Pterosaur - Knowledge

1138: 3219: 892: 4313: 1011: 755: 1158:. 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 1587:

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.

2276: 17169: 17316: 1674: 1607: 2984: 1316:, 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 3278:. 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. 4003: 371: 12510: 12499: 15936: 2692: 4540: 1400:

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

2536: 2171:. 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 1118: 14738: 3211: 975: 1567:. 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 3393: 1045: 124: 177: 2901: 1347: 1305:

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,

1925: 3004: 4294: 1871: 2796: 14744: 1546:, 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 829: 12545: 14121: 4109:

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.

13595: 1438: 3468: 2140: 15944: 2075:). 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. 4512: 2724: 4498: 1502: 2563: 3262: 1800:, 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. 1178: 966:. 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. 1654:

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

455:') 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 1515:(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. 3370:

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

4058:. 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 770:. 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. 1425:, 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 4308:

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.

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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.

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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.

499:, 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 2352:-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 2467:

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.

1288:. 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 10257: 4141:, 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. 1550:

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

546:, 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 467:) 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 2341:. 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. 791:

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

1069:. 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. 7458:

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

3537:), 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 11281:

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.).

1986:. 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. 1026:

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

1714:

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,

3415:(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. 821:. 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 1908:

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

4043:, 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 1699:. 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 888:

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.

1821:. 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 914:

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

4404:

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.

2193:

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

6199:

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".

4450:

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

2304:. 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. 6647:"Pterosaurs evolved a muscular wing–body junction providing multifaceted flight performance benefits: Advanced aerodynamic smoothing, sophisticated wing root control, and wing force generation" 1723:. 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, 1329:

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.

862:

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

9486: 10384: 1232:

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

9262: 6468:

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".

1338:

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.

3597:

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

1523:

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

2444:. 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 6242:

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".

10300:

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)".

1803:

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

924:

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.

10747:

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).

1628:

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

9998:"Evidence for tactile foraging in pterosaurs: a sensitive tip to the beak of Lonchodraco giganteus (Pterosauria, Lonchodectidae) from the Upper Cretaceous of southern England" 2300:

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

4446:. 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 801:, 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 9953:

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".

1229:

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

2097:

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

4133:

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

3330:

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

9207:

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".

1321:

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

6949:

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).

4367:, whose name means "toy finger" for its resemblance to old, inaccurate children's toys. Pterosaurs have sometimes been incorrectly identified as (the ancestors of) 3712:

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

2238:

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

1165:

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

1969:

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

8930:"Does morphology reflect osteohistology-based ontogeny? A case study of Late Cretaceous pterosaur jaw symphyses from Hungary reveals hidden taxonomic diversity" 4124:

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

1421:

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

9359: 9036: 7786: 5482:

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

3980: 904:

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

4790:"Testing pterosaur ingroup relationships through broader sampling of avemetatarsalian taxa and characters and a range of phylogenetic analysis techniques" 3234:

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

3444:

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.

3246:

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

2203:

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

1392:

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.

1375:

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

715:

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

4331:

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

12602: 3614:

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

8869: 1205:("fore membrane"), was the forward-most part of the wing and attached between the wrist and shoulder, creating the "leading edge" during flight. The 1109:, jutted obliquely upwards. The rear edge of the breastbone was the deepest point of the thorax. Clavicles or interclavicles were completely absent. 10392: 10236: 9270: 7778: 1785:

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

910:. 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 9293: 534:

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

11420:

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).

17683: 17217: 5061: 4085:

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

1371:

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

1101:, 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 9059:"Small, immature pterosaurs from the Cretaceous of Africa: implications for taphonomic bias and palaeocommunity structure in flying reptiles" 4563: 4177:

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

2247: 2076: 3968:, embedded in it. The vertebrae are known not to have been eaten and exposed to digestion, as the joints are still articulated. Fossils of 1452:-like filaments known as pycnofibers on the head and torso. The term "pycnofiber", meaning "dense filament", was coined by palaeontologist 4076:

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

4073:, as opposed to the single functional ovary in birds, dismissing the reduction of functional ovaries as a requirement for powered flight. 3304:. 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 1125:

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

7440:

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".

5706:

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

3657:

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

1395:

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

9416: 4433: 2460:(unnatural) group, since the pterodactyloids evolved directly from them and not from a common ancestor, so, with the increasing use of 740:

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

8252:

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

4037:, a pterosaur known by abundant material. This was supported by the description of an additional pterosaur egg belonging to the genus 732:) is debated. Anurognathids were highly specialized. Small flyers with shortened jaws and a wide gape, some had large eyes suggesting 17573: 11578: 9468: 7413:

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

2301: 9893:"A new rhamphorhynchid pterosaur from the Upper Jurassic of Xinjiang, China, and the phylogenetic relationships of basal pterosaurs" 5434:"A new rhamphorhynchid pterosaur from the Upper Jurassic of Xinjiang, China, and the phylogenetic relationships of basal pterosaurs" 17729: 11375: 10601:

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

7086:"The Soft Tissue of Jeholopterus (Pterosauria, Anurognathidae, Batrachognathinae) and the Structure of the Pterosaur Wing Membrane" 1137: 2409:

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

1894:, whose spectacular traits refuted what had become entrenched orthodoxy. In 1970, likewise the description of the furry pterosaur 1850:

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

483:, which covered their bodies and parts of their wings. Pycnofibers grew in several forms, from simple filaments to branching down 11326: 8338:"The phylogenetic relationships of basal archosauromorphs, with an emphasis on the systematics of proterosuchian archosauriforms" 2456:, "advanced" pterosaurs with short tails. However, this traditional division has been largely abandoned. Rhamphorhynchoidea is a 4189:

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

4113:

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

4027:

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

3920:

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

3218: 8874:

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

6950: 1888:

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

3258:

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

1781:. Seeley thought that pterosaurs were warm-blooded and dynamic creatures, closely related to birds. Earlier, the evolutionist 17578: 12581: 11522: 11503: 11484: 9329: 5552: 5327: 5269: 4906: 4856: 4260: 962:, were at the tip tightly fused into a central symphysis. This made the lower jaws function as a single connected whole, the 7133:

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

2348:, indicating a higher diversity of Late Cretaceous pterosaurs than previously accounted for. The recent findings of a small 891: 10258:"An unusual modification of the jaws in cf. Alanqa, a mid-Cretaceous azhdarchid pterosaur from the Kem Kem beds of Morocco" 3334:, seeing as their shoulder-pectoral girdles were too inflexible to move the sternum as in birds, and they possessed strong 2405:

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

1982:. New and old fossils yielded much more information when subjected to modern ultraviolet light or roentgen photography, or 1417:

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

9386: 1486:

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

604:

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

17210: 8776:"Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary" 4144:

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

1913:, who in 1970s laid the foundations of modern pterosaur science. In 1978, he published the first pterosaur textbook, the 3511:

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

17734: 10476:(4 (supplement, abstracts from XX Jornadas Argentinas de Paleontología de Vertebrados, La Plata, 26-28 May 2004)): 40R. 9057:

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

6542: 6381: 1954:

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

3866:, estimating the bite force and potential dietary habits of nine selected species. The study corroborated the view of 1003:. Advanced pterosaurs are unique in possessing special processes projecting adjacent to their condyle and cotyle, the 11496:

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

11347: 11298: 9170: 5715: 4659: 2380: 2284: 17168: 9037:"Pterosaur remains (Archosauria, Ornithodira) from the early Late Cretaceous of "La Buitrera", Río Negro, Argentina" 8504: 5252:(2004). "Origin and relationships of Dinosauria". In Weishampel, David B.; Dodson, Peter; Osmólska, Halszka (eds.). 4312: 580:

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

9591: 9349: 4558: 600:

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

8071: 4152:, the average growth rate during the first year of life was 130% to 173%, slightly faster than the growth rate of 3641:

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

1251:

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

1010: 592:

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

3552:

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

1358:

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

8529: 2255:. Like this archosaur, basal pterosaur lineages have plantigrade hindlimbs that show adaptations for saltation. 17203: 9194: 9147: 8929: 4588: 4318: 1596: 14743: 11128:

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

6446: 5140:(March 1975). "Pterosaur from the Latest Cretaceous of West Texas: Discovery of the Largest Flying Creature". 3932:

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

3771:

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

11571: 11210:

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

10081:(Pterodactyloidea: Azhdarchoidea) from the Early Cretaceous Jehol Biota and its paleoecological implications" 1711: 1031:. Such species also often show a fusion of the front dorsal vertebrae into a rigid whole which is called the 12509: 10186:"Neck biomechanics indicate that giant Transylvanian azhdarchid pterosaurs were short-necked arch predators" 6495:

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

754: 451:

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

4380: 1855: 1716: 24: 15935: 14737: 10848: 8904: 4156:. Growth in these species slowed after sexual maturity, and it would have taken more than three years for 3824:

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

1846:

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

17337: 17315: 10465: 10242: 7811: 4531: 4431:

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

2004: 1474: 870:, and had larger, more extensive, and more bird-like beaks. Some groups had specialised tooth forms. The 12498: 5080:"Discovery of a rare arboreal forest-dwelling flying reptile (Pterosauria, Pterodactyloidea) from China" 4463:, not reflecting their full dietary variation. They are also often shown as aerial predators similar to 176: 17429: 17265: 12548: 10749:"Eggshell and Histology Provide Insight on the Life History of a Pterosaur with Two Functional Ovaries" 4894: 4553: 4440:

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

4385: 3799:

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

1527:. Others were unconvinced, considering the difference with the "quills" found on many of the bird-like 1429:. They covered pads cushioning the impact of walking. Scales are unknown from other parts of the body. 441: 12516: 9997: 4343:, or a fictionalized hybrid of the two. Many children's toys and cartoons feature "pterodactyls" with 17739: 14120: 12574: 11684: 9712:"New pterosaur tracks (Pteraichnidae) from the Late Cretaceous Uhangri Formation, southwestern Korea" 6975: 3884: 2437: 2185:

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

1782: 1689:. Cuvier agreed in 1801, understanding it was an extinct flying reptile. In 1809, he coined the name 1650: 1617: 10241:. The Annual Symposium of Vertebrate Paleontology and Comparative Anatomy. Edinburgh. Archived from 9096:

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

8191:

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

4958: 12678: 12427: 11599: 11564: 11186: 8446:

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

5741: 4568: 4353:-like tails and teeth, a combination that never existed in nature. However, at least one pterosaur 4349: 4324: 3300: 3287: 3283: 1524: 1263:

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

13579: 9609:

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

6010:(Pterosauria, Anurognathidae, Batrachognathinae) and the structure of the pterosaur wing membrane" 14156: 13587: 5037: 4573: 3780: 3338:. Thus, their respiratory system had characteristics comparable to both modern archosaur clades. 3306: 2746: 850: 17701: 14125: 11024:"Prenatal development in pterosaurs and its implications for their postnatal locomotory ability" 8724: 4120:. Due to how underdeveloped the chests of the hatchlings were for flying, it was suggested that 3698:), and shorter, broader wings. These were either terrestrial/aerial predators of vertebrates or 2103:, a "protorosaur" with skin membranes on its hindlimbs likely used for gliding. A 1999 study by 17600: 17568: 17512: 17507: 17492: 17076: 15234: 12121: 10998: 10137:

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

9160: 8978:

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

4942: 4396: 3890: 3875: 3362: 3356: 1930: 1773:, at the time the main English expert on the subject, who also wrote the first pterosaur book, 1612: 1221:

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

999:) was concave and into it fitted a convex extension at the rear of the preceding vertebra, the 17175: 12492: 10238:

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

9487:"The wingtips of the pterosaurs: Anatomy, aeronautical function and 3 ecological implications" 9058: 5317: 5261: 3601:

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

3564:, which presumably still needed to launch from water in case they found themselves in it. The 3447:

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

2345: 944:

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

17696: 17638: 17583: 17181: 12211: 9319: 4390: 4243:, being active throughout the day for short intervals. As a result, the possibly fish-eating 2386: 1851: 1789: 1479: 1159: 17688: 11376:"'Godzilla: King of the Monsters' Trailer Turns Mothra, Rodan, and More Into Epic Spectacle" 10704:

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

10487:

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

9442: 5253: 4388:. They appeared in a number of films and television programs since, including the 1933 film 4251:

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

3820:

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

2275: 17657: 17522: 17449: 17434: 15943: 15329: 13768: 13678: 13594: 12567: 12503: 11949: 11435: 11219: 11084: 10952: 10890: 10801: 10713: 10668: 10610: 10557: 10422: 10340: 10269: 10092: 10009: 9962: 9907: 9821: 9723: 9676: 9216: 9105: 8991: 8944: 8834: 8735: 8562: 8462: 8207: 7905: 7633: 7300: 7186: 7142: 7042: 6803: 6658: 6251: 6208: 6165: 6073: 5965: 5448: 5366: 5207: 5149: 5091: 4844: 4752: 4452: 4425: 4235: 3916: 3238:

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

2164: 1885: 1465: 838: 696: 535: 15951: 3853:

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

2476:

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

507:, which hollowed out their bones to an extreme extent. Pterosaurs spanned a wide range of 8: 17724: 17517: 17444: 16833: 16522: 16209: 15508: 15435: 13203: 12923: 11657: 8505:"Paleontologists find pterosaur precursors that fill a gap in early evolutionary history" 4545: 4098: 4028: 4016: 3928: 3922: 3911: 3569: 3231:

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

2999: 2558: 2356:, and that their diversity might actually have been much larger than previously thought. 2055: 1975: 1770: 1765: 1673: 1655: 1642: 1401: 814: 399: 11439: 11223: 11162: 11129: 11088: 10956: 10894: 10805: 10717: 10672: 10614: 10561: 10426: 10344: 10273: 10096: 10013: 9966: 9911: 9825: 9727: 9680: 9576: 9220: 9109: 8995: 8948: 8838: 8739: 8566: 8466: 8211: 7909: 7637: 7541:"Versuch einer Eintheilung der Säugethiere in 6 Stämme und der Amphibien in 6 Ordnungen" 7304: 7190: 7146: 7046: 7007: 6807: 6662: 6255: 6212: 6169: 6077: 5969: 5452: 5370: 5211: 5153: 5095: 5038:"It's Official: Those Flying Reptiles Called Pterosaurs Were Covered in Fluffy Feathers" 4848: 4756: 3739:

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

3676:, were fish-eaters with long, slender wings, needle-like dentition and long, thin jaws. 2079:

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

608:, which served to stiffen the torso during flight, and provide a stable support for the 17532: 17497: 17250: 16362: 14836: 14679: 13822: 13625: 13612: 13526: 13368: 12999: 12974: 12450: 11243: 11105: 11072: 11048: 11023: 10975: 10936: 10914: 10906: 10827: 10729: 10634: 10466:"Primer reporte de un embrión de pterosaurio (Cretácico inferior, San Luis, Argentina)" 10446: 10366: 10212: 10185: 10115: 10076: 10033: 9978: 9923: 9837: 9777: 9739: 9692: 9637: 9611:"A Jurassic pterosaur from Patagonia and the origin of the pterodactyloid neurocranium" 9610: 9545: 9240: 9121: 9078: 9012: 8979: 8960: 8896: 8850: 8802: 8775: 8751: 8648: 8621: 8486: 8419: 8388: 8364: 8337: 8318: 8269: 8231: 8176: 8159: 8143: 8114: 8024: 7929: 7921: 7803: 7649: 7321: 7288: 7269: 7220: 7158: 7110: 7085: 7066: 6988: 6826: 6791: 6702: 6689: 6646: 6564: 6537: 6450: 6403: 6376: 6316: 6267: 6224: 6181: 6096: 6062:"Respiratory evolution facilitated the origin of pterosaur flight and aerial gigantism" 6061: 6034: 6005: 6004:

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

5981: 5685:

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

5464: 5390: 5231: 5173: 5114: 5079: 4921: 4816: 4789: 4280: 3736: 3314: 3135: 3097: 2983: 2846: 2449: 2243: 2189:

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

1822: 1457: 1285: 1226: 737: 531:, a good oxygen supply and strong muscles made pterosaurs powerful and capable flyers. 500: 488: 387: 171: 10407: 9758: 9661: 8192: 7490:

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

7033:

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

5351: 5192: 4170:, grew to adult size within the first year of life. Additionally, pterodactyloids had 1606: 1007:, and the cotyle also may possess a small prong on its midline called a hypapophysis. 17548: 17527: 17487: 17367: 17285: 16788: 15603: 15532: 14774: 14761: 14704: 14467: 14365: 14267: 14185: 13963: 13940: 13906: 13836: 13377: 13293: 13245: 13219: 12932: 12837: 11518: 11499: 11480: 11453: 11294: 11235: 11167: 11149: 11110: 11053: 10980: 10819: 10770: 10686: 10638: 10626: 10583: 10526: 10518: 10438: 10358: 10217: 10120: 10037: 10025: 9982: 9974: 9892: 9841: 9696: 9642: 9537: 9350:"Pterosaurs breathed in bird-like fashion and had inflatable air sacs in their wings" 9325: 9232: 9190: 9166: 9143: 9125: 9082: 9017: 8807: 8707: 8653: 8490: 8478: 8447: 8424: 8369: 8223: 8193:"A Late Triassic Dinosauromorph Assemblage from New Mexico and the Rise of Dinosaurs" 8016: 8011: 7994: 7881:

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

7326: 7261: 7224: 7212: 7115: 7070: 7058: 6980: 6831: 6706: 6694: 6676: 6569: 6518: 6427: 6408: 6308: 6303: 6286: 6271: 6101: 6039: 5711: 5652: 5548: 5433: 5382: 5323: 5265: 5254: 5249: 5223: 5165: 5137: 5119: 5055: 4902: 4852: 4821: 4770: 4442: 3661: 3620: 3560:-like aerial hawking. These adaptations are also seen in terrestrial pterosaurs like 3522: 2873: 2811: 2759: 2664: 2585: 1991: 1955: 1950: 1935: 1724: 1453: 1350:

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

1290: 1248: 1201:

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

1066: 633: 473: 255: 11247: 10918: 10733: 10657:"Sexually Dimorphic Tridimensionally Preserved Pterosaurs and Their Eggs from China" 9927: 9743: 9549: 9244: 8980:"A small azhdarchoid pterosaur from the latest Cretaceous, the age of flying giants" 8964: 8900: 8854: 8755: 8574: 8322: 8273: 8028: 7933: 7807: 7653: 7540: 7273: 6992: 6513: 6496: 6481: 6454: 6320: 6228: 5985: 5468: 5352:"Neuroanatomy of flying reptiles and implications for flight, posture and behaviour" 5235: 5177: 4105:

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

4002: 3331: 3327: 3214:

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

1862:

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

370: 17558: 17332: 17017: 16900: 16892: 16684: 16613: 16538: 16080: 15983: 15970: 15902: 15782: 15771: 15021: 14878: 14815: 14590: 14572: 14410: 14234: 14050: 14036: 13994: 13500: 13486: 13418: 13097: 12647: 12105: 11865: 11669: 11443: 11227: 11157: 11141: 11100: 11092: 11073:"Powered flight in hatchling pterosaurs: Evidence from wing form and bone strength" 11043: 11035: 10970: 10960: 10898: 10831: 10809: 10760: 10721: 10676: 10618: 10573: 10565: 10508: 10500: 10450: 10430: 10370: 10348: 10309: 10277: 10207: 10197: 10164: 10110: 10100: 10017: 9970: 9915: 9829: 9773: 9731: 9684: 9632: 9622: 9572: 9527: 9224: 9113: 9070: 9007: 8999: 8952: 8888: 8842: 8797: 8787: 8743: 8697: 8643: 8633: 8570: 8470: 8414: 8404: 8359: 8349: 8310: 8300: 8261: 8235: 8215: 8171: 8138: 8130: 8006: 7913: 7795: 7641: 7316: 7308: 7251: 7202: 7194: 7162: 7150: 7105: 7097: 7050: 6970: 6962: 6821: 6811: 6684: 6666: 6559: 6551: 6508: 6477: 6442: 6398: 6390: 6298: 6259: 6216: 6185: 6173: 6091: 6081: 6029: 6021: 5973: 5956:

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

5648: 5456: 5394: 5374: 5215: 5157: 5109: 5099: 5012: 4992: 4811: 4801: 4760: 4713: 4618: 4517: 4174:, meaning that the animals reached a fixed maximum adult size and stopped growing. 4145: 4007: 3946: 3863: 3825: 3821: 3796: 3498: 2966: 2776: 2704: 2453: 2440:

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

2182: 2088: 2060: 2011:, and it has been described as the world's best-preserved skeleton of a pterosaur. 1989:

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

1910: 1836: 1762: 1704: 1426: 988: 885: 875: 798: 601: 561: 452: 445: 237: 10881:

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

10765: 10748: 10281: 10021: 9117: 9074: 7745:

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

6930:

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

6263: 3411:

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

3374:

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

2691: 2535: 2154:" which some controversial studies have posited as a close relative of pterosaurs. 17502: 17406: 17389: 17260: 17106: 16949: 16798: 16583: 16487: 16462: 16455: 16442: 16375: 16261: 16159: 16125: 16021: 16001: 15855: 15713: 15654: 15641: 15612: 15488: 15399: 15258: 15131: 15086: 14958: 14887: 14866: 14692: 14353: 14333: 14305: 14281: 14217: 14209: 14202: 14194: 13949: 13929: 13759: 13735: 13685: 13493: 13476: 13466: 13443: 13399: 13391: 13359: 13348: 13327: 13281: 13273: 13212: 13128: 13068: 12987: 12956: 12846: 12825: 12778: 12663: 12638: 12538: 12416: 12359: 11970: 11748: 11726: 11696: 11587: 11426: 10965: 10941:

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

10569: 10313: 10153:"Azhdarchid pterosaurs: water-trawling pelican mimics or "terrestrial stalkers"?" 10105: 9996:

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

9563:

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

9185:

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

8792: 8289:"The early evolution of archosaurs: relationships and the origin of major clades" 7520:

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

6816: 6086: 5193:"A new giant pterosaur with a robust skull from the latest cretaceous of Romania" 5161: 4926: 4578: 4503: 4472: 4401: 4339: 4225: 3906: 3830: 3785: 3776: 3745: 3732: 3703: 3690: 3666: 3643: 3441: 3381: 3239: 2518: 2360: 2328: 2263: 2132: 2064: 2008: 1876: 1805: 1256: 1143: 1078: 995:. The vertebrae in front of the tail were "procoelous": the cotyle (front of the 950: 897: 871: 863: 728: 684: 639: 418: 309: 221: 43: 11550: 6368: 1630: 1117: 382:

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

17563: 17454: 17377: 17324: 17295: 17255: 17230: 17083: 17056: 17034: 16858: 16772: 16757: 16709: 16648: 16633: 16599: 16424: 16408: 16396: 16382: 16335: 16292: 16252: 16166: 16112: 15840: 15833: 15826: 15798: 15705: 15626: 15553: 15464: 15220: 15112: 15047: 14988: 14928: 14894: 14614: 14541: 14513: 14432: 14382: 14321: 14290: 13890: 13866: 13844: 13790: 13748: 13692: 13660: 13341: 13258: 13235: 13114: 13040: 13008: 12861: 12810: 12758: 12533: 12389: 11890: 11803: 11769: 11266: 11096: 10545: 8747: 7312: 7198: 7054: 4680: 4583: 4481: 4186: 4161: 3990: 3951: 3896: 3867: 3775:

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

3724: 3627: 3594: 3295: 3210: 3154: 3116: 2896: 2687: 2531: 2337: 2323: 2227: 2210:

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

2190: 2104: 1840: 1829:. German studies continued well into the 1930s, describing new species such as 1797: 1761:, also the first non-pterodactyloid pterosaur known. Later in the century, the 1686: 1682: 1547: 1539: 1456:

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

1086: 1053: 996: 980: 974: 784: 749: 708: 690: 660: 517: 508: 464: 350: 151: 19:"Pterodactyl" redirects here. For the genus commonly called "pterodactyl", see 11448: 11421: 10935:

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

10902: 10725: 10681: 10656: 10235:

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

10141:(Pterodactyloidea: Tapejaridae). Flugsaurier 2015 Portsmouth, abstracts, 47–48 9919: 9735: 9688: 9228: 8846: 8827:

Earth and Environmental Science Transactions of the Royal Society of Edinburgh

8686:"Early penguin fossils, plus mitochondrial genomes, calibrate avian evolution" 8474: 8265: 7917: 7896:

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

7256: 7239: 6966: 6792:"A reappraisal of azhdarchid pterosaur functional morphology and paleoecology" 6645:

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

5977: 5460: 5219: 4765: 4740: 3392: 2900: 1970: 1044: 123: 17718: 17623: 17384: 17300: 17290: 17120: 17091: 17067: 17048: 17006: 16991: 16940: 16929: 16822: 16814: 16764: 16701: 16672: 16656: 16640: 16626: 16567: 16545: 16496: 16389: 16150: 16093: 16012: 15847: 15819: 15791: 15662: 15545: 15336: 15266: 15138: 15121: 14977: 14914: 14858: 14623: 14579: 14460: 14453: 14446: 14439: 14424: 14374: 14298: 14255: 14248: 14174: 14021: 14014: 14007: 13856: 13798: 13783: 13776: 13724: 13507: 13176: 13136: 13086: 13055: 13016: 12911: 12903: 12881: 12785: 12769: 12382: 12375: 12331: 12310: 12265: 12128: 11919: 11556: 11153: 10587: 10522: 10029: 6680: 4477: 4197: 4191: 4039: 3849: 3813: 3791: 3728: 3720: 3708: 3684: 3633: 3539: 3527: 3477: 3078: 2979: 2916: 2883: 2790: 2501: 2441: 2396: 2364: 2332: 2312: 2307:

