154-597: Ctenochasmatoidea is a group of early pterosaurs within the suborder Pterodactyloidea . Their remains are usually found in what were once coastal or lake environments. They generally had long wings, long necks, and highly specialized teeth. The earliest known ctenochasmatoid remains date to the Late Jurassic Kimmeridgian age. Previously, a fossil jaw recovered from the Middle Jurassic Stonesfield Slate formation in
308-545: A British geologist. This species however, was reassigned to the genus Camposipterus in 2013, therefore creating the new combination Camposipterus segwickii . Later, in 1861, Owen had uncovered multiple distinctively looking fossil remains yet again in the Cambridge Greensand, these were assigned to a new species named Pterodactylus simus , though the British paleontologist Harry Govier Seeley had created
462-563: A carnivore specializing in small animals. A 2020 study of pterosaur tooth wear supported the hypothesis that Pterodactylus preyed mainly on invertebrates and had a generalist feeding strategy, indicated by a relatively high bite force. Specimens of Pterodactylus have been found mainly in the Solnhofen limestone (geologically known as the Altmühltal Formation) of Bavaria , Germany . The main composition of this formation
616-543: A crest made up of soft tissue instead of bone. Another appearance of Pterodactylus -like creatures is in J. R. R. Tolkien 's Middle-earth legendarium . In this novel, the Nazgûl , introduced as the Black Riders, are nine characters who rode flying monsters that looked similarly built to Pterodactylus . Christopher Tolkien , the son of the author, described the flying monsters as "Nazgûl-birds"; his father described
770-402: 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. 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
924-612: A few isolated, large specimens once assigned to P. kochi that overlap P. antiquus in size. However, all specimens in this third year class also show sign of immaturity. Fully mature Pterodactylus specimens remain unknown, or may have been mistakenly classified as a different genus. The distinct year classes of Pterodactylus antiquus specimens show that this species, like the contemporary Rhamphorhynchus muensteri , likely bred seasonally and grew consistently during its lifetime. A new generation of 1st year class P. antiquus would have been produced seasonally, and reached 2nd-year size by
1078-458: A few millimetres thin transversely. The bony crest base would typically be extended by keratinous or other soft tissue. 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 ,
1232-411: 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
1386-478: A fossilized crustacean, and it was not until 1856 that this species was properly described as a pterosaur by German paleontologist Hermann von Meyer . In his first description of the Mannheim specimen, Collini did not conclude that it was a flying animal. In fact, Collini could not fathom what kind of animal it might have been, rejecting affinities with the birds or the bats. He speculated that it may have been
1540-407: A good wing." Contrary to von Moll's report, the fossil was not missing; it was being studied by Samuel Thomas von Sömmerring , who gave a public lecture about it on December 27, 1810. In January 1811, von Sömmerring wrote a letter to Cuvier deploring the fact that he had only recently been informed of Cuvier's request for information. His lecture was published in 1812, and in it von Sömmerring named
1694-499: A junior synonym. Even if Aerodactylus were valid, at least one specimen with these features is still considered to belong to Pterodactylus , BSP 1929 I 18, which has an occipital lappet similar to the proposed Aerodactylus definition, and also possesses a small triangular soft tissue crest with the peak of the crest positioned above the eyes. Like other pterosaurs (most notably Rhamphorhynchus ), Pterodactylus specimens can vary considerably based on age or level of maturity. Both
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#17328026106151848-457: A larger surface area to catch individual small prey. Most ctenochasmatoids have wing proportions akin to those of modern shorebirds and ducks , and probably possessed a similar frantic, powerful flight style. The exception is Ctenochasma , which appears to have had longer wings and was probably more comparable to modern skuas . Launching varied radically among ctenochasmatoids. In forms like Cycnorhamphus , long limbs and shorter torsos meant
2002-416: A level of relative ease. In forms like Pterodaustro , however, which possessed long torsos and short limbs, launching might have been a more taxing and prolonged affair, only possible in large open areas, just like modern heavy-bodied aquatic birds such as swans , even with the pterosaurian quadrupedal launching. Ctenochasmatoidea was first named as the subfamily Ctenochasmatinae by Franz Nopcsa . Under
2156-445: A limited mobility. These toes were clawed but the claws were smaller than the hand claws. 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
2310-516: 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
2464-425: A new fossil of Tupandactylus cf. imperator was found to have melanosomes in forms that signal an earlier-than-anticipated development of patterns found in extant feathers. The new specimen suggested that pterosaur integumentary melanosomes exhibited a more complex organization than those previously known from other pterosaurs. This indicates the presence of a unique form of melanosomes within pterosaur integument at
2618-427: A new genus name. It has sometimes been placed in the genus Diopecephalus because Harry Govier Seeley based this genus partly on the P. longicollum material. However, it was shown by Bennett that the type specimen later designated for Diopecephalus was a fossil belonging to P. kochi , and no longer thought to be separate from Pterodactylus . Diopecephalus is therefore a synonym of Pterodactylus , and as such
2772-432: A new separate species called Ornithocephalus longicaudus ; the specific name means 'long tail', in reference to the animal's tail size. German paleontologist Hermann von Meyer in 1845 officially emended that the genus Pterodactylus had priority over Ornithocephalus , so he reassigned the species O. münsteri and O. longicaudus into Pterodactylus münsteri and Pterodactylus longicaudus . In 1846, von Meyer created
