title: Pterosaur

text: 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 "Pteranodon". 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 parietal bones 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 Tapejaridae. "Nyctosaurus" 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.


title: Pterosaur

text: 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 "Pterorhynchus" and "Pterodactylus", the true extent of these crests has only been uncovered using ultraviolet photography. While fossil crests used to be restricted to the more advanced Pterodactyloidea, "Pterorhynchus" and "Austriadactylus" show that even some early pterosaurs possessed them.


title: Pterosaur

text: 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 "ossa dentalia", were at the tip tightly fused into a central symphysis. This made the lower jaws function as a single connected whole, the mandible. 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.


title: Pterosaur

text: The vertebral column of pterosaurs numbered between thirty-four and seventy vertebrae. The vertebrae in front of the tail were "procoelous": the cotyle (front of the vertebral body) was concave and into it fitted a convex extension at the rear of the preceding vertebra, the condyle. Advanced pterosaurs are unique in possessing special processes projecting adjacent to their condyle and cotyle, the exapophyses, and the cotyle also may possess a small prong on its midline called a hypapophysis.


title: Pterosaur

text: The neck of pterosaurs was 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 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.


title: Pterosaur

text: 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 sacrum. Such species also often show a fusion of the front dorsal vertebrae into a rigid whole which is called the notarium 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 neural spines 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.


title: Pterosaur

text: The tails of pterosaurs were always rather slender. This means that the caudofemoralis retractor muscle which in most basal Archosauria 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, the zygapophyses, and chevrons. 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.


title: Pterosaur

text: The shoulder girdle was a strong structure that transferred the forces of flapping flight to the thorax. It was probably covered by thick muscle layers. The upper bone, the shoulder blade, was a straight bar. It was connected to a lower bone, the coracoid 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 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.


title: Pterosaur

text: The breastbone, formed by fused paired "sterna", 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 gastralia was present, covering the entire belly. To the front, a long point, the "cristospina", jutted obliquely upwards. The rear edge of the breastbone was the deepest point of the thorax. Clavicles or interclavicles were completely absent.


title: Pterosaur

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


title: Pterosaur

text: 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 fibers called "actinofibrils". 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 camber.


title: Pterosaur

text: 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 air sacs into the wing membrane.


title: Pterosaur

text: The pterosaur wing membrane is divided into three basic units. The first, called the "propatagium" ("fore membrane"), was the forward-most part of the wing and attached between the wrist and shoulder, creating the "leading edge" during flight. The "brachiopatagium" ("arm membrane") was the primary component of the wing, stretching from the highly elongated fourth finger of the hand to the hindlimbs. Finally, at least some pterosaur groups had a membrane that stretched between the legs, possibly connecting to or incorporating the tail, called the uropatagium; the extent of this membrane is not certain, as studies on "Sordes" seem to suggest that it simply connected the legs but did not involve the tail (rendering it a cruropatagium). A common interpretation is that non-pterodactyloid pterosaurs had a broader uro/cruropatagium stretched between their long fifth toes, with pterodactyloids, lacking such toes, only having membranes running along the legs.


title: Pterosaur

text: 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 "Sordes", the anurognathid "Jeholopterus", and a pterodactyloid from the Santana Formation seem to demonstrate that the wing membrane did attach to the hindlimbs, at least in some species. However, modern bats and flying squirrels 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 wing-plans.


title: Pterosaur

text: The bony elements of the arm formed a mechanism to support and extend the wing. Near the body, the humerus 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 ulna and radius, are much longer than the humerus. They were probably incapable of pronation.


title: Pterosaur

text: 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 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 flap. 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 "Changchengopterus pani" and "Darwinopterus linglongtaensis" 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.


title: Pterosaur

text: 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" 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.


title: Pterosaur

text: In derived pterodactyloids like pteranodontians and azhdarchoids, 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 nyctosaurids 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 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 induced drag. The wingfinger is also bent somewhat downwards.


title: Pterosaur

text: 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 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.


title: Pterosaur

text: A laser-simulated fluorescence scan on "Pterodactylus" 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.


title: Pterosaur

text: The pelvis of pterosaurs was of moderate size compared to the body as a whole. Often the three pelvic bones were fused. The ilium 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, pubic bone fused with the broad ischium into an ischiopubic blade. Sometimes, the blades of both sides were also fused, closing the pelvis from below and forming the pelvic canal. The hip joint was not perforated and allowed considerable mobility to the leg. It was directed obliquely upwards, preventing a perfectly vertical position of the leg.


title: Pterosaur

text: 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 chest cavity.


title: Pterosaur

text: 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 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, metatarsus 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.


title: Pterosaur

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


title: Pterosaur

text: 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.


title: Pterosaur

text: The rare conditions that allowed for the fossilisation of pterosaur remains, sometimes also preserved soft tissues. Modern synchrotron or ultraviolet light photography has revealed many traces not visible to the naked eye. These are often imprecisely called "impressions" but mostly consist of petrifications, 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 metatarsals. They covered pads cushioning the impact of walking. Scales are unknown from other parts of the body.


title: Pterosaur

text: Most or all pterosaurs had hair-like filaments known as pycnofibers on the head and torso. The term "pycnofiber", meaning "dense filament", was coined by palaeontologist Alexander Kellner and colleagues in 2009. Pycnofibers were unique structures similar to, but not homologous (sharing a common origin) with, mammalian hair, an example of convergent evolution. A fuzzy integument was first reported from a specimen of "Scaphognathus crassirostris" in 1831 by Georg August Goldfuss, but had been widely doubted. Since the 1990s, pterosaur finds and histological and ultraviolet examination of pterosaur specimens have provided incontrovertible proof: pterosaurs had pycnofiber coats. "Sordes pilosus" (which translates as "hairy demon") and "Jeholopterus ninchengensis" show pycnofibers on the head and body.


title: Pterosaur

text: The presence of pycnofibers strongly indicates that pterosaurs were endothermic (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.


title: Pterosaur

text: 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 proto-feathers. Others were unconvinced, considering the difference with the "quills" found on many of the bird-like maniraptoran specimens too fundamental.


title: Pterosaur

text: A 2018 study of the remains of two small Jurassic-age pterosaurs from Inner Mongolia, China, 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 anurognathids 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 patagium, and the presence of both aktinofibrils and filaments on "Jeholopterus ningchengensis" and "Sordes pilosus". 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 proto-feathers. But Liliana D’Alba points out that the description of the preserved integumentary structures on the two anurogmathid specimens is still based upon gross morphology. She also points out that "Pterorhynchus" 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.


title: Pterosaur

text: In 2022, a new fossil of "Tupandactylus cf. imperator" 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 relatives of "Tupandactylus" 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 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.


title: Pterosaur

text: Pterosaur fossils 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 "Lagerstätten". The pieces from one such "Lagerstätte", the Late Jurassic Solnhofen Limestone in Bavaria, became much sought after by rich collectors. In 1784, the Italian naturalist Cosimo Alessandro Collini was the first scientist in history to describe a pterosaur fossil. At that time the concepts of evolution and extinction were only imperfectly developed. The bizarre build of the pterosaur was therefore shocking, as it could not clearly be assigned to any existing animal group. The discovery of pterosaurs would thus play an 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 the German zoologist Johann Georg Wagler suggested that "Pterodactylus" used its wings as flippers and was affiliated with Ichthyosauria and Plesiosauria.