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

2195: 2151: 2145: 2126: 2109: 2099: 2084: 2080: 1901: 1843: 1811: 1695: 1663: 1639: 1577: 1422: 1346: 1334: 1309: 1296: 1273: 1154: 1081:

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

936: 930: 911: 844: 678: 654: 645: 627: 556: 523: 340: 329: 299: 282: 163: 139: 81: 39: 20: 11231: 10622: 8702: 8685: 8219: 6671: 5104: 4467:, grasping human victims with talons on their feet. However, only the small 3003: 2120: 1924: 817:

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

758:

Size disparity of late Maastrichtian Pterosaurs compared to birds and humans

17113: 17041: 16912: 16807: 16741: 16693: 16503: 16431: 16309: 16277: 16045: 15812: 15740: 15681: 15576: 15213: 15198: 15182: 15173: 15162: 15002: 14965: 14851: 14801: 14644: 14557: 14550: 14530: 14502: 14495: 14481: 14474: 14389: 14346: 14313: 14029: 13971: 13914: 13899: 13879: 13704: 13435: 13427: 13407: 13334: 13313: 13306: 13161: 12949: 12895: 12854: 12404: 12323: 12255: 12200: 12135: 12093: 12086: 12078: 12036: 11927: 11874: 11818: 11794: 11457: 11239: 11171: 11145: 11114: 11057: 11039: 10984: 10849:"Pterosaur hatchlings needed their parents, trove of eggs reveals (Update)" 10823: 10774: 10690: 10630: 10530: 10442: 10362: 10221: 10169: 10152: 10124: 9646: 9541: 9354: 9345: 9236: 9021: 8811: 8711: 8657: 8482: 8448:"Enigmatic dinosaur precursors bridge the gap to the origin of Pterosauria" 8428: 8373: 8227: 8134: 8020: 7330: 7265: 7216: 7119: 7101: 7062: 6984: 6835: 6698: 6573: 6555: 6522: 6497:"New long-tailed pterosaurs (Wukongopteridae) from western Liaoning, China" 6412: 6394: 6312: 6105: 6043: 6025: 5750:

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

5386: 5227: 5169: 5123: 4825: 4774: 4468: 4464: 4416: 4363: 4293: 4259:

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

4256: 4219: 4125: 4033: 3871: 3836: 3801: 3678: 3533: 3494: 3460: 3427: 3397: 3255: 3247: 2795: 2718: 2641: 2473: 2457: 2235: 2043: 2035: 1996: 1905: 1870: 1831: 1758: 1667: 1649:, became much sought after by rich collectors. In 1784, Italian naturalist 1557: 1528: 1494: 1359: 1322: 1243: 1239: 1062: 1037: 880: 867: 724: 720: 543: 512: 145: 8160:"The phylogenetic position of the Pterosauria within the Archosauromorpha" 4997: 4980: 4607:

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

4069:

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

3637:

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

17670: 17649: 17632: 17461: 17411: 17394: 17352: 17347: 17342: 16999: 16959: 16878: 16842: 16576: 16351: 16284: 16227: 16173: 16030: 15698: 15569: 15561: 15521: 15480: 15472: 15347: 15314: 15282: 15274: 15245: 15206: 15190: 15149: 15078: 15070: 15063: 15009: 14950: 14942: 14603: 14565: 14488: 14143: 14076: 14043: 13979: 13320: 13189: 13144: 13121: 12796: 12368: 12347: 12339: 12296: 12149: 12142: 12114: 12057: 12050: 12011: 11905: 11898: 11848: 11825: 11811: 11780: 9516:"Breathing in a box: Constraints on lung ventilation in giant pterosaurs" 4309:

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

4213: 4117: 3985: 3974: 3817: 3763: 3740: 3672: 3606: 3565: 3557: 3553: 3487: 3448: 3432: 3279: 3251: 3173: 2736: 2426: 2259: 2159: 1962: 1890: 1817: 1753: 1748: 1418: 1317: 1313: 1057: 1004: 941: 920: 915: 833: 828: 672: 621: 468: 456: 358: 274: 133: 56: 17195: 11422:"A new darwinopteran pterosaur reveals arborealism and an opposed thumb" 10513: 9003: 6951:"Pterosaur integumentary structures with complex feather-like branching" 5378: 3694:

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

3604:

Interpretations of the habits of basal groups have changed profoundly.

3294:

In 1985, the Smithsonian Institution commissioned aeronautical engineer

1105:

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

1052:

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

17362: 17305: 17275: 17270: 16967: 16922: 16850: 16725: 16717: 16552: 16531: 16476: 16344: 16319: 16269: 16234: 16180: 16068: 16060: 16053: 15805: 15728: 15720: 15619: 15583: 15421: 15322: 15301: 15290: 15227: 14844: 14637: 14630: 14065: 13956: 12963: 12620: 12275: 12245: 12228: 12163: 12156: 12071: 12001: 11991: 11912: 11883: 11832: 11642: 11617: 10910: 10202: 9833: 9711: 9627: 8892: 8638: 8409: 8354: 7925: 7799: 7289:"Pterosaur melanosomes support signalling functions for early feathers" 7207: 4806: 4378:

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

4333: 4301: 4240: 4208: 4166: 4077: 4045: 3879: 3750: 3578: 3561: 3513: 3436:, might have walked or even run bipedally, in addition to flying, like 3371: 3267: 2461: 2219: 2204: 2177: 2093: 2072: 2039: 2030: 1979: 1735: 1720: 1469: 1396: 1389: 1364: 906: 802: 793: 778: 666: 593: 539: 528: 448:

stretching from the ankles to a dramatically lengthened fourth finger.

437: 433: 101: 66: 17675: 11187:"July: Pterosaurs parents | News and features | University of Bristol" 11071:

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

10578: 9140:

Anatomy, Phylogeny and Palaeobiology of Early Archosaurs and Their Kin

8123:

Philosophical Transactions of the Royal Society B: Biological Sciences

6535: 5635:

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

3882:

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

3490:, but they too appear to have been generally efficient on the ground. 2024: 1884:

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

1085:. It was probably covered by thick muscle layers. The upper bone, the 527:, which reached wingspans of at least nine metres. The combination of 17439: 17027: 16469: 16417: 16139: 16132: 16100: 16038: 15764: 15757: 15749: 15669: 15447: 15413: 15363: 15356: 15093: 15031: 14935: 14523: 13922: 13639: 13266: 13106: 12941: 12818: 12629: 12439: 12064: 11626: 11407:

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

11383: 11352: 10486: 9532: 9515: 8867: 8314: 7763:

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

7645: 7492:

Denkschriften der königlichen bayerischen Akademie der Wissenschaften

7154: 6220: 6177: 5291: 4460: 4372: 4230: 4203: 4153: 4138: 4051: 4020: 3964: 3959: 3941: 3901: 3840: 3649: 3615: 3598: 3437: 3419: 3376: 3335: 2860: 2763: 2589: 2469: 2445: 2406: 2353: 2308: 2207: 2114:

and named the group Ornithodira to encompass pterosaurs and dinosaurs

1751:

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

1685:

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

1512: 1483: 1372: 1277: 1102: 992: 822: 810: 733: 702: 613: 496: 460: 188: 106: 50: 17594: 10814: 10789: 10504: 10434: 10353: 10328: 8977: 5408: 1511:

The presence of pycnofibers strongly indicates that pterosaurs were

1482:, but had been widely doubted. Since the 1990s, pterosaur finds and 1276:, are much longer than the humerus. They were probably incapable of 1186:, as depicted here, evidences the possibility that pterosaurs had a 17617: 17471: 17372: 17280: 17099: 16983: 16975: 16749: 16733: 16664: 16241: 15591: 15496: 15428: 15100: 15055: 14828: 14057: 13717: 12614: 12285: 12183: 12022: 11611: 8444: 7995:"The shape of pterosaur evolution: evidence from the fossil record" 7478:

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

7012: 6538:"High lift function of the pteroid bone and forewing of pterosaurs" 6377:"High lift function of the pteroid bone and forewing of pterosaurs" 5260:(2nd ed.). Berkeley: University of California Press. pp. 4129: 4024: 3955: 3862:

A 2021 study reconstructed the adductor musculature of skulls from

3806: 3503: 2920: 2815: 2618: 2505: 2430: 2215: 2168: 2068: 2007:

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

2000: 1983: 1966: 1552: 1535: 1207: 1166: 1090: 1032: 963: 605: 581: 504: 444:. Their wings were formed by a membrane of skin, muscle, and other 429: 425: 208: 96: 91: 76: 71: 61: 11130:"Allometric wing growth links parental care to pterosaur giantism" 10488: 8956: 8305: 8288: 7834:

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

4304:, created by Mark Witton for the Royal Society's 350th anniversary 1917:, and in 1991 the second ever popular science pterosaur book, the 17399: 11538: 10600: 9608: 9514:

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

8440: 8438: 7599:

Ornithosauria – an elementary study of the bones of Pterodactyles

4941:

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

4114: 4070: 3768: 3758: 3754: 3481:

were quadrupeds, and some rather efficient terrestrial predators.

3430:

suggested that smaller pterosaurs with longer hindlimbs, such as

3275: 2296: 2231: 2051: 1945:

This development accelerated through the exploitation of two new

1646: 1437: 1368: 1264: 1000: 859: 806: 767: 609: 484: 111: 86: 17662: 10544:

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

9317: 5315: 3749:), while boreopterids were freshwater diving animals similar to 3467: 3274:

Another issue that has been difficult to understand is how they

2723: 2139: 2107:

found that pterosaurs were avemetatarsalians closely related to

940:, the true extent of these crests has only been uncovered using 550:, particularly in fiction and journalism. However, technically, 436:(228 to 66 million years ago). Pterosaurs are the earliest 17357: 15690: 15457: 15039: 13647: 13514: 12608: 11605: 11541:, multi-authored website about all aspects of pterosaur science 10385:"Prehistoric sharks feasted on flying reptiles, fossil reveals" 9662:"Pterosaur Stance and Gait and the Interpretation of Trackways" 8927: 5319:

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

4525: 4055: 3888:

was specialised in consuming relatively large prey compared to

3855: 3844: 3719:

Among pterodactyloids, a greater variation in diet is present.

3699: 3611: 3574: 3472: 3401: 2613: 2562: 2222:

similarities with pterosaurs and reconstructions of lagerpetid

1939: 1896: 1625: 1563: 1501: 1488: 1461: 1355: 1234: 1217: 1182: 1126: 1082: 1028: 589: 379: 198: 157: 12559: 11544: 9159:

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

8553:

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

8435: 4648: 4627: 4160:

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

3839:

are now understood to be terrestrial predators akin to ground

3573:

may display adaptations for wing-propelled diving like modern

3418:

There was considerable debate whether pterosaurs ambulated as

3261: 2359:

At least some non-pterodactyloid pterosaurs survived into the

2063:. In the 1980s, early cladistic analyses found that they were 14788: 13384: 11327:"The One Born of Fire: a pterosaurological analysis of Rodan" 11070: 8072:"'Superbly preserved' pterosaur fossil unearthed in Scotland" 6428:"Articulation and Function of the Pteroid Bone of Pterosaurs" 4410: 4405: 3910:

proposed to share this feeding habit based on high estimated

3518: 3452: 3423: 3412: 3347: 2953: 2390: 2344:

Small-sized pterosaur species apparently were present in the

2317: 2223: 2211: 1543: 878:

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

597: 414: 11348:"A Monster-Sized Breakdown of Every Insane 'Godzilla' Movie" 10703: 9759:"Pterosaur tracks and the terrestrial ability of pterosaurs" 9324:. Boulder, Colo: Geological Society of America. p. 60. 9206: 7974: 7765:

Sitzungsberichte der Bayerischen Akademie der Wissenschaften

7707: 7697: 7695: 7682: 7680: 7678: 7665: 7663: 6060:

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

6003: 5432:

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

5349: 4636: 3556:

must have foraged while swimming, as they seem incapable of

1215:; the extent of this membrane is not certain, as studies on 1089:, was a straight bar. It was connected to a lower bone, the 503:

had efficient unidirectional "flow-through" breathing using

17553: 17421: 17234: 12456: 10790:"Palaeontology: pterosaur embryo from the Early Cretaceous" 9808:

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

6749: 6747: 6745: 6743: 6741: 6739: 6726: 6724: 6722: 6720: 6718: 6716: 6626: 6616: 6614: 6612: 6610: 6470:

Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen

6334: 6332: 6330: 6241: 5883: 5881: 5879: 5877: 5875: 5862: 5860: 5782: 5780: 5778: 5742:"Taxonomy and systematics of the Late Cretaceous pterosaur 5641:

Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen

5581: 5579: 5035: 4651: 4630: 4368: 4059: 3456: 2292: 1825:

published studies about the wing membranes of specimens of

1449: 1269: 1190:

patagium – a membrane connecting the legs that, unlike the

1177: 1121:

Various configurations proposed for the wings of pterosaurs

585: 492: 10746: 9952: 6059: 5078:

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

4944:

The dinosaur book: the ruling reptiles and their relatives

4897:(2003) , Peter Roach; James Hartmann; Jane Setter (eds.), 3984:

has been interpreted as being attacked or scavenged by an

2335:, several tapejarids and the indeterminate non-azhdarchid 1555:, and the presence of both aktinofibrils and filaments on 17242: 17226: 10326: 10044: 9995: 9158: 9056: 9041:

XXIII Jornadas Argentinas de Paleontología de Vertebrados

8825:

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

8094: 7861:

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

7692: 7675: 7660: 7612:

Dragons of the Air: An account of extinct flying reptiles

7579: 7567: 7555: 7501: 6447:

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

4741:"The Earliest Pterodactyloid and the Origin of the Group" 3716:, in particular, seems to have been a beetle specialist. 2452:, a "primitive" group of long-tailed pterosaurs, and the 2349: 1252: 1191: 1130: 766:

Standing, such giants could reach the height of a modern

569: 479:

Pterosaurs sported coats of hair-like filaments known as

16:

Flying reptiles of the extinct clade or order Pterosauria

11261: 11259: 11257: 9710:

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

9034: 8672:

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

8599:

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

8054: 8052: 7352: 6736: 6713: 6607: 6327: 5872: 5857: 5775: 5721: 5680: 5678: 5676: 5674: 5659: 5576: 5190: 11134:

Proceedings of the Royal Society B: Biological Sciences

11028:

Proceedings of the Royal Society B: Biological Sciences

10697: 10463: 10234: 9513: 9138:

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

8247: 8245: 8190: 7951: 7949: 7947: 7945: 7943: 7175: 7090:

Proceedings of the Royal Society B: Biological Sciences

6948: 6913: 6911: 6909: 6907: 6905: 6785: 6783: 6781: 6779: 6766: 6764: 6762: 6597: 6595: 6593: 6591: 6589: 6587: 6585: 6583: 6374: 6147: 6145: 6132: 6130: 6117: 6115: 5939: 5937: 5912: 5910: 5908: 5847: 5845: 5843: 5830: 5828: 5826: 5813: 5811: 5809: 5807: 5765: 5763: 5630: 5628: 5615: 5613: 5611: 5598: 5596: 5594: 5566: 5564: 5513: 5511: 5509: 5036:

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

3954:

discusses an Early Cretaceous fossil of three cervical

2054:

has been so heavily modified for flight, and immediate

1880:

by John Conway exemplifies the "new look" of pterosaurs

991:

of pterosaurs numbered between thirty-four and seventy

14154: 10934: 10249: 10056: 9934: 9872: 9860: 9848: 9263:"Pterodactyls were too heavy to fly, scientist claims" 8251: 7624:

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

7442:

Journal de Physique, de Chimie et d'Histoire Naturelle

7397: 7395: 7393: 7391: 7342: 7340: 6644: 5739: 5481: 4951: 4459:

In most media appearances, pterosaurs are depicted as

1961:

Even more productive was the Early Cretaceous Chinese

652:

Pterodactyloids include the clades Ornithocheiroidea (

619:

Basal pterosaurs include the clades Dimorphodontidae (

11304: 11254: 11209: 10537: 10457: 9469:"With Wings Flapping, Model Pterodactyl Takes to Air" 9417:"Did giant pterosaurs vault aloft like vampire bats?" 8868:

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

8773: 8684:

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

8622:"Were early pterosaurs inept terrestrial locomotors?" 8581: 8082: 8049: 8040: 8038: 7883:, Ph.D. diss., Department of Biology, Yale University 7376: 6494: 5671: 5077: 4660: 4633: 3972:

have been found with tooth marks from sharks such as

3265:

Skeletal reconstruction of a quadrupedally launching

2267:

is anatomically similar to that of early pterosaurs.

2158:

Two researchers, S. Christopher Bennett in 1996, and

1958:, intercepted the trade and named even more species. 954:

show that even some early pterosaurs possessed them.

612:. Likewise, the sacral vertebrae could form a single 12413: 12307: 12292: 12281: 12271: 12261: 12251: 12241: 12197: 12033: 12018: 12007: 11997: 11987: 11862: 11791: 11776: 11765: 10480: 10408:"Palaeontology: pterosaur egg with a leathery shell" 10406:

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

9709: 9052: 9050: 8769: 8767: 8765: 8242: 7940: 6902: 6890: 6878: 6866: 6854: 6842: 6776: 6759: 6580: 6356: 6344: 6142: 6127: 6112: 6055: 6053: 5934: 5922: 5905: 5893: 5840: 5823: 5804: 5792: 5760: 5625: 5608: 5591: 5561: 5506: 5494: 5350:

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

5345: 5343: 5341: 5339: 4959:"Pterosaur distribution in time and space: an atlas" 4645: 4624: 4493: 4456:(1988), is a notable example from an animated film. 4420:, is portrayed as an enormous irradiated species of 2464:, it has fallen out of favor among most scientists. 2181:. A 2011 archosaur-focused phylogenetic analysis by 355: 347: 336: 325: 317: 305: 295: 287: 279: 271: 11551:"Comments on the phylogeny of the pterodactyloidea" 10075:Wu, Wen-Hao; Zhou, Chang-Fu; Andres, Brian (2017). 9410: 9408: 9321:

Posture, locomotion, and paleoecology of pterosaurs

8818: 7993:Dyke, G.J. McGowan; Nudds, R.L.; Smith, D. (2009). 7787:

Zeitschrift für Anatomie und Entwicklungsgeschichte

7388: 7364: 7337: 6536:Wilkinson M.T.; Unwin D.M.; Ellington C.P. (2006). 6156:and the nature of the pterosaur flight apparatus". 4642: 4621: 3244:

Posture, Locomotion, and Paleoecology of Pterosaurs

3236:

Posture, Locomotion, and Paleoecology of Pterosaurs

1915:

Handbuch der Paläoherptologie, Teil 19: Pterosauria

1662:used its wings as flippers and was affiliated with 13202: 11022:Unwin, David Michael; Deeming, D. Charles (2019). 10999:"First 3D pterosaur eggs found with their parents" 10546:"First occurrence of stomach stones in pterosaurs" 10543: 9381: 9379: 9377: 8774:Longrich, N.R.; Martill, D.M.; Andres, B. (2018). 8293:Bulletin of the American Museum of Natural History 8035: 7726:Marsh, O.C. (1882). "The wings of Pterodactyles". 5999: 5997: 5995: 3653:were likely terrestrial/semiarboreal generalists. 1325:. The wingfinger is also bent somewhat downwards. 14678: 13623: 11127: 10327:Buffetaut E, Martill D, Escuillié F (July 2004). 10299: 9047: 8762: 6529: 6050: 5409:"Pterosaur.net :: Origins and Relationships" 5336: 5191:Buffetaut E, Grigorescu D, Csiki Z (April 2002). 5184: 4255:may have had similar activity patterns to modern 4050:likely pterosaurs buried their eggs, like modern 3847:, eating any prey item they could swallow whole. 3783:obtained food in coastal or freshwater habitats. 3753:, and pteranodonts pelagic plunge-divers akin to 3743:(with the exception of the plunge-diving adapted 3618:, and large insects. Its robust dentition caused 2258:At least one study found that the early Triassic 1839:discovered hair follicles in pterosaur skin, and 515:to the largest known flying creatures, including 17716: 14772: 11861: 10464:Codorniú, L.; Chiappe, L.; Rivarola, D. (2014). 10256:Martill, David M.; Ibrahim, Nizar (March 2015). 10144: 9405: 9132: 9098:Geological Society, London, Special Publications 6244:Geological Society, London, Special Publications 5316:Alexander, David E. & Vogel, Steven (2004). 3997: 3517:had slightly larger feet (47% the length of the 2367:situation for late Cretaceous pterosaur faunas. 2291:It was once thought that competition with early 15981: 15901: 14814: 11515:Pterosaurs: Natural History, Evolution, Anatomy 10930: 10928: 10872: 10870: 10868: 9890: 9374: 7969:Handbuch der Paläoherpetologie XIX. Pterosauria 7415:Acta Theodoro-Palatinae Mannheim 5 Pars Physica 6651:Proceedings of the National Academy of Sciences 6284: 6198: 5992: 5431: 5084:Proceedings of the National Academy of Sciences 5073: 5071: 4875:and its implications for pterosaur phylogeny", 4841:Pterosaurs: Natural History, Evolution, Anatomy 2374: 1693:, "wing-finger". This was in 1815 Latinised to 884:could have over a thousand bristle-like teeth. 12662: 11746: 11586: 11419: 11321: 11319: 10877:Bennett S. C. (1995). "A statistical study of 10399: 10255: 9791: 9789: 9787: 9434: 8683: 8615: 8613: 8611: 8609: 8601:Weltenberger Akademie, Erwin Rutte-Festschrift 7992: 7891: 7889: 7132: 6151: 5710:. The Dinosaur Museum: Blanding, Utah, 15–41. 5639:(Wagner) from the Upper Jurassic of Germany". 5013:"Pterosaur.net :: Terrestrial Locomotion" 4870: 4838: 4734: 4732: 4714:expansion of ecological niches in the Mesozoic 4137:showed possible evidence of their young being 3242:period. However, both Sato and the authors of 572:, some fossils of which have been discovered. 560:, and more broadly to members of the suborder 17211: 12575: 11572: 11413: 10876: 9344: 8389:"Reassessment of the Triassic archosauriform 8106: 7494:, München: mathematisch-physikalische Classe 6790:Witton MP, Naish D (2008). McClain CR (ed.). 6375:Wilkinson MT, Unwin DM, Ellington CP (2006). 6287:"Limb disparity and wing shape in pterosaurs" 4981:"The Extent of the Pterosaur Flight Membrane" 4978: 4940: 4738: 4564:List of pterosaur-bearing stratigraphic units 4375:dinosaurs and not descendants of pterosaurs. 3723:contained many piscivorous taxa, such as the 3525:had very large feet (69% of tibial length in 1048:The shoulder girdle connected to the notarium 16566: 12196: 11021: 10925: 10865: 10074: 9946: 9089: 8928:Prondvai, E.; Bodor, E. R.; Ösi, A. (2014). 7032: 6488: 5708:Feathered Dinosaurs and the Origin of Flight 5684: 5311: 5309: 5068: 3521:), while filter-feeding pterosaurs like the 3451:), in a manner similar to many mammals like 131:Six pterosaurs (top left to bottom right): 15680: 11316: 10594: 10183: 10177: 10150: 9891:Andres, B.; Clark, J. M.; Xing, X. (2010). 9784: 9562: 9387:"Why pterosaurs weren't so scary after all" 9318:Templin, R. J.; Chatterjee, Sankar (2004). 9200: 9179: 8824: 8606: 8119:and the origin of dinosaurs and pterosaurs" 7886: 7407: 6789: 5634: 5060:: CS1 maint: numeric names: authors list ( 4729: 4689: 4683: 3940:Pterosaurs are known to have been eaten by 3320: 2429:more closely related to pterosaurs than to 2014: 1928:The three-dimensionally preserved skull of 773: 17218: 17204: 12738: 12582: 12568: 11579: 11565: 11493: 11409:. Andrews MacMeel Publishing. p. 272. 9750: 9165:. CRC Press Boca Raton, FL. p. 2072. 7980: 7713: 7701: 7686: 7669: 7585: 7573: 7561: 7507: 7433: 7420: 7358: 6942: 6753: 6730: 6632: 6620: 6467: 6461: 6425: 6338: 6285:Dyke GJ, Nudds RL, Rayner JM (July 2006). 5955: 5887: 5866: 5786: 5727: 5665: 5585: 5475: 5130: 4424:. Rodan has appeared in multiple Japanese 4180: 3958:of a pterosaur with the broken tooth of a 3593:Early-on it was recognised that the small 3584: 718:The position of the clade Anurognathidae ( 584:were hollow and air-filled, like those of 369: 122: 17574:Tradeoffs for locomotion in air and water 17225: 11641: 11447: 11284:Evolution and Palaeobiology of Pterosaurs 11161: 11104: 11047: 10974: 10964: 10813: 10764: 10680: 10577: 10512: 10405: 10352: 10302:Zoological Journal of the Linnean Society 10295: 10293: 10291: 10211: 10201: 10168: 10114: 10104: 9636: 9626: 9531: 9440: 9011: 8801: 8791: 8701: 8647: 8637: 8418: 8408: 8363: 8353: 8304: 8175: 8164:Zoological Journal of the Linnean Society 8142: 8010: 7320: 7255: 7206: 7109: 6976:1983/1f7893a1-924d-4cb3-a4bf-c4b1592356e9 6974: 6825: 6815: 6688: 6670: 6563: 6512: 6402: 6302: 6095: 6085: 6033: 5733: 5306: 5289: 5242: 5136: 5113: 5103: 4996: 4972: 4901:, Cambridge: Cambridge University Press, 4815: 4805: 4764: 3978:, and a fossil with tooth marks from the 3859:may have been a specialist molluscivore. 3313:Large-headed species are thought to have 1949:. During the 1970s, the Early Cretaceous 1534:A 2018 study of the remains of two small 1518: 11553:, by Alexander W. A. Kellner (technical) 11345: 10787: 10753:Anais da Academia Brasileira de Ciências 10489:"Argentinian unhatched pterosaur fossil" 10329:"Pterosaurs as part of a spinosaur diet" 9756: 9659: 9565:Annual Review of Ecology and Systematics 9466: 9256: 9254: 7895: 7439: 7286: 6929: 6501:Anais da Academia Brasileira de Ciências 5538: 5536: 5534: 5532: 5530: 5528: 5526: 5285: 5283: 5281: 4414:) which first appeared in the 1956 film 4311: 4292: 4286: 4001: 3466: 3391: 3260: 3217: 3209: 2274: 2138: 2119: 2023: 1923: 1869: 1865: 1734: 1672: 1605: 1500: 1492:(which translates as "hairy demon") and 1436: 1345: 1176: 1136: 1116: 1043: 1009: 973: 890: 832:Reconstruction of crests: three crested 827: 777: 753: 463:of small vertebrates. Later pterosaurs ( 11404: 11275: 9955:Acta Geologica Sinica - English Edition 9807: 9260: 9162:Phylonyms: A Companion to the PhyloCode 9095: 8393:: neither runner nor biped, but hopper" 8386: 8335: 8286: 8157: 7776: 7083: 5702: 5700: 3440:. However, a large number of pterosaur 3298:to build a half-scale working model of 1590: 1498:show pycnofibers on the head and body. 1332:A laser-simulated fluorescence scan on 1097:The breastbone, formed by fused paired 17717: 13611: 11512: 11288:Geological Society Special Publication 10843: 10841: 10650: 10648: 10288: 10062: 10050: 9940: 9884: 9878: 9866: 9854: 9795: 9589: 9362:from the original on February 21, 2009 9291: 8619: 8587: 8280: 8112: 8100: 8088: 8058: 8044: 7955: 7860: 7744: 7623: 7519: 7401: 7382: 7370: 7346: 7237: 7008:"Fur flies over new pterosaur fossils" 7005: 6917: 6896: 6884: 6872: 6860: 6848: 6770: 6601: 6362: 6350: 6136: 6121: 5943: 5928: 5916: 5899: 5851: 5834: 5817: 5798: 5769: 5619: 5602: 5570: 5517: 5500: 5292:"Pterosaurs: Myths and Misconceptions" 5248: 4739:Andres, B.; Clark, J.; Xu, X. (2014). 3914:(BFQ) and absolute bite force values. 2135:theorized to be related to pterosaurs. 1777:, and in 1901 the first popular book, 1472:was first reported from a specimen of 17599: 17598: 17199: 15968: 14760: 14759: 14141: 13610: 12725: 12601: 12563: 11598: 11560: 11474: 11373: 11310: 9414: 9338: 9251: 8552: 7851:, Londen: Penguin Books, 1988, p. 283 7725: 7461:Annales du Musée d'Histoire Naturelle 5542: 5523: 5278: 4947:. New York: McGraw-Hill. p. 153. 4893: 4787: 4010:juvenile from the Solnhofen Limestone 2421:(Sömmerring 1812)". A broader clade, 2302:Cretaceous–Paleogene extinction event 1730: 1056:retractor muscle which in most basal 15969: 10654: 8336:Ezcurra, Martín D. (28 April 2016). 7538: 5697: 4594: 3935: 3828:were likely terrestrial carnivores. 3816:mostly were terrestrial pterosaurs. 3475:trackways show that pterosaurs like 3350:cavities revealed that the animals ( 1172: 874:had recurved teeth for eating meat. 11346:Gonzales, Dave (October 12, 2016). 11184: 10838: 10645: 9590:Anthes, Emily (November 18, 2013). 9577:10.1146/annurev.es.08.110177.002241 9467:Molotsky, Irvin (28 January 1986). 9261:Alleyne, Richard (1 October 2008). 9035:Haluza, A.; Apesteguía, S. (2007). 8530:"Pterosaur Origins Flap into Focus" 8254:Journal of Systematic Palaeontology 5547:. New York: Pi Press. p. 246. 2295:species might have resulted in the 2175:or a branch between the latter and 1904:named a renaissance of pterosaurs. 1225:). A common interpretation is that 969: 825:was relatively large for reptiles. 616:while the pelvic bones fused also. 596:for flight muscles and an enlarged 554:may refer to members of the genus 491:to the down feathers found on both 13: 10550:Journal of Vertebrate Paleontology 9900:Journal of Vertebrate Paleontology 9778:10.1111/j.1502-3931.1996.tb01673.x 9592:"Coldblooded Does Not Mean Stupid" 8177:10.1111/j.1096-3642.1996.tb01267.x 6543:Proceedings of the Royal Society B 6435:Journal of Vertebrate Paleontology 6382:Proceedings of the Royal Society B 6014:Proceedings of the Royal Society B 5441:Journal of Vertebrate Paleontology 4979:Elgin RA, Hone DW, Frey E (2011). 3779:were likely primarily scavengers. 3387: 1147:, preserve the membrane structure 1072: 424:. They existed during most of the 14: 17751: 11532: 11374:Sharf, Zack (December 10, 2018). 10391:. October 3, 2018. Archived from 9294:"Were pterosaurs too big to fly?" 8527: 3341: 2381:List of pterosaur classifications 2285:Arizona Museum of Natural History 17314: 17180: 17174: 17167: 15950: 15942: 15934: 14742: 14736: 14124: 14119: 13593: 13586: 13578: 12544: 12543: 12515: 12508: 12502: 12497: 12491: 11398: 11367: 11339: 11267:"Pterosaurs In Popular Culture." 11203: 11178: 11121: 11064: 11015: 10991: 10781: 10740: 10377: 10320: 10228: 10184:Naish, D.; Witton, M.P. (2017). 10151:Witton, M.P.; Naish, D. (2015). 10131: 10068: 9989: 9975:10.1111/j.1755-6724.2011.00444.x 9801: 9703: 9653: 9602: 9583: 9556: 9507: 9479: 9460: 9415:Hecht, Jeff (16 November 2010). 9311: 9292:Powell, Devin (2 October 2008). 9285: 9152: 9142:. Geological Society of London. 9028: 8971: 8921: 8870:"Systematic reinterpretation of 8861: 8718: 8677: 8664: 8593: 8546: 8521: 8497: 8380: 8329: 8184: 8158:Bennett, S. Christopher (1996). 8151: 8064: 8012:10.1111/j.1420-9101.2008.01682.x 7986: 7961: 7873: 7854: 7841: 7828: 7770: 7757: 7738: 7719: 7617: 7604: 7591: 7532: 7513: 7483: 7470: 7428:Natürliches System der Amphibien 7238:D’Alba, Liliana (January 2019). 6304:10.1111/j.1420-9101.2006.01096.x 6152:Unwin DM, Bakhurina NN (1994). " 5746:(Pterosauria, Pterodactyloidea)" 4617: 4559:Graphical timeline of pterosaurs 4538: 4524: 4510: 4496: 4408:, a fictional giant monster (or 3944:. In the 1 July 2004 edition of 3002: 2982: 2919: 2899: 2814: 2794: 2762: 2722: 2690: 2617: 2588: 2561: 2534: 2504: 1460:(sharing a common origin) with, 1308:In derived pterodactyloids like 1247:, and a pterodactyloid from the 480: 175: 54: 17730:Late Triassic first appearances 14142: 12726: 12589: 9443:"The Great Pterodactyl Project" 8690:Molecular Biology and Evolution 8575:10.1016/j.earscirev.2021.103777 7999:Journal of Evolutionary Biology 7452: 7280: 7231: 7169: 7126: 7077: 7026: 6999: 6932:Nova Acta Academiae Leopoldinae 6923: 6638: 6514:10.1590/s0001-37652010000400024 6419: 6278: 6235: 6192: 5949: 5740:S. Christopher Bennett (1994). 5425: 5401: 5029: 5005: 4706: 4673: 3426:. In the 1980s, paleontologist 3205: 1995:in 2022, was discovered on the 1719:thought pterosaurs were flying 1412: 11517:. Princeton University Press. 7971:, Urban & Fischer, München 7287:Cincotta; et al. (2022). 7244:Nature Ecology & Evolution 7179:Nature Ecology & Evolution 7035:Nature Ecology & Evolution 6955:Nature Ecology & Evolution 5545:The Pterosaurs: From Deep Time 4934: 4927:Merriam-Webster.com Dictionary 4914: 4899:English Pronouncing Dictionary 4887: 4864: 4843:, Princeton University Press, 4832: 4781: 4610: 4601: 4589:Timeline of pterosaur research 4434:Godzilla: King of the Monsters 4319:When Dinosaurs Ruled the Earth 4015:and in Loma del Pterodaustro ( 2472:(family tree) below follows a 2315:, such as the ornithocheirids 1601: 1597:Timeline of pterosaur research 1432: 1197:patagium, leaves the tail free 805:, with the upper jawbone, the 575: 1: 10766:10.1590/0001-3765201520150364 10282:10.1016/j.cretres.2014.11.001 10157:Acta Palaeontologica Polonica 10022:10.1016/j.cretres.2020.104637 9187:The Encyclopedia of Dinosaurs 9118:10.1144/gsl.sp.2003.217.01.10 9075:10.1016/j.cretres.2021.105061 7779:"Das Gehirn der Pterosaurier" 7084:Kellner; et al. (2009). 6264:10.1144/GSL.SP.2003.217.01.14 4985:Acta Palaeontologica Polonica 4722: 3998:Reproduction and life history 3380:show that brain evolution in 3222:Pterosaur flight adaptations. 2270: 2046:similarities with pterosaurs. 1727:coined the term Pterosauria. 1712:Samuel Thomas von Soemmerring 1677:Newman's marsupial pterosaurs 1297:Darwinopterus linglongtaensis 1141:Some specimens, such as this 782:Conical tooth, possibly from 10966:10.1371/journal.pone.0031392 10788:Wang X, Zhou Z (June 2004). 10570:10.1080/02724634.2013.731335 10106:10.1371/journal.pone.0185486 9814:Paläontologische Zeitschrift 8793:10.1371/journal.pbio.2001663 7601:, Cambridge University Press 7006:Briggs, Helen (2018-12-17). 6817:10.1371/journal.pone.0002271 6087:10.1371/journal.pone.0004497 5162:10.1126/science.187.4180.947 3361:) had massive flocculi. The 3346:An X-ray study of pterosaur 3317:in order to better balance. 2375:Classification and phylogeny 1856:David Meredith Seares Watson 1382: 25:Pterodactyl (disambiguation) 7: 11479:. McFarland & Company. 7728:American Journal of Science 6482:10.1127/0077-7749/2011/0131 4532:Evolutionary biology portal 4489: 3795:were piscivores, while the 3546: 3315:forwardly swept their wings 2005:National Museum of Scotland 1634:. The pieces from one such 1578:Tupandactylus cf. imperator 1558:Jeholopterus ningchengensis 1531:specimens too fundamental. 1475:Scaphognathus crassirostris 1448:Most or all pterosaurs had 10: 17756: 17466: 17430:Flying and gliding animals 17266:Fin and flipper locomotion 11588:Avemetatarsalia / Pan-Aves 11494:Wellnhofer, Peter (1991). 11468: 11272:, Accessed 27 August 2010. 11097:10.1038/s41598-021-92499-z 10314:10.1093/zoolinnean/zlaa163 8984:Royal Society Open Science 8748:10.1666/0094-8373-35.3.432 7838:, October 1914, pp. 324–35 7313:10.1038/s41586-022-04622-3 7199:10.1038/s41559-020-01309-8 7055:10.1038/s41559-020-01308-9 5653:10.1127/njgpa/210/1998/421 5322:. JHU Press. p. 191. 4788:Baron, Matthew G. (2020). 4554:Flying and gliding animals 2425:, has been defined as all 2378: 2279:Reconstructed skeleton of 2019: 1919:Encyclopedia of Pterosaurs 1610:Engraving of the original 1594: 1495:Jeholopterus ninchengensis 858:In some cases, fossilized 747: 637:), and Rhamphorhynchidae ( 417:of flying reptiles in the 378:Distribution of pterosaur 18: 17735:Maastrichtian extinctions 17607: 17541: 17480: 17420: 17323: 17312: 17241: 17164: 17066: 17016: 16948: 16939: 16911: 16887: 16876: 16832: 16797: 16787: 16683: 16621: 16612: 16608: 16597: 16562: 16521: 16486: 16450: 16441: 16407: 16370: 16361: 16334: 16330: 16307: 16251: 16222: 16218: 16207: 16149: 16120: 16111: 16088: 16079: 16020: 16011: 15996: 15992: 15977: 15964: 15931: 15897: 15781: 15739: 15649: 15640: 15602: 15540: 15531: 15516: 15507: 15446: 15408: 15397: 15346: 15309: 15300: 15253: 15244: 15172: 15157: 15148: 15120: 15111: 15020: 14997: 14986: 14975: 14923: 14912: 14877: 14823: 14810: 14799: 14783: 14768: 14755: 14733: 14703: 14687: 14674: 14613: 14598: 14589: 14540: 14512: 14419: 14408: 14364: 14341: 14332: 14280: 14276: 14265: 14243: 14232: 14184: 14169: 14165: 14150: 14137: 14117: 14075: 14002: 13993: 13939: 13889: 13874: 13865: 13854: 13831: 13820: 13758: 13743: 13734: 13712: 13703: 13673: 13669: 13658: 13634: 13619: 13606: 13575: 13525: 13475: 13465: 13417: 13367: 13358: 13301: 13292: 13253: 13244: 13233: 13198: 13187: 13171: 13159: 13096: 13081: 13077: 13066: 13050: 13038: 12998: 12982: 12973: 12931: 12922: 12890: 12879: 12836: 12805: 12795: 12768: 12753: 12749: 12734: 12721: 12689: 12671: 12658: 12597: 12528: 12489: 12438: 12412: 12403: 12358: 12318: 12306: 12237: 12226: 12192: 12181: 12104: 12045: 12032: 11983: 11979: 11968: 11937: 11873: 11857: 11846: 11790: 11761: 11757: 11742: 11723: 11695: 11677: 11668: 11650: 11637: 11594: 11449:10.1016/j.cub.2021.03.030 10903:10.1017/S0022336000034946 10726:10.1080/08912960701189800 10682:10.1016/j.cub.2014.04.054 10077:"The toothless pterosaur 9920:10.1080/02724630903409220 9736:10.1017/S0016756802006647 9689:10.1080/10420940390255501 9450:Engineering & Science 9229:10.1007/s00114-008-0397-5 8847:10.1017/S1755691013000303 8475:10.1038/s41586-020-3011-4 8266:10.1017/S1477201907002064 7918:10.1017/S009483730000765X 7257:10.1038/s41559-018-0767-0 6967:10.1038/s41559-018-0728-7 5978:10.1080/10292380009380572 5461:10.1080/02724630903409220 5220:10.1007/s00114-002-0307-1 4766:10.1016/j.cub.2014.03.030 4229:have been inferred to be 4207:have been inferred to be 3885:Tropeognathus mesembrinus 3714:Darwinopterus robustidens 3405:Haenamichnus uhangriensis 3352:Rhamphorhynchus muensteri 3226: 2997: 2977: 2970: 2950: 2943: 2914: 2894: 2887: 2877: 2857: 2850: 2809: 2787: 2780: 2757: 2750: 2740: 2715: 2708: 2685: 2678: 2661: 2654: 2639: 2632: 2610: 2603: 2583: 2576: 2556: 2549: 2529: 2522: 2499: 2492: 1783:St. George Jackson Mivart 1651:Cosimo Alessandro Collini 1575:In 2022, a new fossil of 1341: 1018:was longer than the torso 588:. This provided a higher 393: 386: 377: 368: 268: 263: 172:Scientific classification 170: 130: 121: 34: 11477:The Dinosaur Filmography 11185:Bristol, University of. 10389:Science & Innovation 8674:at 22', Tim Haines, 1999 8620:Witton, Mark P. (2015). 7836:The Aeronautical Journal 7476:Rafinesque, C.S., 1815, 6201:Chinese Science Bulletin 5543:Unwin, David M. (2006). 