2926-534: A new species called "Pterodactylus oweni" , but this was changed to Pterodactylus occidentalis because "P. oweni" was found to have been preoccupied by a pterosaur species described with the same name back in 1864 by Seeley. In 1872, American paleontologist Edward Drinker Cope also found various pterosaur specimens in North America, he assigned these to two new species known as Ornithochirus umbrosus and Ornithochirus harpyia , Cope attempted to assign
3080-439: A new species called Pterodactylus compressirostris . In 1914 however, paleontologist Reginald Hooley redescribed P. compressirostris , to which he erected the genus Lonchodectes ( lit. ' lance biter ' ), and therefore made P. compressirostris the type species, and created the new combination L. compressirostris . In a 2013 review, P. giganteus and P. cuvieri were reassigned to new genera; P. giganteus
3234-453: A new species called Pterodactylus sagittirostris , this species however, was reassigned to the genus Lonchodectes in 1914 by Hooley, which resulted in an L. sagittirostris . This conclusion was revised by Rigal et al. in 2017, who disagreed with Hooley's reassignment, and therefore created the genus Serradraco , which afterwards resulted in a new combination called S. sagittirostris . Assigning new pterosaur species to Pterodactylus
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#17328026106153388-791: A number of changes in classification and thus have acquired a large number of synonyms. Additionally, a number of species assigned to Pterodactylus are based on poor remains that have proven difficult to assign to one species or another and are therefore considered nomina dubia ( lit. ' doubtful names ' ). The following list includes names that were used to identify new pterosaur species that now have been reclassified, or until recently thought to be pertaining to Pterodactylus proper, and names based on other material that has as yet not been assigned to other genera. This list also includes species that are nomina nuda ('naked names'), which are species that were not published formally. Species that are nomina oblita ('forgotten names') are
3542-613: A predominant occurrence in aquatic environments, the exception being the more slender-limbed and short-torsoed gallodactylids . They occupied a wide variety of ecological niches, from generalistic carnivores like Pterodactylus to filter-feeders like Pterodaustro and possible molluscivores like Cycnorhamphus . Most common, however, were straight-jawed, needle-toothed forms, some of the most notable being Ctenochasma and Gnathosaurus ; these possibly occupied an ecological niche akin to that of modern spoonbills , their teeth forming spatula -like jaw profile extensions, allowing them
3696-452: A pterosaur, specifically a species of Pterodactylus . At the time however, most paleontologists incorrectly consider the genus Ornithocephalus ( lit. ' bird-head ' ) to be the valid name for Pterodactylus , and therefore the specimen found was named as Ornithocephalus Münsteri , which was first mentioned by Graf zu Münster himself. Another specimen was found and described by Graf zu Münster in 1839, he assigned this specimen to
3850-411: 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
4004-612: A sea creature, not for any anatomical reason, but because he thought the ocean depths were more likely to have housed unknown types of animals. The idea that pterosaurs were aquatic animals persisted among a minority of scientists as late as 1830, when the German zoologist Johann Georg Wagler published a text on "amphibians" which included an illustration of Pterodactylus using its wings as flippers. Wagler went so far as to classify Pterodactylus , along with other aquatic vertebrates (namely plesiosaurs , ichthyosaurs , and monotremes ), in
4158-549: A separate generic name called Ornithocheirus , and reassigned P. simus as the type species, which created the combination Ornithocheirus simus . Between the years 1869 and 1870, Seeley had reassigned many pterosaur species into Ornithocheirus , while also creating several new species. Many of these species however, are now reclassified to other genera, or considered nomina dubia . In 1874, further specimens were found in England, again by Owen, these ones were assigned to
4312-433: A somewhat longer description, in which he named the animal Petro-Dactyle , this was a typographical error however, and was later corrected by him to Ptéro-Dactyle . He also refuted a hypothesis by Johann Friedrich Blumenbach that it would have been a shore bird. Cuvier remarked: "It is not possible to doubt that the long finger served to support a membrane that, by lengthening the anterior extremity of this animal, formed
4466-449: A supraneural plate that, however, would not contact the notarium. 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,
4620-464: A thousand bristle-like teeth. Dsungaripteridae covered their teeth with jawbone tissue for a crushing function. If teeth were present, they were placed in separate tooth sockets. Replacement teeth were generated behind, not below, the older teeth. 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
4774-416: 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 . 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. The pterosaur wing membrane
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4928-408: A weight of up to 250 kilograms (550 pounds) for the largest species. Compared to the other vertebrate flying groups, the birds and bats, pterosaur skulls were typically quite large. Most pterosaur skulls had elongated jaws. Their skull bones tend to be fused in adult individuals. Early pterosaurs often had heterodont teeth, varying in build, and some still had teeth in the palate. In later groups
5082-453: A wide range of adult sizes , from the very small anurognathids to the largest known flying creatures, including Quetzalcoatlus and Hatzegopteryx , which reached wingspans of at least nine metres. The combination of endothermy , a good oxygen supply and strong muscles made pterosaurs powerful and capable flyers. Pterosaurs are often referred to by popular media or the general public as "flying dinosaurs", but dinosaurs are defined as
5236-482: A wingspan no less than 25 centimetres (10 inches). The most sizeable forms represent the largest known animals ever to fly, with wingspans of up to 10–11 metres (33–36 feet). Standing, such giants could reach the height of a modern giraffe . Traditionally, it was assumed that pterosaurs were extremely light relative to their size. Later, it was understood that this would imply unrealistically low densities of their soft tissues. Some modern estimates therefore extrapolate
