139: 5138: 5128: 5118: 5086: 5077: 5076: 5070: 5062: 5060: 5059: 5044: 5038: 5037: 5035: 5034: 5020: 5014: 5013: 5011: 4987: 4981: 4980: 4974: 4966: 4960: 4959: 4949: 4943: 4942: 4929: 4923: 4922: 4902: 4896: 4895: 4879: 4873: 4872: 4847: 4841: 4840: 4830: 4820: 4796: 4790: 4789: 4779: 4747: 4727: 4721: 4715: 4714: 4711: 4708: 4704: 4698: 4688: 4682: 4678: 4674: 4669: 4668: 4665: 4664: 4661: 4658: 4655: 4650: 4649: 4646: 4643: 4640: 4637: 4634: 4625: 4619: 4616: 4559: 4554: 4553: 4552: 4545: 4540: 4539: 4531: 4529:Dinosaurs portal 4526: 4525: 4524: 4517: 4512: 4511: 4510: 4112:of Germany, and 3837:Thalassodromidae 3833:Dsungaripteridae 3816:was adapted for 3808:Ctenochasmatidae 3534:ctenochasmatoids 3017: 2997: 2984: 2983: 2978:Pteranodontoidea 2957: 2956: 2934: 2914: 2901: 2900: 2891: 2890: 2864: 2863: 2829: 2809: 2794: 2793: 2788:Euctenochasmatia 2777: 2764: 2763: 2754: 2753: 2737: 2722: 2721: 2716:Pterodactyloidea 2705: 2692: 2691: 2668: 2667: 2646: 2645: 2632: 2617: 2616: 2603: 2590: 2589: 2576: 2563: 2562: 2549: 2536: 2535: 2519: 2506: 2505: 2496: 2495: 2465:Pterodactyloidea 2434:Pterosauromorpha 2374:, postulating a 2298:in Mesa Arizona. 2229:showing various 2194:Sterling Nesbitt 2072:Archosauromorpha 1922:Peter Wellnhofer 1885:This drawing of 1852:paleoneurologist 1848:Ferdinand Broili 1774:Early Cretaceous 1716:Pterodactyloidea 1629:Egid Verhelst II 1268:flying squirrels 1000:vertebral column 981:Vertebral column 897:Dsungaripteridae 887:Ctenochasmatidae 865:(drawn to scale) 824:, the nasal and 810:Pterodactyloidea 645:Campyognathoides 573:Pterodactyloidea 439:: from the Late 409: 384: 368: 360: 349: 338: 330: 318: 308: 300: 292: 284: 265: 260: 249:Pterosauromorpha 247: 191: 190: 137: 127: 64: 49:Temporal range: 43: 42: 21: 17766: 17765: 17761: 17760: 17759: 17757: 17756: 17755: 17726: 17725: 17724: 17719: 17711: 17706: 17698: 17693: 17685: 17680: 17672: 17667: 17659: 17657: 17648: 17647: 17642: 17633: 17632: 17627: 17614: 17604: 17599: 17548: 17514:Origin of birds 17487: 17427: 17349:Limb morphology 17330: 17321: 17307:Ray-finned fish 17272:Fish locomotion 17248: 17235: 17205: 17196: 17177: 17137: 17118:Liaoningopterus 17073: 17023: 16961:Tropeognathinae 16946: 16918: 16892: 16875: 16839: 16810:Cimoliopteridae 16790: 16690: 16613: 16600: 16595:Zhenyuanopterus 16569: 16520: 16499:Istiodactylinae 16493: 16474:Longchengpterus 16467:Lingyuanopterus 16454:Istiodactylidae 16448: 16414: 16387:Hongshanopterus 16368: 16337: 16323: 16310: 16273:Hongshanopterus 16258: 16223: 16198: 16171:Barbaridactylus 16156: 16118: 16086: 16033:Pteranodontidae 16018: 16013:Santanadactylus 15997: 15984: 15967: 15960: 15952: 15944: 15928: 15872: 15867:Zhejiangopterus 15788: 15746: 15725:Palaeocursornis 15687: 15666:Montanazhdarcho 15653:Azhdarchiformes 15643: 15624:Chaoyangopterus 15609: 15538: 15514: 15500:Montanazhdarcho 15453: 15413: 15411:Azhdarchomorpha 15380: 15353: 15307: 15270:Aymberedactylus 15251: 15179: 15155: 15143:Aymberedactylus 15118: 15098:Thalassodromeus 15027: 15002: 14991: 14982: 14970:Montanazhdarcho 14928: 14911: 14899:Domeykodactylus 14884: 14878:Tendaguripterus 14815: 14806: 14788: 14775: 14758: 14746: 14730: 14710: 14704:Altmuehlopterus 14661: 14620: 14596: 14547: 14519: 14424: 14407: 14371: 14365:Gallodactylidae 14345:Aurorazhdarchia 14339: 14317:Normannognathus 14293:Gallodactylidae 14281: 14272: 14248: 14235: 14229:Tendaguripterus 14221:Normannognathus 14214:Germanodactylus 14206:Altmuehlopterus 14191: 14170: 14157: 14140: 14102: 14082: 13996: 13961:Batrachognathus 13946: 13941:Vesperopterylus 13896: 13870: 13861: 13836: 13815: 13771:Wukongopterinae 13765: 13747:Wukongopteridae 13741: 13710: 13697:Normannognathus 13674: 13665: 13639: 13626: 13609: 13596: 13588: 13552: 13532: 13505:Jianchangnathus 13488:Scaphognathidae 13478:Digibrevisauria 13460: 13455:Rhamphorhynchus 13424: 13411:Qinglongopterus 13403:Harpactognathus 13371:Rhamphorhynchae 13365: 13360:Qinglongopterus 13339:Harpactognathus 13299: 13293:Scaphognathinae 13285:Parapsicephalus 13249: 13236: 13224:Bergamodactylus 13203: 13194: 13176: 13162: 13140:Parapsicephalus 13103: 13082: 13080:Macronychoptera 13073: 13055: 13033: 13005: 12999:Arcticodactylus 12980: 12968:Raeticodactylus 12929: 12895: 12878: 12858:Arcticodactylus 12843: 12837:Raeticodactylus 12802: 12790:Austriadactylus 12775: 12754: 12741: 12716: 12696: 12690:Avemetatarsalia 12675:Avemetatarsalia 12665: 12650:Avemetatarsalia 12615: 12614: 12604: 12599: 12569: 12564: 12550:Phytodinosauria 12531: 12518: 12474: 12445: 12439:Sauropodomorpha 12428:Sauropodomorpha 12406: 12371:Herrerasauridae 12365: 12313: 12242: 12229: 12197: 12180: 12111: 12039: 11984: 11982:Dinosauromorpha 11967: 11944: 11862: 11849: 11797: 11762: 11760:Avemetatarsalia 11741: 11738:Avemetatarsalia 11722: 11708:Avemetatarsalia 11702: 11675: 11644: 11612: 11611: 11601: 11596: 11546: 11536: 11517: 11498: 11482: 11477: 11476: 11438:Current Biology 11429: 11425: 11414: 11410: 11400: 11398: 11389: 11383: 11379: 11369: 11367: 11355: 11351: 11335: 11328: 11320: 11316: 11291: 11287: 11275: 11266: 11219: 11215: 11206: 11204: 11194: 11190: 11137: 11133: 11080: 11076: 11031: 11027: 11018: 11016: 11008: 11007: 11003: 10950:Rhamphorhynchus 10944: 10937: 10890:Rhamphorhynchus 10886: 10877: 10868: 10866: 10858: 10857: 10850: 10826:10.1038/429621a 10797: 10793: 10756: 10752: 10713: 10709: 10678:(12): 1323–30. 10672:Current Biology 10664: 10657: 10610: 10606: 10553: 10549: 10516:10.1038/432571a 10496: 10492: 10473: 10469: 10446:10.1038/432572a 10421: 10415: 10411: 10394: 10393: 10389: 10365:10.1038/430033a 10336: 10332: 10309: 10300: 10265: 10261: 10244: 10240: 10193: 10189: 10160: 10156: 10147: 10143: 10102:(9): e0185486. 10084: 10080: 10072: 10068: 10060: 10056: 10005: 10001: 9962: 9958: 9950: 9946: 9906: 9900: 9896: 9888: 9884: 9876: 9872: 9864: 9860: 9817: 9813: 9805: 9796: 9772: 9766: 9762: 9719: 9715: 9686:(2–4): 115–26. 9675: 9669: 9665: 9618: 9614: 9599: 9595: 9572: 9568: 9537:(12): 2233–53. 9523: 9519: 9509: 9507: 9505:Qmro.qmul.ac.uk 9500: 9496: 9495: 9491: 9476: 9472: 9456: 9450: 9446: 9436: 9434: 9424: 9417: 9407: 9405: 9396: 9395: 9386: 9376: 9374: 9354: 9350: 9343: 9327: 9323: 9313: 9311: 9301: 9297: 9287: 9285: 9270: 9263: 9216: 9212: 9195: 9191: 9184: 9168: 9164: 9148: 9144: 9105: 9101: 9066: 9059: 9044: 9040: 8987: 8983: 8943: 8937: 8933: 8924: 8922: 8918: 8887: 8877: 8873: 8834: 8830: 8797:(3): e2001663. 8783: 8774: 8734: 8730: 8693: 8689: 8680: 8676: 8629: 8618: 8609: 8605: 8597: 8593: 8562: 8558: 8549: 8547: 8537: 8533: 8524: 8522: 8514: 8513: 8509: 8461: 8454: 8447: 8396: 8392: 8345: 8341: 8296: 8292: 8261: 8254: 8206: 8200: 8196: 8167: 8163: 8122: 8118: 8110: 8106: 8098: 8094: 8081: 8080: 8076: 8068: 8061: 8054: 8047: 8002: 7998: 7992:Wellnhofer 1991 7990: 7986: 7977: 7973: 7965: 7952: 7905: 7898: 7889: 7885: 7870: 7866: 7857: 7853: 7844: 7840: 7831: 7829: 7825: 7805:(1/3): 105–12. 7792: 7786: 7782: 7773: 7769: 7754: 7750: 7735: 7731: 7725:Wellnhofer 1991 7723: 7719: 7713:Wellnhofer 1991 7711: 7704: 7698:Wellnhofer 1991 7696: 7687: 7681:Wellnhofer 1991 7679: 7672: 7643:(615): 309–38. 7633: 7629: 7620: 7616: 7607: 7603: 7597:Wellnhofer 1991 7595: 7591: 7585:Wellnhofer 1991 7583: 7579: 7573:Wellnhofer 1991 7571: 7567: 7548: 7544: 7529: 7525: 7519:Wellnhofer 1991 7517: 7513: 7499: 7495: 7486: 7482: 7468: 7464: 7449: 7445: 7436: 7432: 7423: 7419: 7411: 7400: 7396:, pp. 6–7. 7392: 7388: 7380: 7376: 7370:Wellnhofer 1991 7368: 7364: 7356: 7349: 7296: 7292: 7247: 7243: 7185: 7181: 7152:(6492): 62–64. 7142: 7138: 7093: 7089: 7042: 7038: 7029: 7027: 7015: 7011: 6958: 6954: 6939: 6935: 6927: 6914: 6906: 6902: 6894: 6890: 6882: 6878: 6870: 6866: 6858: 6854: 6799: 6788: 6780: 6771: 6765:Wellnhofer 1991 6763: 6748: 6742:Wellnhofer 1991 6740: 6725: 6654: 6650: 6644:Wellnhofer 1991 6642: 6638: 6632:Wellnhofer 1991 6630: 6619: 6611: 6592: 6545: 6541: 6504: 6500: 6477: 6473: 6441: 6435: 6431: 6384: 6380: 6372: 6368: 6360: 6356: 6350:Wellnhofer 1991 6348: 6339: 6294: 6290: 6251: 6247: 6208: 6204: 6175:(6492): 62–64. 6161: 6154: 6146: 6139: 6131: 6124: 6069: 6062: 6013: 6004: 5965: 5961: 5953: 5946: 5938: 5934: 5926: 5917: 5909: 5905: 5899:Wellnhofer 1991 5897: 5884: 5878:Wellnhofer 1991 5876: 5869: 5861: 5852: 5844: 5835: 5827: 5816: 5808: 5804: 5798:Wellnhofer 1991 5796: 5787: 5779: 5772: 5749: 5745: 5739:Wellnhofer 1991 5737: 5733: 5716: 5709: 5694: 5683: 5677:Wellnhofer 1991 5675: 5671: 5644: 5637: 5629: 5620: 5612: 5603: 5597:Wellnhofer 1991 5595: 5588: 5580: 5573: 5566: 5552: 5535: 5527: 5518: 5510: 5506: 5491: 5487: 5447: 5441: 5437: 5428: 5426: 5418: 5417: 5413: 5365: 5359: 5348: 5341: 5325: 5318: 5308: 5306: 5305:. Pterosaur.net 5301:Naish, Darren. 5299: 5290: 5283: 5258: 5254: 5206: 5200: 5196: 5146: 5142: 5087: 5080: 5064: 5063: 5057: 5055: 5053:livescience.com 5045: 5041: 5032: 5030: 5022: 5021: 5017: 4988: 4984: 4979:: 61–107. 2008. 4972: 4968: 4967: 4963: 4950: 4946: 4931: 4930: 4926: 4920: 4903: 4899: 4880: 4876: 4870: 4848: 4844: 4797: 4793: 4756:Current Biology 4748: 4741: 4736: 4731: 4730: 4722: 4718: 4712: 4709: 4706: 4689: 4685: 4676: 4672: 4652: 4631: 4627: 4626: 4622: 4617: 4613: 4608: 4590:Pterosaur Beach 4555: 4550: 4548: 4541: 4534: 4527: 4522: 4520: 4515:Reptiles portal 4513: 4508: 4506: 4503: 4484:Vesperopterylus 4413:Ray Harryhausen 4361:Rhamphorhynchus 4351:Rhamphorhynchus 4336:Rhamphorhynchus 4300: 4280:Rhamphorhynchus 4260:Rhamphorhynchus 4237:Rhamphorhynchus 4194: 4169:Rhamphorhynchus 4161:Rhamphorhynchus 4106:Rhamphorhynchus 4011: 3999:] (most likely 3949: 3918:Thalassodromeus 3875:pterodactyloids 3842:Thalassodromeus 3797:Germanodactylus 3757:Alcione elainus 3744:Pteranodontidae 3715:Wukongopteridae 3702:Harpactognathus 3678:Rhamphorhynchus 3655:Austriadactylus 3642:and the larger 3598: 3560: 3510:ornithocheirids 3475:in pterosaurs. 