5484:Acta Geoscientica Sinica 4877:Acta Geoscientica Sinica 4871:David M. Unwin (2010), " 4569:List of pterosaur genera 4400:In the latter, animator 4337:or (non-pterodactyloid) 4239:has been inferred to be 4185:Comparisons between the 4046:Hamipterus tianshanensis 3321:Air sacs and respiration 3301:Quetzalcoatlus northropi 3288:Johns Hopkins University 3284:University of Portsmouth 2015:Evolution and extinction 1169:into the wing membrane. 1112: 774:Skull, teeth, and crests 14157:Archaeopterodactyloidea 11475:Berry, Mark F. (2005). 11405:Mansour, David (2005). 11331:Journal of Geek Studies 11232:10.1126/science.1200043 10883:Journal of Paleontology 10623:10.1126/science.1197323 8220:10.1126/science.1143325 7417:, pp. 58–103 (1 plate). 6672:10.1073/pnas.2107631118 5105:10.1073/pnas.0707728105 4839:Mark P. Witton (2013), 4574:Phylogeny of pterosaurs 4361:-like crest and teeth: 4347:-like crests and long, 4181:Daily activity patterns 4097:flaplings found in the 3781:Archaeopterodactyloidea 3585:Diet and feeding habits 3384:was a modular process. 2747:Archaeopterodactyloidea 2199:which argued that both 2167:between pterosaurs and 1858:, but they interpreted 851:Tupandactylus imperator 819:nasoantorbital fenestra 743: 625:), Campylognathididae ( 17569:Terrestrial locomotion 17513:Evolution of cetaceans 17508:Origin of avian flight 17493:Evolution of tetrapods 11545:The Pterosaur Database 11146:10.1098/rspb.2023.1102 11040:10.1098/rspb.2019.0409 10655:Wang, Xiaolin (2014). 10170:10.4202/app.00005.2013 9441:MacCready, P. (1985). 9391:The Observer newspaper 8387:Bennett, S.C. (2020). 8287:Nesbitt, S.J. (2011). 8135:10.1098/rstb.1999.0489 7967:Wellnhofer, P., 1978, 7847:Bakker, Robert, 1986, 7102:10.1098/rspb.2009.0846 6556:10.1098/rspb.2005.3278 6395:10.1098/rspb.2005.3278 6026:10.1098/rspb.2009.0846 4690: 4684: 4397:One Million Years B.C. 4328: 4322:depicting an outsized 4305: 4283:between these genera. 4011: 3900:was corroborated as a 3809:-like filter-feeding. 3493:The forelimb bones of 3482: 3408: 3271: 3223: 3215: 2288: 2246:for incipient flight. 2155: 2136: 2067:(archosaurs closer to 2047: 1942: 1881: 1744: 1678: 1621: 1613:Pterodactylus antiquus 1519:Relation with feathers 1508: 1445: 1351: 1291:Changchengopterus pani 1198: 1148: 1122: 1049: 1019: 984: 901: 855: 845:Tupandactylus navigans 836:. From top to bottom: 788: 759: 694:), and Azhdarchoidea ( 682:), Dsungaripteroidea ( 670:), Ctenochasmatoidea ( 511:, from the very small 440:known to have evolved 23:. For other uses, see 17697:Paleobiology Database 17584:Undulatory locomotion 17533:Homologous structures 11513:Witton, Mark (2013). 11325:Thomas, H.N. (2020). 10139:Thalassodromeus sethi 9520:The Anatomical Record 9348:(February 18, 2009). 8703:10.1093/molbev/msj124 8555:Earth-Science Reviews 8391:Scleromochlus taylori 8117:Scleromochlus taylori 8113:Benton, M.J. (1999). 8076:Associated Press (AP) 7849:The Dinosaur Heresies 4998:10.4202/app.2009.0145 4315: 4296: 4287:Cultural significance 4128:, similar to that of 4005: 3470: 3395: 3366:of total brain mass. 3264: 3221: 3213: 2387:phylogenetic taxonomy 2379:Further information: 2278: 2142: 2123: 2089:basal archosauriforms 2027: 1927: 1873: 1866:Pterosaur renaissance 1852:Ernest Hanbury Hankin 1790:Othniel Charles Marsh 1738: 1676: 1609: 1538:-age pterosaurs from 1504: 1480:Georg August Goldfuss 1444:preserved pycnofibers 1440: 1367:fused with the broad 1349: 1180: 1140: 1120: 1047: 1013: 977: 894: 831: 781: 757: 17528:Analogous structures 17523:Convergent evolution 13769:Archaeoistiodactylus 13679:Archaeoistiodactylus 11434:(11): 2429–2436.e7. 7610:Seeley, H.G., 1901, 7597:Seeley, H.G., 1870, 7430:Munich, 1830: 1–354. 6006:"The soft tissue of 4453:The Land Before Time 4386:1925 film adaptation 4164:pterosaurs, such as 4080:scans of fossilised 3917:Tapejara wellnhoferi 3912:bite force quotients 3268:Pteranodon longiceps 2165:convergent evolution 2143:Life restoration of 2124:Life restoration of 2087:. A placement among 2028:Life restoration of 1886:dinosaur renaissance 1591:History of discovery 1466:convergent evolution 1464:hair, an example of 839:Tapejara wellnhoferi 536:last common ancestor 17579:Rotating locomotion 17518:Comparative anatomy 16834:Targaryendraconidae 16523:Ornithocheiriformes 16210:Ornithocheiromorpha 15509:Neopterodactyloidea 15436:Dsungaripteromorpha 13204:Campylognathoididae 12924:Eudimorphodontoidea 11547:, by Paul Pursglove 11440:2021CBio...31E2429Z 11224:2011Sci...332..705S 11089:2021NatSR..1113130N 10957:2012PLoSO...731392P 10895:1995JPal...69..569B 10806:2004Natur.429..621W 10718:2007HBio...19..273G 10673:2014CBio...24.1323W 10615:2011Sci...331..321L 10562:2013JVPal..33..647C 10427:2004Natur.432..572J 10395:on October 3, 2018. 10345:2004Natur.429...33B 10274:2015CrRes..53...59M 10262:Cretaceous Research 10097:2017PLoSO..1285486W 10014:2021CrRes.11704637M 10002:Cretaceous Research 9967:2011AcGlS..85..507L 9912:2010JVPal..30..163A 9826:2007PalZ...81..376B 9810:Anurognathus ammoni 9728:2002GeoM..139..421H 9716:Geological Magazine 9681:2003Ichno..10..115P 9221:2008NW.....95..891L 9209:Naturwissenschaften 9110:2003GSLSP.217..105K 9063:Cretaceous Research 9004:10.1098/rsos.160333 8996:2016RSOS....360333M 8949:2014Pbio...40..288P 8839:2012EESTR.103..383A 8740:2009Pbio...35..432B 8567:2021ESRv..22103777B 8534:Scientific American 8467:2020Natur.588..445E 8212:2007Sci...317..358I 7910:1983Pbio....9..218P 7777:Edinger, T (1927). 7638:1881Natur..24..335. 7426:Wagler, J. (1830). 7305:2022Natur.604..684C 7240:"Pterosaur plumage" 7191:2020NatEE...4.1592Y 7147:1994Natur.371...62U 7047:2020NatEE...4.1590U 6808:2008PLoSO...3.2271W 6663:2021PNAS..11807631P 6657:(44): e2107631118. 6426:Bennett SC (2007). 6256:2003GSLSP.217..233F 6213:2002ChSBu..47..226W 6170:1994Natur.371...62U 6078:2009PLoSO...4.4497C 5970:2000HBio...14..255B 5637:Pterodactylus kochi 5453:2010JVPal..30..163A 5379:10.1038/nature02048 5371:2003Natur.425..950W 5212:2002NW.....89..180B 5200:Naturwissenschaften 5154:1975Sci...187..947L 5096:2008PNAS..105.1983W 4849:2013pnhe.book.....W 4757:2014CBio...24.1011A 4546:Paleontology portal 4371:, though birds are 4099:Solnhofen limestone 4029:Lagarcito Formation 4017:Lagarcito Formation 3929:Tupuxuara leonardii 3923:Caupedactylus ybaka 3735:and Nyctosauridae. 3702:-like generalists. 3000:Ornithocheiromorpha 2559:Campylognathoididae 2346:Csehbánya Formation 2056:transitional fossil 1976:quantitative method 1931:Anhanguera santanae 1792:in 1870 discovered 1771:Harry Govier Seeley 1766:Cambridge Greensand 1739:Seeley's dynamical 1656:Johann Georg Wagler 1643:Solnhofen Limestone 978:A neck vertebra of 815:antorbital openings 564:of the pterosaurs. 495:and some non-avian 17498:Evolution of birds 17251:Aquatic locomotion 16363:Istiodactyliformes 14837:Lonchognathosaurus 14680:Eupterodactyloidea 13823:Pterodactyliformes 13626:Pterodactylomorpha 13613:Pterodactylomorpha 13527:Pterodactylomorpha 13369:Angustinaripterini 13000:Eudimorphodontinae 12975:Eudimorphodontidae 11498:. Crescent Books. 11077:Scientific Reports 10706:Historical Biology 10203:10.7717/peerj.2908 10079:Jidapterus edentus 10053:, pp. 150–51. 9834:10.1007/bf02990250 9628:10.7717/peerj.2311 9596:The New York Times 9273:on 31 October 2009 9189:. Academic Press. 8893:10.5252/g2012n4a10 8639:10.7717/peerj.1018 8410:10.7717/peerj.8418 8355:10.7717/peerj.1778 8103:, pp. 14, 17. 7879:Padian, K., 1980, 7800:10.1007/bf02117933 5958:Historical Biology 5250:Benton, Michael J. 4930:. Merriam-Webster. 4807:10.7717/peerj.9604 4329: 4306: 4281:niche partitioning 4279:may also indicate 4172:determinate growth 4012: 3981:Toolebuc formation 3767:found clusters of 3737:Niche partitioning 3610:, envisioned as a 3483: 3459:. Footprints from 3409: 3286:and Mike Habib of 3272: 3224: 3216: 3136:Pterodactylomorpha 3098:Pterodactyliformes 2847:Eupterodactyloidea 2450:Rhamphorhynchoidea 2289: 2244:selection pressure 2156: 2137: 2050:Because pterosaur 2048: 1943: 1882: 1823:Karl Alfred Zittel 1779:Dragons of the Air 1745: 1731:Expanding research 1710:In 1812 and 1817, 1703:or members of the 1679: 1622: 1509: 1446: 1352: 1227:non-pterodactyloid 1199: 1149: 1123: 1050: 1020: 985: 902: 856: 789: 760: 501:respiratory system 432:to the end of the 17712: 17711: 17601:Taxon identifiers 17592: 17591: 17549:Animal locomotion 17488:Evolution of fish 17368:facultative biped 17193: 17192: 17189: 17188: 17162: 17161: 17158: 17157: 17154: 17153: 17150: 17149: 17146: 17145: 17142: 17141: 17138: 17137: 17134: 17133: 17130: 17129: 16872: 16871: 16868: 16867: 16789:Targaryendraconia 16783: 16782: 16593: 16592: 16517: 16516: 16513: 16512: 16303: 16302: 16203: 16202: 16199: 16198: 16195: 16194: 16191: 16190: 15960: 15959: 15929: 15928: 15925: 15924: 15921: 15920: 15893: 15892: 15889: 15888: 15885: 15884: 15881: 15880: 15877: 15876: 15873: 15872: 15869: 15868: 15865: 15864: 15636: 15635: 15604:Chaoyangopterinae 15533:Chaoyangopteridae 15393: 15392: 15389: 15388: 15385: 15384: 15381: 15380: 15377: 15376: 15373: 15372: 14908: 14907: 14904: 14903: 14775:Ornithocheiroidea 14762:Ornithocheiroidea 14751: 14750: 14731: 14730: 14727: 14726: 14723: 14722: 14705:Ornithocheiroidea 14670: 14669: 14666: 14665: 14662: 14661: 14658: 14657: 14654: 14653: 14468:Gladocephaloideus 14404: 14403: 14400: 14399: 14366:Aurorazhdarchidae 14268:Ctenochasmatoidea 14228: 14227: 14186:Germanodactylidae 14133: 14132: 14115: 14114: 14111: 14110: 14107: 14106: 14103: 14102: 14099: 14098: 14095: 14094: 13989: 13988: 13964:Dendrorhynchoides 13941:Batrachognathinae 13907:Dendrorhynchoides 13837:Changchengopterus 13816: 13815: 13812: 13811: 13808: 13807: 13602: 13601: 13573: 13572: 13569: 13568: 13565: 13564: 13561: 13560: 13557: 13556: 13553: 13552: 13549: 13548: 13545: 13544: 13461: 13460: 13457: 13456: 13453: 13452: 13378:Angustinaripterus 13294:Rhamphorhynchinae 13246:Rhamphorhynchidae 13229: 13228: 13220:Campylognathoides 13155: 13154: 13034: 13033: 13030: 13029: 13026: 13025: 12933:Raeticodactylidae 12875: 12874: 12871: 12870: 12838:Austriadraconidae 12717: 12716: 12713: 12712: 12709: 12708: 12557: 12556: 12524: 12523: 12487: 12486: 12483: 12482: 12479: 12478: 12475: 12474: 12471: 12470: 12467: 12466: 12399: 12398: 12222: 12221: 12177: 12176: 12173: 12172: 11964: 11963: 11960: 11959: 11842: 11841: 11738: 11737: 11734: 11733: 11719: 11718: 11715: 11714: 11524:978-0-691-15061-1 11505:978-0-517-03701-0 11486:978-0-7864-2453-5 11265:Hone, D. (2010). 11191:www.bristol.ac.uk 10937:"Life history of 10499:(7017): 571–572. 9757:Unwin DM (1997). 9660:Padian K (2003). 9331:978-0-8137-2376-1 8670:BBC Documentary: 8461:(7838): 445–449. 8129:(1388): 1423–46. 7983:, pp. 1–192. 7747:Palaeontographica 7716:, pp. 37–38. 7614:, Londen: Methuen 7539:Kaup, J. (1834). 7299:(7907): 684–688. 7185:(12): 1592–1593. 7041:(12): 1590–1591. 6635:, pp. 53–54. 5554:978-0-13-146308-0 5329:978-0-8018-8059-9 5271:978-0-520-24209-8 5148:(4180): 947–948. 4908:978-3-12-539683-8 4858:978-0-691-15061-1 4595:Explanatory notes 4443:Jurassic Park III 4146:rhamphorhynchoids 4023:). The eggs from 3950:, paleontologist 3936:Natural predators 3761:. An analysis of 3662:Rhamphorhynchidae 3621:Campylognathoides 3201: 3200: 3192: 3191: 3183: 3182: 3164: 3163: 3145: 3144: 3126: 3125: 3107: 3106: 3088: 3087: 3069: 3068: 3060: 3059: 3051: 3050: 3042: 3041: 3033: 3032: 3024: 3023: 3015: 3014: 2932: 2931: 2874:Ornithocheiroidea 2836: 2835: 2827: 2826: 2812:Ctenochasmatoidea 2760:Germanodactylidae 2665:Changchengopterus 2586:Rhamphorhynchidae 2065:Avemetatarsalians 1992:Dearc sgiathanach 1956:Alexander Kellner 1951:Santana Formation 1936:Santana Formation 1725:Johann Jakob Kaup 1454:Alexander Kellner 1249:Santana Formation 1173:Parts of the wing 474:sexual dimorphism 472:head crests with 453:rhamphorhynchoids 408: 407: 403: 259: 17747: 17740:Pterosauromorpha 17705: 17704: 17692: 17691: 17679: 17678: 17666: 17665: 17653: 17652: 17643: 17642: 17641: 17628: 17627: 17626: 17596: 17595: 17559:Robot locomotion 17333:Limb development 17318: 17291:Lobe-finned fish 17220: 17213: 17206: 17197: 17196: 17184: 17178: 17172: 17171: 17018:Coloborhynchinae 16946: 16945: 16901:Ornithocheiridae 16893:Brasileodactylus 16885: 16884: 16795: 16794: 16685:Ornithocheirinae 16619: 16618: 16614:Ornithocheiridae 16610: 16609: 16606: 16605: 16564: 16563: 16560: 16559: 16539:Brasileodactylus 16448: 16447: 16368: 16367: 16336:Lonchodraconidae 16332: 16331: 16328: 16327: 16220: 16219: 16216: 16215: 16118: 16117: 16086: 16085: 16081:Nyctosauromorpha 16018: 16017: 15994: 15993: 15990: 15989: 15984:Pteranodontoidea 15979: 15978: 15971:Pteranodontoidea 15966: 15965: 15955: 15954: 15947: 15946: 15939: 15938: 15903:Pteranodontoidea 15899: 15898: 15783:Quetzalcoatlinae 15772:Mistralazhdarcho 15737: 15736: 15678: 15677: 15647: 15646: 15538: 15537: 15514: 15513: 15406: 15405: 15307: 15306: 15251: 15250: 15170: 15169: 15155: 15154: 15118: 15117: 15022:Thalassodromidae 14995: 14994: 14984: 14983: 14921: 14920: 14879:Dsungaripterinae 14821: 14820: 14816:Dsungaripteridae 14812: 14811: 14808: 14807: 14781: 14780: 14770: 14769: 14757: 14756: 14746: 14741: 14740: 14685: 14684: 14676: 14675: 14596: 14595: 14591:Ctenochasmatinae 14573:Plataleorhynchus 14417: 14416: 14411:Ctenochasmatidae 14339: 14338: 14278: 14277: 14274: 14273: 14241: 14240: 14235:Euctenochasmatia 14167: 14166: 14163: 14162: 14152: 14151: 14139: 14138: 14128: 14123: 14051:Propterodactylus 14037:Ningchengopterus 14000: 13999: 13995:Pterodactyloidea 13872: 13871: 13863: 13862: 13829: 13828: 13741: 13740: 13710: 13709: 13671: 13670: 13667: 13666: 13632: 13631: 13621: 13620: 13608: 13607: 13597: 13591: 13590: 13583: 13582: 13501:Jianchangopterus 13487:Fenghuangopterus 13473: 13472: 13419:Rhamphorhynchini 13365: 13364: 13299: 13298: 13251: 13250: 13242: 13241: 13200: 13199: 13196: 13195: 13169: 13168: 13098:Dimorphodontidae 13079: 13078: 13075: 13074: 13048: 13047: 12980: 12979: 12929: 12928: 12888: 12887: 12803: 12802: 12751: 12750: 12747: 12746: 12736: 12735: 12723: 12722: 12669: 12668: 12660: 12659: 12653: 12652: 12648:Pterosauromorpha 12599: 12598: 12584: 12577: 12570: 12561: 12560: 12547: 12546: 12519: 12513: 12512: 12506: 12501: 12495: 12415: 12410: 12409: 12316: 12315: 12309: 12294: 12283: 12273: 12263: 12253: 12243: 12235: 12234: 12199: 12194: 12193: 12190: 12189: 12106:Sulcimentisauria 12043: 12042: 12035: 12020: 12009: 11999: 11989: 11981: 11980: 11977: 11976: 11871: 11870: 11866:Pterosauromorpha 11864: 11859: 11858: 11855: 11854: 11793: 11778: 11767: 11759: 11758: 11755: 11754: 11744: 11743: 11685:Archosauromorpha 11675: 11674: 11670:Archosauromorpha 11648: 11647: 11639: 11638: 11632: 11631: 11596: 11595: 11581: 11574: 11567: 11558: 11557: 11528: 11509: 11490: 11462: 11461: 11451: 11417: 11411: 11410: 11402: 11396: 11395: 11393: 11391: 11380: 11371: 11365: 11364: 11362: 11360: 11343: 11337: 11323: 11314: 11308: 11302: 11279: 11273: 11263: 11252: 11251: 11218:(6030): 705–08. 11207: 11201: 11200: 11198: 11197: 11182: 11176: 11175: 11165: 11125: 11119: 11118: 11108: 11068: 11062: 11061: 11051: 11019: 11013: 11012: 11010: 11009: 10995: 10989: 10988: 10978: 10968: 10932: 10923: 10922: 10874: 10863: 10862: 10860: 10859: 10845: 10836: 10835: 10817: 10785: 10779: 10778: 10768: 10759:(3): 1599–1609. 10744: 10738: 10737: 10701: 10695: 10694: 10684: 10652: 10643: 10642: 10609:(6015): 321–24. 10598: 10592: 10591: 10581: 10541: 10535: 10534: 10516: 10484: 10478: 10477: 10461: 10455: 10454: 10412: 10403: 10397: 10396: 10381: 10375: 10374: 10356: 10324: 10318: 10317: 10297: 10286: 10285: 10253: 10247: 10246: 10232: 10226: 10225: 10215: 10205: 10181: 10175: 10174: 10172: 10148: 10142: 10135: 10129: 10128: 10118: 10108: 10072: 10066: 10060: 10054: 10048: 10042: 10041: 9993: 9987: 9986: 9950: 9944: 9938: 9932: 9931: 9897: 9888: 9882: 9876: 9870: 9864: 9858: 9852: 9846: 9845: 9805: 9799: 9793: 9782: 9781: 9763: 9754: 9748: 9747: 9707: 9701: 9700: 9666: 9657: 9651: 9650: 9640: 9630: 9606: 9600: 9599: 9587: 9581: 9580: 9560: 9554: 9553: 9535: 9533:10.1002/ar.22839 9511: 9505: 9504: 9502: 9500: 9491: 9483: 9477: 9476: 9464: 9458: 9457: 9447: 9438: 9432: 9431: 9429: 9427: 9412: 9403: 9402: 9400: 9398: 9393:. 11 August 2013 9383: 9372: 9371: 9369: 9367: 9342: 9336: 9335: 9315: 9309: 9308: 9306: 9304: 9289: 9283: 9282: 9280: 9278: 9269:. Archived from 9258: 9249: 9248: 9204: 9198: 9183: 9177: 9176: 9156: 9150: 9136: 9130: 9129: 9093: 9087: 9086: 9054: 9045: 9044: 9032: 9026: 9025: 9015: 8975: 8969: 8968: 8934: 8925: 8919: 8918: 8916: 8915: 8909: 8903:. Archived from 8878: 8872:Piksi barbarulna 8865: 8859: 8858: 8822: 8816: 8815: 8805: 8795: 8771: 8760: 8759: 8722: 8716: 8715: 8705: 8681: 8675: 8668: 8662: 8661: 8651: 8641: 8617: 8604: 8597: 8591: 8585: 8579: 8578: 8550: 8544: 8543: 8541: 8540: 8525: 8519: 8518: 8516: 8515: 8501: 8495: 8494: 8452: 8442: 8433: 8432: 8422: 8412: 8384: 8378: 8377: 8367: 8357: 8333: 8327: 8326: 8308: 8284: 8278: 8277: 8249: 8240: 8239: 8206:(5836): 358–61. 8197: 8188: 8182: 8181: 8179: 8155: 8149: 8148: 8146: 8110: 8104: 8098: 8092: 8086: 8080: 8079: 8068: 8062: 8056: 8047: 8042: 8033: 8032: 8014: 7990: 7984: 7978: 7972: 7965: 7959: 7953: 7938: 7937: 7893: 7884: 7877: 7871: 7870: 7858: 7852: 7845: 7839: 7832: 7826: 7825: 7823: 7822: 7816: 7810:. Archived from 7783: 7774: 7768: 7761: 7755: 7754: 7742: 7736: 7735: 7723: 7717: 7711: 7705: 7699: 7690: 7684: 7673: 7667: 7658: 7657: 7646:10.1038/024335f0 7621: 7615: 7608: 7602: 7595: 7589: 7583: 7577: 7571: 7565: 7559: 7553: 7552: 7536: 7530: 7529: 7517: 7511: 7505: 7499: 7487: 7481: 7474: 7468: 7456: 7450: 7449: 7437: 7431: 7424: 7418: 7411: 7405: 7399: 7386: 7380: 7374: 7368: 7362: 7356: 7350: 7344: 7335: 7334: 7324: 7284: 7278: 7277: 7259: 7235: 7229: 7228: 7210: 7173: 7167: 7166: 7155:10.1038/371062a0 7130: 7124: 7123: 7113: 7096:(1679): 321–29. 7081: 7075: 7074: 7030: 7024: 7023: 7021: 7020: 7003: 6997: 6996: 6978: 6946: 6940: 6939: 6927: 6921: 6915: 6900: 6894: 6888: 6882: 6876: 6870: 6864: 6858: 6852: 6846: 6840: 6839: 6829: 6819: 6787: 6774: 6768: 6757: 6751: 6734: 6728: 6711: 6710: 6692: 6674: 6642: 6636: 6630: 6624: 6618: 6605: 6599: 6578: 6577: 6567: 6550:(1582): 119–26. 6533: 6527: 6526: 6516: 6492: 6486: 6485: 6465: 6459: 6458: 6432: 6423: 6417: 6416: 6406: 6389:(1582): 119–26. 6372: 6366: 6360: 6354: 6348: 6342: 6336: 6325: 6324: 6306: 6282: 6276: 6275: 6239: 6233: 6232: 6221:10.1360/02tb9054 6196: 6190: 6189: 6178:10.1038/371062a0 6149: 6140: 6134: 6125: 6119: 6110: 6109: 6099: 6089: 6057: 6048: 6047: 6037: 6020:(1679): 321–29. 6001: 5990: 5989: 5953: 5947: 5941: 5932: 5926: 5920: 5914: 5903: 5897: 5891: 5885: 5870: 5864: 5855: 5849: 5838: 5832: 5821: 5815: 5802: 5796: 5790: 5784: 5773: 5767: 5758: 5757: 5737: 5731: 5725: 5719: 5704: 5695: 5694: 5682: 5669: 5663: 5657: 5656: 5632: 5623: 5617: 5606: 5600: 5589: 5583: 5574: 5568: 5559: 5558: 5540: 5521: 5515: 5504: 5498: 5492: 5491: 5479: 5473: 5472: 5438: 5429: 5423: 5422: 5420: 5419: 5405: 5399: 5398: 5365:(6961): 950–53. 