title: Pterosaur

text: In 1800, Johann Hermann first suggested that it represented a flying creature in a letter to Georges Cuvier. Cuvier agreed in 1801, understanding it was an extinct flying reptile. In 1809, he coined the name "Ptéro-Dactyle", "wing-finger". This was in 1815 Latinised to "Pterodactylus". 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 "Pterodactylus" or members of the Pterodactyloidea.


title: Pterosaur

text: In 1812 and 1817, Samuel Thomas von Soemmerring 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 very influential during the 19th century. In 1843, Edward Newman thought pterosaurs were flying marsupials. As the bat model correctly depicted pterosaurs as furred and warm-blooded, it better approached the true physiology of pterosaurs than Cuvier's "reptile model". In 1834, Johann Jakob Kaup coined the term Pterosauria.


title: Pterosaur

text: In 1828, Mary Anning found in England the first pterosaur genus outside Germany, named as "Dimorphodon" by Richard Owen, also the first non-pterodactyloid pterosaur known. Later in the century, the Early Cretaceous Cambridge Greensand 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 Harry Govier Seeley, at the time the main English expert on the subject, who also wrote the first pterosaur book, "Ornithosauria", and in 1901 the first popular book, "Dragons of the Air". Seeley thought that pterosaurs were warm-blooded and dynamic creatures, closely related to birds. Earlier, the evolutionist St. George Jackson Mivart had suggested pterosaurs were the direct ancestors of birds. Owen opposed the views of both men, seeing pterosaurs as cold-blooded "true" reptiles.


title: Pterosaur

text: In the US, Othniel Charles Marsh in 1870 discovered "Pteranodon" in the Niobrara Chalk, 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.


title: Pterosaur

text: 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 "Rhamphorhynchus", "Scaphognathus" and "Dorygnathus". 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 Karl Alfred Zittel published studies about the wing membranes of specimens of "Rhamphorhynchus". German studies continued well into the 1930s, describing new species such as "Anurognathus". In 1927, Ferdinand Broili discovered hair follicles in pterosaur skin, and paleoneurologist Tilly Edinger determined that the brains of pterosaurs more resembled those of birds than modern cold-blooded reptiles.


title: Pterosaur

text: 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 themselves from cliffs to achieve a take-off. In 1914, for the first time pterosaur aerodynamics were quantitatively analysed, by Ernest Hanbury Hankin and David Meredith Seares Watson, but they interpreted "Pteranodon" as a pure glider. Little research was done on the group during the 1940s and 1950s.


title: Pterosaur

text: The situation for dinosaurs was comparable. From the 1960s onwards, a dinosaur renaissance took place, a quick increase in the number of studies and critical ideas, influenced by the discovery of additional fossils of "Deinonychus", whose spectacular traits refuted what had become entrenched orthodoxy. In 1970, likewise the description of the furry pterosaur "Sordes" began what Robert Bakker named a renaissance of pterosaurs. Kevin Padian 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 Peter Wellnhofer, who in 1970s laid the foundations of modern pterosaur science. In 1978, he published the first pterosaur textbook, the "Handbuch der Paläoherptologie, Teil 19: Pterosauria", and in 1991 the second ever popular science pterosaur book, the "Encyclopedia of Pterosaurs".


title: Pterosaur

text: This development accelerated through the exploitation of two new "Lagerstätten". During the 1970s, the Early Cretaceous Santana Formation 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 their scientists to describe many new species and revealing a whole new fauna. Soon, Brazilian researchers, among them Alexander Kellner, intercepted the trade and named even more species.


title: Pterosaur

text: Even more productive was the Early Cretaceous Chinese Jehol Biota of Liaoning that since the 1990s has brought forth hundreds of exquisitely preserved two-dimensional fossils, often showing soft tissue remains. Chinese researchers such as Lü Junchang 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 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 allowed to determine their complex relationships through the quantitative method of cladistics. New and old fossils yielded much more information when subjected to modern ultraviolet light or roentgen photography, or CAT-scans. 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.


title: Pterosaur

text: In 2017 a fossil from a 170-million-year-old pterosaur, later named as the species "Dearc sgiathanach" in 2022, was discovered on the Isle of Skye in Scotland. The National Museum of Scotland claims that it the largest of its kind ever discovered from the Jurassic period, and it has been described as the world’s best-preserved skeleton of a pterosaur.


title: Pterosaur

text: Because pterosaur anatomy has been so heavily modified for flight, and immediate transitional fossil 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 called the Archosauromorpha. In the 1980s, early cladistic analyses found that they were Avemetatarsalians (archosaurs closer to dinosaurs than to crocodilians). 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. David Unwin proposed an ancestry among the basal Archosauromorpha, specifically long-necked forms ("protorosaurs") such as tanystropheids. A placement among basal archosauriforms like "Euparkeria" 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 "Sharovipteryx", a "protorosaur" with skin membranes on its hindlimbs likely used for gliding. A 1999 study by Michael Benton found that pterosaurs were avemetatarsalians closely related to "Scleromochlus," and named the group Ornithodira to encompass pterosaurs and dinosaurs"."


title: Pterosaur

text: Two researchers, S. Christopher Bennett in 1996, and paleoartist 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 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 convergent evolution between pterosaurs and dinosaurs. 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 "Scleromochlus" or a branch between the latter and "Lagosuchus". An 2011 archosaur-focused phylogenetic analysis by Sterling Nesbitt benefited from far more data and found strong support for pterosaurs being avemetatarsalians, though "Scleromochlus" was not included due to its poor preservation. A 2016 archosauromorph-focused study by Martin Ezcurra 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 "Scleromochlus" which argued that both "Scleromochlus" and pterosaurs were non-archosaur archosauromorphs, albeit not particularly closely related to each other. By contrast, a later 2020 study proposed that lagerpetid archosaurs were the sister clade to pterosauria. This was based on newly described fossil skulls and forelimbs showing various anatomical similarities with pterosaurs and reconstructions of lagerpetid brains and sensory systems based on CT scans also showing neuroanatomical similarities with pterosaurs. The results of the latter study were subsequently supported by an independent analysis of early pterosauromorph interrelationships.


title: Pterosaur

text: 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 selection pressure for incipient flight. Rupert Wild 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 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 "Scleromochlus". Like this archosaur, basal pterosaur lineages have plantigrade hindlimbs that show adaptations for saltation.


title: Pterosaur

text: At least one study found that the early Triassic ichnofossil "Prorotodactylus" is anatomically similar to that of early pterosaurs.


title: Pterosaur

text: It was once thought that competition with early bird species might have resulted in the extinction 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 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 KT event. At the end of the Cretaceous period, the Cretaceous–Paleogene extinction event, which wiped out all non-avian dinosaurs and most avian dinosaurs as well, and many other animals, seems also to have taken the pterosaurs.