3401: 3399:Ground movement 3393:pterodactyloids 3355: 3334: 3265:. Furthermore, 3251:Late Cretaceous 3240: 3219: 3214: 3213: 3204: 3195: 3190: 3189: 3188: 3187: 3176: 3171: 3170: 3169: 3168: 3157: 3152: 3151: 3150: 3149: 3138: 3133: 3132: 3131: 3130: 3119: 3114: 3113: 3112: 3111: 3100: 3095: 3094: 3093: 3092: 3081: 3072: 3063: 3054: 3045: 3036: 3027: 2944: 2848: 2839: 2530:Macronychoptera 2426:Ornithocephalus 2394: 2388: 2372:Late Cretaceous 2284: 2275:Prorotodactylus 2247:neuroanatomical 2239:sensory systems 2144:archosauromorph 2055:neuroanatomical 2033: 2028: 2020:Jurassic period 1888:Zhejiangopterus 1879: 1838:Rhamphorhynchus 1817:Rhamphorhynchus 1744: 1669:suggested that 1615: 1610: 1604: 1532: 1446: 1426: 1396: 1355: 1321:pteranodontians 1219:brachiopatagium 1186: 1155:Rhamphorhynchus 1126: 1090:shoulder girdle 1086: 1084:Shoulder girdle 983: 962:Austriadactylus 909:Thalassodromeus 883:Istiodactylidae 875:Pteranodontidae 787: 763: 757: 740:Vesperopterylus 696:Germanodactylus 651:Rhamphorhynchus 589: 498:. These may be 476:pterodactyloids 424:are an extinct 332:Macronychoptera 321:Austriadactylus 264: 258: 245: 185: 128: 126: 125: 120: 115: 110: 105: 100: 95: 90: 85: 80: 75: 70: 59: 58: 55:Late Cretaceous 47: 39: 28: 23: 22: 15: 12: 11: 5: 17764: 17754: 17753: 17748: 17743: 17738: 17721: 17720: 17718: 17717: 17704: 17691: 17678: 17665: 17655: 17640: 17624: 17622: 17616: 17615: 17601: 17600: 17598: 17597: 17592: 17587: 17582: 17577: 17572: 17567: 17562: 17556: 17554: 17550: 17549: 17547: 17546: 17541: 17536: 17531: 17526: 17521: 17516: 17511: 17506: 17501: 17495: 17493: 17489: 17488: 17486: 17485: 17480: 17478:Pterosaur wing 17475: 17470: 17469: 17468: 17463: 17458: 17448: 17443: 17437: 17435: 17429: 17428: 17426: 17425: 17420: 17415: 17414: 17413: 17403: 17398: 17393: 17392: 17391: 17386: 17381: 17376: 17371: 17366: 17361: 17356: 17346: 17340: 17338: 17332: 17331: 17324: 17322: 17320: 17319: 17314: 17309: 17304: 17299: 17294: 17289: 17284: 17279: 17274: 17269: 17267:Cephalopod fin 17264: 17258: 17256: 17250: 17249: 17234: 17233: 17226: 17219: 17211: 17202: 17201: 17198: 17197: 17176: 17174: 17171: 17170: 17167: 17166: 17163: 17162: 17159: 17158: 17155: 17154: 17151: 17150: 17147: 17146: 17143: 17142: 17139: 17138: 17136: 17135: 17128: 17121: 17114: 17107: 17099: 17095:Caulkicephalus 17091: 17083: 17081: 17075: 17074: 17072: 17071: 17068:Uktenadactylus 17064: 17056: 17049: 17046:Coloborhynchus 17042: 17034: 17032: 17025: 17024: 17022: 17021: 17014: 17007: 16999: 16991: 16983: 16975: 16966: 16964: 16954: 16948: 16947: 16945: 16944: 16937: 16929: 16927: 16920: 16919: 16917: 16916: 16908: 16899: 16893: 16888: 16885: 16884: 16881: 16880: 16877: 16876: 16874: 16873: 16870:Targaryendraco 16866: 16858: 16849: 16847: 16841: 16840: 16838: 16837: 16830: 16822: 16814: 16812: 16803: 16796: 16795: 16792: 16791: 16789: 16788: 16784:Uktenadactylus 16780: 16772: 16769:Ornithocheirus 16765: 16757: 16749: 16741: 16733: 16725: 16721:Coloborhynchus 16717: 16709: 16700: 16698: 16692: 16691: 16689: 16688: 16680: 16672: 16664: 16660:Caulkicephalus 16656: 16648: 16645:Arthurdactylus 16641: 16633: 16627: 16614: 16611:Ornithocheirae 16609: 16606: 16605: 16602: 16601: 16599: 16598: 16591: 16583: 16581: 16568: 16567: 16560: 16553: 16546: 16538: 16536: 16530: 16529: 16526: 16525: 16522: 16521: 16519: 16518: 16511: 16503: 16501: 16495: 16494: 16492: 16491: 16484: 16477: 16470: 16462: 16456: 16450: 16449: 16447: 16446: 16439: 16436:Linlongopterus 16432: 16424: 16422: 16420:Mimodactylidae 16416: 16415: 16413: 16412: 16408:Lonchodectidae 16404: 16397: 16394:Linlongopterus 16390: 16382: 16376: 16370: 16369: 16367: 16366: 16359: 16351: 16349: 16336: 16335: 16326: 16324: 16319: 16316: 16315: 16312: 16311: 16309: 16308: 16304:Targaryendraco 16300: 16292: 16285: 16277: 16268: 16266: 16264:Lonchodectidae 16260: 16259: 16257: 16256: 16249: 16242: 16234: 16224: 16219: 16216: 16215: 16212: 16211: 16208: 16207: 16204: 16203: 16200: 16199: 16197: 16196: 16188: 16181: 16178:Muzquizopteryx 16174: 16166: 16164: 16158: 16157: 16155: 16154: 16147: 16140: 16132: 16126: 16124:Aponyctosauria 16120: 16119: 16117: 16116: 16108: 16100: 16094: 16088: 16087: 16085: 16084: 16076: 16068: 16061: 16053: 16046: 16037: 16035: 16026: 16020: 16019: 16017: 16016: 16008: 15998: 15993: 15986: 15985: 15973: 15972: 15969: 15968: 15943: 15941: 15938: 15937: 15934: 15933: 15930: 15929: 15927: 15926: 15925: 15924: 15918: 15916: 15906: 15905: 15902: 15901: 15898: 15897: 15894: 15893: 15890: 15889: 15886: 15885: 15882: 15881: 15878: 15877: 15874: 15873: 15871: 15870: 15863: 15855: 15852:Thanatosdrakon 15848: 15845:Quetzalcoatlus 15841: 15838:Phosphatodraco 15834: 15827: 15820: 15813: 15810:Arambourgiania 15806: 15798: 15796: 15790: 15789: 15787: 15786: 15779: 15772: 15765: 15756: 15754: 15745: 15744: 15736: 15728: 15721: 15717:Navajodactylus 15713: 15706: 15697: 15695: 15686: 15685: 15677: 15670: 15661: 15655: 15649: 15648: 15645: 15644: 15642: 15641: 15638:Shenzhoupterus 15634: 15627: 15619: 15617: 15611: 15610: 15608: 15607: 15599: 15591: 15584: 15577: 15569: 15565:Argentinadraco 15561: 15552: 15546: 15540: 15539: 15537: 15536: 15528: 15522: 15516: 15515: 15513: 15512: 15504: 15496: 15488: 15480: 15476:Argentinadraco 15472: 15464: 15462: 15455: 15454: 15452: 15451: 15443: 15436: 15429: 15420: 15414: 15409: 15406: 15405: 15402: 15401: 15398: 15397: 15394: 15393: 15390: 15389: 15386: 15385: 15382: 15381: 15379: 15378: 15371: 15363: 15361: 15355: 15354: 15352: 15351: 15344: 15337: 15330: 15321: 15315: 15309: 15308: 15306: 15305: 15298: 15290: 15282: 15274: 15265: 15259: 15253: 15252: 15250: 15249: 15242: 15235: 15232:Nemicolopterus 15228: 15221: 15214: 15206: 15198: 15189: 15187: 15178: 15177: 15169: 15163: 15157: 15156: 15154: 15153: 15146: 15138: 15136: 15126: 15124:Tapejariformes 15120: 15119: 15117: 15116: 15108: 15101: 15094: 15086: 15078: 15071: 15063: 15059:Argentinadraco 15055: 15047: 15038: 15036: 15029: 15028: 15026: 15025: 15017: 15009: 15003: 15000:Tapejaromorpha 14998: 14992: 14987: 14984: 14983: 14981: 14980: 14973: 14966: 14958: 14950: 14943: 14940:Argentinadraco 14935: 14929: 14924: 14921: 14920: 14917: 14916: 14913: 14912: 14910: 14909: 14906:Dsungaripterus 14902: 14894: 14892: 14886: 14885: 14883: 14882: 14874: 14866: 14859: 14852: 14844: 14835: 14829: 14816: 14811: 14808: 14807: 14805: 14804: 14795: 14789: 14784: 14777: 14776: 14764: 14763: 14760: 14759: 14745: 14743: 14740: 14739: 14736: 14735: 14732: 14731: 14729: 14728: 14727: 14726: 14720: 14718: 14712: 14711: 14709: 14708: 14699: 14693: 14683: 14682: 14679: 14678: 14675: 14674: 14671: 14670: 14667: 14666: 14663: 14662: 14660: 14659: 14652: 14645: 14638: 14630: 14628: 14626:Pterodaustrini 14622: 14621: 14619: 14618: 14610: 14604: 14598: 14597: 14595: 14594: 14587: 14580: 14573: 14565: 14557: 14555: 14553:Gnathosaurinae 14549: 14548: 14546: 14545: 14538: 14530: 14528: 14525:Moganopterinae 14521: 14520: 14518: 14517: 14510: 14503: 14496: 14489: 14482: 14475: 14468: 14461: 14454: 14447: 14444:Balaenognathus 14440: 14431: 14425: 14420: 14417: 14416: 14413: 14412: 14409: 14408: 14406: 14405: 14397: 14394:Aurorazhdarcho 14390: 14381: 14379: 14373: 14372: 14370: 14369: 14361: 14353: 14347: 14341: 14340: 14338: 14337: 14333:Moganopterinae 14329: 14321: 14313: 14306: 14302:Aurorazhdarcho 14297: 14295: 14282: 14277: 14274: 14273: 14271: 14270: 14263: 14255: 14249: 14244: 14241: 14240: 14237: 14236: 14234: 14233: 14225: 14217: 14210: 14201: 14199: 14193: 14192: 14190: 14189: 14181: 14171: 14166: 14159: 14158: 14146: 14145: 14142: 14141: 14129: 14127: 14124: 14123: 14120: 14119: 14116: 14115: 14112: 14111: 14108: 14107: 14104: 14103: 14101: 14100: 14099: 14098: 14092: 14090: 14084: 14083: 14081: 14080: 14073: 14065: 14058: 14051: 14044: 14037: 14029: 14022: 14014: 14008: 14002: 14001: 13998: 13997: 13995: 13994: 13987: 13979: 13971: 13964: 13956: 13954: 13948: 13947: 13945: 13944: 13937: 13930: 13922: 13914: 13906: 13904: 13902:Anurognathinae 13898: 13897: 13895: 13894: 13886: 13880: 13878:Anurognathidae 13871: 13866: 13863: 13862: 13860: 13859: 13856:Douzhanopterus 13852: 13843: 13837: 13832: 13829: 13828: 13825: 13824: 13821: 13820: 13817: 13816: 13814: 13813: 13806: 13802:Douzhanopterus 13798: 13791: 13784: 13775: 13773: 13767: 13766: 13764: 13763: 13760:Kunpengopterus 13755: 13749: 13743: 13742: 13740: 13739: 13732: 13724: 13718: 13712: 13711: 13709: 13708: 13704:Anurognathidae 13700: 13693: 13685: 13675: 13672:Monofenestrata 13670: 13667: 13666: 13664: 13663: 13655: 13646: 13640: 13635: 13628: 13627: 13615: 13614: 13611: 13610: 13587: 13585: 13582: 13581: 13578: 13577: 13574: 13573: 13570: 13569: 13566: 13565: 13562: 13561: 13558: 13557: 13554: 13553: 13551: 13550: 13549: 13548: 13542: 13540: 13534: 13533: 13531: 13530: 13522: 13515: 13508: 13501: 13493: 13491: 13481: 13474: 13473: 13470: 13469: 13466: 13465: 13462: 13461: 13459: 13458: 13451: 13443: 13434: 13432: 13426: 13425: 13423: 13422: 13415: 13407: 13399: 13392: 13384: 13382: 13373: 13367: 13366: 13364: 13363: 13356: 13353:Orientognathus 13349: 13342: 13335: 