5390-999: Is cladogram following a topology recovered by Brian Andres, using the most recent iteration of his data set (Andres, 2021). Andre's analysis found the "aurorazhdarchian" group recovered by the analysis of Vidovic and Martill to mostly be members of the Ctenochasmatidae, with only a small group of gallodactylids falling outside that group. Pterodactylus [REDACTED] Cycnorhamphus Aurorazhdarcho Normannognathus "Pterodactylus" micronyx Liaodactylus Ctenochasma Pterodaustro [REDACTED] Beipiaopterus Gegepterus Kepodactylus Elanodactylus Feilongus Moganopterus Ardeadactylus Huanhepterus Plataleorhynchus [REDACTED] Gnathosaurus [REDACTED] [REDACTED] [REDACTED] Pterosaurs Ornithosauria Seeley , 1870 Pterosaurs are an extinct clade of flying reptiles in
5544-425: Is characterized by skulls of around 55 to 95 millimeters (2.2 to 3.7 in) long, but are still immature however. These first two size groups were once classified as juveniles and adults of the species P. kochi , until further study showed that even the supposed "adults" were immature, and possibly belong to a distinct genus. A third year class is represented by specimens of the "traditional" P. antiquus , as well as
5698-555: 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. 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
5852-422: Is curved to behind, resulting in a rounded wing tip, which reduces induced drag . The wingfinger is also bent somewhat downwards. 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
6006-421: 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
6160-419: Is fine-grained limestone that originated mainly from the nearby towns Solnhofen and Eichstätt, which is formed by mud silt deposits. The Solnhofen Limestone is a diverse Lagerstätte that contains a wide range of different creatures, including highly detailed fossilized imprints of soft bodied organisms such as jellyfishes . Abundant specimens of pterosaurs similar to Pterodactylus were also found within
6314-418: 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 . A bone unique to pterosaurs, known as
Ctenochasmatoidea - Misplaced Pages Continue
6468-475: Is similar to modern crocodilians , rather than the rapid growth of modern birds . Comparisons between the scleral rings of Pterodactylus antiquus and modern birds and reptiles suggest that it may have been diurnal . This may also indicate niche partitioning with contemporary pterosaurs inferred to be nocturnal , such as Ctenochasma and Rhamphorhynchus . Based on the shape, size, and arrangement of its teeth, Pterodactylus has long been recognized as
6622-489: Is sometimes used to refer to any kind of animal belonging to the order Pterosauria , though most of the time to Pterodactylus itself and the distantly-related Pteranodon , the most well-known members of the group. The popular aspect of Pterodactylus consists of an elongated head crest, and potentially large wings. Studies of Pterodactylus , however, conclude that it may even lack a bony cranial crest, though several analysis have proven that Pterodactylus may in fact have
6776-746: Is thought to contain only a single species , Pterodactylus antiquus , which was the first pterosaur to be named and identified as a flying reptile and one of the first prehistoric reptiles to ever be discovered. Fossil remains of Pterodactylus have primarily been found in the Solnhofen limestone of Bavaria , Germany , which dates from the Late Jurassic period ( Tithonian stage), about 150.8 to 148.5 million years ago. More fragmentary remains of Pterodactylus have tentatively been identified from elsewhere in Europe and in Africa . Pterodactylus
6930-409: Is truly valid. The skulls of adult Pterodactylus were long and thin, with about 90 narrow and conical teeth. The teeth extended back from the tips of both jaws, and became smaller farther away from the jaw tips. This was unlike the ones seen in most relatives, where teeth were absent in the upper jaw tip and were relatively uniform in size. The teeth of Pterodactylus also extended farther back into
7084-550: Is unavailable for use as a new genus for "P." longicollum . "P." longicollum was eventually made the type species of a separate genus Ardeadactylus . The only well-known and well-supported species left by the first decades of the 21st century were P. antiquus and P. kochi . However, most studies between 1995 and 2010 found little reason to separate even these two species, and treated them as synonymous. More recent studies of pterosaur relationships have found anurognathids and pterodactyloids to be sister groups, which would limit
7238-535: The International Code of Zoological Nomenclature , this makes Nopcsa the author of Ctenochasmatidae and Ctenochasmatoidea as well. The modern clade Ctenochasmatoidea was defined by David Unwin in 2003 as the clade containing Cycnorhamphus suevicus , Pterodaustro guinazui , their most recent common ancestor, and all its descendants. Below is a cladogram showing the results of a phylogenetic analysis presented by Steven Vidovic and David Martill, using
7392-477: The United Kingdom , was considered the oldest known. This specimen supposedly represented a member of the family Ctenochasmatidae, though further examination suggested it belonged to a teleosaurid stem-crocodilian instead of a pterosaur. Most ctenochasmatoids were aquatic or semi-aquatic pterosaurs, possessing large webbed hindfeet and long torsos - both adaptations for swimming and floating -, as well as
7546-570: The ctenochasmatoids may have been more closely related to the more advanced dsungaripteroids , or in some cases, fall outside both groups. Their conclusion was published in 2017, in which they placed Pterodactylus as a basal member of the suborder Pterodactyloidea . Eosipterus yangi Pterodactylus antiquus [REDACTED] Diopecephalus kochi Gallodactylidae [REDACTED] Ctenochasmatidae [REDACTED] Altmuehlopterus ramphastinus [REDACTED] Dsungaripteroidea [REDACTED] [REDACTED] As illustrated below,
7700-526: The dinosaurs Archaeopteryx and Compsognathus were also found within the limestone, these specimens were related to early evolution of feathers , since they were some of the only ones that had them during the Jurassic period. Various lizard remains were also found alongside those of Pterodactylus , with several specimens assigned to Ardeosaurus , Bavarisaurus and Eichstaettisaurus . Crocodylomorph specimens were widely distributed within
7854-831: The order Pterosauria . They existed during most of the Mesozoic : from the Late Triassic to the end of the Cretaceous (228 to 66 million years ago). Pterosaurs are the earliest vertebrates known to have evolved powered flight . Their wings were formed by a membrane of skin, muscle, and other tissues stretching from the ankles to a dramatically lengthened fourth finger. There were two major types of pterosaurs. Basal pterosaurs (also called 'non-pterodactyloid pterosaurs' or ' rhamphorhynchoids ') were smaller animals with fully toothed jaws and, typically, long tails. Their wide wing membranes probably included and connected