title: Pterosaur

text: In the early 2010s, several new pterosaur taxa were discovered dating to the Campanian/Maastrichtian, such as the ornithocheirids "Piksi" and ""Ornithocheirus"", possible pteranodontids and nyctosaurids, several tapejarids and the indeterminate non-azhdarchid "Navajodactylus". 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.


title: Pterosaur

text: Small-sized pterosaur species apparently were present in the Csehbánya Formation, indicating a higher diversity of Late Cretaceous pterosaurs than previously accounted for. The recent findings of a small cat-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 dinosaurs, and that their diversity might actually have been much larger than previously thought.


title: Pterosaur

text: At least some non-pterodactyloid pterosaurs survived into the Late Cretaceous, postulating a Lazarus taxa situation for late Cretaceous pterosaur faunas.


title: Pterosaur

text: 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 outcompeted by birds, pterosaurs essentially specialized a trend already occurring in previous eras of the Mesozoic.


title: Pterosaur

text: In phylogenetic taxonomy, the clade 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, "Preondactylus" and "Quetzalcoatlus" 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 presence of an enlarged fourth finger that supports a wing membrane. This "apomorophy-based" definition was adopted by the PhyloCode in 2020 as "[T]he clade characterized by the apomorphy fourth manual digit hypertrophied to support a wing membrane, as inherited by "Pterodactylus" (originally "Ornithocephalus") "antiquus" (Sömmerring 1812)". A broader clade, Pterosauromorpha, has been defined as all ornithodirans more closely related to pterosaurs than to dinosaurs.


title: Pterosaur

text: The internal classification of pterosaurs has historically been difficult, because there were many gaps in the fossil record. 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 suborders: the Rhamphorhynchoidea, a "primitive" group of long-tailed pterosaurs, and the Pterodactyloidea, "advanced" pterosaurs with short tails. However, this traditional division has been largely abandoned. Rhamphorhynchoidea is a paraphyletic (unnatural) group, since the pterodactyloids evolved directly from them and not from a common ancestor, so, with the increasing use of cladistics, it has fallen out of favor among most scientists.


title: Pterosaur

text: 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 cladogram (family tree) below follows a phylogenetic analysis presented by Longrich, Martill and Andres in 2018, with clade names after Andres "et al." (2014).


title: Pterosaur

text: 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 "Posture, Locomotion, and Paleoecology of Pterosaurs" it is theorized that they were able to fly due to the oxygen-rich, dense atmosphere of the Late Cretaceous period. However, both Sato and the authors of "Posture, Locomotion, and Paleoecology of Pterosaurs" 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 azhdarchids and tapejarids. Furthermore, Darren Naish concluded that atmospheric differences between the present and the Mesozoic were not needed for the giant size of pterosaurs.


title: Pterosaur

text: Another issue that has been difficult to understand is how they took off. 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. Mark Witton of the University of Portsmouth and Mike Habib of Johns Hopkins University 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 and travel thousands of kilometres.


title: Pterosaur

text: In 1985, the Smithsonian Institution commissioned aeronautical engineer Paul MacCready to build a half-scale working model of "Quetzalcoatlus northropi". 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 "On the Wing".


title: Pterosaur

text: 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 subcutaneous 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 hepatic piston, seeing as their shoulder-pectoral girdles were too inflexible to move the sternum as in birds, and they possessed strong gastralia. Thus, their respiratory system had characteristics comparable to both modern archosaur clades.


title: Pterosaur

text: An X-ray study of pterosaur brain cavities revealed that the animals ("Rhamphorhynchus muensteri" and "Anhanguera santanae") had massive flocculi. The flocculus 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% of total brain mass.


title: Pterosaur

text: 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 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 sauropsids to achieve high intelligence levels with small brains. Studies on the endocast of "Allkaruen" show that brain evolution in pterodactyloids was a modular process.


title: Pterosaur

text: Pterosaurs' hip sockets are oriented facing slightly upwards, and the head of the femur (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.


title: Pterosaur

text: There was considerable debate whether pterosaurs ambulated as quadrupeds or as bipeds. In the 1980s, paleontologist Kevin Padian suggested that smaller pterosaurs with longer hindlimbs, such as "Dimorphodon", might have walked or even run bipedally, in addition to flying, like road runners. However, a large number of pterosaur trackways 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.


title: Pterosaur

text: Fossil footprints show that pterosaurs stood with the entire foot in contact with the ground (plantigrade), in a manner similar to many mammals like humans and bears. Footprints from azhdarchids 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 in pterosaurs.


title: Pterosaur

text: 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 cruropatagia, but they too appear to have been generally efficient on the ground.


title: Pterosaur

text: The forelimb bones of azhdarchids and ornithocheirids 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 ungulate 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.


title: Pterosaur

text: 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 length of the lower leg. This suggests that azhdarchids were better adapted to walking on dry, relatively solid ground. "Pteranodon" had slightly larger feet (47% the length of the tibia), while filter-feeding pterosaurs like the ctenochasmatoids had very large feet (69% of tibial length in "Pterodactylus", 84% in "Pterodaustro"), 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 "Scleromochlus".


title: Pterosaur

text: 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 interpreted to have been suggestive of a water-based version of the typical quadrupedal launch, and several like boreopterids must have foraged while swimming, as they seem incapable of frigatebird-like aerial hawking. These adaptations are also seen in terrestrial pterosaurs like azhdarchids, which presumably still needed to launch from water in case they found themselves in it. The nyctosaurid "Alcione" may display adaptations for wing-propelled diving like modern gannets and tropicbirds.


title: Pterosaur

text: 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.


title: Pterosaur

text: Early-on it was recognised that the small Anurognathidae 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 nightjars or extant insectivorous bats, being capable of high manoeuvrability at relatively low speeds.


title: Pterosaur

text: Interpretations of the habits of basal groups have changed profoundly. "Dimorphodon", envisioned as a puffin analogue in the past, is indicated by its jaw structure, gait, and poor flight capabilities, as a terrestrial/semiarboreal predator of small mammals, squamates, and large insects. Its robust dentition caused "Campylognathoides" 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 "Carniadactylus" and the larger "Eudimorphodon" were highly aerial animals and fast, agile flyers with long robust wings. "Eudimorphodon" has been found with fish remains in its stomach, but its dentition suggests an opportunistic diet. Slender-winged "Austriadactylus" and "Caviramus" were likely terrestrial/semiarboreal generalists. "Caviramus" likely had a strong bite force, indicating an adaptation towards hard food items that might have been chewed in view of the tooth wear.


title: Pterosaur

text: Some Rhamphorhynchidae, such as "Rhamphorhynchus" itself or "Dorygnathus", were fish-eaters with long, slender wings, needle-like dentition and long, thin jaws. "Sericipterus", "Scaphognathus" and "Harpactognathus" had more robust jaws and teeth (which were ziphodont, dagger-shaped, in " Sericipterus"), and shorter, broader wings. These were either terrestrial/aerial predators of vertebrates or corvid-like generalists. Wukongopteridae like "Darwinopterus" were first considered aerial predators. Lacking a robust jaw structure or powerful flying muscles, they are now seen as arboreal or semiterrestrial insectivores. "Darwinopterus robustidens", in particular, seems to have been a beetle specialist.