13328: 13321: 13313: 13307: 13301: 13300: 13298: 13297: 13289: 13281: 13274: 13270:Dolicorhamphus 13265: 13259: 13250: 13247:Breviquartossa 13245: 13242: 13241: 13238: 13237: 13235: 13234: 13227: 13219: 13217: 13204: 13199: 13196: 13195: 13193: 13192: 13183: 13177: 13171: 13168: 13167: 13164: 13163: 13161: 13160: 13152: 13144: 13136: 13129: 13126:Caelestiventus 13122: 13113: 13111: 13105: 13104: 13102: 13101: 13093: 13083: 13078: 13075: 13074: 13072: 13071: 13062: 13056: 13052:Zambellisauria 13050: 13047: 13046: 13043: 13042: 13039: 13038: 13035: 13034: 13032: 13031: 13024: 13020:Carniadactylus 13015: 13013: 13007: 13006: 13004: 13003: 12994: 12988: 12982: 12981: 12979: 12978: 12971: 12964: 12957: 12948: 12946: 12937: 12931: 12930: 12928: 12927: 12919: 12911: 12902: 12896: 12891: 12888: 12887: 12884: 12883: 12880: 12879: 12877: 12876: 12873:Seazzadactylus 12869: 12862: 12853: 12851: 12845: 12844: 12842: 12841: 12833: 12826: 12822:Carniadactylus 12817: 12811: 12804: 12803: 12801: 12800: 12793: 12785: 12783: 12777: 12776: 12774: 12773: 12770:Daohugoupterus 12765: 12755: 12750: 12743: 12742: 12730: 12729: 12726: 12725: 12722: 12721: 12718: 12717: 12715: 12714: 12713: 12712: 12706: 12704: 12698: 12697: 12695: 12694: 12693: 12692: 12683: 12677: 12667: 12666: 12662: 12661: 12652: 12643: 12634: 12628: 12622: 12606: 12605: 12598: 12597: 12590: 12583: 12575: 12566: 12565: 12563: 12562: 12552: 12547: 12545:Ornithoscelida 12540: 12537: 12536: 12533: 12532: 12501: 12499: 12496: 12495: 12492: 12491: 12488: 12487: 12484: 12483: 12480: 12479: 12476: 12475: 12473: 12472: 12471: 12470: 12464: 12455: 12453: 12447: 12446: 12444: 12443: 12442: 12441: 12432: 12430: 12418: 12412: 12411: 12408: 12407: 12405: 12404: 12401:Staurikosaurus 12397: 12390: 12383: 12375: 12373: 12367: 12366: 12364: 12363: 12355: 12347: 12339: 12330: 12324: 12315: 12314: 12312: 12311: 12301: 12290: 12280: 12270: 12260: 12249: 12243: 12238: 12235: 12234: 12231: 12230: 12228: 12227: 12226: 12225: 12216: 12214: 12198: 12193: 12190: 12189: 12186: 12185: 12182: 12181: 12179: 12178: 12171: 12164: 12157: 12150: 12143: 12136: 12129: 12121: 12119: 12113: 12112: 12110: 12109: 12101: 12094: 12086: 12079: 12072: 12065: 12057: 12051: 12041: 12040: 12038: 12037: 12027: 12016: 12006: 11995: 11985: 11980: 11977: 11976: 11973: 11972: 11969: 11968: 11966: 11965: 11964: 11963: 11954: 11952: 11946: 11945: 11943: 11942: 11935: 11927: 11920: 11913: 11906: 11902:Faxinalipterus 11898: 11890: 11888: 11879: 11863: 11858: 11855: 11854: 11851: 11850: 11848: 11847: 11840: 11833: 11826: 11819: 11815:Incertovenator 11810: 11808: 11799: 11798: 11796: 11795: 11785: 11781:Incertovenator 11773: 11763: 11758: 11751: 11750: 11747: 11746: 11743: 11742: 11735: 11732: 11731: 11728: 11727: 11724: 11723: 11721: 11720: 11719: 11718: 11712: 11710: 11704: 11703: 11701: 11700: 11699: 11698: 11689: 11683: 11677: 11676: 11674: 11673: 11672: 11671: 11662: 11656: 11646: 11645: 11641: 11640: 11631: 11625: 11619: 11603: 11602: 11595: 11594: 11587: 11580: 11572: 11566: 11565: 11559: 11553: 11545: 11544:External links 11542: 11541: 11540: 11534: 11521: 11515: 11502: 11496: 11481: 11478: 11475: 11474: 11423: 11408: 11377: 11349: 11326: 11324:, p. 452. 11314: 11285: 11264: 11213: 11188: 11131: 11074: 11025: 11001: 10935: 10875: 10848: 10791: 10750: 10707: 10655: 10604: 10567:(3): 647–654. 10547: 10490: 10467: 10409: 10387: 10330: 10319:(2): 602–635. 10298: 10259: 10256:on 2016-04-06. 10238: 10187: 10154: 10141: 10078: 10076:, p. 199. 10066: 10054: 9999: 9956: 9954:, p. 134. 9944: 9894: 9892:, p. 122. 9882: 9880:, p. 121. 9870: 9868:, p. 103. 9858: 9811: 9794: 9760: 9713: 9663: 9612: 9593: 9566: 9517: 9489: 9484:New York Times 9470: 9444: 9415: 9384: 9348: 9341: 9321: 9295: 9261: 9210: 9189: 9182: 9162: 9142: 9099: 9057: 9038: 8981: 8954:(2): 288–321. 8931: 8871: 8828: 8772: 8728: 8707:(6): 1144–55. 8687: 8674: 8616: 8603: 8591: 8556: 8539:Black, Riley. 8531: 8507: 8445: 8390: 8339: 8290: 8252: 8194: 8181:(3): 261–308. 8161: 8116: 8104: 8092: 8089:. 22 Feb 2022. 8074: 8059: 8045: 7996: 7984: 7971: 7950: 7896: 7883: 7864: 7851: 7838: 7780: 7767: 7748: 7729: 7717: 7702: 7685: 7670: 7627: 7614: 7601: 7589: 7577: 7565: 7542: 7523: 7511: 7493: 7480: 7462: 7443: 7430: 7417: 7398: 7386: 7374: 7362: 7347: 7290: 7241: 7179: 7136: 7087: 7036: 7009: 6952: 6933: 6912: 6900: 6888: 6876: 6864: 6852: 6786: 6769: 6746: 6723: 6648: 6636: 6617: 6590: 6539: 6518:(4): 1045–62. 6498: 6471: 6429: 6378: 6366: 6354: 6337: 6308:(4): 1339–42. 6288: 6245: 6202: 6165:Sordes pilosus 6152: 6137: 6122: 6060: 6002: 5959: 5944: 5932: 5915: 5903: 5882: 5867: 5850: 5833: 5814: 5802: 5785: 5770: 5743: 5731: 5707: 5681: 5669: 5635: 5618: 5601: 5586: 5571: 5564: 5533: 5516: 5504: 5485: 5458:(1): 163–187. 5435: 5411: 5346: 5339: 5316: 5288: 5281: 5267:The Dinosauria 5252: 5194: 5140: 5101:(6): 1983–87. 5078: 5039: 5015: 4982: 4961: 4944: 4924: 4918: 4897: 4874: 4868: 4842: 4791: 4762:(9): 1011–16. 4738: 4737: 4735: 4732: 4729: 4728: 4716: 4683: 4620: 4610: 4609: 4607: 4604: 4603: 4602: 4597: 4595:Pterosaur size 4592: 4587: 4582: 4577: 4572: 4567: 4561: 4560: 4546: 4532: 4518: 4502: 4499: 4493:Kunpengopterus 4392:The Lost World 4368:have both the 4309:Quetzalcoatlus 4299: 4296: 4193: 4190: 4173:pterodactyloid 4137:superprecocial 4042:was laid by a 4019:pterodactyloid 4010: 4007: 4002:Platypterygius 3973:, most likely 3963:Eric Buffetaut 3948: 3945: 3908:Dsungaripterus 3879:pteranodontids 3788:istiodactylids 3736:Ornithocheirae 3639:Carniadactylus 3606:Anurognathidae 3597: 3594: 3559: 3556: 3400: 3397: 3354: 3353:Nervous system 3351: 3343:hepatic piston 3333: 3330: 3307:Paul MacCready 3239: 3236: 3218: 3215: 3210: 3209: 3206: 3205: 3201: 3200: 3197: 3196: 3192: 3191: 3183: 3182: 3181: 3178: 3177: 3173: 3172: 3166:Breviquartossa 3164: 3163: 3162: 3159: 3158: 3154: 3153: 3145: 3144: 3143: 3140: 3139: 3135: 3134: 3128:Monofenestrata 3126: 3125: 3124: 3121: 3120: 3116: 3115: 3107: 3106: 3105: 3102: 3101: 3097: 3096: 3088: 3087: 3086: 3083: 3082: 3078: 3077: 3074: 3073: 3069: 3068: 3065: 3064: 3060: 3059: 3056: 3055: 3051: 3050: 3047: 3046: 3042: 3041: 3038: 3037: 3033: 3032: 3029: 3028: 3024: 3023: 3020: 3019: 3007: 3004: 3003: 3000: 2999: 2987: 2982: 2980: 2974: 2973: 2970: 2969: 2960: 2955: 2953: 2950: 2949: 2946: 2945: 2941: 2940: 2937: 2936: 2924: 2921: 2920: 2917: 2916: 2908:Tapejaromorpha 2904: 2899: 2897: 2889: 2887: 2881: 2880: 2877: 2876: 2867: 2862: 2860: 2854: 2853: 2850: 2849: 2845: 2844: 2841: 2840: 2836: 2835: 2832: 2831: 2819: 2816: 2815: 2812: 2811: 2797: 2792: 2790: 2784: 2783: 2780: 2779: 2767: 2762: 2760: 2752: 2750: 2744: 2743: 2740: 2739: 2725: 2720: 2718: 2712: 2711: 2708: 2707: 2699:Anurognathidae 2695: 2690: 2688: 2685: 2684: 2681: 2680: 2671: 2666: 2664: 2661: 2660: 2657: 2656: 2649: 2644: 2642: 2639: 2638: 2635: 2634: 2620: 2615: 2613: 2610: 2609: 2606: 2605: 2593: 2588: 2586: 2583: 2582: 2579: 2578: 2566: 2561: 2559: 2556: 2555: 2552: 2551: 2543:Dimorphodontia 2539: 2534: 2532: 2526: 2525: 2522: 2521: 2509: 2504: 2502: 2494: 2493: 2449:classification 2413:Quetzalcoatlus 2387: 2384: 2349:Navajodactylus 2340:pteranodontids 2335:Ornithocheirus 2292:Quetzalcoatlus 2283: 2280: 2202:Martin Ezcurra 2116:Michael Benton 2096:tanystropheids 2032: 2029: 2027: 2024: 1878: 1875: 1809:Niobrara Chalk 1754:reconstruction 1743: 1740: 1698:Georges Cuvier 1694:Johann Hermann 1614: 1611: 1603: 1600: 1575:Sordes pilosus 1551:Inner Mongolia 1536:proto-feathers 1531: 1528: 1500:Sordes pilosus 1445: 1442: 1434:petrifications 1425: 1422: 1395: 1392: 1390:chest cavity. 1354: 1351: 1185: 1182: 1163:fibers called 1125: 1122: 1098:shoulder blade 1085: 1082: 1065:caudofemoralis 1008:vertebral body 992:Arambourgiania 982: 979: 923:parietal bones 820:. Unlike most 796:Coloborhynchus 786: 783: 761:Pterosaur size 759:Main article: 756: 753: 720:Quetzalcoatlus 702:Dsungaripterus 672:Ornithocheirus 621:shoulder blade 588: 585: 529:Quetzalcoatlus 453:powered flight 417: 416: 402: 401: 395: 394: 386: 385: 377: 376: 375: 374: 373: 372: 364: 362:Dimorphodontia 356: 345: 326: 325: 324: 314: 296: 288: 277: 276: 272: 271: 256: 252: 251: 243: 236: 235: 230: 223: 222: 217: 213: 212: 207: 203: 202: 197: 193: 192: 179: 178: 163:Quetzalcoatlus 139: 138: 130: 129: 121: 116: 111: 106: 101: 96: 91: 86: 81: 76: 71: 66: 48: 26: 9: 6: 4: 3: 2: 17763: 17752: 17749: 17747: 17744: 17742: 17739: 17737: 17734: 17733: 17731: 17714: 17709: 17705: 17701: 17696: 17692: 17688: 17683: 17679: 17675: 17670: 17666: 17662: 17656: 17651: 17645: 17641: 17636: 17630: 17626: 17625: 17623: 17621: 17617: 17613: 17608: 17596: 17593: 17591: 17588: 17586: 17583: 17581: 17578: 17576: 17573: 17571: 17568: 17566: 17563: 17561: 17558: 17557: 17555: 17551: 17545: 17542: 17540: 17537: 17535: 17532: 17530: 17527: 17525: 17522: 17520: 17517: 17515: 17512: 17510: 17507: 17505: 17502: 17500: 17497: 17496: 17494: 17490: 17484: 17481: 17479: 17476: 17474: 17471: 17467: 17464: 17462: 17459: 17457: 17454: 17453: 17452: 17449: 17447: 17444: 17442: 17439: 17438: 17436: 17434: 17430: 17424: 17421: 17419: 17416: 17412: 17409: 17408: 17407: 17404: 17402: 17399: 17397: 17394: 17390: 17387: 17385: 17382: 17380: 17377: 17375: 17372: 17370: 17367: 17365: 17362: 17360: 17357: 17355: 17352: 17351: 17350: 17347: 17345: 17342: 17341: 17339: 17337: 17333: 17328: 17318: 17315: 17313: 17312:Pectoral fins 17310: 17308: 17305: 17303: 17300: 17298: 17295: 17293: 17290: 17288: 17285: 17283: 17280: 17278: 