5356: 5347: 5334: 5333: 5313: 5304: 5303: 5301: 5299: 5287: 5276: 5275: 5259: 5246: 5240: 5239: 5197: 5188: 5182: 5181: 5134: 5128: 5127: 5117: 5107: 5075: 5066: 5065: 5059: 5051: 5049: 5048: 5033: 5027: 5026: 5024: 5023: 5009: 5003: 5002: 5000: 4976: 4970: 4969: 4963: 4955: 4949: 4948: 4938: 4932: 4931: 4918: 4912: 4911: 4891: 4885: 4884: 4868: 4862: 4861: 4836: 4830: 4829: 4819: 4809: 4785: 4779: 4778: 4768: 4736: 4716: 4710: 4704: 4703: 4700: 4697: 4693: 4687: 4677: 4671: 4667: 4663: 4658: 4657: 4654: 4653: 4650: 4647: 4644: 4639: 4638: 4635: 4632: 4629: 4626: 4623: 4614: 4608: 4605: 4548: 4543: 4542: 4541: 4534: 4529: 4528: 4520: 4518:Dinosaurs portal 4515: 4514: 4513: 4506: 4501: 4500: 4499: 4101:of Germany, and 3826:Thalassodromidae 3822:Dsungaripteridae 3805:was adapted for 3797:Ctenochasmatidae 3523:ctenochasmatoids 3006: 2986: 2973: 2972: 2967:Pteranodontoidea 2946: 2945: 2923: 2903: 2890: 2889: 2880: 2879: 2853: 2852: 2818: 2798: 2783: 2782: 2777:Euctenochasmatia 2766: 2753: 2752: 2743: 2742: 2726: 2711: 2710: 2705:Pterodactyloidea 2694: 2681: 2680: 2657: 2656: 2635: 2634: 2621: 2606: 2605: 2592: 2579: 2578: 2565: 2552: 2551: 2538: 2525: 2524: 2508: 2495: 2494: 2485: 2484: 2454:Pterodactyloidea 2423:Pterosauromorpha 2363:, postulating a 2287:in Mesa Arizona. 2218:showing various 2183:Sterling Nesbitt 2061:Archosauromorpha 1911:Peter Wellnhofer 1874:This drawing of 1841:paleoneurologist 1837:Ferdinand Broili 1763:Early Cretaceous 1705:Pterodactyloidea 1618:Egid Verhelst II 1257:flying squirrels 989:vertebral column 970:Vertebral column 886:Dsungaripteridae 876:Ctenochasmatidae 854:(drawn to scale) 813:, the nasal and 799:Pterodactyloidea 634:Campyognathoides 562:Pterodactyloidea 428:: from the Late 398: 373: 357: 349: 338: 327: 319: 307: 297: 289: 281: 273: 254: 249: 238:Pterosauromorpha 236: 180: 179: 126: 116: 53: 38:Temporal range: 32: 31: 17755: 17754: 17750: 17749: 17748: 17746: 17745: 17744: 17715: 17714: 17713: 17708: 17700: 17695: 17687: 17682: 17674: 17669: 17661: 17656: 17648: 17646: 17637: 17636: 17631: 17622: 17621: 17616: 17603: 17593: 17588: 17537: 17503:Origin of birds 17476: 17416: 17338:Limb morphology 17319: 17310: 17296:Ray-finned fish 17261:Fish locomotion 17237: 17224: 17194: 17185: 17166: 17126: 17107:Liaoningopterus 17062: 17012: 16950:Tropeognathinae 16935: 16907: 16881: 16864: 16828: 16799:Cimoliopteridae 16779: 16679: 16602: 16589: 16584:Zhenyuanopterus 16558: 16509: 16488:Istiodactylinae 16482: 16463:Longchengpterus 16456:Lingyuanopterus 16443:Istiodactylidae 16437: 16403: 16376:Hongshanopterus 16357: 16326: 16312: 16299: 16262:Hongshanopterus 16247: 16212: 16187: 16160:Barbaridactylus 16145: 16107: 16075: 16022:Pteranodontidae 16007: 16002:Santanadactylus 15986: 15973: 15956: 15949: 15941: 15933: 15917: 15861: 15856:Zhejiangopterus 15777: 15735: 15714:Palaeocursornis 15676: 15655:Montanazhdarcho 15642:Azhdarchiformes 15632: 15613:Chaoyangopterus 15598: 15527: 15503: 15489:Montanazhdarcho 15442: 15402: 15400:Azhdarchomorpha 15369: 15342: 15296: 15259:Aymberedactylus 15240: 15168: 15144: 15132:Aymberedactylus 15107: 15087:Thalassodromeus 15016: 14991: 14980: 14971: 14959:Montanazhdarcho 14917: 14900: 14888:Domeykodactylus 14873: 14867:Tendaguripterus 14804: 14795: 14777: 14764: 14747: 14735: 14719: 14699: 14693:Altmuehlopterus 14650: 14609: 14585: 14536: 14508: 14413: 14396: 14360: 14354:Gallodactylidae 14334:Aurorazhdarchia 14328: 14306:Normannognathus 14282:Gallodactylidae 14270: 14261: 14237: 14224: 14218:Tendaguripterus 14210:Normannognathus 14203:Germanodactylus 14195:Altmuehlopterus 14180: 14159: 14146: 14129: 14091: 14071: 13985: 13950:Batrachognathus 13935: 13930:Vesperopterylus 13885: 13859: 13850: 13825: 13804: 13760:Wukongopterinae 13754: 13736:Wukongopteridae 13730: 13699: 13686:Normannognathus 13663: 13654: 13628: 13615: 13598: 13585: 13577: 13541: 13521: 13494:Jianchangnathus 13477:Scaphognathidae 13467:Digibrevisauria 13449: 13444:Rhamphorhynchus 13413: 13400:Qinglongopterus 13392:Harpactognathus 13360:Rhamphorhynchae 13354: 13349:Qinglongopterus 13328:Harpactognathus 13288: 13282:Scaphognathinae 13274:Parapsicephalus 13238: 13225: 13213:Bergamodactylus 13192: 13183: 13165: 13151: 13129:Parapsicephalus 13092: 13071: 13069:Macronychoptera 13062: 13044: 13022: 12994: 12988:Arcticodactylus 12969: 12957:Raeticodactylus 12918: 12884: 12867: 12847:Arcticodactylus 12832: 12826:Raeticodactylus 12791: 12779:Austriadactylus 12764: 12743: 12730: 12705: 12685: 12679:Avemetatarsalia 12664:Avemetatarsalia 12654: 12639:Avemetatarsalia 12604: 12603: 12593: 12588: 12558: 12553: 12539:Phytodinosauria 12520: 12507: 12463: 12434: 12428:Sauropodomorpha 12417:Sauropodomorpha 12395: 12360:Herrerasauridae 12354: 12302: 12231: 12218: 12186: 12169: 12100: 12028: 11973: 11971:Dinosauromorpha 11956: 11933: 11851: 11838: 11786: 11751: 11749:Avemetatarsalia 11730: 11727:Avemetatarsalia 11711: 11697:Avemetatarsalia 11691: 11664: 11633: 11601: 11600: 11590: 11585: 11535: 11525: 11506: 11487: 11471: 11466: 11465: 11427:Current Biology 11418: 11414: 11403: 11399: 11389: 11387: 11378: 11372: 11368: 11358: 11356: 11344: 11340: 11324: 11317: 11309: 11305: 11280: 11276: 11264: 11255: 11208: 11204: 11195: 11193: 11183: 11179: 11126: 11122: 11069: 11065: 11020: 11016: 11007: 11005: 10997: 10996: 10992: 10939:Rhamphorhynchus 10933: 10926: 10879:Rhamphorhynchus 10875: 10866: 10857: 10855: 10847: 10846: 10839: 10815:10.1038/429621a 10786: 10782: 10745: 10741: 10702: 10698: 10667:(12): 1323–30. 10661:Current Biology 10653: 10646: 10599: 10595: 10542: 10538: 10505:10.1038/432571a 10485: 10481: 10462: 10458: 10435:10.1038/432572a 10410: 10404: 10400: 10383: 10382: 10378: 10354:10.1038/430033a 10325: 10321: 10298: 10289: 10254: 10250: 10233: 10229: 10182: 10178: 10149: 10145: 10136: 10132: 10091:(9): e0185486. 10073: 10069: 10061: 10057: 10049: 10045: 9994: 9990: 9951: 9947: 9939: 9935: 9895: 9889: 9885: 9877: 9873: 9865: 9861: 9853: 9849: 9806: 9802: 9794: 9785: 9761: 9755: 9751: 9708: 9704: 9675:(2–4): 115–26. 9664: 9658: 9654: 9607: 9603: 9588: 9584: 9561: 9557: 9526:(12): 2233–53. 9512: 9508: 9498: 9496: 9494:Qmro.qmul.ac.uk 9489: 9485: 9484: 9480: 9465: 9461: 9445: 9439: 9435: 9425: 9423: 9413: 9406: 9396: 9394: 9385: 9384: 9375: 9365: 9363: 9343: 9339: 9332: 9316: 9312: 9302: 9300: 9290: 9286: 9276: 9274: 9259: 9252: 9205: 9201: 9184: 9180: 9173: 9157: 9153: 9137: 9133: 9094: 9090: 9055: 9048: 9033: 9029: 8976: 8972: 8932: 8926: 8922: 8913: 8911: 8907: 8876: 8866: 8862: 8823: 8819: 8786:(3): e2001663. 8772: 8763: 8723: 8719: 8682: 8678: 8669: 8665: 8618: 8607: 8598: 8594: 8586: 8582: 8551: 8547: 8538: 8536: 8526: 8522: 8513: 8511: 8503: 8502: 8498: 8450: 8443: 8436: 8385: 8381: 8334: 8330: 8285: 8281: 8250: 8243: 8195: 8189: 8185: 8156: 8152: 8111: 8107: 8099: 8095: 8087: 8083: 8070: 8069: 8065: 8057: 8050: 8043: 8036: 7991: 7987: 7981:Wellnhofer 1991 7979: 7975: 7966: 7962: 7954: 7941: 7894: 7887: 7878: 7874: 7859: 7855: 7846: 7842: 7833: 7829: 7820: 7818: 7814: 7794:(1/3): 105–12. 7781: 7775: 7771: 7762: 7758: 7743: 7739: 7724: 7720: 7714:Wellnhofer 1991 7712: 7708: 7702:Wellnhofer 1991 7700: 7693: 7687:Wellnhofer 1991 7685: 7676: 7670:Wellnhofer 1991 7668: 7661: 7632:(615): 309–38. 7622: 7618: 7609: 7605: 7596: 7592: 7586:Wellnhofer 1991 7584: 7580: 7574:Wellnhofer 1991 7572: 7568: 7562:Wellnhofer 1991 7560: 7556: 7537: 7533: 7518: 7514: 7508:Wellnhofer 1991 7506: 7502: 7488: 7484: 7475: 7471: 7457: 7453: 7438: 7434: 7425: 7421: 7412: 7408: 7400: 7389: 7385:, pp. 6–7. 7381: 7377: 7369: 7365: 7359:Wellnhofer 1991 7357: 7353: 7345: 7338: 7285: 7281: 7236: 7232: 7174: 7170: 7141:(6492): 62–64. 7131: 7127: 7082: 7078: 7031: 7027: 7018: 7016: 7004: 7000: 6947: 6943: 6928: 6924: 6916: 6903: 6895: 6891: 6883: 6879: 6871: 6867: 6859: 6855: 6847: 6843: 6788: 6777: 6769: 6760: 6754:Wellnhofer 1991 6752: 6737: 6731:Wellnhofer 1991 6729: 6714: 6643: 6639: 6633:Wellnhofer 1991 6631: 6627: 6621:Wellnhofer 1991 6619: 6608: 6600: 6581: 6534: 6530: 6493: 6489: 6466: 6462: 6430: 6424: 6420: 6373: 6369: 6361: 6357: 6349: 6345: 6339:Wellnhofer 1991 6337: 6328: 6283: 6279: 6240: 6236: 6197: 6193: 6164:(6492): 62–64. 6150: 6143: 6135: 6128: 6120: 6113: 6058: 6051: 6002: 5993: 5954: 5950: 5942: 5935: 5927: 5923: 5915: 5906: 5898: 5894: 5888:Wellnhofer 1991 5886: 5873: 5867:Wellnhofer 1991 5865: 5858: 5850: 5841: 5833: 5824: 5816: 5805: 5797: 5793: 5787:Wellnhofer 1991 5785: 5776: 5768: 5761: 5738: 5734: 5728:Wellnhofer 1991 5726: 5722: 5705: 5698: 5683: 5672: 5666:Wellnhofer 1991 5664: 5660: 5633: 5626: 5618: 5609: 5601: 5592: 5586:Wellnhofer 1991 5584: 5577: 5569: 5562: 5555: 5541: 5524: 5516: 5507: 5499: 5495: 5480: 5476: 5436: 5430: 5426: 5417: 5415: 5407: 5406: 5402: 5354: 5348: 5337: 5330: 5314: 5307: 5297: 5295: 5294:. Pterosaur.net 5290:Naish, Darren. 5288: 5279: 5272: 5247: 5243: 5195: 5189: 5185: 5135: 5131: 5076: 5069: 5053: 5052: 5046: 5044: 5042:livescience.com 5034: 5030: 5021: 5019: 5011: 5010: 5006: 4977: 4973: 4968:: 61–107. 2008. 4961: 4957: 4956: 4952: 4939: 4935: 4920: 4919: 4915: 4909: 4892: 4888: 4869: 4865: 4859: 4837: 4833: 4786: 4782: 4745:Current Biology 4737: 4730: 4725: 4720: 4719: 4711: 4707: 4701: 4698: 4695: 4678: 4674: 4665: 4661: 4641: 4620: 4616: 4615: 4611: 4606: 4602: 4597: 4579:Pterosaur Beach 4544: 4539: 4537: 4530: 4523: 4516: 4511: 4509: 4504:Reptiles portal 4502: 4497: 4495: 4492: 4473:Vesperopterylus 4402:Ray Harryhausen 4350:Rhamphorhynchus 4340:Rhamphorhynchus 4325:Rhamphorhynchus 4289: 4269:Rhamphorhynchus 4249:Rhamphorhynchus 4226:Rhamphorhynchus 4183: 4158:Rhamphorhynchus 4150:Rhamphorhynchus 4095:Rhamphorhynchus 4000: 3988:] (most likely 3938: 3907:Thalassodromeus 3864:pterodactyloids 3831:Thalassodromeus 3786:Germanodactylus 3746:Alcione elainus 3733:Pteranodontidae 3704:Wukongopteridae 3691:Harpactognathus 3667:Rhamphorhynchus 3644:Austriadactylus 3631:and the larger 3587: 3549: 3499:ornithocheirids 3464:in pterosaurs. 3390: 3388:Ground movement 3382:pterodactyloids 3344: 3323: 3254:. Furthermore, 3240:Late Cretaceous 3229: 3208: 3203: 3202: 3193: 3184: 3179: 3178: 3177: 3176: 3165: 3160: 3159: 3158: 3157: 3146: 3141: 3140: 3139: 3138: 3127: 3122: 3121: 3120: 3119: 3108: 3103: 3102: 3101: 3100: 3089: 3084: 3083: 3082: 3081: 3070: 3061: 3052: 3043: 3034: 3025: 3016: 2933: 2837: 2828: 2519:Macronychoptera 2415:Ornithocephalus 2383: 2377: 2361:Late Cretaceous 2273: 2264:Prorotodactylus 2236:neuroanatomical 2228:sensory systems 2133:archosauromorph 2044:neuroanatomical 2022: 2017: 2009:Jurassic period 1877:Zhejiangopterus 1868: 1827:Rhamphorhynchus 1806:Rhamphorhynchus 1733: 1658:suggested that 1604: 1599: 1593: 1521: 1435: 1415: 1385: 1344: 1310:pteranodontians 1208:brachiopatagium 1175: 1144:Rhamphorhynchus 1115: 1079:shoulder girdle 1075: 1073:Shoulder girdle 972: 951:Austriadactylus 898:Thalassodromeus 872:Istiodactylidae 864:Pteranodontidae 776: 752: 746: 729:Vesperopterylus 685:Germanodactylus 640:Rhamphorhynchus 578: 487:. These may be 465:pterodactyloids 413:are an extinct 321:Macronychoptera 310:Austriadactylus 253: 247: 234: 174: 117: 115: 114: 109: 104: 99: 94: 89: 84: 79: 74: 69: 64: 59: 48: 47: 44:Late Cretaceous 36: 28: 17: 12: 11: 5: 17753: 17743: 17742: 17737: 17732: 17727: 17710: 17709: 17707: 17706: 17693: 17680: 17667: 17654: 17644: 17629: 17613: 17611: 17605: 17604: 17590: 17589: 17587: 17586: 17581: 17576: 17571: 17566: 17561: 17556: 17551: 17545: 17543: 17539: 17538: 17536: 17535: 17530: 17525: 17520: 17515: 17510: 17505: 17500: 17495: 17490: 17484: 17482: 17478: 17477: 17475: 17474: 17469: 17467:Pterosaur wing 17464: 17459: 17458: 17457: 17452: 17447: 17437: 17432: 17426: 17424: 17418: 17417: 17415: 17414: 17409: 17404: 17403: 17402: 17392: 17387: 17382: 17381: 17380: 17375: 17370: 17365: 17360: 17355: 17350: 17345: 17335: 17329: 17327: 17321: 17320: 17313: 17311: 17309: 17308: 17303: 17298: 17293: 17288: 17283: 17278: 17273: 17268: 17263: 17258: 17256:Cephalopod fin 17253: 17247: 17245: 17239: 17238: 17223: 17222: 17215: 17208: 17200: 17191: 17190: 17187: 17186: 17165: 17163: 17160: 17159: 17156: 17155: 17152: 17151: 17148: 17147: 17144: 17143: 17140: 17139: 17136: 17135: 17132: 17131: 17128: 17127: 17125: 17124: 17117: 17110: 17103: 17096: 17088: 17084:Caulkicephalus 17080: 17072: 17070: 17064: 17063: 17061: 17060: 17057:Uktenadactylus 17053: 17045: 17038: 17035:Coloborhynchus 17031: 17023: 17021: 17014: 17013: 17011: 17010: 17003: 16996: 16988: 16980: 16972: 16964: 16955: 16953: 16943: 16937: 16936: 16934: 16933: 16926: 16918: 16916: 16909: 16908: 16906: 16905: 16897: 16888: 16882: 16877: 16874: 16873: 16870: 16869: 16866: 16865: 16863: 16862: 16859:Targaryendraco 16855: 16847: 16838: 16836: 16830: 16829: 16827: 16826: 16819: 16811: 16803: 16801: 16792: 16785: 16784: 16781: 16780: 16778: 16777: 16773:Uktenadactylus 16769: 16761: 16758:Ornithocheirus 16754: 16746: 16738: 16730: 16722: 16714: 16710:Coloborhynchus 16706: 16698: 16689: 16687: 16681: 16680: 16678: 16677: 16669: 16661: 16653: 16649:Caulkicephalus 16645: 16637: 16634:Arthurdactylus 16630: 16622: 16616: 16603: 16600:Ornithocheirae 16598: 16595: 16594: 16591: 16590: 16588: 16587: 16580: 16572: 16570: 16557: 16556: 16549: 16542: 16535: 16527: 16525: 16519: 16518: 16515: 16514: 16511: 16510: 16508: 16507: 16500: 16492: 16490: 16484: 16483: 16481: 16480: 16473: 16466: 16459: 16451: 16445: 16439: 16438: 16436: 16435: 16428: 16425:Linlongopterus 16421: 16413: 16411: 16409:Mimodactylidae 16405: 16404: 16402: 16401: 16397:Lonchodectidae 16393: 16386: 16383:Linlongopterus 16379: 16371: 16365: 16359: 16358: 16356: 16355: 16348: 16340: 16338: 16325: 16324: 16315: 16313: 16308: 16305: 16304: 16301: 16300: 16298: 16297: 16293:Targaryendraco 16289: 16281: 16274: 16266: 16257: 16255: 16253:Lonchodectidae 16249: 16248: 16246: 16245: 16238: 16231: 16223: 16213: 16208: 16205: 16204: 16201: 16200: 16197: 16196: 16193: 16192: 16189: 16188: 16186: 16185: 16177: 16170: 16167:Muzquizopteryx 16163: 16155: 16153: 16147: 16146: 16144: 16143: 16136: 16129: 16121: 16115: 16113:Aponyctosauria 16109: 16108: 16106: 16105: 16097: 16089: 16083: 16077: 16076: 16074: 16073: 16065: 16057: 16050: 16042: 16035: 16026: 16024: 16015: 16009: 16008: 16006: 16005: 15997: 15987: 15982: 15975: 15974: 15962: 15961: 15958: 15957: 15932: 15930: 15927: 15926: 15923: 15922: 15919: 15918: 15916: 15915: 15914: 15913: 15907: 15905: 15895: 15894: 15891: 15890: 15887: 15886: 15883: 15882: 15879: 15878: 15875: 15874: 15871: 15870: 15867: 15866: 15863: 15862: 15860: 15859: 15852: 15844: 15841:Thanatosdrakon 15837: 15834:Quetzalcoatlus 15830: 15827:Phosphatodraco 15823: 15816: 15809: 15802: 15799:Arambourgiania 15795: 15787: 15785: 15779: 15778: 15776: 15775: 15768: 15761: 15754: 15745: 15743: 15734: 15733: 15725: 15717: 15710: 15706:Navajodactylus 15702: 15695: 15686: 15684: 15675: 15674: 15666: 15659: 15650: 15644: 15638: 15637: 15634: 15633: 15631: 15630: 15627:Shenzhoupterus 15623: 15616: 15608: 15606: 15600: 15599: 15597: 15596: 15588: 15580: 15573: 15566: 15558: 15554:Argentinadraco 15550: 15541: 15535: 15529: 15528: 15526: 15525: 15517: 15511: 15505: 15504: 15502: 15501: 15493: 15485: 15477: 15469: 15465:Argentinadraco 15461: 15453: 15451: 15444: 15443: 15441: 15440: 15432: 15425: 15418: 15409: 15403: 15398: 15395: 15394: 15391: 15390: 15387: 15386: 15383: 15382: 15379: 15378: 15375: 15374: 15371: 15370: 15368: 15367: 15360: 15352: 15350: 15344: 15343: 15341: 15340: 15333: 15326: 15319: 15310: 15304: 15298: 15297: 15295: 15294: 15287: 15279: 15271: 15263: 15254: 15248: 15242: 15241: 15239: 15238: 15231: 15224: 15221:Nemicolopterus 15217: 15210: 15203: 15195: 15187: 15178: 15176: 15167: 15166: 15158: 15152: 15146: 15145: 15143: 15142: 15135: 15127: 15125: 15115: 15113:Tapejariformes 15109: 15108: 15106: 15105: 15097: 15090: 15083: 15075: 15067: 15060: 15052: 15048:Argentinadraco 15044: 15036: 15027: 15025: 15018: 15017: 15015: 15014: 15006: 14998: 14992: 14989:Tapejaromorpha 14987: 14981: 14976: 14973: 14972: 14970: 14969: 14962: 14955: 14947: 14939: 14932: 14929:Argentinadraco 14924: 14918: 14913: 14910: 14909: 14906: 14905: 14902: 14901: 14899: 14898: 14895:Dsungaripterus 14891: 14883: 14881: 14875: 14874: 14872: 14871: 14863: 14855: 14848: 14841: 14833: 14824: 14818: 14805: 14800: 14797: 14796: 14794: 14793: 14784: 14778: 14773: 14766: 14765: 14753: 14752: 14749: 14748: 14734: 14732: 14729: 14728: 14725: 14724: 14721: 14720: 14718: 14717: 14716: 14715: 14709: 14707: 14701: 14700: 14698: 14697: 14688: 14682: 14672: 14671: 14668: 14667: 14664: 14663: 14660: 14659: 14656: 14655: 14652: 14651: 14649: 14648: 14641: 14634: 14627: 14619: 14617: 14615:Pterodaustrini 14611: 14610: 14608: 14607: 14599: 14593: 14587: 14586: 14584: 14583: 14576: 14569: 14562: 14554: 14546: 14544: 14542:Gnathosaurinae 14538: 14537: 14535: 14534: 14527: 14519: 14517: 14514:Moganopterinae 14510: 14509: 14507: 14506: 14499: 14492: 14485: 14478: 14471: 14464: 14457: 14450: 14443: 14436: 14433:Balaenognathus 14429: 14420: 14414: 14409: 14406: 14405: 14402: 14401: 14398: 14397: 14395: 14394: 14386: 14383:Aurorazhdarcho 14379: 14370: 14368: 14362: 14361: 14359: 14358: 14350: 14342: 14336: 14330: 14329: 14327: 14326: 14322:Moganopterinae 14318: 14310: 14302: 14295: 14291:Aurorazhdarcho 14286: 14284: 14271: 14266: 14263: 14262: 14260: 14259: 14252: 14244: 14238: 14233: 14230: 14229: 14226: 14225: 14223: 14222: 14214: 14206: 14199: 14190: 14188: 14182: 14181: 14179: 14178: 14170: 14160: 14155: 14148: 14147: 14135: 14134: 14131: 14130: 14118: 14116: 14113: 14112: 14109: 14108: 14105: 14104: 14101: 14100: 14097: 14096: 14093: 14092: 14090: 14089: 14088: 14087: 14081: 14079: 14073: 14072: 14070: 14069: 14062: 14054: 14047: 14040: 14033: 14026: 14018: 14011: 14003: 13997: 13991: 13990: 13987: 13986: 13984: 13983: 13976: 13968: 13960: 13953: 13945: 13943: 13937: 13936: 13934: 13933: 13926: 13919: 13911: 13903: 13895: 13893: 13891:Anurognathinae 13887: 13886: 13884: 13883: 13875: 13869: 13867:Anurognathidae 13860: 13855: 13852: 13851: 13849: 13848: 13845:Douzhanopterus 13841: 13832: 13826: 13821: 13818: 13817: 13814: 13813: 13810: 13809: 13806: 13805: 13803: 13802: 13795: 13791:Douzhanopterus 13787: 13780: 13773: 13764: 13762: 13756: 13755: 13753: 13752: 13749:Kunpengopterus 13744: 13738: 13732: 13731: 13729: 13728: 13721: 13713: 13707: 13701: 13700: 13698: 13697: 13693:Anurognathidae 13689: 13682: 13674: 13664: 13661:Monofenestrata 13659: 13656: 13655: 13653: 13652: 13644: 13635: 13629: 13624: 13617: 13616: 13604: 13603: 13600: 13599: 13576: 13574: 13571: 13570: 13567: 13566: 13563: 13562: 13559: 13558: 13555: 13554: 13551: 13550: 13547: 13546: 13543: 13542: 13540: 13539: 13538: 13537: 13531: 13529: 13523: 13522: 13520: 13519: 13511: 13504: 13497: 13490: 13482: 13480: 13470: 13463: 13462: 13459: 13458: 13455: 13454: 