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#17328026106158008-477: 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 protofeathers. But Liliana D'Alba points out that
8162-427: 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
8316-414: The "bat model" depicted pterosaurs as warm-blooded and furred, it would turn out to be more correct in certain aspects than Cuvier's "reptile model" in the long run. In 1834, Johann Jakob Kaup coined the term Pterosauria. Pterodactylus Pterodactylus (from Ancient Greek : πτεροδάκτυλος , romanized : pterodáktylos ' winged finger ' ) is a genus of extinct pterosaurs . It
8470-536: The 1990s it was understood that this was even true for part of the remaining species. P. elegans , for example, was found by numerous studies to be an immature Ctenochasma . Another species of Pterodactylus originally based on small, immature specimens was P. micronyx . However, it has been difficult to determine exactly of what genus and species P. micronyx might be the juvenile form. Stéphane Jouve, Christopher Bennett and others had once suggested that it probably belonged either to Gnathosaurus subulatus or one of
8624-428: 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. The presence of pycnofibers strongly indicates that pterosaurs were endothermic (warm-blooded). They aided thermoregulation, as
8778-480: The 2010s, a large body of research had been developed based on the idea that P. kochi and P. scolopaciceps were early growth stages of P. antiquus . However, in 2014, two scientists began publishing research that challenged this paradigm. Steven Vidovic and David Martill concluded that differences between specimens of P. kochi , P. scolopaciceps , and P. antiquus , such as different lengths of neck vertebrae, thinner or thicker teeth, more rounded skulls, and how far
8932-587: The Late Jurassic period, while both Pteranodon and Quetzalcoatlus lived during the Late Cretaceous . Pterodactylus lived alongside other small pterosaurs such as the well-known Rhamphorhynchus , as well as other genera such as Scaphognathus , Anurognathus and Ctenochasma . Pterodactylus is classified as an early-branching member of the ctenochasmatid lineage , within the pterosaur clade Pterodactyloidea . The type specimen of
9086-593: The animal now known as Pterodactylus antiquus was the first pterosaur fossil ever to be identified. The first Pterodactylus specimen was described by the Italian scientist Cosimo Alessandro Collini in 1784, based on a fossil skeleton that had been unearthed from the Solnhofen limestone of Bavaria. Collini was the curator of the Naturalienkabinett , or nature cabinet of curiosities (a precursor to
9240-428: 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. There was a clear difference between early pterosaurs and advanced species regarding
9394-484: The ankles. The exact curvature of the trailing edge, however, is still equivocal. 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
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#17328026106159548-427: 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. 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,
9702-486: The back edge of the antorbital fenestra and the back of the skull. In at least one specimen, the crest had a short bony base, also seen in related pterosaurs like Germanodactylus . Solid crests have only been found on large, fully adult specimens of Pterodactylus , indicating that this was a display structure that became larger and more well developed as individuals reached maturity. In 2013, pterosaur researcher S. Christopher Bennett noted that other authors claimed that
9856-413: 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
10010-447: 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. The front of the pubic bones articulated with a unique structure, the paired prepubic bones. Together these formed
10164-474: The clade Anurognathidae ( Anurognathus , Jeholopterus , Vesperopterylus ) is debated. Anurognathids were highly specialized. Small flyers with shortened jaws and a wide gape, some had large eyes suggesting nocturnal or crepuscular habits, mouth bristles, and feet adapted for clinging. Parallel adaptations are seen in birds and bats that prey on insects in flight. Pterosaurs had a wide range of sizes, though they were generally large. The smallest species had
10318-445: The clades Ornithocheiroidea ( Istiodactylus , Ornithocheirus , Pteranodon ), Ctenochasmatoidea ( Ctenochasma , Pterodactylus ), Dsungaripteroidea ( Germanodactylus , Dsungaripterus ), and Azhdarchoidea ( Tapejara , Tupuxuara , Quetzalcoatlus ). The two groups overlapped in time, but the earliest pterosaurs in the fossil record are basal pterosaurs, and the latest pterosaurs are pterodactyloids. The position of
10472-658: The class Gryphi, between birds and mammals. The German/French scientist Johann Hermann was the one who first stated that Pterodactylus used its long fourth finger to support a wing membrane. Back in March 1800, Hermann alerted the prominent French scientist Georges Cuvier to the existence of Collini's fossil, believing that it had been captured by the invading forces of the French Consulate and sent to collections in Paris (and perhaps to Cuvier himself) as war booty; at
10626-502: The descendants of the last common ancestor of the Saurischia and Ornithischia , which excludes the pterosaurs. Pterosaurs are nonetheless more closely related to birds and other dinosaurs than to crocodiles or any other living reptile, though they are not bird ancestors. Pterosaurs are also colloquially referred to as pterodactyls , particularly in fiction and journalism. However, technically, pterodactyl may refer to members of
10780-421: The description of Rhamphorhynchus ; fossil collector Georg Graf zu Münster alerted the German paleontologist Samuel Thomas von Sömmerring about several distinct fossil specimens, Sömmerring thought that they belonged to an ancient bird. Further fossil preparations had uncovered teeth, to which Graf zu Münster created a skull cast. He later sent the cast to Professor Georg August Goldfuss , who recognized it as
10934-431: The description of the preserved integumentary structures on the two anurognathid 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. In 2022,