17275: 17273: 17270: 17268: 17265: 17263: 17260: 17259: 17257: 17255: 17251: 17247: 17243: 17239: 17232: 17227: 17225: 17220: 17218: 17213: 17212: 17209: 17194: 17190: 17188: 17181: 17134: 17133: 17132:Maaradactylus 17129: 17127: 17126: 17122: 17120: 17119: 17115: 17113: 17112: 17108: 17105: 17104: 17103:Cearadactylus 17100: 17097: 17096: 17092: 17090: 17089: 17085: 17084: 17082: 17080: 17079:Anhanguerinae 17076: 17070: 17069: 17065: 17062: 17061: 17060:Siroccopteryx 17057: 17055: 17054: 17050: 17048: 17047: 17043: 17041: 17040: 17036: 17035: 17033: 17030: 17026: 17020: 17019: 17018:Tropeognathus 17015: 17013: 17012: 17008: 17005: 17004: 17003:Siroccopteryx 17000: 16997: 16996: 16992: 16989: 16988: 16984: 16981: 16980: 16976: 16973: 16972: 16968: 16967: 16965: 16962: 16958: 16955: 16953: 16952:Anhangueridae 16949: 16943: 16942: 16941:Iberodactylus 16938: 16936: 16935: 16931: 16930: 16928: 16925: 16921: 16914: 16913: 16909: 16906: 16905: 16901: 16900: 16897: 16894: 16891: 16886: 16872: 16871: 16867: 16864: 16863: 16859: 16856: 16855: 16851: 16850: 16848: 16846: 16842: 16836: 16835: 16834:Cimoliopterus 16831: 16828: 16827: 16826:Camposipterus 16823: 16821: 16820: 16816: 16815: 16813: 16811: 16807: 16804: 16801: 16797: 16786: 16785: 16781: 16778: 16777: 16776:Siroccopteryx 16773: 16771: 16770: 16766: 16763: 16762: 16758: 16755: 16754: 16750: 16747: 16746: 16742: 16739: 16738: 16734: 16731: 16730: 16726: 16723: 16722: 16718: 16715: 16714: 16713:Camposipterus 16710: 16707: 16706: 16702: 16701: 16699: 16697: 16693: 16686: 16685: 16684:Tropeognathus 16681: 16678: 16677: 16673: 16670: 16669: 16668:Cimoliopterus 16665: 16662: 16661: 16657: 16654: 16653: 16652:Camposipterus 16649: 16647: 16646: 16642: 16640: 16639: 16638:Araripesaurus 16635: 16634: 16631: 16628: 16626: 16622: 16618: 16615: 16612: 16607: 16597: 16596: 16592: 16590: 16589: 16585: 16584: 16582: 16580: 16579:Boreopteridae 16576: 16572: 16566: 16565: 16561: 16559: 16558: 16557:Cearadactylus 16554: 16552: 16551: 16547: 16545: 16544: 16540: 16539: 16537: 16535: 16531: 16517: 16516: 16512: 16510: 16509: 16508:Istiodactylus 16505: 16504: 16502: 16500: 16496: 16490: 16489: 16485: 16483: 16482: 16478: 16476: 16475: 16471: 16469: 16468: 16464: 16463: 16460: 16457: 16455: 16451: 16445: 16444: 16440: 16438: 16437: 16433: 16431: 16430: 16426: 16425: 16423: 16421: 16417: 16410: 16409: 16405: 16403: 16402: 16401:Yixianopterus 16398: 16396: 16395: 16391: 16389: 16388: 16384: 16383: 16380: 16377: 16375: 16371: 16365: 16364: 16360: 16358: 16357: 16353: 16352: 16350: 16348: 16344: 16340: 16333: 16332: 16328: 16327: 16325: 16322: 16317: 16306: 16305: 16301: 16298: 16297: 16293: 16291: 16290: 16286: 16283: 16282: 16278: 16275: 16274: 16270: 16269: 16267: 16265: 16261: 16255: 16254: 16250: 16248: 16247: 16243: 16241: 16240: 16236: 16235: 16232: 16228: 16225: 16222: 16217: 16194: 16193: 16189: 16187: 16186: 16182: 16180: 16179: 16175: 16173: 16172: 16168: 16167: 16165: 16163: 16162:Nyctosauridae 16159: 16153: 16152: 16148: 16146: 16145: 16141: 16139: 16138: 16134: 16133: 16130: 16127: 16125: 16121: 16114: 16113: 16109: 16107: 16106: 16105:Alamodactylus 16102: 16101: 16098: 16095: 16093: 16089: 16082: 16081: 16077: 16074: 16073: 16069: 16067: 16066: 16062: 16059: 16058: 16054: 16052: 16051: 16047: 16044: 16043: 16039: 16038: 16036: 16034: 16030: 16027: 16025: 16024:Pteranodontia 16021: 16015: 16014: 16010: 16009: 16006: 16002: 15999: 15996: 15991: 15987: 15983: 15978: 15974: 15964: 15959: 15956: 15948: 15922: 15921: 15920: 15919: 15917: 15915: 15911: 15907: 15869: 15868: 15864: 15861: 15860: 15859:Wellnhopterus 15856: 15854: 15853: 15849: 15847: 15846: 15842: 15840: 15839: 15835: 15833: 15832: 15831:Hatzegopteryx 15828: 15826: 15825: 15821: 15819: 15818: 15814: 15812: 15811: 15807: 15805: 15804: 15803:Aralazhdarcho 15800: 15799: 15797: 15795: 15791: 15785: 15784: 15780: 15778: 15777: 15773: 15771: 15770: 15766: 15763: 15762: 15758: 15757: 15755: 15753: 15749: 15742: 15741: 15737: 15734: 15733: 15729: 15727: 15726: 15722: 15719: 15718: 15714: 15712: 15711: 15707: 15704: 15703: 15699: 15698: 15696: 15694: 15690: 15683: 15682: 15678: 15676: 15675: 15674:Radiodactylus 15671: 15668: 15667: 15663: 15662: 15659: 15656: 15654: 15650: 15640: 15639: 15635: 15633: 15632: 15628: 15626: 15625: 15621: 15620: 15618: 15616: 15612: 15605: 15604: 15600: 15597: 15596: 15592: 15590: 15589: 15585: 15583: 15582: 15578: 15575: 15574: 15570: 15567: 15566: 15562: 15559: 15558: 15557:Apatorhamphus 15554: 15553: 15550: 15547: 15545: 15541: 15535: 15534: 15530: 15529: 15526: 15523: 15521: 15517: 15510: 15509: 15505: 15502: 15501: 15497: 15494: 15493: 15489: 15486: 15485: 15481: 15478: 15477: 15473: 15471: 15470: 15466: 15465: 15463: 15460: 15456: 15449: 15448: 15444: 15442: 15441: 15437: 15435: 15434: 15430: 15427: 15426: 15422: 15421: 15418: 15415: 15412: 15407: 15377: 15376: 15372: 15370: 15369: 15365: 15364: 15362: 15360: 15356: 15350: 15349: 15348:Tupandactylus 15345: 15343: 15342: 15338: 15336: 15335: 15331: 15328: 15327: 15323: 15322: 15319: 15316: 15314: 15310: 15304: 15303: 15299: 15296: 15295: 15291: 15288: 15287: 15283: 15280: 15279: 15278:Caupedactylus 15275: 15272: 15271: 15267: 15266: 15263: 15260: 15258: 15254: 15248: 15247: 15243: 15241: 15240: 15236: 15234: 15233: 15229: 15227: 15226: 15222: 15220: 15219: 15215: 15212: 15211: 15207: 15204: 15203: 15199: 15196: 15195: 15191: 15190: 15188: 15186: 15182: 15176: 15175: 15171: 15170: 15167: 15164: 15162: 15158: 15152: 15151: 15150:Caupedactylus 15147: 15145: 15144: 15140: 15139: 15137: 15134: 15133:Caupedactylia 15130: 15127: 15125: 15121: 15114: 15113: 15109: 15107: 15106: 15102: 15100: 15099: 15095: 15092: 15091: 15087: 15084: 15083: 15079: 15077: 15076: 15072: 15069: 15068: 15064: 15061: 15060: 15056: 15053: 15052: 15048: 15045: 15044: 15040: 15039: 15037: 15034: 15030: 15023: 15022: 15018: 15016: 15015: 15011: 15010: 15007: 15004: 15001: 14996: 14993: 14990: 14989:Neoazhdarchia 14985: 14979: 14978: 14974: 14972: 14971: 14967: 14964: 14963: 14959: 14956: 14955: 14951: 14949: 14948: 14944: 14942: 14941: 14937: 14936: 14933: 14930: 14927: 14926:Azhdarchoidea 14922: 14908: 14907: 14903: 14901: 14900: 14896: 14895: 14893: 14891: 14887: 14880: 14879: 14875: 14872: 14871: 14870:Puntanipterus 14867: 14865: 14864: 14860: 14858: 14857: 14853: 14850: 14849: 14845: 14842: 14841: 14837: 14836: 14833: 14830: 14828: 14824: 14820: 14817: 14814: 14809: 14802: 14801: 14797: 14796: 14793: 14790: 14787: 14782: 14778: 14774: 14769: 14765: 14756: 14750: 14724: 14723: 14722: 14721: 14719: 14717: 14713: 14706: 14705: 14701: 14700: 14697: 14694: 14692: 14688: 14684: 14658: 14657: 14653: 14651: 14650: 14646: 14644: 14643: 14639: 14637: 14636: 14635:Beipiaopterus 14632: 14631: 14629: 14627: 14623: 14617: 14616: 14612: 14611: 14608: 14605: 14603: 14599: 14593: 14592: 14591:Tacuadactylus 14588: 14586: 14585: 14581: 14579: 14578: 14574: 14571: 14570: 14566: 14564: 14563: 14559: 14558: 14556: 14554: 14550: 14544: 14543: 14539: 14537: 14536: 14532: 14531: 14529: 14526: 14522: 14516: 14515: 14511: 14509: 14508: 14504: 14502: 14501: 14497: 14495: 14494: 14490: 14488: 14487: 14483: 14481: 14480: 14476: 14474: 14473: 14472:Forfexopterus 14469: 14467: 14466: 14465:Elanodactylus 14462: 14460: 14459: 14458:Cratonopterus 14455: 14453: 14452: 14451:Cathayopterus 14448: 14446: 14445: 14441: 14438: 14437: 14436:Ardeadactylus 14433: 14432: 14429: 14426: 14423: 14418: 14403: 14402: 14398: 14396: 14395: 14391: 14388: 14387: 14386:Ardeadactylus 14383: 14382: 14380: 14378: 14374: 14367: 14366: 14362: 14360: 14359: 14355: 14354: 14351: 14348: 14346: 14342: 14335: 14334: 14330: 14327: 14326: 14322: 14319: 14318: 14314: 14312: 14311: 14310:Cycnorhamphus 14307: 14304: 14303: 14299: 14298: 14296: 14294: 14290: 14286: 14283: 14280: 14275: 14269: 14268: 14267:Pterodactylus 14264: 14262: 14261: 14260:Diopecephalus 14257: 14256: 14253: 14250: 14247: 14242: 14231: 14230: 14226: 14223: 14222: 14218: 14216: 14215: 14211: 14208: 14207: 14203: 14202: 14200: 14198: 14194: 14188: 14187: 14186:Prejanopterus 14183: 14182: 14179: 14175: 14172: 14169: 14164: 14160: 14156: 14151: 14147: 14138: 14133: 14096: 14095: 14094: 14093: 14091: 14089: 14085: 14079: 14078: 14074: 14071: 14070: 14066: 14064: 14063: 14059: 14057: 14056: 14052: 14050: 14049: 14045: 14043: 14042: 14038: 14035: 14034: 14033:Herbstosaurus 14030: 14028: 14027: 14026:Eurolimnornis 14023: 14021: 14020: 14019:Dermodactylus 14016: 14015: 14012: 14009: 14007: 14003: 13993: 13992: 13988: 13985: 13984: 13980: 13977: 13976: 13972: 13970: 13969: 13965: 13963: 13962: 13958: 13957: 13955: 13953: 13949: 13943: 13942: 13938: 13936: 13935: 13931: 13928: 13927: 13923: 13920: 13919: 13915: 13913: 13912: 13908: 13907: 13905: 13903: 13899: 13893: 13892: 13888: 13887: 13884: 13881: 13879: 13875: 13872: 13869: 13868:Caelidracones 13864: 13858: 13857: 13853: 13850: 13849: 13845: 13844: 13841: 13838: 13835: 13830: 13812: 13811: 13810:Wukongopterus 13807: 13804: 13803: 13799: 13797: 13796: 13795:Darwinopterus 13792: 13790: 13789: 13788:Cuspicephalus 13785: 13782: 13781: 13777: 13776: 13774: 13772: 13768: 13762: 13761: 13757: 13756: 13753: 13750: 13748: 13744: 13738: 13737: 13736:Pterorhynchus 13733: 13731: 13730: 13726: 13725: 13722: 13719: 13717: 13713: 13706: 13705: 13701: 13699: 13698: 13694: 13692: 13691: 13687: 13686: 13683: 13679: 13676: 13673: 13668: 13661: 13660: 13656: 13653: 13652: 13648: 13647: 13644: 13641: 13638: 