title: Pterosaur

text: Among pterodactyloids, a greater variation in diet is present. Pteranodontia contained many piscivorous taxa, such as the Ornithocheirae, Boreopteridae, Pteranodontidae and Nyctosauridae. Niche partitioning caused ornithocheirans and the later nyctosaurids to be aerial dip-feeders like today's frigatebirds (with the exception of the plunge-diving adapted "Alcione elainus"), while boreopterids were freshwater diving animals similar to cormorants, and pteranodonts pelagic plunge-divers akin to boobies and gannets. An analysis of "Lonchodraco" found clusters of foramina at the tip of its beak; birds with similarly numerous foramina have sensitive beaks used to feel for food, so "Lonchodraco" may have used its beak to feel for fish or invertebrates in shallow water. The istiodactylids were likely primarily scavengers. Archaeopterodactyloidea obtained food in coastal or freshwater habitats. "Germanodactylus" and "Pterodactylus" were piscivores, while the Ctenochasmatidae were suspension feeders, using their numerous fine teeth to filter small organisms from shallow water. "Pterodaustro" was adapted for flamingo-like filter-feeding.


title: Pterosaur

text: In contrast, Azhdarchoidea mostly were terrestrial pterosaurs. Tapejaridae were arboreal omnivores, supplementing seeds and fruits with small insects and vertebrates. Dsungaripteridae were specialist molluscivores, using their powerful jaws to crush the shells of molluscs and crustaceans. Thalassodromidae were likely terrestrial carnivores. "Thalassodromeus" 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. Azhdarchidae are now understood to be terrestrial predators akin to ground hornbills or some storks, eating any prey item they could swallow whole. "Hatzegopteryx" was a robustly built predator of relatively large prey, including medium-sized dinosaurs. "Alanqa" may have been a specialist molluscivore.


title: Pterosaur

text: A 2021 study reconstructed the adductor musculature of skulls from pterodactyloids, estimating the bite force and potential dietary habits of nine selected species. The study corroborated the view of pteranodontids, nyctosaurids and anhanuerids as piscivores based on them being relatively weak but fast biters, and suggest that "Tropeognathus mesembrinus" was specialised in consuming relatively large prey compared to "Anhanguera". "Dsungaripterus" was corroborated as a durophage, with "Thalassodromeus" proposed to share this feeding habit based on high estimated bite force quotients (BFQ) and absolute bite force values. "Tapejara wellnhoferi" was corroborated as a specialised consumer of hard plant material with a relatively high BFQ and high mechanical advantage, and "Caupedactylus ybaka" and "Tupuxuara leonardii" were proposed to be ground-feeding generalists with intermediate bite force values and less specialised jaws.


title: Pterosaur

text: Pterosaurs are known to have been eaten by theropods. In the 1 July 2004 edition of "Nature", paleontologist Eric Buffetaut discusses an Early Cretaceous fossil of three cervical vertebrae of a pterosaur with the broken tooth of a spinosaur, most likely "Irritator", embedded in it. The vertebrae are known not to have been eaten and exposed to digestion, as the joints are still articulated.


title: Pterosaur

text: 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 egg was found in the quarries of Liaoning, the same place that yielded feathered dinosaurs. The egg was squashed flat with no signs of cracking, so evidently the eggs had leathery shells, as in modern lizards. This was supported by the description of an additional pterosaur egg belonging to the genus "Darwinopterus", 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 "Hamipterus tianshanensis" 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 likely pterosaurs buried their eggs, like modern crocodiles and turtles. 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 birds.


title: Pterosaur

text: A "Darwinopterus" specimen showcases that at least some pterosaurs had a pair of functional ovaries, as opposed to the single functional ovary in birds, dismissing the reduction of functional ovaries as a requirement for powered flight.


title: Pterosaur

text: Wing membranes preserved in pterosaur embryos are well developed, suggesting that pterosaurs were ready to fly soon after birth. However, tomography scans of fossilised "Hamipterus" 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, including pterodactylids, rhamphorhinchids, ctenochasmatids and azhdarchids. All preserve bones that show a relatively high degree of hardening ("ossification") for their age, and wing proportions similar to adults. In fact, many pterosaur flaplings have been considered adults and placed in separate species in the past. Additionally, flaplings are normally found in the same sediments as adults and juveniles of the same species, such as the "Pterodactylus" and "Rhamphorhynchus" flaplings found in the Solnhofen limestone of Germany, and "Pterodaustro" flaplings from Argentina. All are found in deep aquatic environment far from shore.


title: Pterosaur

text: 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 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 "Hamipterus" nests were shown preserving many male and female pterosaurs together with their eggs in a manner to a similar to that of modern seabird colonies. Due to how underdeveloped the chests of the hatchlings were for flying, it was suggested that "Hamipterus" may have practiced some form of parental care. However, this study has since been criticised. Most evidence currently leans towards pterosaur hatchlings being superprecocial, similar to that of megapode 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.


title: Pterosaur

text: Growth rates of pterosaurs once they hatched varied across different groups. In more primitive, long-tailed pterosaurs ("rhamphorhynchoids"), such as "Rhamphorhynchus", the average growth rate during the first year of life was 130% to 173%, slightly faster than the growth rate of alligators. Growth in these species slowed after sexual maturity, and it would have taken more than three years for "Rhamphorhynchus" to attain maximum size. In contrast, the more advanced, large pterodactyloid pterosaurs, such as "Pteranodon", grew to adult size within the first year of life. Additionally, pterodactyloids had "determinate growth", meaning that the animals reached a fixed maximum adult size and stopped growing.


title: Pterosaur

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


title: Pterosaur

text: Comparisons between the scleral rings of pterosaurs and modern birds and reptiles have been used to infer daily activity patterns of pterosaurs. The pterosaur genera "Pterodactylus", "Scaphognathus", and "Tupuxuara" have been inferred to be diurnal, "Ctenochasma", "Pterodaustro", and "Rhamphorhynchus" have been inferred to be nocturnal, and "Tapejara" has been inferred to be cathemeral, being active throughout the day for short intervals. As a result, the possibly fish-eating "Ctenochasma" and "Rhamphorhynchus" may have had similar activity patterns to modern nocturnal seabirds, and the filter-feeding "Pterodaustro" may have had similar activity patterns to modern anseriform birds that feed at night. The differences between activity patterns of the Solnhofen pterosaurs "Ctenochasma", "Rhamphorhynchus", "Scaphognathus", and "Pterodactylus" may also indicate niche partitioning between these genera.


title: Pterosaur

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


title: Pterosaur

text: The vague generic term "pterodactyl" is often used for these creatures. The animals depicted in fiction and pop culture frequently represent either the "Pteranodon" or (non-pterodactyloid) "Rhamphorhynchus", or a fictionalized hybrid of the two. Many children's toys and cartoons feature "pterodactyls" with "Pteranodon"-like crests and long, "Rhamphorhynchus"-like tails and teeth, a combination that never existed in nature. However, at least one pterosaur "did" have both the "Pteranodon"-like crest and teeth: "Ludodactylus", whose name means "toy finger" for its resemblance to old, inaccurate children's toys. Pterosaurs have sometimes been incorrectly identified as (the ancestors of) birds, though birds are theropod dinosaurs and not descendants of pterosaurs.