11088-544: The difference with the "quills" found on many of the bird-like maniraptoran specimens too fundamental. 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
11242-457: The distinct species P. kochi ) have a small number of teeth, as few as 15 in some, and the teeth have a relatively broad base. The teeth of other P. antiquus specimens are both narrower and more numerous (up to 90 teeth are present in several specimens). Pterodactylus specimens can be divided into two distinct year classes. In the first year class, the skulls are only 15 to 45 millimeters (0.59 to 1.77 in) in length. The second year class
11396-410: The down feathers found on both avian and some non-avian dinosaurs , suggesting that early feathers evolved in the common ancestor of pterosaurs and dinosaurs, possibly as insulation. They were warm-blooded (endothermic), active animals. The respiratory system had efficient unidirectional "flow-through" breathing using air sacs , which hollowed out their bones to an extreme extent. Pterosaurs spanned
11550-405: The earliest available definitions for each clade name. Diopecephalus Aerodactylus [REDACTED] Huanhepterus Ardeadactylus Aurorazhdarcho Cycnorhamphus Feilongus Moganopterus Gladocephaloideus Pterodaustro [REDACTED] Beipiaopterus Gegepterus Ctenochasma Gnathosaurus Plataleorhynchus [REDACTED] Below
11704-475: The estimated total length of its skull) and has a maximum height of 0.9 mm (0.035 inches) above the orbit. Several specimens previously referred to P. antiquus preserved evidence of the soft tissue extensions of these crests, including an "occipital lappet", a flexible, tab-like structure extending from the back of the skull. Most of these specimens have been reclassified in the related species Aerodactylus scolopaciceps , which may however be nothing more than
11858-404: 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. The bony elements of the arm formed a mechanism to support and extend the wing. Near the body, the humerus or upper arm bone
12012-676: The feather-specific melanosome signaling found in extant birds are possibly homologous with those found in pterosaurs. 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, Italian naturalist Cosimo Alessandro Collini
12166-432: 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
12320-520: The first was named in 1846 as Pterodactylus giganteus ; the specific name means 'the gigantic one' in Latin, in reference to the large size of the remains, and the second species was named in 1851 as Pterodactylus cuvieri , in honor of the French scientist Georges Cuvier. Later in 1851, Owen named and described new pterosaur specimens that have been found yet again in England. He assigned these specimens to
12474-450: 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
12628-400: 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
12782-417: 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
12936-417: The formation, these include the rhamphorhynchids Rhamphorhynchus and Scaphognathus , several gallodactylids such as Aerodactylus , Ardeadactylus , Aurorazhdarcho and Cycnorhamphus , the ctenochasmatids Ctenochasma and Gnathosaurus , the anurognathid Anurognathus , the germanodactylid Germanodactylus , as well as the basal euctenochasmatian Diopecephalus . Fossil remains of
13090-402: The fossil site, most were assigned to the metriorhynchid genera Cricosaurus , Dakosaurus , Geosaurus and Rhacheosaurus . These genera are colloquially called as marine or sea crocodiles due to their similar built. The turtle genera Eurysternum and Paleomedusa were also found within the formation. Fossils of the ichthyosaur Aegirosaurus also appeared to be present in
13244-452: The fossilization process. Bennett pointed in particular to the data used to distinguish Aerodactylus , which was so different from the data for related species, it might be due to an unnatural assemblage of specimens. As a result, Bennett continued to consider Diopecephalus and Aerodactylus simply as year-classes of immature Pterodactylus antiquus . During its over-200-year history, the various species of Pterodactylus have gone through
13398-415: 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
13552-473: 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
13706-429: The genus Pterodactylus , and more broadly to members of the suborder Pterodactyloidea of the pterosaurs. Pterosaurs had a variety of lifestyles. Traditionally seen as fish-eaters, the group is now understood to have also included hunters of land animals, insectivores, fruit eaters and even predators of other pterosaurs. They reproduced by eggs , some fossils of which have been discovered. The anatomy of pterosaurs
13860-431: The ground, furred, warmblooded and had a wing membrane reaching the ankle. Some of these elements have been confirmed, some refuted by modern research, while others remain disputed. In 1815, the generic name Ptéro-Dactyle was latinized to Pterodactylus by Constantine Samuel Rafinesque . Unaware of Rafinesque's publication however, Cuvier himself in 1819 latinized the name Ptéro-Dactyle again to Pterodactylus , but
14014-683: The ground, they walked well on all four limbs with an upright posture, standing plantigrade on the hind feet and folding the wing finger upward to walk on the three-fingered "hand". They could take off from the ground, and fossil trackways show that at least some species were able to run, wade, and/or swim. Their jaws had horny beaks, and some groups lacked teeth. Some groups developed elaborate head crests with sexual dimorphism . Pterosaurs sported coats of hair-like filaments known as pycnofibers , which covered their bodies and parts of their wings. Pycnofibers grew in several forms, from simple filaments to branching down feathers . These may be homologous to
14168-464: 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
14322-412: 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
14476-478: The hind legs. On the ground, they would have had an awkward sprawling posture, but the anatomy of their joints and strong claws would have made them effective climbers, and some may have even lived in trees. Basal pterosaurs were insectivores or predators of small vertebrates. Later pterosaurs ( pterodactyloids ) evolved many sizes, shapes, and lifestyles. Pterodactyloids had narrower wings with free hind limbs, highly reduced tails, and long necks with large heads. On
14630-480: The jaw compared to close relatives, and some were present below the front of the nasoantorbital fenestra , which is the largest opening in the skull. Another autapomorphy that Pterodactylus has is that the skull and jaws were straight, which are unlike the upwardly curved jaws seen in the related ctenochasmatids . Pterodactylus , like related pterosaurs, had a crest on its skull composed mainly of soft tissues. In adult Pterodactylus , this crest extended between