13633: 13629: 13625: 13620: 13616: 13607: 13603: 13600: 13592: 13546: 13545: 13544: 13543: 13541: 13539: 13535: 13528: 13527: 13523: 13521: 13520: 13519:Scaphognathus 13516: 13514: 13513: 13509: 13507: 13506: 13502: 13500: 13499: 13495: 13494: 13492: 13489: 13485: 13482: 13479: 13475: 13457: 13456: 13452: 13449: 13448: 13444: 13441: 13440: 13436: 13435: 13433: 13431: 13427: 13421: 13420: 13416: 13413: 13412: 13408: 13405: 13404: 13400: 13398: 13397: 13393: 13391: 13390: 13386: 13385: 13383: 13381: 13377: 13374: 13372: 13368: 13362: 13361: 13357: 13355: 13354: 13350: 13348: 13347: 13343: 13341: 13340: 13336: 13334: 13333: 13329: 13327: 13326: 13322: 13320: 13319: 13315: 13314: 13311: 13308: 13306: 13302: 13295: 13294: 13290: 13287: 13286: 13282: 13280: 13279: 13275: 13272: 13271: 13267: 13266: 13263: 13260: 13258: 13254: 13251: 13248: 13243: 13233: 13232: 13228: 13226: 13225: 13221: 13220: 13218: 13216: 13212: 13208: 13205: 13202: 13197: 13190: 13189: 13188:Eudimorphodon 13185: 13184: 13181: 13178: 13174: 13169: 13158: 13157: 13153: 13150: 13149: 13148:Peteinosaurus 13145: 13142: 13141: 13137: 13135: 13134: 13130: 13128: 13127: 13123: 13120: 13119: 13115: 13114: 13112: 13110: 13106: 13100: 13099: 13098:Herbstosaurus 13095: 13094: 13091: 13087: 13084: 13081: 13076: 13069: 13068: 13067:Peteinosaurus 13064: 13063: 13060: 13057: 13053: 13048: 13030: 13029: 13028:Eudimorphodon 13025: 13022: 13021: 13017: 13016: 13014: 13012: 13008: 13001: 13000: 12996: 12995: 12992: 12989: 12987: 12983: 12977: 12976: 12972: 12970: 12969: 12965: 12963: 12962: 12958: 12955: 12954: 12950: 12949: 12947: 12945: 12941: 12938: 12936: 12932: 12925: 12924: 12923:Preondactylia 12920: 12917: 12916: 12915:Peteinosaurus 12912: 12909: 12908: 12904: 12903: 12900: 12897: 12894: 12893:Eopterosauria 12889: 12875: 12874: 12870: 12868: 12867: 12863: 12860: 12859: 12855: 12854: 12852: 12850: 12846: 12839: 12838: 12834: 12832: 12831: 12827: 12824: 12823: 12819: 12818: 12815: 12812: 12809: 12805: 12799: 12798: 12797:Preondactylus 12794: 12792: 12791: 12787: 12786: 12784: 12782: 12781:Preondactylia 12778: 12772: 12771: 12767: 12766: 12763: 12759: 12756: 12753: 12748: 12744: 12740: 12735: 12731: 12710: 12709: 12708: 12707: 12705: 12703: 12699: 12691: 12687: 12686: 12685: 12684: 12681: 12678: 12676: 12672: 12668: 12660: 12656: 12653: 12651: 12647: 12644: 12642: 12638: 12635: 12633: 12629: 12627: 12623: 12621: 12617: 12616: 12611: 12607: 12603: 12596: 12591: 12589: 12584: 12582: 12577: 12576: 12573: 12561: 12553: 12551: 12548: 12546: 12542: 12541: 12538: 12529: 12525: 12522: 12516: 12511: 12505: 12469: 12465: 12463: 12459: 12458: 12457: 12456: 12454: 12452: 12448: 12440: 12436: 12435: 12434: 12433: 12431: 12429: 12422: 12419: 12417: 12413: 12403: 12402: 12398: 12396: 12395: 12394:Sanjuansaurus 12391: 12389: 12388: 12387:Herrerasaurus 12384: 12382: 12381: 12377: 12376: 12374: 12372: 12368: 12361: 12360: 12356: 12353: 12352: 12348: 12345: 12344: 12343:Daemonosaurus 12340: 12337: 12336: 12332: 12331: 12328: 12325: 12323: 12322:Herrerasauria 12316: 12310: 12309: 12302: 12299: 12298: 12291: 12289: 12288: 12281: 12279: 12278: 12277:Daemonosaurus 12271: 12269: 12268: 12261: 12259: 12258: 12251: 12250: 12247: 12244: 12241: 12236: 12224: 12220: 12219: 12218: 12217: 12215: 12213: 12206: 12202: 12199: 12196: 12191: 12177: 12176: 12172: 12170: 12169: 12165: 12163: 12162: 12158: 12156: 12155: 12151: 12149: 12148: 12144: 12142: 12141: 12140:Eucoelophysis 12137: 12135: 12134: 12130: 12128: 12127: 12123: 12122: 12120: 12118: 12114: 12107: 12106: 12102: 12100: 12099: 12095: 12092: 12091: 12087: 12085: 12084: 12080: 12078: 12077: 12073: 12071: 12070: 12066: 12064: 12063: 12059: 12058: 12055: 12052: 12049: 12042: 12036: 12035: 12028: 12025: 12024: 12017: 12015: 12014: 12007: 12005: 12004: 11997: 11996: 11993: 11989: 11986: 11983: 11978: 11962: 11958: 11957: 11956: 11955: 11953: 11951: 11947: 11941: 11940: 11936: 11933: 11932: 11931:Scleromochlus 11928: 11926: 11925: 11921: 11919: 11918: 11914: 11912: 11911: 11907: 11904: 11903: 11899: 11897: 11896: 11892: 11891: 11889: 11887: 11883: 11880: 11878: 11871: 11867: 11864: 11861: 11856: 11846: 11845: 11841: 11839: 11838: 11834: 11832: 11831: 11827: 11825: 11824: 11820: 11817: 11816: 11812: 11811: 11809: 11807: 11800: 11794: 11793: 11786: 11783: 11782: 11775: 11774: 11771: 11767: 11764: 11761: 11756: 11752: 11740: 11739: 11733: 11716: 11715: 11714: 11713: 11711: 11709: 11705: 11697: 11693: 11692: 11691: 11690: 11687: 11684: 11682: 11678: 11670: 11666: 11665: 11664: 11663: 11660: 11657: 11655: 11651: 11647: 11639: 11635: 11632: 11630: 11626: 11624: 11620: 11618: 11614: 11613: 11608: 11604: 11600: 11593: 11588: 11586: 11581: 11579: 11574: 11573: 11570: 11563: 11560: 11557: 11554: 11551: 11550:Pterosaur.net 11548: 11547: 11537: 11531: 11527: 11522: 11518: 11512: 11508: 11503: 11499: 11493: 11489: 11484: 11483: 11470: 11466: 11461: 11456: 11452: 11448: 11444: 11440: 11439: 11434: 11427: 11419: 11412: 11397: 11396: 11388: 11381: 11366: 11365: 11360: 11353: 11346: 11343: 11339: 11333: 11331: 11323: 11318: 11311: 11310:1-86239-143-2 11307: 11303: 11300: 11296: 11289: 11282: 11281:Pterosaur.net 11279: 11273: 11271: 11269: 11260: 11256: 11252: 11248: 11244: 11240: 11236: 11232: 11228: 11224: 11217: 11203: 11199: 11192: 11184: 11180: 11175: 11170: 11166: 11162: 11158: 11154: 11150: 11146: 11142: 11135: 11127: 11123: 11118: 11113: 11109: 11105: 11101: 11097: 11093: 11089: 11085: 11078: 11070: 11066: 11061: 11056: 11052: 11048: 11044: 11040: 11036: 11029: 11015: 11011: 11005: 10997: 10993: 10988: 10983: 10978: 10973: 10969: 10965: 10962:(2): e31392. 10961: 10957: 10953: 10951: 10942: 10940: 10931: 10927: 10923: 10919: 10915: 10911: 10907: 10903: 10900:(3): 569–80. 10899: 10895: 10891: 10884: 10882: 10880: 10865: 10861: 10855: 10853: 10844: 10840: 10836: 10832: 10827: 10822: 10818: 10814: 10811:(6992): 621. 10810: 10806: 10802: 10795: 10787: 10783: 10778: 10773: 10769: 10765: 10761: 10754: 10746: 10742: 10738: 10734: 10730: 10726: 10723:(4): 273–77. 10722: 10718: 10711: 10703: 10699: 10694: 10689: 10685: 10681: 10677: 10673: 10669: 10662: 10660: 10651: 10647: 10643: 10639: 10635: 10631: 10627: 10623: 10619: 10615: 10608: 10600: 10596: 10591: 10586: 10582: 10578: 10574: 10570: 10566: 10562: 10558: 10551: 10543: 10539: 10535: 10531: 10526: 10521: 10517: 10513: 10509: 10505: 10501: 10494: 10486: 10482: 10478: 10471: 10463: 10459: 10455: 10451: 10447: 10443: 10439: 10435: 10432:(7017): 572. 10431: 10427: 10420: 10413: 10405: 10401: 10397: 10391: 10383: 10379: 10375: 10371: 10366: 10361: 10357: 10353: 10349: 10345: 10341: 10334: 10326: 10322: 10318: 10314: 10307: 10305: 10303: 10294: 10290: 10286: 10282: 10278: 10274: 10270: 10263: 10255: 10251: 10250: 10242: 10234: 10230: 10225: 10220: 10215: 10210: 10206: 10202: 10198: 10191: 10182: 10177: 10173: 10169: 10165: 10158: 10151: 10145: 10137: 10133: 10128: 10123: 10118: 10113: 10109: 10105: 10101: 10097: 10093: 10091: 10082: 10075: 10070: 10063: 10058: 10050: 10046: 10042: 10038: 10034: 10030: 10026: 10022: 10018: 10014: 10010: 10003: 9995: 9991: 9987: 9983: 9979: 9975: 9972:(3): 507–14. 9971: 9967: 9960: 9953: 9948: 9940: 9936: 9932: 9928: 9924: 9920: 9917:(1): 163–87. 9916: 9912: 9905: 9898: 9891: 9886: 9879: 9874: 9867: 9862: 9854: 9850: 9846: 9842: 9838: 9834: 9831:(4): 376–98. 9830: 9826: 9822: 9815: 9808: 9803: 9801: 9799: 9790: 9786: 9783:(4): 373–86. 9782: 9778: 9771: 9764: 9756: 9752: 9748: 9744: 9740: 9736: 9733:(4): 421–35. 9732: 9728: 9724: 9717: 9709: 9705: 9701: 9697: 9693: 9689: 9685: 9681: 9674: 9667: 9659: 9655: 9650: 9645: 9640: 9635: 9631: 9627: 9623: 9616: 9608: 9604: 9597: 9589: 9585: 9581: 9577: 9570: 9562: 9558: 9554: 9550: 9545: 9540: 9536: 9532: 9528: 9521: 9506: 9499: 9493: 9485: 9481: 9474: 9466: 9462: 9455: 9448: 9433: 9429: 9422: 9420: 9403: 9399: 9393: 9391: 9389: 9372: 9368: 9367: 9362: 9358: 9357:Naish, Darren 9352: 9344: 9338: 9334: 9333: 9325: 9310: 9306: 9299: 9283: 9279: 9278:The Telegraph 9275: 9268: 9266: 9257: 9253: 9249: 9245: 9241: 9237: 9233: 9229: 9226:(9): 891–97. 9225: 9221: 9214: 9207: 9203: 9199: 9193: 9185: 9183:9780429821202 9179: 9175: 9174: 9166: 9160: 9156: 9152: 9146: 9138: 9134: 9130: 9126: 9122: 9118: 9115:(1): 105–37. 9114: 9110: 9103: 9095: 9091: 9087: 9083: 9079: 9075: 9071: 9064: 9062: 9053: 9049: 9042: 9034: 9030: 9025: 9020: 9016: 9012: 9008: 9004: 9001:(8): 160333. 9000: 8996: 8992: 8985: 8977: 8973: 8969: 8968:10.1666/13030 8965: 8961: 8957: 8953: 8949: 8942: 8935: 8921:on 2013-01-15 8917: 8913: 8909: 8905: 8901: 8898:(4): 883–94. 8897: 8893: 8892:Geodiversitas 8886: 8884: 8875: 8867: 8863: 8859: 8855: 8851: 8847: 8843: 8839: 8832: 8824: 8820: 8815: 8810: 8805: 8800: 8796: 8792: 8788: 8781: 8779: 8777: 8768: 8764: 8760: 8756: 8752: 8748: 8745:(3): 432–46. 8744: 8740: 8732: 8724: 8720: 8715: 8710: 8706: 8702: 8698: 8691: 8684: 8678: 8670: 8666: 8661: 8656: 8651: 8646: 8642: 8638: 8634: 8627: 8625: 8623: 8621: 8613: 8607: 8601:, p. 18. 8600: 8595: 8587: 8583: 8579: 8575: 8571: 8567: 8560: 8546: 8542: 8535: 8521: 8517: 8511: 8503: 8499: 8495: 8491: 8487: 8483: 8479: 8475: 8471: 8467: 8460: 8452: 8450: 8441: 8437: 8432: 8427: 8422: 8417: 8413: 8409: 8405: 8403: 8394: 8386: 8382: 8377: 8372: 8367: 8362: 8358: 8354: 8350: 8343: 8335: 8331: 8327: 8323: 8318: 8317:10.1206/352.1 8313: 8309: 8305: 8301: 8294: 8286: 8282: 8278: 8274: 8271:(4): 465–69. 