13451: 13450: 13448: 13447: 13440: 13432: 13423: 13421: 13415: 13414: 13412: 13411: 13404: 13396: 13388: 13381: 13373: 13371: 13362: 13356: 13355: 13353: 13352: 13345: 13342:Orientognathus 13338: 13331: 13324: 13317: 13310: 13302: 13296: 13290: 13289: 13287: 13286: 13278: 13270: 13263: 13259:Dolicorhamphus 13254: 13248: 13239: 13236:Breviquartossa 13234: 13231: 13230: 13227: 13226: 13224: 13223: 13216: 13208: 13206: 13193: 13188: 13185: 13184: 13182: 13181: 13172: 13166: 13160: 13157: 13156: 13153: 13152: 13150: 13149: 13141: 13133: 13125: 13118: 13115:Caelestiventus 13111: 13102: 13100: 13094: 13093: 13091: 13090: 13082: 13072: 13067: 13064: 13063: 13061: 13060: 13051: 13045: 13041:Zambellisauria 13039: 13036: 13035: 13032: 13031: 13028: 13027: 13024: 13023: 13021: 13020: 13013: 13009:Carniadactylus 13004: 13002: 12996: 12995: 12993: 12992: 12983: 12977: 12971: 12970: 12968: 12967: 12960: 12953: 12946: 12937: 12935: 12926: 12920: 12919: 12917: 12916: 12908: 12900: 12891: 12885: 12880: 12877: 12876: 12873: 12872: 12869: 12868: 12866: 12865: 12862:Seazzadactylus 12858: 12851: 12842: 12840: 12834: 12833: 12831: 12830: 12822: 12815: 12811:Carniadactylus 12806: 12800: 12793: 12792: 12790: 12789: 12782: 12774: 12772: 12766: 12765: 12763: 12762: 12759:Daohugoupterus 12754: 12744: 12739: 12732: 12731: 12719: 12718: 12715: 12714: 12711: 12710: 12707: 12706: 12704: 12703: 12702: 12701: 12695: 12693: 12687: 12686: 12684: 12683: 12682: 12681: 12672: 12666: 12656: 12655: 12651: 12650: 12641: 12632: 12623: 12617: 12611: 12595: 12594: 12587: 12586: 12579: 12572: 12564: 12555: 12554: 12552: 12551: 12541: 12536: 12534:Ornithoscelida 12529: 12526: 12525: 12522: 12521: 12490: 12488: 12485: 12484: 12481: 12480: 12477: 12476: 12473: 12472: 12469: 12468: 12465: 12464: 12462: 12461: 12460: 12459: 12453: 12444: 12442: 12436: 12435: 12433: 12432: 12431: 12430: 12421: 12419: 12407: 12401: 12400: 12397: 12396: 12394: 12393: 12390:Staurikosaurus 12386: 12379: 12372: 12364: 12362: 12356: 12355: 12353: 12352: 12344: 12336: 12328: 12319: 12313: 12304: 12303: 12301: 12300: 12290: 12279: 12269: 12259: 12249: 12238: 12232: 12227: 12224: 12223: 12220: 12219: 12217: 12216: 12215: 12214: 12205: 12203: 12187: 12182: 12179: 12178: 12175: 12174: 12171: 12170: 12168: 12167: 12160: 12153: 12146: 12139: 12132: 12125: 12118: 12110: 12108: 12102: 12101: 12099: 12098: 12090: 12083: 12075: 12068: 12061: 12054: 12046: 12040: 12030: 12029: 12027: 12026: 12016: 12005: 11995: 11984: 11974: 11969: 11966: 11965: 11962: 11961: 11958: 11957: 11955: 11954: 11953: 11952: 11943: 11941: 11935: 11934: 11932: 11931: 11924: 11916: 11909: 11902: 11895: 11891:Faxinalipterus 11887: 11879: 11877: 11868: 11852: 11847: 11844: 11843: 11840: 11839: 11837: 11836: 11829: 11822: 11815: 11808: 11804:Incertovenator 11799: 11797: 11788: 11787: 11785: 11784: 11774: 11770:Incertovenator 11762: 11752: 11747: 11740: 11739: 11736: 11735: 11732: 11731: 11724: 11721: 11720: 11717: 11716: 11713: 11712: 11710: 11709: 11708: 11707: 11701: 11699: 11693: 11692: 11690: 11689: 11688: 11687: 11678: 11672: 11666: 11665: 11663: 11662: 11661: 11660: 11651: 11645: 11635: 11634: 11630: 11629: 11620: 11614: 11608: 11592: 11591: 11584: 11583: 11576: 11569: 11561: 11555: 11554: 11548: 11542: 11534: 11533:External links 11531: 11530: 11529: 11523: 11510: 11504: 11491: 11485: 11470: 11467: 11464: 11463: 11412: 11397: 11366: 11338: 11315: 11313:, p. 452. 11303: 11274: 11253: 11202: 11177: 11120: 11063: 11014: 10990: 10924: 10864: 10837: 10780: 10739: 10696: 10644: 10593: 10556:(3): 647–654. 10536: 10479: 10456: 10398: 10376: 10319: 10308:(2): 602–635. 10287: 10248: 10245:on 2016-04-06. 10227: 10176: 10143: 10130: 10067: 10065:, p. 199. 10055: 10043: 9988: 9945: 9943:, p. 134. 9933: 9883: 9881:, p. 122. 9871: 9869:, p. 121. 9859: 9857:, p. 103. 9847: 9800: 9783: 9749: 9702: 9652: 9601: 9582: 9555: 9506: 9478: 9473:New York Times 9459: 9433: 9404: 9373: 9337: 9330: 9310: 9284: 9250: 9199: 9178: 9171: 9151: 9131: 9088: 9046: 9027: 8970: 8943:(2): 288–321. 8920: 8860: 8817: 8761: 8717: 8696:(6): 1144–55. 8676: 8663: 8605: 8592: 8580: 8545: 8528:Black, Riley. 8520: 8496: 8434: 8379: 8328: 8279: 8241: 8183: 8170:(3): 261–308. 8150: 8105: 8093: 8081: 8078:. 22 Feb 2022. 8063: 8048: 8034: 7985: 7973: 7960: 7939: 7885: 7872: 7853: 7840: 7827: 7769: 7756: 7737: 7718: 7706: 7691: 7674: 7659: 7616: 7603: 7590: 7578: 7566: 7554: 7531: 7512: 7500: 7482: 7469: 7451: 7432: 7419: 7406: 7387: 7375: 7363: 7351: 7336: 7279: 7230: 7168: 7125: 7076: 7025: 6998: 6941: 6922: 6901: 6889: 6877: 6865: 6853: 6841: 6775: 6758: 6735: 6712: 6637: 6625: 6606: 6579: 6528: 6507:(4): 1045–62. 6487: 6460: 6418: 6367: 6355: 6343: 6326: 6297:(4): 1339–42. 6277: 6234: 6191: 6154:Sordes pilosus 6141: 6126: 6111: 6049: 5991: 5948: 5933: 5921: 5904: 5892: 5871: 5856: 5839: 5822: 5803: 5791: 5774: 5759: 5732: 5720: 5696: 5670: 5658: 5624: 5607: 5590: 5575: 5560: 5553: 5522: 5505: 5493: 5474: 5447:(1): 163–187. 5424: 5400: 5335: 5328: 5305: 5277: 5270: 5256:The Dinosauria 5241: 5183: 5129: 5090:(6): 1983–87. 5067: 5028: 5004: 4971: 4950: 4933: 4913: 4907: 4886: 4863: 4857: 4831: 4780: 4751:(9): 1011–16. 4727: 4726: 4724: 4721: 4718: 4717: 4705: 4672: 4609: 4599: 4598: 4596: 4593: 4592: 4591: 4586: 4584:Pterosaur size 4581: 4576: 4571: 4566: 4561: 4556: 4550: 4549: 4535: 4521: 4507: 4491: 4488: 4482:Kunpengopterus 4381:The Lost World 4357:have both the 4298:Quetzalcoatlus 4288: 4285: 4182: 4179: 4162:pterodactyloid 4126:superprecocial 4031:was laid by a 4008:pterodactyloid 3999: 3996: 3991:Platypterygius 3962:, most likely 3952:Eric Buffetaut 3937: 3934: 3897:Dsungaripterus 3868:pteranodontids 3777:istiodactylids 3725:Ornithocheirae 3628:Carniadactylus 3595:Anurognathidae 3586: 3583: 3548: 3545: 3389: 3386: 3343: 3342:Nervous system 3340: 3332:hepatic piston 3322: 3319: 3296:Paul MacCready 3228: 3225: 3207: 3204: 3199: 3198: 3195: 3194: 3190: 3189: 3186: 3185: 3181: 3180: 3172: 3171: 3170: 3167: 3166: 3162: 3161: 3155:Breviquartossa 3153: 3152: 3151: 3148: 3147: 3143: 3142: 3134: 3133: 3132: 3129: 3128: 3124: 3123: 3117:Monofenestrata 3115: 3114: 3113: 3110: 3109: 3105: 3104: 3096: 3095: 3094: 3091: 3090: 3086: 3085: 3077: 3076: 3075: 3072: 3071: 3067: 3066: 3063: 3062: 3058: 3057: 3054: 3053: 3049: 3048: 3045: 3044: 3040: 3039: 3036: 3035: 3031: 3030: 3027: 3026: 3022: 3021: 3018: 3017: 3013: 3012: 3009: 3008: 2996: 2993: 2992: 2989: 2988: 2976: 2971: 2969: 2963: 2962: 2959: 2958: 2949: 2944: 2942: 2939: 2938: 2935: 2934: 2930: 2929: 2926: 2925: 2913: 2910: 2909: 2906: 2905: 2897:Tapejaromorpha 2893: 2888: 2886: 2878: 2876: 2870: 2869: 2866: 2865: 2856: 2851: 2849: 2843: 2842: 2839: 2838: 2834: 2833: 2830: 2829: 2825: 2824: 2821: 2820: 2808: 2805: 2804: 2801: 2800: 2786: 2781: 2779: 2773: 2772: 2769: 2768: 2756: 2751: 2749: 2741: 2739: 2733: 2732: 2729: 2728: 2714: 2709: 2707: 2701: 2700: 2697: 2696: 2688:Anurognathidae 2684: 2679: 2677: 2674: 2673: 2670: 2669: 2660: 2655: 2653: 2650: 2649: 2646: 2645: 2638: 2633: 2631: 2628: 2627: 2624: 2623: 2609: 2604: 2602: 2599: 2598: 2595: 2594: 2582: 2577: 2575: 2572: 2571: 2568: 2567: 2555: 2550: 2548: 2545: 2544: 2541: 2540: 2532:Dimorphodontia 2528: 2523: 2521: 2515: 2514: 2511: 2510: 2498: 2493: 2491: 2483: 2482: 2438:classification 2402:Quetzalcoatlus 2376: 2373: 2338:Navajodactylus 2329:pteranodontids 2324:Ornithocheirus 2281:Quetzalcoatlus 2272: 2269: 2191:Martin Ezcurra 2105:Michael Benton 2085:tanystropheids 2021: 2018: 2016: 2013: 1867: 1864: 1798:Niobrara Chalk 1743:reconstruction 1732: 1729: 1687:Georges Cuvier 1683:Johann Hermann 1603: 1600: 1592: 1589: 1564:Sordes pilosus 1540:Inner Mongolia 1525:proto-feathers 1520: 1517: 1489:Sordes pilosus 1434: 1431: 1423:petrifications 1414: 1411: 1384: 1381: 1379:chest cavity. 1343: 1340: 1174: 1171: 1152:fibers called 1114: 1111: 1087:shoulder blade 1074: 1071: 1054:caudofemoralis 997:vertebral body 981:Arambourgiania 971: 968: 912:parietal bones 809:. Unlike most 785:Coloborhynchus 775: 772: 750:Pterosaur size 748:Main article: 745: 742: 709:Quetzalcoatlus 691:Dsungaripterus 661:Ornithocheirus 610:shoulder blade 577: 574: 518:Quetzalcoatlus 442:powered flight 406: 405: 391: 390: 384: 383: 375: 374: 366: 365: 364: 363: 362: 361: 353: 351:Dimorphodontia 345: 334: 315: 314: 313: 303: 285: 277: 266: 265: 261: 260: 245: 241: 240: 232: 225: 224: 219: 212: 211: 206: 202: 201: 196: 192: 191: 186: 182: 181: 168: 167: 152:Quetzalcoatlus 128: 127: 119: 118: 110: 105: 100: 95: 90: 85: 80: 75: 70: 65: 60: 55: 37: 15: 9: 6: 4: 3: 2: 17752: 17741: 17738: 17736: 17733: 17731: 17728: 17726: 17723: 17722: 17720: 17703: 17698: 17694: 17690: 17685: 17681: 17677: 17672: 17668: 17664: 17659: 17655: 17651: 17645: 17640: 17634: 17630: 17625: 17619: 17615: 17614: 17612: 17610: 17606: 17602: 17597: 17585: 17582: 17580: 17577: 17575: 17572: 17570: 17567: 17565: 17562: 17560: 17557: 17555: 17552: 17550: 17547: 17546: 17544: 17540: 17534: 17531: 17529: 17526: 17524: 17521: 17519: 17516: 17514: 17511: 17509: 17506: 17504: 17501: 17499: 17496: 17494: 17491: 17489: 17486: 17485: 17483: 17479: 17473: 17470: 17468: 17465: 17463: 17460: 17456: 17453: 17451: 17448: 17446: 17443: 17442: 17441: 17438: 17436: 17433: 17431: 17428: 17427: 17425: 17423: 17419: 17413: 17410: 17408: 17405: 17401: 17398: 17397: 17396: 17393: 17391: 17388: 17386: 17383: 17379: 17376: 17374: 17371: 17369: 17366: 17364: 17361: 17359: 17356: 17354: 17351: 17349: 17346: 17344: 17341: 17340: 17339: 17336: 17334: 17331: 17330: 17328: 17326: 17322: 17317: 17307: 17304: 17302: 17301:Pectoral fins 17299: 17297: 17294: 17292: 17289: 17287: 17284: 17282: 17279: 17277: 17274: 17272: 17269: 17267: 17264: 17262: 17259: 17257: 17254: 17252: 17249: 17248: 17246: 17244: 17240: 17236: 17232: 17228: 17221: 17216: 17214: 17209: 17207: 17202: 17201: 17198: 17183: 17179: 17177: 17170: 17123: 17122: 17121:Maaradactylus 17118: 17116: 17115: 17111: 17109: 17108: 17104: 17102: 17101: 17097: 17094: 17093: 17092:Cearadactylus 17089: 17086: 17085: 17081: 17079: 17078: 17074: 17073: 17071: 17069: 17068:Anhanguerinae 17065: 17059: 17058: 17054: 17051: 17050: 17049:Siroccopteryx 17046: 17044: 17043: 17039: 17037: 17036: 17032: 17030: 17029: 17025: 17024: 17022: 17019: 17015: 17009: 17008: 17007:Tropeognathus 17004: 17002: 17001: 16997: 16994: 16993: 16992:Siroccopteryx 16989: 16986: 16985: 16981: 16978: 16977: 16973: 16970: 16969: 16965: 16962: 16961: 16957: 16956: 16954: 16951: 16947: 16944: 16942: 16941:Anhangueridae 16938: 16932: 16931: 16930:Iberodactylus 16927: 16925: 16924: 16920: 16919: 16917: 16914: 16910: 16903: 16902: 16898: 16895: 16894: 16890: 16889: 16886: 16883: 16880: 16875: 16861: 16860: 16856: 16853: 16852: 16848: 16845: 16844: 16840: 16839: 16837: 16835: 16831: 16825: 16824: 16823:Cimoliopterus 16820: 16817: 16816: 16815:Camposipterus 16812: 16810: 16809: 16805: 16804: 16802: 16800: 16796: 16793: 16790: 16786: 16775: 16774: 16770: 16767: 16766: 16765:Siroccopteryx 16762: 16760: 16759: 16755: 16752: 16751: 16747: 16744: 16743: 16739: 16736: 16735: 16731: 16728: 16727: 16723: 16720: 16719: 16715: 16712: 16711: 16707: 16704: 16703: 16702:Camposipterus 16699: 16696: 16695: 16691: 16690: 16688: 16686: 16682: 16675: 16674: 16673:Tropeognathus 16670: 16667: 16666: 16662: 16659: 16658: 16657:Cimoliopterus 16654: 16651: 16650: 16646: 16643: 16642: 16641:Camposipterus 16638: 16636: 16635: 16631: 16629: 16628: 16627:Araripesaurus 16624: 16623: 16620: 16617: 16615: 16611: 16607: 16604: 16601: 16596: 16586: 16585: 16581: 16579: 16578: 16574: 16573: 16571: 16569: 16568:Boreopteridae 16565: 16561: 16555: 16554: 16550: 16548: 16547: 16546:Cearadactylus 16543: 16541: 16540: 16536: 16534: 16533: 16529: 16528: 16526: 16524: 16520: 16506: 16505: 16501: 16499: 16498: 16497:Istiodactylus 16494: 16493: 16491: 16489: 16485: 16479: 16478: 16474: 16472: 16471: 16467: 16465: 16464: 16460: 16458: 16457: 16453: 16452: 16449: 16446: 16444: 16440: 16434: 16433: 16429: 16427: 16426: 16422: 16420: 16419: 16415: 16414: 16412: 16410: 16406: 16399: 16398: 16394: 16392: 16391: 16390:Yixianopterus 16387: 16385: 16384: 16380: 16378: 16377: 16373: 16372: 16369: 16366: 16364: 16360: 16354: 16353: 16349: 16347: 16346: 16342: 16341: 16339: 16337: 16333: 16329: 16322: 16321: 16317: 16316: 16314: 16311: 16306: 16295: 16294: 16290: 16287: 16286: 16282: 16280: 16279: 16275: 16272: 16271: 16267: 16264: 16263: 16259: 16258: 16256: 16254: 16250: 16244: 16243: 16239: 16237: 16236: 16232: 16230: 16229: 16225: 16224: 16221: 16217: 16214: 16211: 16206: 16183: 16182: 16178: 16176: 16175: 16171: 16169: 16168: 16164: 16162: 16161: 16157: 16156: 16154: 16152: 16151:Nyctosauridae 16148: 16142: 16141: 16137: 16135: 16134: 16130: 16128: 16127: 16123: 16122: 16119: 16116: 16114: 16110: 16103: 16102: 16098: 16096: 16095: 16094:Alamodactylus 16091: 16090: 16087: 16084: 16082: 16078: 16071: 16070: 16066: 16063: 16062: 16058: 16056: 16055: 16051: 16048: 16047: 16043: 16041: 16040: 16036: 16033: 16032: 16028: 16027: 16025: 16023: 16019: 16016: 16014: 16013:Pteranodontia 16010: 16004: 16003: 15999: 15998: 15995: 15991: 15988: 15985: 15980: 15976: 15972: 15967: 15963: 15953: 15948: 15945: 15937: 15911: 15910: 15909: 15908: 15906: 15904: 15900: 15896: 15858: 15857: 15853: 15850: 15849: 15848:Wellnhopterus 15845: 15843: 15842: 15838: 15836: 15835: 15831: 15829: 15828: 15824: 15822: 15821: 15820:Hatzegopteryx 15817: 15815: 15814: 15810: 15808: 15807: 15803: 15801: 15800: 15796: 15794: 15793: 15792:Aralazhdarcho 15789: 15788: 15786: 15784: 15780: 15774: 15773: 15769: 15767: 15766: 15762: 15760: 15759: 15755: 15752: 15751: 15747: 15746: 15744: 15742: 15738: 15731: 15730: 15726: 15723: 15722: 15718: 15716: 15715: 15711: 15708: 15707: 15703: 15701: 15700: 15696: 15693: 15692: 15688: 15687: 15685: 15683: 15679: 15672: 15671: 15667: 15665: 15664: 15663:Radiodactylus 15660: 15657: 15656: 15652: 15651: 15648: 15645: 15643: 15639: 15629: 15628: 15624: 15622: 15621: 15617: 15615: 15614: 15610: 15609: 15607: 15605: 15601: 15594: 15593: 15589: 15586: 15585: 15581: 15579: 15578: 15574: 15572: 15571: 15567: 15564: 15563: 15559: 15556: 15555: 15551: 15548: 15547: 15546:Apatorhamphus 15543: 15542: 15539: 15536: 15534: 15530: 15524: 15523: 15519: 15518: 15515: 15512: 15510: 15506: 15499: 15498: 15494: 15491: 15490: 15486: 15483: 15482: 15478: 15475: 15474: 15470: 15467: 15466: 15462: 15460: 15459: 15455: 15454: 15452: 15449: 15445: 15438: 15437: 15433: 15431: 15430: 15426: 15424: 15423: 15419: 15416: 15415: 15411: 15410: 15407: 15404: 15401: 15396: 15366: 15365: 15361: 15359: 15358: 15354: 15353: 15351: 15349: 15345: 15339: 15338: 15337:Tupandactylus 15334: 15332: 15331: 15327: 15325: 15324: 15320: 15317: 15316: 15312: 15311: 15308: 15305: 15303: 15299: 15293: 15292: 15288: 15285: 15284: 15280: 15277: 15276: 15272: 15269: 15268: 15267:Caupedactylus 15264: 15261: 15260: 15256: 15255: 15252: 15249: 15247: 15243: 15237: 15236: 15232: 15230: 15229: 15225: 15223: 15222: 15218: 15216: 15215: 15211: 15209: 15208: 15204: 15201: 15200: 15196: 15193: 15192: 15188: 15185: 15184: 15180: 15179: 15177: 15175: 15171: 15165: 15164: 15160: 15159: 15156: 15153: 15151: 15147: 15141: 15140: 15139:Caupedactylus 15136: 15134: 15133: 15129: 15128: 15126: 15123: 15122:Caupedactylia 15119: 15116: 15114: 15110: 15103: 15102: 15098: 15096: 15095: 15091: 15089: 15088: 15084: 15081: 15080: 15076: 15073: 15072: 15068: 15066: 15065: 15061: 15058: 15057: 15053: 15050: 15049: 15045: 15042: 15041: 15037: 15034: 15033: 15029: 15028: 15026: 15023: 15019: 15012: 15011: 15007: 15005: 15004: 15000: 14999: 14996: 14993: 14990: 14985: 14982: 14979: 14978:Neoazhdarchia 14974: 14968: 14967: 14963: 14961: 14960: 14956: 14953: 14952: 14948: 14945: 14944: 14940: 14938: 14937: 14933: 14931: 14930: 14926: 14925: 14922: 14919: 14916: 14915:Azhdarchoidea 14911: 14897: 14896: 14892: 14890: 14889: 14885: 14884: 14882: 14880: 14876: 14869: 14868: 14864: 14861: 14860: 14859:Puntanipterus 14856: 14854: 14853: 14849: 14847: 14846: 14842: 14839: 14838: 14834: 14831: 14830: 14826: 14825: 14822: 14819: 14817: 14813: 14809: 14806: 14803: 14798: 14791: 14790: 14786: 14785: 14782: 14779: 14776: 14771: 14767: 14763: 14758: 14754: 14745: 14739: 14713: 14712: 14711: 14710: 14708: 14706: 14702: 14695: 14694: 14690: 14689: 14686: 14683: 14681: 14677: 14673: 14647: 14646: 14642: 14640: 14639: 14635: 14633: 14632: 14628: 14626: 14625: 14624:Beipiaopterus 14621: 14620: 14618: 14616: 14612: 14606: 14605: 14601: 14600: 14597: 14594: 14592: 14588: 14582: 14581: 14580:Tacuadactylus 14577: 14575: 14574: 14570: 14568: 14567: 14563: 14560: 14559: 14555: 14553: 14552: 14548: 14547: 14545: 14543: 14539: 14533: 14532: 14528: 14526: 14525: 14521: 14520: 14518: 14515: 14511: 14505: 14504: 14500: 14498: 14497: 14493: 14491: 14490: 14486: 14484: 14483: 14479: 14477: 14476: 14472: 14470: 14469: 14465: 14463: 14462: 14461:Forfexopterus 14458: 14456: 14455: 14454:Elanodactylus 14451: 14449: 14448: 14447:Cratonopterus 14444: 14442: 14441: 14440:Cathayopterus 14437: 14435: 14434: 14430: 14427: 14426: 14425:Ardeadactylus 14422: 14421: 14418: 14415: 14412: 14407: 14392: 14391: 14387: 14385: 14384: 14380: 14377: 14376: 14375:Ardeadactylus 14372: 14371: 14369: 14367: 14363: 14356: 14355: 14351: 14349: 14348: 14344: 14343: 14340: 14337: 14335: 14331: 14324: 14323: 14319: 14316: 14315: 14311: 14308: 14307: 14303: 14301: 14300: 14299:Cycnorhamphus 14296: 14293: 14292: 14288: 14287: 14285: 14283: 14279: 14275: 14272: 14269: 14264: 14258: 14257: 14256:Pterodactylus 14253: 14251: 14250: 14249:Diopecephalus 14246: 14245: 14242: 14239: 14236: 14231: 14220: 14219: 14215: 14212: 14211: 14207: 14205: 14204: 14200: 14197: 14196: 14192: 14191: 14189: 14187: 14183: 14177: 14176: 14175:Prejanopterus 14172: 14171: 14168: 14164: 14161: 14158: 14153: 14149: 14145: 14140: 14136: 14127: 14122: 14085: 14084: 14083: 14082: 14080: 14078: 14074: 14068: 14067: 14063: 14060: 14059: 14055: 14053: 14052: 14048: 14046: 14045: 14041: 14039: 14038: 14034: 14032: 14031: 14027: 14024: 14023: 14022:Herbstosaurus 14019: 14017: 14016: 14015:Eurolimnornis 14012: 14010: 14009: 14008:Dermodactylus 14005: 14004: 14001: 13998: 13996: 13992: 13982: 13981: 13977: 13974: 13973: 13969: 13966: 13965: 13961: 13959: 13958: 13954: 13952: 13951: 13947: 13946: 13944: 13942: 13938: 13932: 13931: 13927: 13925: 13924: 13920: 13917: 13916: 13912: 13909: 13908: 13904: 13902: 13901: 13897: 13896: 13894: 13892: 13888: 13882: 13881: 13877: 13876: 13873: 13870: 13868: 13864: 13861: 13858: 13857:Caelidracones 13853: 13847: 13846: 13842: 13839: 13838: 13834: 13833: 13830: 13827: 13824: 13819: 13801: 13800: 13799:Wukongopterus 13796: 13793: 13792: 13788: 13786: 13785: 13784:Darwinopterus 13781: 13779: 13778: 13777:Cuspicephalus 13774: 13771: 13770: 13766: 13765: 13763: 13761: 13757: 13751: 13750: 13746: 13745: 13742: 13739: 13737: 13733: 13727: 13726: 13725:Pterorhynchus 13722: 