title: Pterosaur

text: Pterosaurs were used in fiction in Sir Arthur Conan Doyle's 1912 novel "The Lost World" and its 1925 film adaptation. They appeared in a number of films and television programs since, including the 1933 film "King Kong", and 1966's "One Million Years B.C." In the latter, animator Ray Harryhausen 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. Rodan, a fictional giant monster (or "kaiju") which first appeared in the 1956 film "Rodan", is portrayed as an enormous irradiated species of "Pteranodon". Rodan has appeared in multiple Japanese "Godzilla" films released during the 1960s, 1970s, 1990s, and 2000s, and also appeared in the 2019 American-produced film "".


title: Pterosaur

text: After the 1960s, pterosaurs remained mostly absent from notable American film appearances until 2001's "Jurassic Park III". 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 "Pteranodon" 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 "The Land Before Time" (1988), is a notable example from an animated film.


title: Pterosaur

text: In most media appearances, pterosaurs are depicted as piscivores, not reflecting their full dietary variation. They are also often shown as aerial predators similar to birds of prey, grasping human victims with talons on their feet. However, only the small anurognathid "Vesperopterylus" and small wukongopterid "Kunpengopterus" 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.


title: Ranma ½

text: "Ranma ½" has a comedic formula and a sex-changing main character, who often willfully transforms into a girl to advance his goals. The series also contains many other characters, whose intricate relationships with each other, unusual characteristics, and eccentric personalities drive most of the stories. Although the characters and their relationships are complicated, they rarely change once they are firmly introduced and settled into the series.


title: Ranma ½

text: The manga has been adapted into two anime series created by Studio Deen: "Ranma ½" and , which together were broadcast on Fuji Television from 1989 to 1992. In addition, they developed 12 original video animations and three films. In 2011, a live-action television special was produced and aired on Nippon Television. The manga and anime series were licensed by Viz Media for English-language releases in North America. Madman Entertainment released the manga, part of the anime series and the first two films in Australasia, while MVM Films released the first two films in the United Kingdom.


title: Ranma ½

text: The "Ranma ½" manga has over 55 million copies in circulation, making it one of the best-selling manga series. Both the manga and anime are cited as among the first in their respective media to have become popular in the United States.


title: Ranma ½

text: On a training journey in the Bayankala Mountain Range in the Qinghai Province of China, Ranma Saotome and his father Genma fall into the cursed springs at . The cursed spring causes any afflicted to assume the physical form of whatever drowned there hundreds or thousands of years ago whenever they come into contact with cold water, which reverts on contact with hot water but resumes with exposure to cold water. Genma fell into the spring of a drowned panda while Ranma fell into the spring of a drowned girl.


title: Ranma ½

text: Soun Tendo is a fellow practitioner of or "Anything-Goes School" of martial arts and owner of a dojo. Genma and Soun agreed years ago that their children would marry and carry on the Tendo Dojo. Soun has three teenaged daughters: the polite and easygoing Kasumi, the greedy and indifferent Nabiki and the short-tempered, martial arts practicing Akane. Akane, who is Ranma's age, is appointed for bridal duty by her sisters with the reasoning that they are the older sisters and can dump the duty on her, and that they all dislike the arranged engagement and think Akane's dislike of men is the right way to express it to the fathers. At the appointed time they are surprised when a panda comes in and puts a girl in front of their father. The Tendo girls all laugh. It takes several more pages for the situation to be explained to Soun Tendo and his daughters. Both Ranma and Akane refuse the engagement initially, having not been consulted on the decision, but the fathers are insistent and they are generally treated as betrothed and end up helping or saving each other on some occasions. They are frequently found in each other's company and are constantly arguing in their trademark awkward love-hate manner that is a franchise focus.


title: Ranma ½

text: Ranma goes to school with Akane at , where he meets his recurring opponent Tatewaki Kuno, the conceited kendo team captain who aggressively pursues Akane, but also falls in love with Ranma's female form without ever discovering his curse (despite most other characters eventually knowing it). Nerima serves as a backdrop for more martial arts mayhem with the introduction of Ranma's regular rivals, such as the eternally lost Ryoga Hibiki who traveled halfway across Japan getting from the front of his house to the back, where Ranma spent three days waiting for him. Ryoga, seeking revenge on Ranma, followed him to Jusenkyo where he ultimately fell into the Spring of the Drowned Piglet. Now when splashed with cold water he takes the form of a little black pig. Not knowing this, Akane takes the piglet as a pet and names it P-chan, but Ranma knows and hates him for keeping this secret and taking advantage of the situation. Another rival is the nearsighted Mousse, who also fell into a cursed spring and becomes a duck when he gets wet, and finally, there is Genma and Soun's impish grand master, Happosai, who spends his time stealing the underwear of schoolgirls.


title: Ranma ½

text: Ranma's prospective paramours include the martial arts rhythmic gymnastics champion (and Tatewaki's sister) Kodachi Kuno, and his second fiancée and childhood friend Ukyo Kuonji the okonomiyaki vendor, along with the Chinese Amazon Shampoo, supported by her great-grandmother Cologne. As the series progresses, the school becomes more eccentric with the return of the demented, Hawaii-obsessed Principal Kuno and the placement of the power-leeching alternating child/adult Hinako Ninomiya as Ranma's English teacher. Ranma's indecision in choosing his true love causes chaos in his romantic and school life.


title: Ranma ½

text: Rumiko Takahashi stated that "Ranma ½" was conceived to be a martial arts manga that connects all aspects of everyday life to martial arts. Because her previous series had female protagonists, the author decided that she wanted a male this time. However, she was worried about writing a male main character, and therefore decided to make him half-female. Before deciding on water for initiating his changes, she considered Ranma changing every time he was punched. It was after deciding this that she felt Jusenkyo had to be set in China, as it is the only place that could have such mysterious springs. She drew inspiration for "Ranma ½" from a variety of real-world objects. Some of the places frequently seen in the series are modeled after actual locations in Nerima, Tokyo (both the home of Takahashi and the setting of "Ranma ½").


title: Ranma ½

text: In a 1990 interview with "Amazing Heroes", Takahashi stated that she had four assistants that draw the backgrounds, panel lines and tone, while she creates the story and layout, and pencils and inks the characters. All her assistants are female; Takahashi stated that "I don't use male assistants so that the girls will work more seriously if they aren't worried about boys." In 1992, she explained her process as beginning with laying out the chapter in the evening so as to finish it by dawn, and resting for a day before calling her assistants. They finish it in two or three nights, usually utilizing five days for a chapter.


title: Ranma ½

text: Takahashi purposefully aimed the series to be popular with women and children. In 1993, an "Animerica" interviewer talking with Takahashi asked her if she intended the sex-changing theme "as an effort to enlighten a male-dominated society." Takahashi said that she does not think in terms of societal agendas and that she created the "Ranma ½" concept from simply wanting "a simple, fun idea". She added that she, as a woman and while recalling what manga she liked to read as a child, felt that "humans turning into animals might also be fun and ... you know, like a fairy tale." In 2013, she revealed that at the start of "Ranma" her editor told her to make it more dramatic, but she felt that was something she could not do. However, she admitted that drama did start to appear at the end. She also sat in on the voice actor auditions for the anime, where she insisted that male and female Ranma be voiced by different actors whose gender corresponded to that of the part.