8270: 8266: 8259: 8257: 8248: 8244: 8240: 8236: 8232: 8228: 8224: 8220: 8216: 8212: 8205: 8198: 8189: 8184: 8180: 8176: 8172: 8165: 8156: 8151: 8147: 8143: 8139: 8135: 8131: 8129: 8120: 8113: 8108: 8102:, p. 13. 8101: 8096: 8088: 8084: 8078: 8072:, p. 10. 8071: 8066: 8064: 8057: 8052: 8050: 8041: 8037: 8033: 8029: 8024: 8019: 8016:(4): 890–98. 8015: 8011: 8007: 8000: 7993: 7988: 7981: 7975: 7968: 7963: 7961: 7959: 7957: 7955: 7946: 7942: 7938: 7934: 7930: 7926: 7922: 7918: 7915:(3): 218–39. 7914: 7910: 7903: 7901: 7893: 7887: 7879: 7875: 7868: 7861: 7855: 7848: 7842: 7828:on 2020-07-28 7824: 7820: 7816: 7812: 7808: 7804: 7800: 7799: 7791: 7784: 7777: 7771: 7763: 7759: 7752: 7744: 7740: 7733: 7726: 7721: 7715:, p. 31. 7714: 7709: 7707: 7700:, p. 36. 7699: 7694: 7692: 7690: 7683:, p. 35. 7682: 7677: 7675: 7666: 7662: 7658: 7654: 7650: 7646: 7642: 7638: 7631: 7624: 7618: 7611: 7605: 7599:, p. 33. 7598: 7593: 7587:, p. 29. 7586: 7581: 7575:, p. 28. 7574: 7569: 7561: 7557: 7556:Isis von Oken 7553: 7546: 7538: 7534: 7527: 7521:, p. 27. 7520: 7515: 7508: 7504: 7497: 7490: 7484: 7477: 7473: 7466: 7458: 7454: 7447: 7440: 7434: 7427: 7421: 7414: 7409: 7407: 7405: 7403: 7395: 7390: 7383: 7378: 7372:, p. 22. 7371: 7366: 7359: 7354: 7352: 7343: 7339: 7334: 7329: 7325: 7321: 7317: 7313: 7309: 7305: 7301: 7294: 7286: 7282: 7278: 7274: 7269: 7264: 7260: 7256: 7252: 7245: 7237: 7233: 7229: 7225: 7220: 7215: 7211: 7207: 7203: 7199: 7195: 7191: 7183: 7175: 7171: 7167: 7163: 7159: 7155: 7151: 7147: 7140: 7132: 7128: 7123: 7118: 7114: 7110: 7106: 7102: 7098: 7091: 7083: 7079: 7075: 7071: 7067: 7063: 7059: 7055: 7051: 7047: 7040: 7026: 7025: 7020: 7013: 7005: 7001: 6997: 6993: 6988: 6983: 6979: 6975: 6971: 6967: 6963: 6956: 6948: 6944: 6937: 6931:, p. 51. 6930: 6925: 6923: 6921: 6919: 6917: 6910:, p. 48. 6909: 6904: 6898:, p. 47. 6897: 6892: 6886:, p. 43. 6885: 6880: 6874:, p. 39. 6873: 6868: 6862:, p. 37. 6861: 6856: 6848: 6844: 6839: 6834: 6829: 6824: 6820: 6816: 6812: 6808: 6804: 6797: 6795: 6793: 6791: 6784:, p. 36. 6783: 6778: 6776: 6774: 6767:, p. 57. 6766: 6761: 6759: 6757: 6755: 6753: 6751: 6744:, p. 56. 6743: 6738: 6736: 6734: 6732: 6730: 6728: 6719: 6715: 6711: 6707: 6702: 6697: 6693: 6689: 6684: 6679: 6675: 6671: 6667: 6663: 6659: 6652: 6645: 6640: 6634:, p. 55. 6633: 6628: 6626: 6624: 6622: 6615:, p. 35. 6614: 6609: 6607: 6605: 6603: 6601: 6599: 6597: 6595: 6586: 6582: 6577: 6572: 6568: 6564: 6560: 6556: 6555: 6550: 6543: 6535: 6531: 6526: 6521: 6517: 6513: 6509: 6502: 6494: 6490: 6487:(3): 265–75. 6486: 6482: 6475: 6467: 6463: 6459: 6455: 6452:(4): 881–91. 6451: 6447: 6440: 6433: 6425: 6421: 6416: 6411: 6407: 6403: 6399: 6395: 6394: 6389: 6382: 6376:, p. 34. 6375: 6370: 6364:, p. 33. 6363: 6358: 6352:, p. 53. 6351: 6346: 6344: 6342: 6333: 6329: 6325: 6321: 6316: 6311: 6307: 6303: 6302:J. Evol. Biol 6299: 6292: 6284: 6280: 6276: 6272: 6268: 6264: 6261:(1): 233–66. 6260: 6256: 6249: 6241: 6237: 6233: 6229: 6225: 6221: 6217: 6213: 6206: 6198: 6194: 6190: 6186: 6182: 6178: 6174: 6170: 6166: 6159: 6157: 6150:, p. 55. 6149: 6144: 6142: 6135:, p. 52. 6134: 6129: 6127: 6118: 6114: 6109: 6104: 6099: 6094: 6090: 6086: 6082: 6078: 6074: 6067: 6065: 6056: 6052: 6047: 6042: 6038: 6034: 6030: 6026: 6022: 6020: 6011: 6009: 6007: 5998: 5994: 5990: 5986: 5982: 5978: 5975:(4): 255–84. 5974: 5970: 5963: 5957:, p. 53. 5956: 5951: 5949: 5942:, p. 54. 5941: 5936: 5930:, p. 32. 5929: 5924: 5922: 5920: 5913:, p. 44. 5912: 5907: 5901:, p. 52. 5900: 5895: 5893: 5891: 5889: 5887: 5880:, p. 51. 5879: 5874: 5872: 5865:, p. 31. 5864: 5859: 5857: 5855: 5848:, p. 30. 5847: 5842: 5840: 5838: 5831:, p. 46. 5830: 5825: 5823: 5821: 5819: 5812:, p. 45. 5811: 5806: 5800:, p. 50. 5799: 5794: 5792: 5790: 5783:, p. 28. 5782: 5777: 5775: 5766: 5762: 5758: 5756: 5747: 5741:, p. 49. 5740: 5735: 5728: 5727:1-932075-01-1 5724: 5720: 5714: 5712: 5703: 5699: 5692: 5690: 5688: 5686: 5679:, p. 48. 5678: 5673: 5665: 5661: 5658:(3): 421–41. 5657: 5653: 5649: 5642: 5640: 5633:, p. 24. 5632: 5627: 5625: 5623: 5616:, p. 26. 5615: 5610: 5608: 5606: 5599:, p. 47. 5598: 5593: 5591: 5584:, p. 27. 5583: 5578: 5576: 5567: 5561: 5557: 5550: 5548: 5546: 5544: 5542: 5540: 5538: 5531:, p. 23. 5530: 5525: 5523: 5521: 5514:, p. 58. 5513: 5508: 5500: 5496: 5489: 5481: 5477: 5473: 5469: 5465: 5461: 5457: 5453: 5446: 5439: 5425: 5424:pterosaur.net 5421: 5415: 5407: 5403: 5399: 5395: 5391: 5387: 5383: 5379: 5375: 5371: 5364: 5357: 5355: 5353: 5351: 5342: 5336: 5332: 5331: 5323: 5321: 5304: 5297: 5295: 5293: 5284: 5278: 5274: 5269: 5268: 5262: 5256: 5248: 5244: 5240: 5236: 5232: 5228: 5224: 5220: 5217:(4): 180–84. 5216: 5212: 5205: 5198: 5190: 5186: 5182: 5178: 5174: 5170: 5166: 5162: 5158: 5154: 5150: 5144: 5136: 5132: 5127: 5122: 5117: 5112: 5108: 5104: 5100: 5096: 5092: 5085: 5083: 5074: 5068: 5054: 5050: 5043: 5029: 5028:pterosaur.net 5025: 5019: 5010: 5005: 5002:(1): 99–111. 5001: 4997: 4993: 4986: 4978: 4971: 4965: 4957: 4956: 4948: 4940: 4939: 4934: 4928: 4921: 4915: 4911: 4907: 4906:Jones, Daniel 4901: 4893: 4889: 4885: 4884:Darwinopterus 4878: 4871: 4865: 4861: 4857: 4853: 4846: 4838: 4834: 4829: 4824: 4819: 4814: 4810: 4806: 4802: 4795: 4787: 4783: 4778: 4773: 4769: 4765: 4761: 4757: 4753: 4746: 4744: 4739: 4726: 4720: 4703: 4697: 4693: 4687: 4681: 4680: 4667: 4624: 4615: 4611: 4601: 4598: 4596: 4593: 4591: 4588: 4586: 4583: 4581: 4578: 4576: 4573: 4571: 4568: 4566: 4563: 4562: 4558: 4547: 4544: 4538: 4533: 4530: 4519: 4516: 4505: 4498: 4495: 4494: 4490: 4489:wukongopterid 4486: 4485: 4481: 4477: 4476:birds of prey 4473: 4468: 4466: 4465: 4460: 4456: 4455: 4449: 4447: 4446: 4441: 4439: 4434: 4430: 4429: 4424: 4423: 4418: 4414: 4410: 4409: 4405:, and 1966's 4404: 4403: 4398: 4394: 4393: 4387: 4385: 4381: 4377: 4376: 4371: 4367: 4363: 4362: 4357: 4353: 4352: 4347: 4346: 4338: 4337: 4332: 4331: 4325: 4321: 4314: 4310: 4306: 4302: 4295: 4293: 4289: 4288:Pterodactylus 4285: 4284:Scaphognathus 4281: 4277: 4273: 4269: 4265: 4261: 4257: 4253: 4249: 4248: 4243: 4239: 4238: 4233: 4232: 4227: 4226: 4221: 4217: 4216: 4211: 4210: 4209:Scaphognathus 4205: 4204: 4203:Pterodactylus 4199: 4198:scleral rings 4189: 4186: 4184: 4180: 4179: 4174: 4170: 4166: 4162: 4158: 4153: 4151: 4147: 4142: 4138: 4134: 4130: 4127: 4123: 4117: 4115: 4111: 4107: 4103: 4102:Pterodactylus 4099: 4094: 4090: 4085: 4083: 4079: 4078:Darwinopterus 4074: 4072: 4068: 4064: 4059: 4058: 4053: 4052: 4051:Darwinopterus 4047: 4046: 4041: 4037: 4033: 4029: 4020: 4015: 4006: 4004: 4003: 3998: 3994: 3993: 3988: 3987: 3982: 3978: 3977: 3972: 3968: 3964: 3960: 3959: 3954: 3944: 3942: 3941: 3936: 3935: 3930: 3929: 3924: 3920: 3919: 3914: 3910: 3909: 3904: 3903: 3898: 3897: 3892: 3888: 3884: 3880: 3876: 3871: 3869: 3868: 3863: 3862: 3861:Hatzegopteryx 3857: 3853: 3849: 3844: 3843: 3838: 3834: 3830: 3826: 3825:Azhdarchoidea 3823:In contrast, 3821: 3819: 3815: 3814: 3809: 3805: 3804: 3803:Pterodactylus 3799: 3798: 3793: 3789: 3785: 3781: 3777: 3776: 3771: 3767: 3763: 3759: 3758: 3753: 3749: 3745: 3741: 3740:Boreopteridae 3737: 3733: 3732:Pteranodontia 3728: 3726: 3722: 3721: 3720:Darwinopterus 3716: 3712: 3708: 3704: 3703: 3698: 3697: 3696:Scaphognathus 3692: 3691: 3686: 3685: 3680: 3679: 3674: 3669: 3667: 3663: 3662: 3657: 3656: 3651: 3650:Eudimorphodon 3647: 3646: 3645:Eudimorphodon 3641: 3640: 3634: 3633: 3628: 3624: 3620: 3619: 3613: 3611: 3607: 3602: 3593: 3591: 3587: 3583: 3582: 3578: 3574: 3570: 3566: 3555: 3553: 3552: 3551:Scleromochlus 3547: 3546: 3541: 3540: 3539:Pterodactylus 3535: 3531: 3527: 3526: 3519: 3516: 3511: 3507: 3502: 3500: 3491: 3490: 3489:Hatzegopteryx 3485: 3480: 3476: 3473: 3469: 3465: 3461: 3456: 3454: 3450: 3446: 3445: 3440: 3436: 3432: 3427: 3425: 3417: 3414: 3410: 3407:The probable 3405: 3396: 3394: 3390: 3389: 3384: 3378: 3375: 3371: 3369: 3364: 3360: 3350: 3348: 3344: 3340: 3329: 3327: 3322: 3320: 3319: 3314: 3313: 3308: 3303: 3300: 3296: 3292: 3288: 3281: 3280: 3274: 3270: 3268: 3264: 3260: 3256: 3252: 3248: 3243: 3231: 3223: 3208: 3207: 3199: 3198: 3186: 3180: 3179: 3167: 3161: 3160: 3148: 3142: 3141: 3129: 3123: 3122: 3110: 3104: 3103: 3091: 3090:Caelidracones 3085: 3084: 3076: 3075: 3067: 3066: 3058: 3057: 3049: 3048: 3040: 3039: 3031: 3030: 3022: 3021: 3018: 3016: 3012: 3006: 3005: 3002: 3001: 2998: 2996: 2992: 2991:Pteranodontia 2986: 2985: 2979: 2976: 2975: 2972: 2971: 2968: 2967: 2966: 2959: 2958: 2952: 2951: 2948: 2947: 2939: 2938: 2935: 2933: 2929: 2928:Neoazhdarchia 2923: 2922: 2919: 2918: 2915: 2913: 2909: 2903: 2902: 2896: 2895:Azhdarchoidea 2893: 2892: 2886: 2883: 2882: 2879: 2878: 2875: 2874: 2873: 2866: 2865: 2859: 2856: 2855: 2852: 2851: 2843: 2842: 2834: 2833: 2830: 2828: 2824: 2818: 2817: 2814: 2813: 2810: 2808: 2804: 2803: 2802:Pterodactylus 2796: 2795: 2789: 2786: 2785: 2782: 2781: 2778: 2776: 2772: 2766: 2765: 2759: 2756: 2755: 2749: 2746: 2745: 2742: 2741: 2738: 2736: 2732: 2731: 2724: 2723: 2717: 2714: 2713: 2710: 2709: 2706: 2704: 2700: 2694: 2693: 2687: 2686: 2683: 2682: 2679: 2678: 2677: 2670: 2669: 2663: 2662: 2659: 2658: 2655: 2654: 2648: 2647: 2641: 2640: 2637: 2636: 2633: 2631: 2627: 2626: 2619: 2618: 2612: 2611: 2608: 2607: 2604: 2602: 2598: 2592: 2591: 2585: 2584: 2581: 2580: 2577: 2575: 2571: 2565: 2564: 2558: 2557: 2554: 2553: 2550: 2548: 2544: 2538: 2537: 2531: 2528: 2527: 2524: 2523: 2520: 2518: 2514: 2513:Eopterosauria 2508: 2507: 2501: 2498: 2497: 2492: 2490: 2486: 2482: 2476: 2474: 2470: 2466: 2462: 2458: 2454: 2453:fossil record 2450: 2447:The internal 2445: 2443: 2439: 2438:ornithodirans 2435: 