13720: 13719: 13715: 13714: 13711: 13708: 13706: 13702: 13695: 13694: 13690: 13688: 13687: 13683: 13681: 13680: 13676: 13675: 13672: 13668: 13665: 13662: 13657: 13650: 13649: 13645: 13642: 13641: 13637: 13636: 13633: 13630: 13627: 13622: 13618: 13614: 13609: 13605: 13596: 13592: 13589: 13581: 13535: 13534: 13533: 13532: 13530: 13528: 13524: 13517: 13516: 13512: 13510: 13509: 13508:Scaphognathus 13505: 13503: 13502: 13498: 13496: 13495: 13491: 13489: 13488: 13484: 13483: 13481: 13478: 13474: 13471: 13468: 13464: 13446: 13445: 13441: 13438: 13437: 13433: 13430: 13429: 13425: 13424: 13422: 13420: 13416: 13410: 13409: 13405: 13402: 13401: 13397: 13394: 13393: 13389: 13387: 13386: 13382: 13380: 13379: 13375: 13374: 13372: 13370: 13366: 13363: 13361: 13357: 13351: 13350: 13346: 13344: 13343: 13339: 13337: 13336: 13332: 13330: 13329: 13325: 13323: 13322: 13318: 13316: 13315: 13311: 13309: 13308: 13304: 13303: 13300: 13297: 13295: 13291: 13284: 13283: 13279: 13276: 13275: 13271: 13269: 13268: 13264: 13261: 13260: 13256: 13255: 13252: 13249: 13247: 13243: 13240: 13237: 13232: 13222: 13221: 13217: 13215: 13214: 13210: 13209: 13207: 13205: 13201: 13197: 13194: 13191: 13186: 13179: 13178: 13177:Eudimorphodon 13174: 13173: 13170: 13167: 13163: 13158: 13147: 13146: 13142: 13139: 13138: 13137:Peteinosaurus 13134: 13131: 13130: 13126: 13124: 13123: 13119: 13117: 13116: 13112: 13109: 13108: 13104: 13103: 13101: 13099: 13095: 13089: 13088: 13087:Herbstosaurus 13084: 13083: 13080: 13076: 13073: 13070: 13065: 13058: 13057: 13056:Peteinosaurus 13053: 13052: 13049: 13046: 13042: 13037: 13019: 13018: 13017:Eudimorphodon 13014: 13011: 13010: 13006: 13005: 13003: 13001: 12997: 12990: 12989: 12985: 12984: 12981: 12978: 12976: 12972: 12966: 12965: 12961: 12959: 12958: 12954: 12952: 12951: 12947: 12944: 12943: 12939: 12938: 12936: 12934: 12930: 12927: 12925: 12921: 12914: 12913: 12912:Preondactylia 12909: 12906: 12905: 12904:Peteinosaurus 12901: 12898: 12897: 12893: 12892: 12889: 12886: 12883: 12882:Eopterosauria 12878: 12864: 12863: 12859: 12857: 12856: 12852: 12849: 12848: 12844: 12843: 12841: 12839: 12835: 12828: 12827: 12823: 12821: 12820: 12816: 12813: 12812: 12808: 12807: 12804: 12801: 12798: 12794: 12788: 12787: 12786:Preondactylus 12783: 12781: 12780: 12776: 12775: 12773: 12771: 12770:Preondactylia 12767: 12761: 12760: 12756: 12755: 12752: 12748: 12745: 12742: 12737: 12733: 12729: 12724: 12720: 12699: 12698: 12697: 12696: 12694: 12692: 12688: 12680: 12676: 12675: 12674: 12673: 12670: 12667: 12665: 12661: 12657: 12649: 12645: 12642: 12640: 12636: 12633: 12631: 12627: 12624: 12622: 12618: 12616: 12612: 12610: 12606: 12605: 12600: 12596: 12592: 12585: 12580: 12578: 12573: 12571: 12566: 12565: 12562: 12550: 12542: 12540: 12537: 12535: 12531: 12530: 12527: 12518: 12514: 12511: 12505: 12500: 12494: 12458: 12454: 12452: 12448: 12447: 12446: 12445: 12443: 12441: 12437: 12429: 12425: 12424: 12423: 12422: 12420: 12418: 12411: 12408: 12406: 12402: 12392: 12391: 12387: 12385: 12384: 12383:Sanjuansaurus 12380: 12378: 12377: 12376:Herrerasaurus 12373: 12371: 12370: 12366: 12365: 12363: 12361: 12357: 12350: 12349: 12345: 12342: 12341: 12337: 12334: 12333: 12332:Daemonosaurus 12329: 12326: 12325: 12321: 12320: 12317: 12314: 12312: 12311:Herrerasauria 12305: 12299: 12298: 12291: 12288: 12287: 12280: 12278: 12277: 12270: 12268: 12267: 12266:Daemonosaurus 12260: 12258: 12257: 12250: 12248: 12247: 12240: 12239: 12236: 12233: 12230: 12225: 12213: 12209: 12208: 12207: 12206: 12204: 12202: 12195: 12191: 12188: 12185: 12180: 12166: 12165: 12161: 12159: 12158: 12154: 12152: 12151: 12147: 12145: 12144: 12140: 12138: 12137: 12133: 12131: 12130: 12129:Eucoelophysis 12126: 12124: 12123: 12119: 12117: 12116: 12112: 12111: 12109: 12107: 12103: 12096: 12095: 12091: 12089: 12088: 12084: 12081: 12080: 12076: 12074: 12073: 12069: 12067: 12066: 12062: 12060: 12059: 12055: 12053: 12052: 12048: 12047: 12044: 12041: 12038: 12031: 12025: 12024: 12017: 12014: 12013: 12006: 12004: 12003: 11996: 11994: 11993: 11986: 11985: 11982: 11978: 11975: 11972: 11967: 11951: 11947: 11946: 11945: 11944: 11942: 11940: 11936: 11930: 11929: 11925: 11922: 11921: 11920:Scleromochlus 11917: 11915: 11914: 11910: 11908: 11907: 11903: 11901: 11900: 11896: 11893: 11892: 11888: 11886: 11885: 11881: 11880: 11878: 11876: 11872: 11869: 11867: 11860: 11856: 11853: 11850: 11845: 11835: 11834: 11830: 11828: 11827: 11823: 11821: 11820: 11816: 11814: 11813: 11809: 11806: 11805: 11801: 11800: 11798: 11796: 11789: 11783: 11782: 11775: 11772: 11771: 11764: 11763: 11760: 11756: 11753: 11750: 11745: 11741: 11729: 11728: 11722: 11705: 11704: 11703: 11702: 11700: 11698: 11694: 11686: 11682: 11681: 11680: 11679: 11676: 11673: 11671: 11667: 11659: 11655: 11654: 11653: 11652: 11649: 11646: 11644: 11640: 11636: 11628: 11624: 11621: 11619: 11615: 11613: 11609: 11607: 11603: 11602: 11597: 11593: 11589: 11582: 11577: 11575: 11570: 11568: 11563: 11562: 11559: 11552: 11549: 11546: 11543: 11540: 11539:Pterosaur.net 11537: 11536: 11526: 11520: 11516: 11511: 11507: 11501: 11497: 11492: 11488: 11482: 11478: 11473: 11472: 11459: 11455: 11450: 11445: 11441: 11437: 11433: 11429: 11428: 11423: 11416: 11408: 11401: 11386: 11385: 11377: 11370: 11355: 11354: 11349: 11342: 11335: 11332: 11328: 11322: 11320: 11312: 11307: 11300: 11299:1-86239-143-2 11296: 11292: 11289: 11285: 11278: 11271: 11270:Pterosaur.net 11268: 11262: 11260: 11258: 11249: 11245: 11241: 11237: 11233: 11229: 11225: 11221: 11217: 11213: 11206: 11192: 11188: 11181: 11173: 11169: 11164: 11159: 11155: 11151: 11147: 11143: 11139: 11135: 11131: 11124: 11116: 11112: 11107: 11102: 11098: 11094: 11090: 11086: 11082: 11078: 11074: 11067: 11059: 11055: 11050: 11045: 11041: 11037: 11033: 11029: 11025: 11018: 11004: 11000: 10994: 10986: 10982: 10977: 10972: 10967: 10962: 10958: 10954: 10951:(2): e31392. 10950: 10946: 10942: 10940: 10931: 10929: 10920: 10916: 10912: 10908: 10904: 10900: 10896: 10892: 10889:(3): 569–80. 10888: 10884: 10880: 10873: 10871: 10869: 10854: 10850: 10844: 10842: 10833: 10829: 10825: 10821: 10816: 10811: 10807: 10803: 10800:(6992): 621. 10799: 10795: 10791: 10784: 10776: 10772: 10767: 10762: 10758: 10754: 10750: 10743: 10735: 10731: 10727: 10723: 10719: 10715: 10712:(4): 273–77. 10711: 10707: 10700: 10692: 10688: 10683: 10678: 10674: 10670: 10666: 10662: 10658: 10651: 10649: 10640: 10636: 10632: 10628: 10624: 10620: 10616: 10612: 10608: 10604: 10597: 10589: 10585: 10580: 10575: 10571: 10567: 10563: 10559: 10555: 10551: 10547: 10540: 10532: 10528: 10524: 10520: 10515: 10510: 10506: 10502: 10498: 10494: 10490: 10483: 10475: 10471: 10467: 10460: 10452: 10448: 10444: 10440: 10436: 10432: 10428: 10424: 10421:(7017): 572. 10420: 10416: 10409: 10402: 10394: 10390: 10386: 10380: 10372: 10368: 10364: 10360: 10355: 10350: 10346: 10342: 10338: 10334: 10330: 10323: 10315: 10311: 10307: 10303: 10296: 10294: 10292: 10283: 10279: 10275: 10271: 10267: 10263: 10259: 10252: 10244: 10240: 10239: 10231: 10223: 10219: 10214: 10209: 10204: 10199: 10195: 10191: 10187: 10180: 10171: 10166: 10162: 10158: 10154: 10147: 10140: 10134: 10126: 10122: 10117: 10112: 10107: 10102: 10098: 10094: 10090: 10086: 10082: 10080: 10071: 10064: 10059: 10052: 10047: 10039: 10035: 10031: 10027: 10023: 10019: 10015: 10011: 10007: 10003: 9999: 9992: 9984: 9980: 9976: 9972: 9968: 9964: 9961:(3): 507–14. 9960: 9956: 9949: 9942: 9937: 9929: 9925: 9921: 9917: 9913: 9909: 9906:(1): 163–87. 9905: 9901: 9894: 9887: 9880: 9875: 9868: 9863: 9856: 9851: 9843: 9839: 9835: 9831: 9827: 9823: 9820:(4): 376–98. 9819: 9815: 9811: 9804: 9797: 9792: 9790: 9788: 9779: 9775: 9772:(4): 373–86. 9771: 9767: 9760: 9753: 9745: 9741: 9737: 9733: 9729: 9725: 9722:(4): 421–35. 9721: 9717: 9713: 9706: 9698: 9694: 9690: 9686: 9682: 9678: 9674: 9670: 9663: 9656: 9648: 9644: 9639: 9634: 9629: 9624: 9620: 9616: 9612: 9605: 9597: 9593: 9586: 9578: 9574: 9570: 9566: 9559: 9551: 9547: 9543: 9539: 9534: 9529: 9525: 9521: 9517: 9510: 9495: 9488: 9482: 9474: 9470: 9463: 9455: 9451: 9444: 9437: 9422: 9418: 9411: 9409: 9392: 9388: 9382: 9380: 9378: 9361: 9357: 9356: 9351: 9347: 9346:Naish, Darren 9341: 9333: 9327: 9323: 9322: 9314: 9299: 9295: 9288: 9272: 9268: 9267:The Telegraph 9264: 9257: 9255: 9246: 9242: 9238: 9234: 9230: 9226: 9222: 9218: 9215:(9): 891–97. 9214: 9210: 9203: 9196: 9192: 9188: 9182: 9174: 9172:9780429821202 9168: 9164: 9163: 9155: 9149: 9145: 9141: 9135: 9127: 9123: 9119: 9115: 9111: 9107: 9104:(1): 105–37. 9103: 9099: 9092: 9084: 9080: 9076: 9072: 9068: 9064: 9060: 9053: 9051: 9042: 9038: 9031: 9023: 9019: 9014: 9009: 9005: 9001: 8997: 8993: 8990:(8): 160333. 8989: 8985: 8981: 8974: 8966: 8962: 8958: 8957:10.1666/13030 8954: 8950: 8946: 8942: 8938: 8931: 8924: 8910:on 2013-01-15 8906: 8902: 8898: 8894: 8890: 8887:(4): 883–94. 8886: 8882: 8881:Geodiversitas 8875: 8873: 8864: 8856: 8852: 8848: 8844: 8840: 8836: 8832: 8828: 8821: 8813: 8809: 8804: 8799: 8794: 8789: 8785: 8781: 8777: 8770: 8768: 8766: 8757: 8753: 8749: 8745: 8741: 8737: 8734:(3): 432–46. 8733: 8729: 8721: 8713: 8709: 8704: 8699: 8695: 8691: 8687: 8680: 8673: 8667: 8659: 8655: 8650: 8645: 8640: 8635: 8631: 8627: 8623: 8616: 8614: 8612: 8610: 8602: 8596: 8590:, p. 18. 8589: 8584: 8576: 8572: 8568: 8564: 8560: 8556: 8549: 8535: 8531: 8524: 8510: 8506: 8500: 8492: 8488: 8484: 8480: 8476: 8472: 8468: 8464: 8460: 8456: 8449: 8441: 8439: 8430: 8426: 8421: 8416: 8411: 8406: 8402: 8398: 8394: 8392: 8383: 8375: 8371: 8366: 8361: 8356: 8351: 8347: 8343: 8339: 8332: 8324: 8320: 8316: 8312: 8307: 8306:10.1206/352.1 8302: 8298: 8294: 8290: 8283: 8275: 8271: 8267: 8263: 8260:(4): 465–69. 8259: 8255: 8248: 8246: 8237: 8233: 8229: 8225: 8221: 8217: 8213: 8209: 8205: 8201: 8194: 8187: 8178: 8173: 8169: 8165: 8161: 8154: 8145: 8140: 8136: 8132: 8128: 8124: 8120: 8118: 8109: 8102: 8097: 8091:, p. 13. 8090: 8085: 8077: 8073: 8067: 8061:, p. 10. 8060: 8055: 8053: 8046: 8041: 8039: 8030: 8026: 8022: 8018: 8013: 8008: 8005:(4): 890–98. 8004: 8000: 7996: 7989: 7982: 7977: 7970: 7964: 7957: 7952: 7950: 7948: 7946: 7944: 7935: 7931: 7927: 7923: 7919: 7915: 7911: 7907: 7904:(3): 218–39. 7903: 7899: 7892: 7890: 7882: 7876: 7868: 7864: 7857: 7850: 7844: 7837: 7831: 7817:on 2020-07-28 7813: 7809: 7805: 7801: 7797: 7793: 7789: 7788: 7780: 7773: 7766: 7760: 7752: 7748: 7741: 7733: 7729: 7722: 7715: 7710: 7704:, p. 31. 7703: 7698: 7696: 7689:, p. 36. 7688: 7683: 7681: 7679: 7672:, p. 35. 7671: 7666: 7664: 7655: 7651: 7647: 7643: 7639: 7635: 7631: 7627: 7620: 7613: 7607: 7600: 7594: 7588:, p. 33. 7587: 7582: 7576:, p. 29. 7575: 7570: 7564:, p. 28. 7563: 7558: 7550: 7546: 7545:Isis von Oken 7542: 7535: 7527: 7523: 7516: 7510:, p. 27. 7509: 7504: 7497: 7493: 7486: 7479: 7473: 7466: 7462: 7455: 7447: 7443: 7436: 7429: 7423: 7416: 7410: 7403: 7398: 7396: 7394: 7392: 7384: 7379: 7372: 7367: 7361:, p. 22. 7360: 7355: 7348: 7343: 7341: 7332: 7328: 7323: 7318: 7314: 7310: 7306: 7302: 7298: 7294: 7290: 7283: 7275: 7271: 7267: 7263: 7258: 7253: 7249: 7245: 7241: 7234: 7226: 7222: 7218: 7214: 7209: 7204: 7200: 7196: 7192: 7188: 7184: 7180: 7172: 7164: 7160: 7156: 7152: 7148: 7144: 7140: 7136: 7129: 7121: 7117: 7112: 7107: 7103: 7099: 7095: 7091: 7087: 7080: 7072: 7068: 7064: 7060: 7056: 7052: 7048: 7044: 7040: 7036: 7029: 7015: 7014: 7009: 7002: 6994: 6990: 6986: 6982: 6977: 6972: 6968: 6964: 6960: 6956: 6952: 6945: 6937: 6933: 6926: 6920:, p. 51. 6919: 6914: 6912: 6910: 6908: 6906: 6899:, p. 48. 6898: 6893: 6887:, p. 47. 6886: 6881: 6875:, p. 43. 6874: 6869: 6863:, p. 39. 6862: 6857: 6851:, p. 37. 6850: 6845: 6837: 6833: 6828: 6823: 6818: 6813: 6809: 6805: 6801: 6797: 6793: 6786: 6784: 6782: 6780: 6773:, p. 36. 6772: 6767: 6765: 6763: 6756:, p. 57. 6755: 6750: 6748: 6746: 6744: 6742: 6740: 6733:, p. 56. 6732: 6727: 6725: 6723: 6721: 6719: 6717: 6708: 6704: 6700: 6696: 6691: 6686: 6682: 6678: 6673: 6668: 6664: 6660: 6656: 6652: 6648: 6641: 6634: 6629: 6623:, p. 55. 6622: 6617: 6615: 6613: 6611: 6604:, p. 35. 6603: 6598: 6596: 6594: 6592: 6590: 6588: 6586: 6584: 6575: 6571: 6566: 6561: 6557: 6553: 6549: 6545: 6544: 6539: 6532: 6524: 6520: 6515: 6510: 6506: 6502: 6498: 6491: 6483: 6479: 6476:(3): 265–75. 6475: 6471: 6464: 6456: 6452: 6448: 6444: 6441:(4): 881–91. 6440: 6436: 6429: 6422: 6414: 6410: 6405: 6400: 6396: 6392: 6388: 6384: 6383: 6378: 6371: 6365:, p. 34. 6364: 6359: 6353:, p. 33. 6352: 6347: 6341:, p. 53. 6340: 6335: 6333: 6331: 6322: 6318: 6314: 6310: 6305: 6300: 6296: 6292: 6291:J. Evol. Biol 6288: 6281: 6273: 6269: 6265: 6261: 6257: 6253: 6250:(1): 233–66. 6249: 6245: 6238: 6230: 6226: 6222: 6218: 6214: 6210: 6206: 6202: 6195: 6187: 6183: 6179: 6175: 6171: 6167: 6163: 6159: 6155: 6148: 6146: 6139:, p. 55. 6138: 6133: 6131: 6124:, p. 52. 6123: 6118: 6116: 6107: 6103: 6098: 6093: 6088: 6083: 6079: 6075: 6071: 6067: 6063: 6056: 6054: 6045: 6041: 6036: 6031: 6027: 6023: 6019: 6015: 6011: 6009: 6000: 5998: 5996: 5987: 5983: 5979: 5975: 5971: 5967: 5964:(4): 255–84. 5963: 5959: 5952: 5946:, p. 53. 5945: 5940: 5938: 5931:, p. 54. 5930: 5925: 5919:, p. 32. 5918: 5913: 5911: 5909: 5902:, p. 44. 5901: 5896: 5890:, p. 52. 5889: 5884: 5882: 5880: 5878: 5876: 5869:, p. 51. 5868: 5863: 5861: 5854:, p. 31. 5853: 5848: 5846: 5844: 5837:, p. 30. 5836: 5831: 5829: 5827: 5820:, p. 46. 5819: 5814: 5812: 5810: 5808: 5801:, p. 45. 5800: 5795: 5789:, p. 50. 5788: 5783: 5781: 5779: 5772:, p. 28. 5771: 5766: 5764: 5755: 5751: 5747: 5745: 5736: 5730:, p. 49. 5729: 5724: 5717: 5716:1-932075-01-1 5713: 5709: 5703: 5701: 5692: 5688: 5681: 5679: 5677: 5675: 5668:, p. 48. 5667: 5662: 5654: 5650: 5647:(3): 421–41. 5646: 5642: 5638: 5631: 5629: 5622:, p. 24. 5621: 5616: 5614: 5612: 5605:, p. 26. 5604: 5599: 5597: 5595: 5588:, p. 47. 5587: 5582: 5580: 5573:, p. 27. 5572: 5567: 5565: 5556: 5550: 5546: 5539: 5537: 5535: 5533: 5531: 5529: 5527: 5520:, p. 23. 5519: 5514: 5512: 5510: 5503:, p. 58. 5502: 5497: 5489: 5485: 5478: 5470: 5466: 5462: 5458: 5454: 5450: 5446: 5442: 5435: 5428: 5414: 5413:pterosaur.net 5410: 5404: 5396: 5392: 5388: 5384: 5380: 5376: 5372: 5368: 5364: 5360: 5353: 5346: 5344: 5342: 5340: 5331: 5325: 5321: 5320: 5312: 5310: 5293: 5286: 5284: 5282: 5273: 5267: 5263: 5258: 5257: 5251: 5245: 5237: 5233: 5229: 5225: 5221: 5217: 5213: 5209: 5206:(4): 180–84. 5205: 5201: 5194: 5187: 5179: 5175: 5171: 5167: 5163: 5159: 5155: 5151: 5147: 5143: 5139: 5133: 5125: 5121: 5116: 5111: 5106: 5101: 5097: 5093: 5089: 5085: 5081: 5074: 5072: 5063: 5057: 5043: 5039: 5032: 5018: 5017:pterosaur.net 5014: 5008: 4999: 4994: 4991:(1): 99–111. 4990: 4986: 4982: 4975: 4967: 4960: 4954: 4946: 4945: 4937: 4929: 4928: 4923: 4917: 4910: 4904: 4900: 4896: 4895:Jones, Daniel 4890: 4882: 4878: 4874: 4873:Darwinopterus 4867: 4860: 4854: 4850: 4846: 4842: 4835: 4827: 4823: 4818: 4813: 4808: 4803: 4799: 4795: 4791: 4784: 4776: 4772: 4767: 4762: 4758: 4754: 4750: 4746: 4742: 4735: 4733: 4728: 4715: 4709: 4692: 4686: 4682: 4676: 4670: 4669: 4656: 4613: 4604: 4600: 4590: 4587: 4585: 4582: 4580: 4577: 4575: 4572: 4570: 4567: 4565: 4562: 4560: 4557: 4555: 4552: 4551: 4547: 4536: 4533: 4527: 4522: 4519: 4508: 4505: 4494: 4487: 4484: 4483: 4479: 4478:wukongopterid 4475: 4474: 4470: 4466: 4465:birds of prey 4462: 4457: 4455: 4454: 4449: 4445: 4444: 4438: 4436: 4435: 4430: 4428: 4423: 4419: 4418: 4413: 4412: 4407: 4403: 4399: 4398: 4394:, and 1966's 4393: 4392: 4387: 4383: 4382: 4376: 4374: 4370: 4366: 4365: 4360: 4356: 4352: 4351: 4346: 4342: 4341: 4336: 4335: 4327: 4326: 4321: 4320: 4314: 4310: 4303: 4299: 4295: 4291: 4284: 4282: 4278: 4277:Pterodactylus 4274: 4273:Scaphognathus 4270: 4266: 4262: 4258: 4254: 4250: 4246: 4242: 4238: 4237: 4232: 4228: 4227: 4222: 4221: 4216: 4215: 4210: 4206: 4205: 4200: 4199: 4198:Scaphognathus 4194: 4193: 4192:Pterodactylus 4188: 4187:scleral rings 4178: 4175: 4173: 4169: 4168: 4163: 4159: 4155: 4151: 4147: 4142: 4140: 4136: 4131: 4127: 4123: 4119: 4116: 4112: 4106: 4104: 4100: 4096: 4092: 4091:Pterodactylus 4088: 4083: 4079: 4074: 4072: 4068: 4067:Darwinopterus 4063: 4061: 4057: 4053: 4048: 4047: 4042: 4041: 4040:Darwinopterus 4036: 4035: 4030: 4026: 4022: 4018: 4009: 4004: 3995: 3993: 3992: 3987: 3983: 3982: 3977: 3976: 3971: 3967: 3966: 3961: 3957: 3953: 3949: 3948: 3943: 3933: 3931: 3930: 3925: 3924: 3919: 3918: 3913: 3909: 3908: 3903: 3899: 3898: 3893: 3892: 3887: 3886: 3881: 3877: 3873: 3869: 3865: 3860: 3858: 3857: 3852: 3851: 3850:Hatzegopteryx 3846: 3842: 3838: 3833: 3832: 3827: 3823: 3819: 3815: 3814:Azhdarchoidea 3812:In contrast, 3810: 3808: 3804: 3803: 3798: 3794: 3793: 3792:Pterodactylus 3788: 3787: 3782: 3778: 3774: 3770: 3766: 3765: 3760: 3756: 3752: 3748: 3747: 3742: 3738: 3734: 3730: 3729:Boreopteridae 3726: 3722: 3721:Pteranodontia 3717: 3715: 3711: 3710: 3709:Darwinopterus 3705: 3701: 3697: 3693: 3692: 3687: 3686: 3685:Scaphognathus 3681: 3680: 3675: 3674: 3669: 3668: 3663: 3658: 3656: 3652: 3651: 3646: 3645: 3640: 3639:Eudimorphodon 3636: 3635: 3634:Eudimorphodon 3630: 3629: 3623: 3622: 3617: 3613: 3609: 3608: 3602: 3600: 3596: 3591: 3582: 3580: 3576: 3572: 3571: 3567: 3563: 3559: 3555: 3544: 3542: 3541: 3540:Scleromochlus 3536: 3535: 3530: 3529: 3528:Pterodactylus 3524: 3520: 3516: 3515: 3508: 3505: 3500: 3496: 3491: 3489: 3480: 3479: 3478:Hatzegopteryx 3474: 3469: 3465: 3462: 3458: 3454: 3450: 3445: 3443: 3439: 3435: 3434: 3429: 3425: 3421: 3416: 3414: 3406: 3403: 3399: 3396:The probable 3394: 3385: 3383: 3379: 3378: 3373: 3367: 3364: 3360: 3358: 3353: 3349: 3339: 3337: 3333: 3329: 3318: 3316: 3311: 3309: 3308: 3303: 3302: 3297: 3292: 3289: 3285: 3281: 3277: 3270: 3269: 3263: 3259: 3257: 3253: 3249: 3245: 3241: 3237: 3232: 3220: 3212: 3197: 3196: 3188: 3187: 3175: 3169: 3168: 3156: 3150: 3149: 3137: 3131: 3130: 3118: 3112: 3111: 3099: 3093: 3092: 3080: 3079:Caelidracones 3074: 3073: 3065: 3064: 3056: 3055: 3047: 3046: 3038: 3037: 3029: 3028: 3020: 3019: 3011: 3010: 3007: 3005: 3001: 2995: 2994: 2991: 2990: 2987: 2985: 2981: 2980:Pteranodontia 2975: 2974: 2968: 2965: 2964: 2961: 2960: 2957: 2956: 2955: 2948: 2947: 2941: 2940: 2937: 2936: 2928: 2927: 2924: 2922: 2918: 2917:Neoazhdarchia 2912: 2911: 2908: 2907: 2904: 2902: 2898: 2892: 2891: 2885: 2884:Azhdarchoidea 2882: 2881: 2875: 2872: 2871: 2868: 2867: 2864: 2863: 2862: 2855: 2854: 2848: 2845: 2844: 2841: 2840: 2832: 2831: 2823: 2822: 2819: 2817: 2813: 2807: 2806: 2803: 2802: 2799: 2797: 2793: 2792: 2791:Pterodactylus 2785: 2784: 2778: 2775: 2774: 2771: 2770: 2767: 2765: 2761: 2755: 2754: 2748: 2745: 2744: 2738: 2735: 2734: 2731: 2730: 2727: 2725: 2721: 2720: 2713: 2712: 2706: 2703: 2702: 2699: 2698: 2695: 2693: 2689: 2683: 2682: 2676: 2675: 2672: 2671: 2668: 2667: 2666: 2659: 2658: 2652: 2651: 2648: 2647: 2644: 2643: 2637: 2636: 2630: 2629: 2626: 2625: 2622: 2620: 2616: 2615: 2608: 2607: 2601: 2600: 2597: 2596: 2593: 2591: 2587: 2581: 2580: 2574: 2573: 2570: 2569: 2566: 2564: 2560: 2554: 2553: 2547: 2546: 2543: 2542: 2539: 2537: 2533: 2527: 