title: Ranma ½

text: Written and illustrated by Rumiko Takahashi, "Ranma ½" began publication in the "shōnen" manga anthology "Weekly Shōnen Sunday" issue #36 published on August 19, 1987, following the ending of her series "Urusei Yatsura". From August 1987 until March 1996, the manga was published on a near weekly basis with the occasional colored page to spruce up the usually black and white stories. After nearly a decade of storylines, the final chapter was published in "Weekly Shōnen Sunday" issue #12 on March 6, 1996. The 407 chapters were periodically collected and published by Shogakukan into a total of 38 black and white "tankōbon" volumes from 1988 to 1996. They were reassembled into 38 "shinsōban" from April 2002 to October 2003.


title: Ranma ½

text: North American publisher Viz Media originally released "Ranma ½" in a monthly comic book format that contained two chapters each from 1992 to 2003, and had the images "flipped" to read left-to-right, causing the art to be mirrored. These were periodically collected into graphic novels. On March 18, 2004, after releasing 21 volumes, Viz announced that it would reprint a number of its graphic novels. The content remained the same, but the novels moved to a smaller format with different covers and a price drop. Each volume covers roughly the same amount of material as the Japanese volumes, but retained its left-to-right format and had minor differences in grouping so that it spans 36 volumes rather than the original 38. The final volume was released in stores on November 14, 2006, thus making it Viz's longest running manga, spanning over 13 years. At Anime Expo on July 7, 2013, Viz Media announced re-release of the manga in a format that combines two individual volumes into a single large one, and restores the original right-to-left reading order (a first in North America for this series). The first 2-in-1 book (volumes 1-2) was published on March 11, 2014; the final (volumes 35-36) in January, 2017. As of July 27, 2021, VIZ has released all 19 2-in-1 books digitally. Madman Entertainment publishes the two-in-one version in Australasia.


title: Ranma ½

text: An anime television series was created by Studio Deen and aired weekly between April 15, 1989, and September 16, 1989, on Fuji TV for 18 episodes, before being canceled due to low ratings. The series was then reworked by most of the same staff, retitled and launched in a different time slot, running for 143 episodes from October 20, 1989, to September 25, 1992. The anime stays true to the original manga but does differ by keeping Ranma's gender transformation a secret from the high school students, at least throughout most of its length. It also does not introduce Hikaru Gosunkugi until very late in the series, instead, Sasuke Sarugakure, the diminutive ninja retainer of the Kuno family fills a number of Gosunkugi's roles in early storylines but is a major character in his own right. The anime also alters the placement of many story arcs and contains numerous original episodes and characters not adapted from the manga.


title: Ranma ½

text: Viz Media licensed both anime series in 1993, making "Ranma ½" one of the first anime titles licensed by Viz. The English dub produced for the series was recorded by The Ocean Group in Vancouver, British Columbia. They released the series on VHS from their own "Viz Video" label, and on DVD a few years later in association with Pioneer Home Entertainment. Their releases collected both anime series as one, separated episodes into what they call "seasons", and changed the ordering of many of the episodes. Viz themselves re-released it on DVD in 2007 using their own DVD production company. At Otakon 2013, Viz announced that they re-acquired the TV series for Blu-ray and DVD release in 2014. The show is streamed on their anime channel service Neon Alley since Autumn 2013. In September 2020, Toonami co-creator Jason DeMarco revealed that he had previously tried to get "Ranma ½" aired on the American TV programming block, but "it's something we never were able to figure out, because, frankly, there's too much nudity." Madman Entertainment licensed some of the series for release in Australasia, although their rights expired after releasing only the first four "seasons" as one series.


title: Ranma ½

text: Studio Deen also created three theatrical films; "The Battle of Nekonron, China! A Battle to Defy the Rules!" on November 2, 1991; "Battle at Togenkyo! Get Back the Brides" on August 1, 1992; and "Super Indiscriminate Decisive Battle! Team Ranma vs. the Legendary Phoenix" on August 20, 1994. The first two films are feature length, but the third was originally shown in theaters with two other films: "Ghost Sweeper Mikami" and "Heisei Dog Stories: Bow".


title: Ranma ½

text: Following the ending of the TV series, 11 original video animations were released directly to home video, the earliest on December 7, 1993, and the eleventh on June 4, 1996. All but one are based on stories originally in the manga. Twelve years later, a "Ranma" animation was created for the "It's a Rumic World" exhibition of Rumiko Takahashi's artwork. Based on the "Nightmare! Incense of Deep Sleep" manga story from volume 34, it was shown on odd numbered days at the exhibition in Tokyo from July 30 to August 11, 2008. But it was not released until January 29, 2010, when it was put in a DVD box set with the "Urusei Yatsura" and "Inuyasha" specials that premiered at the same exhibit. It was then released on DVD and Blu-ray by itself on October 20, 2010. Viz Media also licensed all three films, and the original 11 OVAs for distribution in North America (however they released the third film as an OVA). MVM Films has released the first two films in the United Kingdom, while Madman Entertainment released them in Australasia.


title: Ranma ½

text: There have been fifteen video games based on the "Ranma ½" franchise. While most are fighting games, there have been several RPGs and puzzle games. Only two have been released in Western countries. "Ranma ½: Chōnai Gekitōhen" was released in the US as "Street Combat"; the characters were Americanized, having their appearances completely changed, and the music was changed as well. However, "" was released in both North America and Europe unaltered.


title: Ranma ½

text: A live action television adaption of "Ranma ½" aired on Nippon TV, in a two-hour time-slot, on December 9, 2011. Although it was initially reported that the special would contain an original story, the film does take its main plot from one of the manga's early stories with several other early scenes mixed in. The special stars Yui Aragaki as Akane, with Kento Kaku and Natsuna Watanabe playing male and female Ranma respectively. Ryōsei Tayama is cast as the antagonist, the new original character Okamada. The all-girl pop group 9nine contribute "Chikutaku☆2Nite" as the theme song. It was released on both DVD and Blu-ray on March 21, 2012.


title: Ranma ½

text: "The Ranma ½ Memorial Book" was published just as the manga ended in 1996. Acting as an end-cap to the series, it collects various illustrations from the series, features an interview with Takahashi, and includes tidbits about Ranma: summaries of his battles, his daily schedule, trivia, and a few exclusive illustrations.


title: Ranma ½

text: A "Movie + OVA Visual Comic" was released to illustrate the theatrical film "Super Indiscriminate Decisive Battle! Team Ranma vs. the Legendary Phoenix" and the OVA episodes "The One to Carry On" (both parts). It also included information on the voice actors, character designs, and a layout of the Tendo dojo.


title: Ranma ½

text: Additionally, guidebooks were released for three of the "Ranma ½" video games; these included not only strategies, but also interviews. Two books including interviews with the cast of the live-action TV drama, and some select stories, were released in 2011.