2431: 2427: 2423: 2422:Pterodactylus 2419: 2414: 2410: 2409: 2408:Preondactylus 2403: 2399: 2393: 2383: 2379: 2377: 2373: 2368: 2366: 2362: 2358: 2353: 2351: 2350: 2345: 2341: 2337: 2336: 2331: 2330: 2325: 2324:Maastrichtian 2321: 2316: 2314: 2309: 2305: 2297: 2293: 2288: 2279: 2277: 2276: 2272: 2267: 2265: 2264:Scleromochlus 2260: 2256: 2250: 2248: 2245:also showing 2244: 2240: 2236: 2232: 2228: 2224: 2220: 2217: 2213: 2212:Scleromochlus 2209: 2208: 2207:Scleromochlus 2203: 2199: 2198:Scleromochlus 2195: 2191: 2190: 2185: 2184:Scleromochlus 2181: 2177: 2172: 2164: 2160: 2158: 2157:Sharovipteryx 2152: 2145: 2141: 2139: 2138:Scleromochlus 2133: 2129: 2128: 2124: 2122: 2121:Scleromochlus 2117: 2113: 2112: 2111:Sharovipteryx 2107: 2106: 2101: 2097: 2093: 2089: 2085: 2081: 2077: 2073: 2068: 2064: 2056: 2052: 2048: 2044: 2043: 2037: 2023: 2021: 2017: 2013: 2009: 2005: 2004: 1998: 1996: 1992: 1988: 1983: 1979: 1975: 1970: 1968: 1963: 1959: 1952: 1948: 1944: 1943: 1937: 1933: 1931: 1927: 1923: 1918: 1914: 1913:Robert Bakker 1910: 1909: 1904: 1903: 1898: 1890: 1889: 1883: 1874: 1872: 1868: 1864: 1858: 1856: 1855:Tilly Edinger 1853: 1849: 1845: 1844: 1839: 1835: 1831: 1830: 1825: 1824: 1823:Scaphognathus 1819: 1818: 1812: 1810: 1806: 1802: 1797: 1795: 1791: 1787: 1786:Ornithosauria 1783: 1778: 1775: 1771: 1767: 1766: 1761: 1753: 1748: 1739: 1737: 1733: 1729: 1728:Edward Newman 1724: 1719: 1717: 1713: 1712:Pterodactylus 1709: 1708: 1707:Pterodactylus 1703: 1702:Ptéro-Dactyle 1699: 1695: 1686: 1682: 1680: 1676: 1675:Ichthyosauria 1672: 1671:Pterodactylus 1668: 1663: 1659: 1655: 1652: 1651:Late Jurassic 1648: 1644: 1643: 1638: 1630: 1626: 1625: 1619: 1609: 1599: 1596: 1595:Tupandactylus 1593:relatives of 1591: 1590: 1584: 1581: 1580:Pterorhynchus 1577: 1576: 1571: 1570: 1565: 1560: 1559:anurognathids 1556: 1552: 1548: 1543: 1541: 1537: 1527: 1525: 1518: 1514: 1510: 1508: 1507: 1502: 1501: 1496: 1492: 1488: 1487: 1482: 1478: 1474: 1470: 1466: 1462: 1454: 1450: 1441: 1439: 1435: 1431: 1421: 1417: 1414: 1409: 1404: 1402: 1391: 1387: 1385: 1381: 1377: 1372: 1368: 1359: 1350: 1348: 1347: 1346:Pterodactylus 1341: 1337: 1335: 1330: 1326: 1322: 1317: 1313: 1310: 1309: 1304: 1303: 1298: 1292: 1290: 1286: 1282: 1277: 1272: 1269: 1265: 1261: 1257: 1256: 1252: 1248: 1247: 1241: 1239: 1235: 1234:cruropatagium 1231: 1230: 1225: 1221: 1220: 1215: 1207: 1204: 1200: 1196: 1195: 1190: 1181: 1179: 1174: 1172: 1168: 1167: 1166:actinofibrils 1157: 1156: 1150: 1146: 1143: 1139: 1130: 1121: 1119: 1115: 1111: 1106: 1103: 1099: 1095: 1091: 1081: 1079: 1075: 1070: 1066: 1057: 1053: 1050: 1049:neural spines 1045: 1041: 1035: 1028: 1023: 1019: 1017: 1013: 1009: 1005: 1001: 994: 993: 987: 978: 976: 972: 971:ossa dentalia 966: 964: 963: 958: 957:Pterorhynchus 954: 950: 949: 948:Pterodactylus 944: 943: 942:Pterorhynchus 936: 934: 933: 928: 924: 920: 919: 911: 910: 906:The skull of 904: 900: 898: 894: 893: 888: 884: 880: 876: 872: 864: 863: 858: 857: 852: 851: 846: 841: 837: 835: 831: 827: 823: 819: 815: 811: 806: 798: 797: 791: 782: 780: 775: 767: 762: 752: 750: 746: 742: 741: 737: 733: 727: 724: 722: 721: 716: 715: 710: 709: 704: 703: 698: 697: 692: 691: 690:Pterodactylus 686: 685: 680: 679: 674: 673: 668: 667: 666:Istiodactylus 661: 659: 658: 657:Scaphognathus 653: 652: 647: 646: 641: 640: 639:Eudimorphodon 635: 634: 628: 626: 622: 618: 614: 610: 606: 602: 598: 594: 584: 582: 576: 574: 570: 569: 568:Pterodactylus 564: 560: 556: 552: 548: 543: 541: 537: 536: 535:Hatzegopteryx 531: 530: 525: 524:anurognathids 521: 517: 513: 509: 505: 501: 497: 493: 488: 486: 481: 477: 473: 469: 465: 460: 458: 454: 450: 446: 442: 438: 434: 431: 427: 423: 415: 412: 408: 407:Ornithosauria 403: 400: 396: 392: 387: 383: 378: 371: 365: 363: 357: 354: 353: 352:Peteinosaurus 346: 343: 342: 341:Eudimorphodon 335: 334: 333: 327: 323: 322: 315: 313: 312: 311:Preondactylus 305: 304: 303: 302:Preondactylia 297: 295: 294:Eopterosauria 289: 287: 281: 280: 278: 273: 268: 263: 257: 254: 253: 250: 244: 241: 238: 237: 234: 231: 228: 225: 224: 221: 218: 215: 214: 211: 208: 205: 204: 201: 198: 195: 194: 189: 184: 180: 177: 176: 175:Tropeognathus 171: 170: 165: 164: 159: 158: 153: 152: 151:Pterodactylus 147: 146: 140: 136: 131: 124: 119: 114: 109: 104: 99: 94: 89: 84: 79: 74: 69: 63: 56: 52: 51:Late Triassic 44: 41: 37: 33: 32:Pterodactylus 19: 17619: 17184: 17130: 17125:Ludodactylus 17123: 17116: 17109: 17101: 17093: 17086: 17066: 17058: 17053:Nicorhynchus 17051: 17044: 17037: 17016: 17009: 17001: 16993: 16985: 16977: 16969: 16939: 16932: 16924:Hamipteridae 16910: 16902: 16868: 16860: 16852: 16832: 16824: 16819:Aetodactylus 16817: 16782: 16774: 16767: 16759: 16753:Ludodactylus 16751: 16743: 16735: 16727: 16719: 16711: 16705:Aetodactylus 16703: 16682: 16674: 16666: 16658: 16650: 16643: 16636: 16593: 16586: 16562: 16555: 16548: 16541: 16515:Liaoxipterus 16513: 16506: 16486: 16479: 16472: 16465: 16443:Mimodactylus 16441: 16434: 16427: 16406: 16399: 16392: 16385: 16361: 16354: 16329: 16321:Lanceodontia 16302: 16294: 16289:Lonchodectes 16287: 16279: 16271: 16251: 16244: 16237: 16190: 16183: 16176: 16169: 16149: 16142: 16135: 16110: 16103: 16078: 16070: 16063: 16057:Ornithostoma 16055: 16048: 16040: 16011: 15951: 15865: 15857: 15850: 15843: 15836: 15829: 15824:Eurazhdarcho 15822: 15815: 15808: 15801: 15781: 15774: 15767: 15759: 15752:Azhdarchinae 15738: 15730: 15723: 15715: 15708: 15700: 15693:Azhdarchidae 15679: 15672: 15664: 15636: 15629: 15622: 15601: 15593: 15588:Meilifeilong 15586: 15579: 15571: 15563: 15555: 15531: 15506: 15498: 15490: 15482: 15474: 15467: 15445: 15438: 15431: 15423: 15373: 15366: 15346: 15339: 15332: 15324: 15300: 15292: 15284: 15276: 15268: 15244: 15237: 15230: 15225:Huaxiapterus 15223: 15216: 15210:Eopteranodon 15208: 15200: 15194:Afrotapejara 15192: 15185:Sinopterinae 15174:Afrotapejara 15172: 15148: 15141: 15110: 15103: 15096: 15088: 15080: 15073: 15065: 15057: 15049: 15041: 15019: 15014:Bennettazhia 15012: 14977:Ornithostoma 14975: 14968: 14960: 14952: 14945: 14938: 14904: 14897: 14876: 14868: 14863:Ordosipterus 14861: 14854: 14846: 14838: 14813:Tapejaroidea 14798: 14702: 14656:Pterodaustro 14654: 14647: 14640: 14633: 14613: 14589: 14582: 14575: 14569:Huanhepterus 14567: 14562:Gnathosaurus 14560: 14542:Moganopterus 14540: 14533: 14514:Pterofiltrus 14512: 14507:Petrodactyle 14505: 14498: 14493:Liaodactylus 14491: 14486:Kepodactylus 14484: 14477: 14470: 14463: 14456: 14449: 14442: 14434: 14401:Huanhepterus 14399: 14392: 14384: 14363: 14358:Aerodactylus 14356: 14331: 14325:Petrodactyle 14323: 14315: 14308: 14300: 14265: 14258: 14227: 14219: 14212: 14204: 14184: 14075: 14067: 14060: 14053: 14046: 14041:Kryptodrakon 14039: 14031: 14024: 14017: 13989: 13983:Jeholopterus 13981: 13973: 13966: 13959: 13939: 13932: 13926:Jeholopterus 13924: 13916: 13911:Anurognathus 13909: 13891:Mesadactylus 13889: 13854: 13846: 13808: 13800: 13793: 13786: 13778: 13758: 13734: 13727: 13716:Darwinoptera 13702: 13695: 13688: 13657: 13649: 13595: 13524: 13517: 13510: 13503: 13496: 13453: 13447:Nesodactylus 13445: 13439:Cacibupteryx 13437: 13419:Sericipterus 13417: 13409: 13401: 13394: 13387: 13358: 13351: 13346:Nesodactylus 13344: 13337: 13330: 13325:Cacibupteryx 13323: 13318:Bellubrunnus 13316: 13291: 13283: 13276: 13268: 13229: 13222: 13186: 13173:Lonchognatha 13154: 13146: 13138: 13131: 13124: 13116: 13096: 13065: 13026: 13018: 12997: 12973: 12966: 12961:Pachagnathus 12959: 12951: 12921: 12913: 12907:Austriadraco 12905: 12871: 12866:Austriadraco 12864: 12856: 12835: 12828: 12820: 12795: 12788: 12768: 12751: 12738: 12701: 12654: 12645: 12636: 12601: 12507: 12416:Eusaurischia 12399: 12392: 12385: 12378: 12357: 12349: 12341: 12335:Chindesaurus 12333: 12306: 12295: 12285: 12275: 12267:Chindesaurus 12265: 12255: 12223:Ornithischia 12212:Ornithischia 12173: 12166: 12159: 12152: 12147:Ignotosaurus 12145: 12138: 12131: 12124: 12105:Technosaurus 12103: 12098:Soumyasaurus 12096: 12090:Pisanosaurus 12088: 12081: 12074: 12067: 12060: 12048:Silesauridae 12032: 12021: 12011: 12001: 11949: 11939:Venetoraptor 11937: 11929: 11922: 11915: 11908: 11900: 11893: 11886:Lagerpetidae 11842: 11835: 11830:Spondylosoma 11828: 11821: 11813: 11806:Aphanosauria 11790: 11779: 11736: 11633: 11525: 11506: 11487: 11442: 11436: 11426: 11417: 11411: 11399:. Retrieved 11393: 11380: 11368:. Retrieved 11362: 11352: 11344: 11341: 11317: 11301: 11298: 11294: 11288: 11280: 11226: 11222: 11216: 11205:. Retrieved 11201: 11191: 11148: 11144: 11134: 11094:(1): 13130. 11091: 11087: 11077: 11042: 11038: 11028: 11017:. Retrieved 11013: 11004: 10959: 10955: 10949: 10897: 10893: 10889: 10867:. Retrieved 10863: 10808: 10804: 10794: 10767: 10763: 10753: 10720: 10716: 10710: 10675: 10671: 10617: 10613: 10607: 10564: 10560: 10550: 10525:11336/156308 10507: 10503: 10493: 10484: 10480: 10470: 10429: 10425: 10412: 10404:the original 10399: 10390: 10350:(6995): 33. 