14784-401: 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,
14938-547: The legs. 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
15092-465: The long metacarpals for the bones of the lower arm, the lower arm for the humerus , this upper arm for the breast bone and this sternum again for the shoulder blades . Sömmerring did not change his opinion that these forms were bats and this "bat model" for interpreting pterosaurs would remain influential long after a consensus had been reached around 1860 that they were reptiles. The standard assumptions were that pterosaurs were quadrupedal, clumsy on
15246-471: The long fourth finger to the ankle and a covering of fur (neither wing membranes nor fur had been preserved in the specimen). Hermann also added a membrane between the neck and wrist, as is the condition in bats . Cuvier agreed with this interpretation, and at Hermann's suggestion, Cuvier became the first to publish these ideas in December 1800 in a very short description. However, contrary to Hermann, Cuvier
15400-402: 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. 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
15554-421: 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. The shoulder girdle was a strong structure that transferred the forces of flapping flight to
15708-469: The modern concept of the natural history museum), in the palace of Charles Theodore, Elector of Bavaria at Mannheim . The specimen had been given to the collection by Count Friedrich Ferdinand zu Pappenheim around 1780, having been recovered from a lithographic limestone quarry in Eichstätt . The actual date of the specimen's discovery and entry into the collection is unknown however, and it
15862-426: The more inclusive group Caelidracones to just two clades. In 1996, Bennett suggested that the differences between specimens of P. kochi and P. antiquus could be explained by differences in age, with P. kochi (including specimens alternately classified in the species P. scolopaciceps ) representing an immature growth stage of P. antiquus . In a 2004 paper, Jouve used a different method of analysis and recovered
16016-519: The name Pterodactyli to contain Pterodactylus and other pterosaurs known at the time. This was emended to the family Pterodactylidae by Prince Charles Lucien Bonaparte in 1838. However, this group has more recently been given several competing definitions. Beginning in 2014, researchers Steven Vidovic and David Martill constructed an analysis in which several pterosaurs traditionally thought of as archaeopterodactyloids closely related to
16170-575: 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. The torso
16324-557: The new genus Aerodactylus for P. scolopaciceps as well. So, what Bennett considered early growth stages of one species, Vidovic and Martill considered representatives of new species. In 2017, Bennett challenged this hypothesis, he claimed that while Vidovic and Martill had identified real differences between these three groups of specimens, they had not provided any rationale that the differences were enough to distinguish them as species, rather than just individual variation, growth changes, or simply due to crushing and distortion during
16478-425: The new species Pterodactylus gemmingi based on long-tailed remains; the specific name honors the fossil collector Carl Eming von Gemming . Later, in 1847, von Meyer finally erected the generic name Rhamphorhynchus ( lit. ' beak snout ' ) due to the distinctively long tails seen in the specimens found, which are much longer than those seen in Pterodactylus . He assigned the species P. longicaudus as
16632-401: The ones that have been disused, and species that are nomina rejecta ('rejected names') are the ones that have been rejected because a more preferable name had been accepted instead. Pterodactylus is regarded as one of the most iconic prehistoric creatures, with multiple appearances in books, movies, as well as television series and several videogames. The informal name " pterodactyl "
16786-458: 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. Most or all pterosaurs had hair -like filaments known as pycnofibers on
16940-572: The proportions of the limb bones, size and shape of the skull, and size and number of teeth changed as the animals grew. Historically, this has led to various growth stages (including growth stages of related pterosaurs) being mistaken for new species of Pterodactylus . Several detailed studies using various methods to measure growth curves among known specimens have suggested that there is actually only one valid species of Pterodactylus , P. antiquus . The youngest immature specimens of Pterodactylus antiquus (alternately interpreted as young specimens of
17094-405: 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
17248-402: 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. 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
17402-441: 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
17556-488: 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
17710-549: The results of a different topology are based on a phylogenetic analysis made by Longrich, Martill, and Andres in 2018. Unlike the previous results above, they placed Pterodactylus within the clade Euctenochasmatia , resulting in a more derived position. Germanodactylus cristatus Germanodactylus rhamphastinus [REDACTED] Pterodactylus antiquus [REDACTED] Cycnorhamphus suevicus Normannognathus wellnhoferi Ctenochasmatidae [REDACTED] Numerous species have been assigned to Pterodactylus in
17864-416: The same result, showing that the "distinctive" features of P. kochi were age-related, and using mathematical comparison to show that the two forms are different growth stages of the same species. An additional review of the specimens published in 2013 demonstrated that some of the supposed differences between P. kochi and P. antiquus were due to measurement errors, further supporting their synonymy. By
18018-508: The site, as well as fish remains, with many specimens assigned to ray-finned fishes such as the halecomorphs Lepidotes , Propterus , Gyrodus , Mesturus , Proscinetes , Caturus , Ophiopsis and Ophiopsiella , the pachycormids Asthenocormus , Hypsocormus and Orthocormus , as well as the aspidorhynchid Aspidorhynchus , and the ichthyodectid Thrissops . Initial classifications for Pterodactylus started when paleontologist Hermann von Meyer used