title: Ranma ½

text: The music from the "Ranma ½" TV series, films and OVAs have been released on various CDs. Four from the TV series, two from the first film, one from the second, one from the third film and OVAs, and three compiling the music by DoCo used in the OVAs. DoCo is a pop group composed of the anime's main female characters' voice actresses. Several compilation albums were also released, some composed of the opening and closing theme songs and others of image songs. Many of the image songs were first released as singles.


title: Ranma ½

text: By November 2006, it was reported that "Ranma ½" had sold over 49 million manga volumes in Japan. Shogakukan has printed 53 million copies as of November 2011. And by April 2021 it had 55 million copies in circulation.


title: Ranma ½

text: The "Ranma ½" anime was ranked number 17 on "Anime Insider"s 2001 list of the Top 50 Anime, although the list was limited to series that were released in North America. It ranked 36th on TV Asahi's 2006 list of Japan's 100 favorite animated TV series, which is based on an online poll of the Japanese people, up from the previous year's list where it ranked 45th. In November 2006, the New York Comic Con announced that it would host the first-ever American Anime Awards. Fans had the chance to vote for their favorite anime online during the month of January 2007. Only the five nominees receiving the most votes for each category were announced on February 5. Among the 12 different categories, "Ranma ½" was voted into the "Best Comedy Anime" category, and the "Ranma ½" OVAs were voted into the "Best Short Series" category. A 2019 NHK poll of 210,061 people saw "Ranma ½" and "Ranma ½ Nettōhen" named Takahashi's second best-animated work. Shampoo and Ranma were voted fourth and fifth place respectively in her characters category.


title: Ranma ½

text: Although Lum from Takahashi's first series "Urusei Yatsura" is often cited as the first "tsundere" character in anime and manga, Theron Martin of "Anime News Network" stated that "Ranma ½"s Akane Tendo is closer to how they would later typically be portrayed in the 2000s. He also suggested that one could argue "Ranma" is an early example of a harem or reverse harem series, due to the main character attracting suitors in both genders. The series's publication in North America proved highly successful as well, being many Americans' first introduction to manga and its anime adaptation one of the first Japanese animation shows to achieve popularity in the US. In an overview of the series, Jason Thompson called "Ranma ½" "the direct ancestor of all comedy-action manga, like "Sumomomo Momomo" and "History's Strongest Disciple Kenichi"", although noted that it was not the first, but only spanned the period when manga and anime sales were at their height. Relating it to Takahashi's other works, he summed the series up as "At the start, the fighting is minimal and it's almost a semi-serious relationship comedy, like "Maison Ikkoku"; then it turns completely ridiculous; and by the climax, when Ranma fights the evil bird-people of Phoenix Mountain in an excessively long and un-funny fight scene, it's like a warmup for "Inuyasha"." Reviewing the final volume of the manga, "Anime News Network" remarked that "Every dimension of Rumiko Takahashi's storytelling skills come into play here: comedy, romance and introspection, and of course, high-flying fantasy martial-arts action." However, they felt some of the action scenes were hard to follow and noted that the mirroring to left-to-right format caused errors with the art.


title: Ranma ½

text: In their review of Viz Media's season five DVD box set, "Anime News Network" praised the Japanese cast's performance and the animation, but criticized the English version's slight script changes and minor voice actors while praising its main cast. They also remarked that while "Ranma ½" is a classic, after a hundred episodes, the same jokes are just not funny anymore. "THEM Anime Reviews"' Raphael See called the television series and the OVAs "one of the funniest things [he's] ever seen, anime or otherwise" and also praised the English dub as some of the best. However, he was much more critical of the first two films particularly for both using the same damsel in distress plot. Mike Toole of "Anime News Network" included "Big Trouble in Nekronon, China" at number 83 on The Other 100 Best Anime Movies of All Time, a list of "lesser-known, lesser-loved classics," calling it "a solid action-comedy and a good, well-rounded example of the appeal of "Ranma ½""


title: Ranma ½

text: Hiroshi Aro admitted that he created "Futaba-kun Change!" based on "Ranma ½". Western comic book artists that have cited "Ranma ½" as an influence include Canadian Bryan Lee O'Malley on his series "Scott Pilgrim" and American Colleen Coover on her erotic series "Small Favors".


title: Ranma ½

text: Film director Makoto Shinkai mentioned that "Ranma ½" served as an inspiration for the 2016 animation film "Your Name". Matt Bozon, creator of the "Shantae" video game series, cited "Ranma ½" as a big influence on his work. The title of the fourth game, "", is also a tribute to the series.


title: Internet pornography

text: Internet pornography is any pornography that is accessible over the internet, primarily via websites, FTP servers peer-to-peer file sharing, or Usenet newsgroups. The availability of widespread public access to the World Wide Web in late 1990s led to the growth of internet pornography.


title: Internet pornography

text: A 2015 study finds "a big jump" in pornography viewing over the past few decades, with the largest increase occurring between people born in the 1970s and those born in the 1980s. While the study's authors note this increase is "smaller than conventional wisdom might predict," it's still quite significant. Those who were born in the 1980s onward are also the first to grow up in a world where they have access to the internet beginning in their teenage years, and this early exposure and access to internet pornography may be the primary driver of the increase. The sex and tech conference series Arse Elektronika dedicated their 2007 conference to what they call "pr0nnovation". The con presented a keynote by culture theorist Mark Dery and published a reader about the subject.


title: Internet pornography

text: , a single company, MindGeek, owns and operates many popular pornographic websites, including video sharing services Pornhub, RedTube, and YouPorn, as well as adult film producers Brazzers, Digital Playground, Men.com, Reality Kings, and Sean Cody, among others, but does not own the websites xHamster and XVideos. It has been alleged to be a monopoly.


title: Internet pornography

text: Pornography is regarded by some as one of the driving forces behind the expansion of the World Wide Web, like camcorders, VCRs and cable television before it. Pornographic images had been transmitted over the internet as ASCII porn but to send images over network needed computers with graphics capability and also higher network bandwidth. This was possible in the late 1980s and early 1990s through the use of anonymous FTP servers and through the Gopher protocol. At this time the internet had widespread ever since the late 1970s. One of the early Gopher/FTP sites was at Tudelft and was called the Digital Archive on the 17th Floor, This small image archive contained some low quality scanned pornographic images that were initially available to anyone anonymously, but the site soon became restricted to Netherlands only access, Pornographic videos started appearing on FTP and Gopher servers as well.


title: Internet pornography

text: Usenet newsgroups provided an early way of sharing images over the narrow bandwidth available in the early 1990s. Because of the network restrictions of the time, images had to be encoded as ascii text and then broken into sections before being posted to the Alt.binaries of the usenet. These files could then be downloaded and then reassembled before being decoded back to an image. Automated software such as Aub ("Assemble Usenet Binaries") allowed the automatic download and assembly of the images from a newsgroup. There was a rapid growth in the number of posts in the early 1990s but image quality was restricted by the size of files that could be posted.