2526: 2520: 2517: 2516: 2513: 2512: 2509: 2507: 2503: 2502:Eopterosauria 2497: 2496: 2490: 2487: 2486: 2481: 2479: 2475: 2471: 2465: 2463: 2459: 2455: 2451: 2447: 2443: 2442:fossil record 2439: 2436:The internal 2434: 2432: 2428: 2427:ornithodirans 2424: 2420: 2416: 2412: 2411:Pterodactylus 2408: 2403: 2399: 2398: 2397:Preondactylus 2392: 2388: 2382: 2372: 2368: 2366: 2362: 2357: 2355: 2351: 2347: 2342: 2340: 2339: 2334: 2330: 2326: 2325: 2320: 2319: 2314: 2313:Maastrichtian 2310: 2305: 2303: 2298: 2294: 2286: 2282: 2277: 2268: 2266: 2265: 2261: 2256: 2254: 2253:Scleromochlus 2249: 2245: 2239: 2237: 2234:also showing 2233: 2229: 2225: 2221: 2217: 2213: 2209: 2206: 2202: 2201:Scleromochlus 2198: 2197: 2196:Scleromochlus 2192: 2188: 2187:Scleromochlus 2184: 2180: 2179: 2174: 2173:Scleromochlus 2170: 2166: 2161: 2153: 2149: 2147: 2146:Sharovipteryx 2141: 2134: 2130: 2128: 2127:Scleromochlus 2122: 2118: 2117: 2113: 2111: 2110:Scleromochlus 2106: 2102: 2101: 2100:Sharovipteryx 2096: 2095: 2090: 2086: 2082: 2078: 2074: 2070: 2066: 2062: 2057: 2053: 2045: 2041: 2037: 2033: 2032: 2026: 2012: 2010: 2006: 2002: 1998: 1994: 1993: 1987: 1985: 1981: 1977: 1972: 1968: 1964: 1959: 1957: 1952: 1948: 1941: 1937: 1933: 1932: 1926: 1922: 1920: 1916: 1912: 1907: 1903: 1902:Robert Bakker 1899: 1898: 1893: 1892: 1887: 1879: 1878: 1872: 1863: 1861: 1857: 1853: 1847: 1845: 1844:Tilly Edinger 1842: 1838: 1834: 1833: 1828: 1824: 1820: 1819: 1814: 1813: 1812:Scaphognathus 1808: 1807: 1801: 1799: 1795: 1791: 1786: 1784: 1780: 1776: 1775:Ornithosauria 1772: 1767: 1764: 1760: 1756: 1755: 1750: 1742: 1737: 1728: 1726: 1722: 1718: 1717:Edward Newman 1713: 1708: 1706: 1702: 1701:Pterodactylus 1698: 1697: 1696:Pterodactylus 1692: 1691:Ptéro-Dactyle 1688: 1684: 1675: 1671: 1669: 1665: 1664:Ichthyosauria 1661: 1660:Pterodactylus 1657: 1652: 1648: 1644: 1641: 1640:Late Jurassic 1637: 1633: 1632: 1627: 1619: 1615: 1614: 1608: 1598: 1588: 1585: 1584:Tupandactylus 1582:relatives of 1580: 1579: 1573: 1570: 1569:Pterorhynchus 1566: 1565: 1560: 1559: 1554: 1549: 1548:anurognathids 1545: 1541: 1537: 1532: 1530: 1526: 1516: 1514: 1507: 1503: 1499: 1497: 1496: 1491: 1490: 1485: 1481: 1477: 1476: 1471: 1467: 1463: 1459: 1455: 1451: 1443: 1439: 1430: 1428: 1424: 1420: 1410: 1406: 1403: 1398: 1393: 1391: 1380: 1376: 1374: 1370: 1366: 1361: 1357: 1348: 1339: 1337: 1336: 1335:Pterodactylus 1330: 1326: 1324: 1319: 1315: 1311: 1306: 1302: 1299: 1298: 1293: 1292: 1287: 1281: 1279: 1275: 1271: 1266: 1261: 1258: 1254: 1250: 1246: 1245: 1241: 1237: 1236: 1230: 1228: 1224: 1223:cruropatagium 1220: 1219: 1214: 1210: 1209: 1204: 1196: 1193: 1189: 1185: 1184: 1179: 1170: 1168: 1163: 1161: 1157: 1156: 1155:actinofibrils 1146: 1145: 1139: 1135: 1132: 1128: 1119: 1110: 1108: 1104: 1100: 1095: 1092: 1088: 1084: 1080: 1070: 1068: 1064: 1059: 1055: 1046: 1042: 1039: 1038:neural spines 1034: 1030: 1024: 1017: 1012: 1008: 1006: 1002: 998: 994: 990: 983: 982: 976: 967: 965: 961: 960:ossa dentalia 955: 953: 952: 947: 946:Pterorhynchus 943: 939: 938: 937:Pterodactylus 933: 932: 931:Pterorhynchus 925: 923: 922: 917: 913: 909: 908: 900: 899: 895:The skull of 893: 889: 887: 883: 882: 877: 873: 869: 865: 861: 853: 852: 847: 846: 841: 840: 835: 830: 826: 824: 820: 816: 812: 808: 804: 800: 795: 787: 786: 780: 771: 769: 764: 756: 751: 741: 739: 735: 731: 730: 726: 722: 716: 713: 711: 710: 705: 704: 699: 698: 693: 692: 687: 686: 681: 680: 679:Pterodactylus 675: 674: 669: 668: 663: 662: 657: 656: 655:Istiodactylus 650: 648: 647: 646:Scaphognathus 642: 641: 636: 635: 630: 629: 628:Eudimorphodon 624: 623: 617: 615: 611: 607: 603: 599: 595: 591: 587: 583: 573: 571: 565: 563: 559: 558: 557:Pterodactylus 553: 549: 545: 541: 537: 532: 530: 526: 525: 524:Hatzegopteryx 520: 519: 514: 513:anurognathids 510: 506: 502: 498: 494: 490: 486: 482: 477: 475: 470: 466: 462: 458: 454: 449: 447: 443: 439: 435: 431: 427: 423: 420: 416: 412: 404: 401: 397: 396:Ornithosauria 392: 389: 385: 381: 376: 372: 367: 360: 354: 352: 346: 343: 342: 341:Peteinosaurus 335: 332: 331: 330:Eudimorphodon 324: 323: 322: 316: 312: 311: 304: 302: 301: 300:Preondactylus 294: 293: 292: 291:Preondactylia 286: 284: 283:Eopterosauria 278: 276: 270: 269: 267: 262: 257: 252: 246: 243: 242: 239: 233: 230: 227: 226: 223: 220: 217: 214: 213: 210: 207: 204: 203: 200: 197: 194: 193: 190: 187: 184: 183: 178: 173: 169: 166: 165: 164:Tropeognathus 160: 159: 154: 153: 148: 147: 142: 141: 140:Pterodactylus 136: 135: 129: 125: 120: 113: 108: 103: 98: 93: 88: 83: 78: 73: 68: 63: 58: 52: 45: 41: 40:Late Triassic 33: 30: 26: 22: 21:Pterodactylus 17608: 17173: 17119: 17114:Ludodactylus 17112: 17105: 17098: 17090: 17082: 17075: 17055: 17047: 17042:Nicorhynchus 17040: 17033: 17026: 17005: 16998: 16990: 16982: 16974: 16966: 16958: 16928: 16921: 16913:Hamipteridae 16899: 16891: 16857: 16849: 16841: 16821: 16813: 16808:Aetodactylus 16806: 16771: 16763: 16756: 16748: 16742:Ludodactylus 16740: 16732: 16724: 16716: 16708: 16700: 16694:Aetodactylus 16692: 16671: 16663: 16655: 16647: 16639: 16632: 16625: 16582: 16575: 16551: 16544: 16537: 16530: 16504:Liaoxipterus 16502: 16495: 16475: 16468: 16461: 16454: 16432:Mimodactylus 16430: 16423: 16416: 16395: 16388: 16381: 16374: 16350: 16343: 16318: 16310:Lanceodontia 16291: 16283: 16278:Lonchodectes 16276: 16268: 16260: 16240: 16233: 16226: 16179: 16172: 16165: 16158: 16138: 16131: 16124: 16099: 16092: 16067: 16059: 16052: 16046:Ornithostoma 16044: 16037: 16029: 16000: 15940: 15854: 15846: 15839: 15832: 15825: 15818: 15813:Eurazhdarcho 15811: 15804: 15797: 15790: 15770: 15763: 15756: 15748: 15741:Azhdarchinae 15727: 15719: 15712: 15704: 15697: 15689: 15682:Azhdarchidae 15668: 15661: 15653: 15625: 15618: 15611: 15590: 15582: 15577:Meilifeilong 15575: 15568: 15560: 15552: 15544: 15520: 15495: 15487: 15479: 15471: 15463: 15456: 15434: 15427: 15420: 15412: 15362: 15355: 15335: 15328: 15321: 15313: 15289: 15281: 15273: 15265: 15257: 15233: 15226: 15219: 15214:Huaxiapterus 15212: 15205: 15199:Eopteranodon 15197: 15189: 15183:Afrotapejara 15181: 15174:Sinopterinae 15163:Afrotapejara 15161: 15137: 15130: 15099: 15092: 15085: 15077: 15069: 15062: 15054: 15046: 15038: 15030: 15008: 15003:Bennettazhia 15001: 14966:Ornithostoma 14964: 14957: 14949: 14941: 14934: 14927: 14893: 14886: 14865: 14857: 14852:Ordosipterus 14850: 14843: 14835: 14827: 14802:Tapejaroidea 14787: 14691: 14645:Pterodaustro 14643: 14636: 14629: 14622: 14602: 14578: 14571: 14564: 14558:Huanhepterus 14556: 14551:Gnathosaurus 14549: 14531:Moganopterus 14529: 14522: 14503:Pterofiltrus 14501: 14496:Petrodactyle 14494: 14487: 14482:Liaodactylus 14480: 14475:Kepodactylus 14473: 14466: 14459: 14452: 14445: 14438: 14431: 14423: 14390:Huanhepterus 14388: 14381: 14373: 14352: 14347:Aerodactylus 14345: 14320: 14314:Petrodactyle 14312: 14304: 14297: 14289: 14254: 14247: 14216: 14208: 14201: 14193: 14173: 14064: 14056: 14049: 14042: 14035: 14030:Kryptodrakon 14028: 14020: 14013: 14006: 13978: 13972:Jeholopterus 13970: 13962: 13955: 13948: 13928: 13921: 13915:Jeholopterus 13913: 13905: 13900:Anurognathus 13898: 13880:Mesadactylus 13878: 13843: 13835: 13797: 13789: 13782: 13775: 13767: 13747: 13723: 13716: 13705:Darwinoptera 13691: 13684: 13677: 13646: 13638: 13584: 13513: 13506: 13499: 13492: 13485: 13442: 13436:Nesodactylus 13434: 13428:Cacibupteryx 13426: 13408:Sericipterus 13406: 13398: 13390: 13383: 13376: 13347: 13340: 13335:Nesodactylus 13333: 13326: 13319: 13314:Cacibupteryx 13312: 13307:Bellubrunnus 13305: 13280: 13272: 13265: 13257: 13218: 13211: 13175: 13162:Lonchognatha 13143: 13135: 13127: 13120: 13113: 13105: 13085: 13054: 13015: 13007: 12986: 12962: 12955: 12950:Pachagnathus 12948: 12940: 12910: 12902: 12896:Austriadraco 12894: 12860: 12855:Austriadraco 12853: 12845: 12824: 12817: 12809: 12784: 12777: 12757: 12740: 12727: 12690: 12643: 12634: 12625: 12590: 12496: 12405:Eusaurischia 12388: 12381: 12374: 12367: 12346: 12338: 12330: 12324:Chindesaurus 12322: 12295: 12284: 12274: 12264: 12256:Chindesaurus 12254: 12244: 12212:Ornithischia 12201:Ornithischia 12162: 12155: 12148: 12141: 12136:Ignotosaurus 12134: 12127: 12120: 12113: 12094:Technosaurus 12092: 12087:Soumyasaurus 12085: 12079:Pisanosaurus 12077: 12070: 12063: 12056: 12049: 12037:Silesauridae 12021: 12010: 12000: 11990: 11938: 11928:Venetoraptor 11926: 11918: 11911: 11904: 11897: 11889: 11882: 11875:Lagerpetidae 11831: 11824: 11819:Spondylosoma 11817: 11810: 11802: 11795:Aphanosauria 11779: 11768: 11725: 11622: 11514: 11495: 11476: 11431: 11425: 11415: 11406: 11400: 11388:. Retrieved 11382: 11369: 11357:. Retrieved 11351: 11341: 11333: 11330: 11306: 11290: 11287: 11283: 11277: 11269: 11215: 11211: 11205: 11194:. Retrieved 11190: 11180: 11137: 11133: 11123: 11083:(1): 13130. 11080: 11076: 11066: 11031: 11027: 11017: 11006:. Retrieved 11002: 10993: 10948: 10944: 10938: 10886: 10882: 10878: 10856:. Retrieved 10852: 10797: 10793: 10783: 10756: 10752: 10742: 10709: 10705: 10699: 10664: 10660: 10606: 10602: 10596: 10553: 10549: 10539: 10514:11336/156308 10496: 10492: 10482: 10473: 10469: 10459: 10418: 10414: 10401: 10393:the original 10388: 10379: 10339:(6995): 33. 10336: 10332: 10322: 10305: 10301: 10265: 10261: 10251: 10243:the original 10237: 10230: 10193: 10189: 10179: 10160: 10156: 10146: 10138: 10133: 10088: 10084: 10078: 10070: 10058: 10046: 10005: 10001: 9991: 9958: 9954: 9948: 9936: 9903: 9899: 9886: 9874: 9862: 9850: 9817: 9813: 9809: 9803: 9798:, p. 51 9769: 9765: 9752: 9719: 9715: 9705: 9672: 9668: 9655: 9618: 9614: 9604: 9595: 9585: 9568: 9564: 9558: 9523: 9519: 9509: 9497:. Retrieved 9493: 9481: 9472: 9462: 9453: 9449: 9436: 9424:. Retrieved 9421:NewScientist 9420: 9395:. Retrieved 9390: 9364:. Retrieved 9355:ScienceBlogs 9353: 9340: 9320: 9313: 9301:. Retrieved 9298:NewScientist 9297: 9287: 9275:. Retrieved 9271:the original 9266: 9212: 9208: 9202: 9186: 9181: 9161: 9154: 9139: 9134: 9101: 9097: 9091: 9066: 9062: 9040: 9030: 8987: 8983: 8973: 8940: 8937:Paleobiology 8936: 8923: 8912:. Retrieved 8905:the original 8884: 8880: 8871: 8863: 8830: 8826: 8820: 8783: 8780:PLOS Biology 8779: 8731: 8728:Paleobiology 8727: 8720: 8693: 8689: 8679: 8671: 8666: 8629: 8625: 8603:, pp. 231–38 8600: 8595: 8583: 8558: 8554: 8548: 8537:. Retrieved 8533: 8523: 8512:. Retrieved 8508: 8499: 8458: 8454: 8400: 8396: 8390: 8382: 8345: 8341: 8331: 8296: 8292: 8282: 8257: 8253: 8203: 8199: 8186: 8167: 8163: 8153: 8126: 8122: 8116: 8108: 8096: 8084: 8075: 8066: 8002: 7998: 7988: 7976: 7968: 7963: 7958:, p. 9. 7901: 7898:Paleobiology 7897: 7880: 7875: 7866: 7862: 7856: 7848: 7843: 7835: 7830: 7819:. Retrieved 7812:the original 7791: 7785: 7772: 7764: 7759: 7750: 7746: 7740: 7731: 7727: 7721: 7709: 7629: 7625: 7619: 7611: 7606: 7598: 7593: 7581: 7569: 7557: 7548: 7544: 7534: 7525: 7521: 7515: 7503: 7495: 7491: 7485: 7477: 7472: 7464: 7460: 7454: 7445: 7441: 7435: 7427: 7422: 7414: 7409: 7404:, p. 7. 7378: 7373:, p. 6. 7366: 7354: 7349:, p. 5. 7296: 7292: 7282: 7250:(1): 12–13. 7247: 7243: 7233: 7182: 7178: 7171: 7138: 7134: 7128: 7093: 7089: 7079: 7038: 7034: 7028: 7017:. Retrieved 7011: 7001: 6961:(1): 24–30. 6958: 6954: 6944: 6935: 6931: 6925: 6892: 6880: 6868: 6856: 6844: 6802:(5): e2271. 6799: 6795: 6654: 6650: 6640: 6628: 6547: 6541: 6531: 6504: 6500: 6490: 6473: 6469: 6463: 6438: 6434: 6421: 6386: 6380: 6370: 6358: 6346: 6294: 6290: 6280: 6247: 6243: 6237: 6204: 6200: 6194: 6161: 6157: 6153: 6072:(2): e4497. 6069: 6065: 6017: 6013: 6008:Jeholopterus 6007: 5961: 5957: 5951: 5924: 5895: 5794: 5753: 5749: 5743: 5735: 5723: 5707: 5693:(5): 213–16. 5690: 5686: 5661: 5644: 5640: 5636: 5544: 5496: 5487: 5483: 5477: 5444: 5440: 5427: 5416:. Retrieved 5412: 5403: 5362: 5358: 5318: 5296:. Retrieved 5255: 5244: 5203: 5199: 5186: 5145: 5141: 5132: 5087: 5083: 5045:. Retrieved 5041: 5031: 5020:. Retrieved 5016: 5007: 4988: 4984: 4974: 4965: 4953: 4943: 4936: 4925: 4916: 4898: 4889: 4880: 4876: 4872: 4866: 4840: 4834: 4797: 4793: 4783: 4748: 4744: 4708: 4675: 4612: 4603: 4480: 4471: 4469:anurognathid 4458: 4451: 4447: 4441: 4439: 4432: 4426: 4421: 4415: 4409: 4395: 4389: 4379: 4377: 4364:Ludodactylus 4362: 4358: 4354: 4348: 4344: 4338: 4332: 4330: 4323: 4317: 4307: 4297: 4290: 4276: 4272: 4268: 4264: 4253:Pterodaustro 4252: 4248: 4244: 4234: 4224: 4220:Pterodaustro 4218: 4212: 4202: 4196: 4190: 4184: 4176: 4171: 4165: 4157: 4149: 4148:"), such as 4143: 4134: 4121: 4110: 4107: 4103:Pterodaustro 4102: 4094: 4090: 4087:ossification 4086: 4081: 4075: 4066: 4064: 4044: 4038: 4034:Pterodaustro 4032: 4013: 3989: 3979: 3973: 3969: 3963: 3945: 3939: 3927: 3921: 3915: 3905: 3895: 3889: 3883: 3872:nyctosaurids 3861: 3854: 3848: 3837:Azhdarchidae 3829: 3811: 3802:Pterodaustro 3800: 3790: 3784: 3772: 3762: 3744: 3741:frigatebirds 3718: 3713: 3707: 3696:Sericipterus 3695: 3689: 3683: 3679:Sericipterus 3677: 3671: 3665: 3659: 3654: 3648: 3642: 3638: 3632: 3626: 3619: 3605: 3603: 3592: 3588: 3568: 3554:boreopterids 3550: 3538: 3534:Pterodaustro 3532: 3526: 3512: 3509: 3492: 3488:cruropatagia 3484: 3476: 3446: 3438:road runners 3431: 3428:Kevin Padian 3417: 3410: 3404: 3375: 3368: 3355: 3351: 3345: 3328:subcutaneous 3324: 3312: 3305: 3299: 3293: 3273: 3266: 3256:Darren Naish 3243: 3235: 3233: 3230: 3206:Paleobiology 2998: 2978: 2952: 2951: 2915: 2895: 2859: 2858: 2810: 2789: 2788: 2758: 2719:Kryptodrakon 2717: 2716: 2686: 2663: 2662: 2642:Darwinoptera 2640: 2612: 2611: 2584: 2557: 2530: 2500: 2488: 2477: 2474:phylogenetic 2466: 2458:paraphyletic 2435: 2422: 2418: 2414: 2413:(originally 2410: 2401: 2395: 2384: 2369: 2365:Lazarus taxa 2358: 2343: 2336: 2333:nyctosaurids 2327:", possible 2322: 2316: 2306: 2290: 2280: 2262: 2257: 2252: 2240: 2200: 2194: 2186: 2176: 2172: 2157: 2144: 2125: 2115: 2108: 2098: 2092: 2081:protorosaurs 2073:crocodilians 2049: 2029: 1997:Isle of Skye 1990: 1988: 1960: 1947:Lagerstätten 1946: 1944: 1929: 1918: 1914: 1906:Kevin Padian 1895: 1889: 1883: 1875: 1859: 1848: 1832:Anurognathus 1830: 1826: 1816: 1810: 1804: 1802: 1793: 1787: 1778: 1774: 1759:Richard Owen 1752: 1746: 1740: 1709: 1700: 1694: 1690: 1680: 1668:Plesiosauria 1659: 1635: 1631:Lagerstätten 1629: 1623: 1616:specimen by 1611: 1583: 1576: 1574: 1568: 1562: 1556: 1533: 1529:maniraptoran 1522: 1510: 1506:Jeholopterus 1505: 1493: 1487: 1484:histological 1473: 1447: 1441: 1416: 1413:Soft tissues 1407: 1394: 1386: 1377: 1353: 1333: 1331: 1327: 1323:induced drag 1318:nyctosaurids 1314:azhdarchoids 1307: 1303: 1295: 1289: 1282: 1262: 1260:wing-plans. 1244:Jeholopterus 1242: 1240:anurognathid 1233: 1231: 1222: 1216: 1212: 1206: 1202: 1200: 1194: 1187: 1181: 1164: 1153: 1150: 1142: 1124: 1106: 1098: 1096: 1076: 1063:zygapophyses 1051: 1025: 1021: 1015: 1014:The neck of 986: 979: 959: 956: 949: 945: 935: 929: 926: 919: 905: 903: 896: 881:Pterodaustro 879: 868:Azhdarchidae 857: 849: 843: 837: 818: 790: 783: 765: 761: 725:Jeholopterus 721:Anurognathus 719: 717: 714: 707: 701: 695: 689: 683: 677: 671: 665: 659: 653: 651: 644: 638: 632: 626: 620: 618: 579: 566: 555: 551: 548:pterodactyls 547: 544:Ornithischia 533: 522: 516: 478: 457:insectivores 450: 421: 410: 409: 395: 394: 339: 328: 320: 308: 298: 290: 250: 228: 215: 162: 156: 150: 146:Anurognathus 144: 138: 132: 49:228–66 29: 17671:iNaturalist 17639:Pterosauria 17633:Wikispecies 17609:Pterosauria 17462:Insect wing 17412:Webbed foot 17353:unguligrade 17348:plantigrade 17343:digitigrade 17000:Thapunngaka 16960:Amblydectes 16879:Anhangueria 16843:Aussiedraco 16577:Boreopterus 16352:Lonchodraco 16285:Lonchodraco 16228:Aussiedraco 16174:Nyctosaurus 16031:Bogolubovia 15699:Bogolubovia 15570:Lacusovagus 15562:Eoazhdarcho 15522:Eoazhdarcho 15481:Leptostomia 15473:Keresdrakon 15348:Caiuajarina 15315:Bakonydraco 15283:Lacusovagus 15275:Keresdrakon 15246:Tapejarinae 15207:Huaxiadraco 15191:Bakonydraco 15150:Tapejaridae 15079:Leptostomia 15071:Lacusovagus 15064:Kariridraco 15010:Keresdrakon 14951:Leptostomia 14943:Keresdrakon 14604:Ctenochasma 14566:Lusognathus 14489:Otogopterus 14144:Lophocratia 14077:Lophocratia 14044:Pangupterus 13980:Sinomacrops 13321:Dorygnathus 13190:Novialoidea 13145:Rhamphinion 13122:Dimorphodon 12797:Caviramidae 12741:Pterosauria 12728:Pterosauria 12691:Pterosauria 12630:Archosauria 12591:Pterosauria 12369:Gnathovorax 12348:Caseosaurus 12150:Lutungutali 12143:Kwanasaurus 12115:Amanasaurus 12058:Asilisaurus 12051:Agnosphitys 12012:Nyasasaurus 11950:Pterosauria 11939:Pterosauria 11906:Kongonaphon 11899:Ixalerpeton 11849:Ornithodira 11826:Teleocrater 11812:Dongusuchus 11781:Mambachiton 11627:Archosauria 10470:Ameghiniana 10063:Witton 2013 10051:Witton 2013 9941:Witton 2013 9879:Witton 2013 9867:Witton 2013 9855:Witton 2013 9796:Witton 2013 9456:(2): 18–24. 8588:Witton 2013 8101:Witton 2013 8089:Witton 2013 8059:Witton 2013 8045:Witton 2013 7956:Witton 2013 7402:Witton 2013 7383:Witton 2013 7371:Witton 2013 7347:Witton 2013 7208:10468/11874 6918:Witton 2013 6897:Witton 2013 6885:Witton 2013 6873:Witton 2013 6861:Witton 2013 6849:Witton 2013 6771:Witton 2013 6602:Witton 2013 6363:Witton 2013 6351:Witton 2013 6137:Witton 2013 6122:Witton 2013 5944:Witton 2013 5929:Witton 2013 5917:Witton 2013 5900:Witton 2013 5852:Witton 2013 5835:Witton 2013 5818:Witton 2013 5799:Witton 2013 5770:Witton 2013 5620:Witton 2013 5603:Witton 2013 5571:Witton 2013 5518:Witton 2013 5501:Witton 2013 4922:"Pterosaur" 4699:wing lizard 4316:Scene from 4265:Ctenochasma 4263:pterosaurs 4245:Ctenochasma 4214:Ctenochasma 3986:ichthyosaur 3975:Squalicorax 3876:anhanuerids 3818:Tapejaridae 3773:Lonchodraco 3764:Lonchodraco 3673:Dorygnathus 3607:Dimorphodon 3579:tropicbirds 3566:nyctosaurid 3562:azhdarchids 3558:frigatebird 3495:azhdarchids 3461:azhdarchids 3449:plantigrade 3433:Dimorphodon 3307:On the Wing 3280:Mark Witton 3248:azhdarchids 3174:Novialoidea 2737:Lophocratia 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A fuzzy 1458:homologous 1397:metatarsal 1390:metatarsus 1365:pubic bone 1016:Anhanguera 907:Pteranodon 860:keratinous 834:tapejarids 811:archosaurs 803:premaxilla 794:heterodont 667:Pteranodon 594:breastbone 540:Saurischia 529:endothermy 489:homologous 434:Cretaceous 411:Pterosaurs 264:Subgroups 17481:Evolution 17440:Bird wing 17385:Arthropod 17378:quadruped 17028:Aerodraco 16470:Luchibang 16418:Haopterus 16140:Simurghia 16133:Epapatelo 16101:Cretornis 16039:Dawndraco 15765:Azhdarcho 15758:Albadraco 15750:Aerotitan 15670:Alanqidae 15448:Alanqidae 15414:Cretornis 15364:Torukjara 15357:Caiuajara 15094:Tupuxuara 15032:Aerotitan 14936:Inabtanin 14524:Feilongus 13923:Luopterus 13640:Allkaruen 13267:Klobiodon 13107:Allkaruen 12942:Caviramus 12819:Caviramus 12607:Kingdom: 12455:includes 12451:Theropoda 12440:Theropoda 12065:Gamatavus 11604:Kingdom: 11384:IndieWire 11353:Thrillist 11293:: 56–63. 11154:0962-8452 10639:206529739 10588:0272-4634 10523:1476-4687 10268:: 59–67. 10196:: e2908. 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