18172-586: The skull, the sutures between elements disappeared. In some later pterosaurs, the backbone over the shoulders fused into a structure known as a notarium , which served to stiffen the torso during flight, and provide a stable support for the shoulder blade . Likewise, the sacral vertebrae could form a single synsacrum while the pelvic bones fused also. Basal pterosaurs include the clades Dimorphodontidae ( Dimorphodon ), Campylognathididae ( Eudimorphodon , Campyognathoides ), and Rhamphorhynchidae ( Rhamphorhynchus , Scaphognathus ). Pterodactyloids include
18326-480: The soft tissue crest of Pterodactylus extended backward behind the skull; Bennett himself, however, didn't find any evidence for the crest extending past the back of the skull. Two specimens of P. antiquus (the holotype specimen BSP AS I 739 and the incomplete skull BMMS 7, the largest known skull of P. antiquus ) have a low bony crest on their skulls; in BMMS 7 it is 47.5 mm long (1.87 inches, more or less 24% of
18480-465: The species Ornithocephalus antiquus . The animal was described as being both a bat, and a form in between mammals and birds, i.e. not intermediate in descent but in "affinity" or archetype . Cuvier disagreed, and the same year in his Ossemens fossiles provided a lengthy description in which he restated that the animal was a reptile. It was not until 1817 that a second specimen of Pterodactylus came to light, again from Solnhofen . This tiny specimen
18634-482: The species belonging to Ctenochasma , though after additional research Bennett assigned it to the genus Aurorazhdarcho . Another species with a complex history is P. longicollum , named by von Meyer in 1854, based on a large specimen with a long neck and fewer teeth. Many researchers, including David Unwin , have found P. longicollum to be distinct from P. kochi and P. antiquus . Unwin found P. longicollum to be closer to Germanodactylus and therefore requiring
18788-512: The specific name he then gave, longirostris , has to give precedence to von Sömmerring's antiquus . In 1888, English naturalist Richard Lydekker designated Pterodactylus antiquus as the type species of Pterodactylus , and considered Ornithocephalus antiquus a synonym. He also designated specimen BSP AS.I.739 as the holotype of the genus. Pterodactylus is known from over 30 fossil specimens, and though most belong to juveniles, many preserve complete skeletons. Pterodactylus antiquus
18942-409: The specimens he found to the genus Ornithocheirus , but misspelled forgetting the 'e'. In 1875 however, Cope reassigned the species O. umbrosus and O. harpyia into Pterodactylus umbrosus and Pterodactylus harpyia , though these species had been considered nomina dubia ever since. Paleontologist Samuel Wendell Williston unearthed the first skull of the pterosaur, and found that the animal
19096-400: 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
19250-405: 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. 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 )
19404-452: The teeth extended back in the jaws, were significant enough to separate them into three distinct species. Vidovic and Martill also performed a phylogenetic analysis which treated all relevant specimens as distinct units, and found that the P. kochi type specimen did not form a natural group with that of P. antiquus . They concluded that the genus Diopecephalus could be returned to use to distinguish "P". kochi from P. antiquus . They named
19558-405: The teeth mostly became conical. Front teeth were often longer, forming a "prey grab" in transversely expanded jaw tips, but size and position were very variable among species. With the derived Pterodactyloidea , the skulls became even more elongated, sometimes surpassing the combined neck and torso in length. This was caused by a stretching and fusion of the front snout bone, the premaxilla , with
19712-480: The term to the genus Pterodactylus or members of the Pterodactyloidea . 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 influential during the 19th century. In 1843, Edward Newman thought pterosaurs were flying marsupials . Ironically, as
19866-399: The time special French political commissars systematically seized art treasures and objects of scientific interest. Hermann sent Cuvier a letter containing his own interpretation of the specimen (though he had not examined it personally), which he believed to be a mammal , including the first known life restoration of a pterosaur. Hermann restored the animal with wing membranes extending from
20020-408: The time the next generation hatched, creating distinct 'clumps' of similarly-sized and aged individuals in the fossil record. The smallest size class probably consisted of individuals that had just begun to fly and were less than one year old. The second year class represents individuals one to two years old, and the rare third year class is composed of specimens over two years old. This growth pattern
20174-453: The time, distinct from previously known contemporary integumentary structures and more similar to those reported from mammalian hair and avian feathers. The feather fossils obtained from this specimen also suggest the presence of Stage IIIa feathers, a new discovery that indicates more complex feather structures were present in pterosaurs. The study describing this specimen further clarifies the timeline of avian feather evolution and suggests that
20328-422: 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. 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
20482-640: The type species of Rhamphorhynchus , which resulted in a new combination called Rhamphorhynchus longicaudus . The species R. münsteri was later changed to R. muensteri by Lydekker in 1888, due to the ICZN rule that prohibits non-standard Latin characters, such as ü , in scientific names. Beginning in 1846, many pterosaur specimens were found near the village of Burham in Kent , England by British paleontologists James Scott Bowerbank and Sir Richard Owen . Bowerbank had assigned fossil remains to two new species;
20636-416: The upper jawbone, the maxilla . Unlike most archosaurs , the nasal and antorbital openings of pterodactyloid pterosaurs merged into a single large opening, called the nasoantorbital fenestra . This feature likely evolved to lighten the skull for flight. In contrast, the bones behind the eye socket contracted and rotated, strongly inclining the rear skull and bringing the jaw joint forward. The braincase
20790-416: 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
20944-402: The wing. It faced sideways and somewhat upwards. 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