title: Internet pornography

text: The method was also used to disseminate pornographic images, which were usually scanned from adult magazines. This type of distribution was generally free (apart from fees for internet access), and provided a great deal of anonymity. The anonymity made it safe and easy to ignore copyright restrictions, as well as protecting the identity of uploaders and downloaders. Around this time frame, pornography was also distributed via pornographic Bulletin Board Systems such as Rusty n Edie's. These BBSes could charge users for access, leading to the first commercial online pornography.


title: Internet pornography

text: A 1995 article written in "The Georgetown Law Journal" titled "Marketing Pornography on the Information Superhighway: A Survey of 917,410 Images, Description, Short Stories and Animations Downloaded 8.5 Million Times by Consumers in Over 2000 Cities in Forty Countries, Provinces and Territories" by Martin Rimm, a Carnegie Mellon University graduate student, claimed that (as of 1994) 83.5% of the images on Usenet newsgroups where images were stored were pornographic in nature. Before publication, Philip Elmer-DeWitt used the research in a "Time" magazine article, "On a Screen Near You: Cyberporn." The findings were attacked by journalists and civil liberties advocates who insisted the findings were seriously flawed. "Rimm's implication that he might be able to determine 'the percentage of all images available on the Usenet that are pornographic on any given day' was sheer fantasy" wrote Mike Godwin in HotWired. The research was cited during a session of U.S. Congress. The student changed his name and disappeared from public view. Godwin recounts the episode in "Fighting a Cyberporn Panic" in his book "".


title: Internet pornography

text: The invention of the World Wide Web spurred both commercial and non-commercial distribution of pornography. The rise of pornography websites offering photos, video clips and streaming media including live webcam access allowed greater access to pornography.


title: Internet pornography

text: On the Web, there are both commercial and free pornography sites. The bandwidth usage of a pornography site is relatively high, and the income a free site can earn through advertising may not be sufficient to cover the costs of that bandwidth. One recent entry into the free pornography website market are Thumbnail gallery post sites. These are free websites that post links to commercial sites, providing a sampling of the commercial site in the form of thumbnail images, or in the form of Free Hosted Galleries—samplings of full-sized content provided and hosted by the commercial sites to promote their site. Some free websites primarily serve as portals by keeping up-to-date indexes of these smaller sampler sites. These intents to create directories about adult content and websites were followed by the creation of adult wikis where the user can contribute their knowledge and recommend quality resources and links. When a user purchases a subscription to a commercial site after clicking through from a free thumbnail gallery site, the commercial site makes a payment to the owner of the free site. There are several forms of sites delivering adult content.


title: Internet pornography

text: The most common form of adult content is a categorized list (more often a table) of small pictures (called "thumbnails") linked to galleries. These sites are called a thumbnail gallery post (TGP). As a rule, these sites sort thumbs by category and type of content available on a linked gallery. Sites containing thumbs that lead to galleries with video content are called MGP (movie gallery post). The main benefit of TGP/MGP is that the surfer can get a first impression of the content provided by a gallery without actually visiting it.


title: Internet pornography

text: However, TGP sites are open to abuse, with the most abusive form being the so-called CJ (abbreviation for circlejerk), that contains links that mislead the surfer to sites he or she actually did not wish to see. This is also called a redirect.


title: Internet pornography

text: Linklists, unlike TGP/MGP sites, do not display a huge number of pictures. A linklist is a (frequently) categorised web list of links to so-called "freesites*", but unlike TGPs, links are provided in a form of text, not thumbs. It's still a question which form is more descriptive to a surfer, but many webmasters cite a trend that thumbs are much more productive, and simplify searching. On the other hand, linklists have a larger amount of unique text, which helps them improve their positions in search engine listings. TopLists are linklists whose internal ranking of freesites is based on incoming traffic from those freesites, except that freesites designed for TopLists have many more galleries.


title: Internet pornography

text: Another free source of pornography on the internet are the Usenet newsgroups that were the first home to such material. Newsgroups tend to be poorly organized and flooded with content that is off-topic or spam. Commercial software and websites are available that allow browsing the images or videos on newsgroups, sometimes with galleries of thumbnail images.


title: Internet pornography

text: Peer-to-peer file sharing networks provide another form of free access to pornography. While such networks have been associated largely with the illegal sharing of copyrighted music and movies, the sharing of pornography has also been a popular use for file sharing. Many commercial sites have recognized this trend and have begun distributing free samples of their content on peer-to-peer networks.


title: Internet pornography

text: As of 2011, the majority of viewers of online pornography were men; women tended to prefer romance novels and erotic fan fiction. Women comprised about one quarter to one third of visitors to popular pornography websites, but were only 2% of subscribers to pay sites. Subscribers with female names were flagged as signs of potential credit card fraud, because "so many of these charges result in an angry wife or mother demanding a refund for the misuse of her card." Nonetheless, women spend more time on pornography websites, particularly Pornhub, than men and were more interested in pornography upon marriage.


title: Internet pornography

text: Pornographic images may be either scanned into the computer from photographs or magazines, produced with a digital camera or a frame from a video before being uploading onto a pornographic website. The JPEG format is one of the most common formats for these images. Another format is GIF which may provide an animated image where the people in the picture move. It can last for only a second or two up to a few minutes and then reruns (repeats) indefinitely. If the position of the objects in the last frame is about the same as the first frame, there is the illusion of continuous action.


title: Internet pornography

text: Pornographic video clips may be distributed in a number of formats, including MPEG, WMV, and QuickTime. More recently, VCD and DVD image files allow the distribution of whole VCDs and DVDs. Many commercial porn sites exist that allow one to view pornographic streaming video. As of 2020, some internet pornography sites have begun offering 5K resolution content, while 1080p and 4K resolution are still more common.


title: Internet pornography

text: Since mid-2006, advertising-supported free pornographic video sharing websites based on the YouTube format have appeared. Referred to as Porn 2.0, these sites generally use Flash technology to distribute videos that were uploaded by users; these include user-generated content as well as scenes from commercial porn movies and advertising clips from pornographic websites.


title: Internet pornography

text: Another format of adult content that emerged with the advent of the internet is live webcams. Webcam content can generally be divided into two categories: group shows offered to members of an adult paysite, and one-on-one private sessions usually sold on a pay-per-view basis.


title: Internet pornography

text: Server-based webcam sex shows spur unique international economics: adult models in various countries perform live webcam shows and chat for clients in affluent countries. This kind of activity is sometimes mediated by companies that will set up websites and manage finances. They may maintain "office" space for the models to perform from, or they provide the interface for models to work at home, with their own computer with webcam. As of 2020, most so-called cam hosts stream directly from their home, due to the fast internet lines and cheap HD webcams, that are available at low-cost. The models get paid via tips or by selling exclusive content to their viewers through live cam sites, which can reach more than 20,000 viewers at once. Live cam sites are very popular. Big sites like Chaturbate or LiveJasmin are among the 100 most popular websites according to Alexa Internet.


title: Internet pornography

text: Other formats include text and audio files. While pornographic and erotic stories, distributed as text files, web pages, and via message boards and newsgroups, have been semi-popular, audio porn, via formats like MP3 and FLV, have increased in popularity. Audio porn can include recordings of people having sex or merely reading erotic stories. (Pornographic magazines are available in Zinio format, which provides a reader program to enable access.)