10347: 10343: 10333: 10316: 10312: 10276: 10272: 10262: 10254:the original 10248: 10241: 10204: 10200: 10190: 10171: 10167: 10157: 10149: 10144: 10099: 10095: 10089: 10081: 10069: 10057: 10016: 10012: 10002: 9969: 9965: 9959: 9947: 9914: 9910: 9897: 9885: 9873: 9861: 9828: 9824: 9820: 9814: 9809:, p. 51 9780: 9776: 9763: 9730: 9726: 9716: 9683: 9679: 9666: 9629: 9625: 9615: 9606: 9596: 9579: 9575: 9569: 9534: 9530: 9520: 9508:. Retrieved 9504: 9492: 9483: 9473: 9464: 9460: 9447: 9435:. Retrieved 9432:NewScientist 9431: 9406:. Retrieved 9401: 9375:. Retrieved 9366:ScienceBlogs 9364: 9351: 9331: 9324: 9312:. Retrieved 9309:NewScientist 9308: 9298: 9286:. Retrieved 9282:the original 9277: 9223: 9219: 9213: 9197: 9192: 9172: 9165: 9150: 9145: 9112: 9108: 9102: 9077: 9073: 9051: 9041: 8998: 8994: 8984: 8951: 8948:Paleobiology 8947: 8934: 8923:. Retrieved 8916:the original 8895: 8891: 8882: 8874: 8841: 8837: 8831: 8794: 8791:PLOS Biology 8790: 8742: 8739:Paleobiology 8738: 8731: 8704: 8700: 8690: 8682: 8677: 8640: 8636: 8614:, pp. 231–38 8611: 8606: 8594: 8569: 8565: 8559: 8548:. Retrieved 8544: 8534: 8523:. Retrieved 8519: 8510: 8469: 8465: 8411: 8407: 8401: 8393: 8356: 8352: 8342: 8307: 8303: 8293: 8268: 8264: 8214: 8210: 8197: 8178: 8174: 8164: 8137: 8133: 8127: 8119: 8107: 8095: 8086: 8077: 8013: 8009: 7999: 7987: 7979: 7974: 7969:, p. 9. 7912: 7909:Paleobiology 7908: 7891: 7886: 7877: 7873: 7867: 7859: 7854: 7846: 7841: 7830:. Retrieved 7823:the original 7802: 7796: 7783: 7775: 7770: 7761: 7757: 7751: 7742: 7738: 7732: 7720: 7640: 7636: 7630: 7622: 7617: 7609: 7604: 7592: 7580: 7568: 7559: 7555: 7545: 7536: 7532: 7526: 7514: 7506: 7502: 7496: 7488: 7483: 7475: 7471: 7465: 7456: 7452: 7446: 7438: 7433: 7425: 7420: 7415:, p. 7. 7389: 7384:, p. 6. 7377: 7365: 7360:, p. 5. 7307: 7303: 7293: 7261:(1): 12–13. 7258: 7254: 7244: 7193: 7189: 7182: 7149: 7145: 7139: 7104: 7100: 7090: 7049: 7045: 7039: 7028:. Retrieved 7022: 7012: 6972:(1): 24–30. 6969: 6965: 6955: 6946: 6942: 6936: 6903: 6891: 6879: 6867: 6855: 6813:(5): e2271. 6810: 6806: 6665: 6661: 6651: 6639: 6558: 6552: 6542: 6515: 6511: 6501: 6484: 6480: 6474: 6449: 6445: 6432: 6397: 6391: 6381: 6369: 6357: 6305: 6301: 6291: 6258: 6254: 6248: 6215: 6211: 6205: 6172: 6168: 6164: 6083:(2): e4497. 6080: 6076: 6028: 6024: 6019:Jeholopterus 6018: 5972: 5968: 5962: 5935: 5906: 5805: 5764: 5760: 5754: 5746: 5734: 5718: 5704:(5): 213–16. 5701: 5697: 5672: 5655: 5651: 5647: 5555: 5507: 5498: 5494: 5488: 5455: 5451: 5438: 5427:. Retrieved 5423: 5414: 5373: 5369: 5329: 5307:. Retrieved 5266: 5255: 5214: 5210: 5197: 5156: 5152: 5143: 5098: 5094: 5056:. Retrieved 5052: 5042: 5031:. Retrieved 5027: 5018: 4999: 4995: 4985: 4976: 4964: 4954: 4947: 4936: 4927: 4909: 4900: 4891: 4887: 4883: 4877: 4851: 4845: 4808: 4804: 4794: 4759: 4755: 4719: 4686: 4623: 4614: 4491: 4482: 4480:anurognathid 4469: 4462: 4458: 4452: 4450: 4443: 4437: 4432: 4426: 4420: 4406: 4400: 4390: 4388: 4375:Ludodactylus 4373: 4369: 4365: 4359: 4355: 4349: 4343: 4341: 4334: 4328: 4318: 4308: 4301: 4287: 4283: 4279: 4275: 4264:Pterodaustro 4263: 4259: 4255: 4245: 4235: 4231:Pterodaustro 4229: 4223: 4213: 4207: 4201: 4195: 4187: 4182: 4176: 4168: 4160: 4159:"), such as 4154: 4145: 4132: 4121: 4118: 4114:Pterodaustro 4113: 4105: 4101: 4098:ossification 4097: 4092: 4086: 4077: 4075: 4055: 4049: 4045:Pterodaustro 4043: 4024: 4000: 3990: 3984: 3980: 3974: 3956: 3950: 3938: 3932: 3926: 3916: 3906: 3900: 3894: 3883:nyctosaurids 3872: 3865: 3859: 3848:Azhdarchidae 3840: 3822: 3813:Pterodaustro 3811: 3801: 3795: 3783: 3773: 3755: 3752:frigatebirds 3729: 3724: 3718: 3707:Sericipterus 3706: 3700: 3694: 3690:Sericipterus 3688: 3682: 3676: 3670: 3665: 3659: 3653: 3649: 3643: 3637: 3630: 3616: 3614: 3603: 3599: 3579: 3565:boreopterids 3561: 3549: 3545:Pterodaustro 3543: 3537: 3523: 3520: 3503: 3499:cruropatagia 3495: 3487: 3457: 3449:road runners 3442: 3439:Kevin Padian 3428: 3421: 3415: 3386: 3379: 3366: 3362: 3356: 3339:subcutaneous 3335: 3323: 3316: 3310: 3304: 3284: 3277: 3267:Darren Naish 3254: 3246: 3244: 3241: 3217:Paleobiology 3009: 2989: 2963: 2962: 2926: 2906: 2870: 2869: 2821: 2800: 2799: 2769: 2730:Kryptodrakon 2728: 2727: 2697: 2674: 2673: 2653:Darwinoptera 2651: 2623: 2622: 2595: 2568: 2541: 2511: 2499: 2488: 2485:phylogenetic 2477: 2469:paraphyletic 2446: 2433: 2429: 2425: 2424:(originally 2421: 2412: 2406: 2395: 2380: 2376:Lazarus taxa 2369: 2354: 2347: 2344:nyctosaurids 2338:", possible 2333: 2327: 2317: 2301: 2291: 2273: 2268: 2263: 2251: 2211: 2205: 2197: 2187: 2183: 2168: 2155: 2136: 2126: 2119: 2109: 2103: 2092:protorosaurs 2084:crocodilians 2060: 2040: 2008:Isle of Skye 2001: 1999: 1971: 1958:Lagerstätten 1957: 1955: 1940: 1929: 1925: 1917:Kevin Padian 1906: 1900: 1894: 1886: 1870: 1859: 1843:Anurognathus 1841: 1837: 1827: 1821: 1815: 1813: 1804: 1798: 1789: 1785: 1770:Richard Owen 1763: 1757: 1751: 1720: 1711: 1705: 1701: 1691: 1679:Plesiosauria 1670: 1646: 1642:Lagerstätten 1640: 1634: 1627:specimen by 1622: 1594: 1587: 1585: 1579: 1573: 1567: 1544: 1540:maniraptoran 1533: 1521: 1517:Jeholopterus 1516: 1504: 1498: 1495:histological 1484: 1458: 1452: 1427: 1424:Soft tissues 1418: 1405: 1397: 1388: 1364: 1344: 1342: 1338: 1334:induced drag 1329:nyctosaurids 1325:azhdarchoids 1318: 1314: 1306: 1300: 1293: 1273: 1271:wing-plans. 1255:Jeholopterus 1253: 1251:anurognathid 1244: 1242: 1233: 1227: 1223: 1217: 1213: 1211: 1205: 1198: 1192: 1175: 1164: 1161: 1153: 1135: 1117: 1109: 1107: 1087: 1074:zygapophyses 1062: 1036: 1032: 1026: 1025:The neck of 997: 990: 970: 967: 960: 956: 946: 940: 937: 930: 916: 914: 907: 892:Pterodaustro 890: 879:Azhdarchidae 868: 860: 854: 848: 829: 801: 794: 776: 772: 736:Jeholopterus 732:Anurognathus 730: 728: 725: 718: 712: 706: 700: 694: 688: 682: 676: 670: 664: 662: 655: 649: 643: 637: 631: 629: 590: 577: 566: 562: 559:pterodactyls 558: 555:Ornithischia 544: 533: 527: 489: 468:insectivores 461: 432: 421: 420: 406: 405: 350: 339: 331: 319: 309: 301: 261: 239: 226: 173: 167: 161: 157:Anurognathus 155: 149: 143: 60:228–66 40: 17682:iNaturalist 17650:Pterosauria 17644:Wikispecies 17620:Pterosauria 17473:Insect wing 17423:Webbed foot 17364:unguligrade 17359:plantigrade 17354:digitigrade 17011:Thapunngaka 16971:Amblydectes 16890:Anhangueria 16854:Aussiedraco 16588:Boreopterus 16363:Lonchodraco 16296:Lonchodraco 16239:Aussiedraco 16185:Nyctosaurus 16042:Bogolubovia 15710:Bogolubovia 15581:Lacusovagus 15573:Eoazhdarcho 15533:Eoazhdarcho 15492:Leptostomia 15484:Keresdrakon 15359:Caiuajarina 15326:Bakonydraco 15294:Lacusovagus 15286:Keresdrakon 15257:Tapejarinae 15218:Huaxiadraco 15202:Bakonydraco 15161:Tapejaridae 15090:Leptostomia 15082:Lacusovagus 15075:Kariridraco 15021:Keresdrakon 14962:Leptostomia 14954:Keresdrakon 14615:Ctenochasma 14577:Lusognathus 14500:Otogopterus 14155:Lophocratia 14088:Lophocratia 14055:Pangupterus 13991:Sinomacrops 13332:Dorygnathus 13201:Novialoidea 13156:Rhamphinion 13133:Dimorphodon 12808:Caviramidae 12752:Pterosauria 12739:Pterosauria 12702:Pterosauria 12641:Archosauria 12602:Pterosauria 12380:Gnathovorax 12359:Caseosaurus 12161:Lutungutali 12154:Kwanasaurus 12126:Amanasaurus 12069:Asilisaurus 12062:Agnosphitys 12023:Nyasasaurus 11961:Pterosauria 11950:Pterosauria 11917:Kongonaphon 11910:Ixalerpeton 11860:Ornithodira 11837:Teleocrater 11823:Dongusuchus 11792:Mambachiton 11638:Archosauria 10481:Ameghiniana 10074:Witton 2013 10062:Witton 2013 9952:Witton 2013 9890:Witton 2013 9878:Witton 2013 9866:Witton 2013 9807:Witton 2013 9467:(2): 18–24. 8599:Witton 2013 8112:Witton 2013 8100:Witton 2013 8070:Witton 2013 8056:Witton 2013 7967:Witton 2013 7413:Witton 2013 7394:Witton 2013 7382:Witton 2013 7358:Witton 2013 7219:10468/11874 6929:Witton 2013 6908:Witton 2013 6896:Witton 2013 6884:Witton 2013 6872:Witton 2013 6860:Witton 2013 6782:Witton 2013 6613:Witton 2013 6374:Witton 2013 6362:Witton 2013 6148:Witton 2013 6133:Witton 2013 5955:Witton 2013 5940:Witton 2013 5928:Witton 2013 5911:Witton 2013 5863:Witton 2013 5846:Witton 2013 5829:Witton 2013 5810:Witton 2013 5781:Witton 2013 5631:Witton 2013 5614:Witton 2013 5582:Witton 2013 5529:Witton 2013 5512:Witton 2013 4933:"Pterosaur" 4710:wing lizard 4327:Scene from 4276:Ctenochasma 4274:pterosaurs 4256:Ctenochasma 4225:Ctenochasma 3997:ichthyosaur 3986:Squalicorax 3887:anhanuerids 3829:Tapejaridae 3784:Lonchodraco 3775:Lonchodraco 3684:Dorygnathus 3618:Dimorphodon 3590:tropicbirds 3577:nyctosaurid 3573:azhdarchids 3569:frigatebird 3506:azhdarchids 3472:azhdarchids 3460:plantigrade 3444:Dimorphodon 3318:On the Wing 3291:Mark Witton 3259:azhdarchids 3185:Novialoidea 2748:Lophocratia 2500:Pterosauria 2271:ichnofossil 2259:Rupert Wild 2171:paleoartist 2163:protorosaur 2161:a gliding " 2094:") such as 2088:David Unwin 2070:called the 2049:share many 2047:Lagerpetids 1982:Lü Junchang 1974:Jehol Biota 1945:, from the 1911:began what 1902:Deinonychus 1846:. 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A fuzzy 1469:homologous 1408:metatarsal 1401:metatarsus 1376:pubic bone 1027:Anhanguera 918:Pteranodon 871:keratinous 845:tapejarids 822:archosaurs 814:premaxilla 805:heterodont 678:Pteranodon 605:breastbone 551:Saurischia 540:endothermy 500:homologous 445:Cretaceous 422:Pterosaurs 275:Subgroups 17492:Evolution 17451:Bird wing 17396:Arthropod 17389:quadruped 17039:Aerodraco 16481:Luchibang 16429:Haopterus 16151:Simurghia 16144:Epapatelo 16112:Cretornis 16050:Dawndraco 15776:Azhdarcho 15769:Albadraco 15761:Aerotitan 15681:Alanqidae 15459:Alanqidae 15425:Cretornis 15375:Torukjara 15368:Caiuajara 15105:Tupuxuara 15043:Aerotitan 14947:Inabtanin 14535:Feilongus 13934:Luopterus 13651:Allkaruen 13278:Klobiodon 13118:Allkaruen 12953:Caviramus 12830:Caviramus 12618:Kingdom: 12466:includes 12462:Theropoda 12451:Theropoda 12076:Gamatavus 11615:Kingdom: 11395:IndieWire 11364:Thrillist 11304:: 56–63. 11165:0962-8452 10650:206529739 10599:0272-4634 10534:1476-4687 10279:: 59–67. 10207:: e2908. 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