21098-475: 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. 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. The pelvis of pterosaurs
21252-489: The years since its discovery. In the first half of the 19th century any new pterosaur species would be named Pterodactylus , which thus became a " wastebasket taxon ". Even after clearly different forms had later been given their own generic name, new species would be created from the very productive sites, throughout Europe and North America, often based on only slightly different material. The earliest reassignments of pterosaur species to Pterodactylus started in 1825, with
21406-479: Was a generalist carnivore that probably fed on a variety of invertebrates and vertebrates. Like all pterosaurs, Pterodactylus had wings formed by a skin and muscle membrane stretching from its elongated fourth finger to its hind limbs. It was supported internally by collagen fibres and externally by keratinous ridges. Pterodactylus was a small pterosaur compared to other famous genera such as Pteranodon and Quetzalcoatlus , and it also lived earlier, during
21560-515: Was a relatively small pterosaur, with an estimated adult wingspan of about 1.04 meters (3 ft 5 in), based on the only known adult specimen, which is represented by an isolated skull. Other "species" were once thought to have been smaller. However, these smaller specimens have been shown to represent juveniles of Pterodactylus , as well as its contemporary relatives including Ctenochasma , Germanodactylus , Aurorazhdarcho , Gnathosaurus , and hypothetically Aerodactylus if this genus
21714-401: 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. The necks of pterosaurs were relatively long and straight. In pterodactyloids,
21868-467: Was convinced the animal was a reptile . The specimen had not in fact been seized by the French. Rather, in 1802, following the death of Charles Theodore, it was brought to Munich , where Baron Johann Paul Carl von Moll had obtained a general exemption of confiscation for the Bavarian collections. Cuvier asked von Moll to study the fossil but was informed it could not be found. In 1809 Cuvier published
22022-451: Was highly modified from their reptilian ancestors by the adaptation to flight. Pterosaur bones were hollow and air-filled, like those of birds . This provided a higher muscle attachment surface for a given skeletal weight. The bone walls were often paper-thin. They had a large and keeled breastbone for flight muscles and an enlarged brain able to coordinate complex flying behaviour. Pterosaur skeletons often show considerable fusion. In
22176-403: Was not mentioned in a catalogue of the collection taken in 1767, so it must have been acquired at some point between that date and its 1784 description by Collini. This makes it potentially the earliest documented pterosaur find; the "Pester Exemplar" of the genus Aurorazhdarcho was described in 1779 and possibly discovered earlier than the Mannheim specimen, but it was at first considered to be
22330-477: Was not only common in Europe, but also in North America; paleontologists such as Othniel Charles Marsh in 1871 for example, described several toothless pterosaur specimens, which were accompanied by teeth that belonged to the fish Xiphactinus , which Marsh assumed that these teeth belonged to the pterosaur specimens he found, since all pterosaurs discovered at the time had teeth. He then assigned these specimens to
22484-407: 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
22638-485: Was reassigned to a genus called Lonchodraco ('lance dragon'), which resulted in a new combination called L. giganteus , and P. cuvieri was reassigned to the new genus Cimoliopterus ('chalk wing'), creating C. cuvieri . Back in 1859, Owen had found remains the front part of a snout in the Cambridge Greensand , and assigned it into the species Pterodactylus segwickii ; in honor of Adam Sedgwick ,
22792-643: Was relatively large for reptiles. In some cases, fossilized keratinous beak tissue has been preserved, though in toothed forms, the beak is small and restricted to the jaw tips and does not involve the teeth. Some advanced beaked forms were toothless, such as the Pteranodontidae and Azhdarchidae , and had larger, more extensive, and more bird-like beaks. Some groups had specialised tooth forms. The Istiodactylidae had recurved teeth for eating meat. Ctenochasmatidae used combs of numerous needle-like teeth for filter feeding; Pterodaustro could have over
22946-403: 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
23100-482: Was that if such creatures were still alive, only the sea was a credible habitat; Collini suggested it might be a swimming animal that used its long front limbs as paddles. A few scientists continued to support the aquatic interpretation even until 1830, when German zoologist Johann Georg Wagler suggested that Pterodactylus used its wings as flippers and was affiliated with Ichthyosauria and Plesiosauria . In 1800, Johann Hermann first suggested that it represented
23254-410: Was that year described by von Sömmerring as Ornithocephalus brevirostris , named for its short snout, now understood to be a juvenile character (this specimen is now thought to represent a juvenile specimen of a different genus, probably Ctenochasma ). He provided a restoration of the skeleton, the first one published for any pterosaur. This restoration was very inaccurate, von Sömmerring mistaking
23408-456: Was the deepest point of the thorax. Clavicles or interclavicles were completely absent. 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
23562-400: Was the first scientist to describe a pterosaur fossil. At that time the concepts of evolution and extinction were imperfectly developed. The bizarre build of the pterosaur was shocking, as it could not clearly be assigned to any existing animal group. The discovery of pterosaurs would thus play an important role in the progress of modern paleontology and geology. Scientific opinion at the time
23716-579: Was toothless, this made Marsh create the genus Pteranodon (lit. 'toothless wing'), and therefore reassigned all the American pterosaur species, including the ones that he named, from Pterodactylus to Pteranodon . Later, in the 1980s, subsequent revisions by Peter Wellnhofer had reduced the number of recognized species to about half a dozen. Many species assigned to Pterodactylus had been based on juvenile specimens, and subsequently been recognized as immature individuals of other species or genera. By
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