This timeline of pterosaur research is a chronologically ordered list of important fossil discoveries, controversies of interpretation, and taxonomic revisions of pterosaurs, the famed flying reptiles of the Mesozoic era. Although pterosaurs went extinct millions of years before humans evolved, humans have coexisted with pterosaur fossils for millennia. Before the development of paleontology as a formal science, these remains would have been interpreted through a mythological lens. Myths about thunderbirds told by the Native Americans of the modern Western United States may have been influenced by observations of Pteranodon fossils. These thunderbirds were said to have warred with water monsters, which agrees well with the co-occurrence of Pteranodon and the ancient marine reptiles of the seaway over which it flew.^[1]

The formal study of pterosaurs began in the late 18th century when naturalist Cosimo Alessandro Collini of Mannheim, Germany published a description of an unusual animal with long arms, each bearing an elongated finger. He recognized that this long finger could support a membrane like that of a bat wing, but because the unnamed creature was found in deposits that preserve marine life he concluded that these strange arms were used as flippers.^[2] The creature was restudied again in the very early 19th century by French anatomist Georges Cuvier, who recognized both that the creature was a reptile and that its "flippers" were wings. He called the creature the Ptero-dactyle, a name since revised to Pterodactylus.^[3]

Although Cuvier's interpretation later became the consensus, it was just one of many early interpretations of the creature and its relatives, including that they were bats, strange birds, or the primordial handiwork of Satan himself.^[4] Similar animals like the long-tailed Rhamphorhynchus and Gnathosaurus were soon discovered around Europe and it became obvious that earth was once home to a diverse group of flying reptiles.^[5] The British anatomist Sir Richard Owen dubbed this vanished order the Pterosauria. Soon after, he described Britain's own first pterosaur, Dimorphodon.^[6] Later in the 19th century pterosaurs were discovered in North America as well, the first of which was a spectacular animal named Pteranodon by paleontologist Othniel Charles Marsh.^[7]

Various aspects of pterosaur biology invited controversy from the beginning. Samuel Thomas von Soemmering ignited a multi-century debate over how pterosaurs walked on the ground by suggesting they crawled on all fours like bats. August Quenstedt, by contrast, argued that they walked on their hind limbs.^[8] In the early 20th century, Hankin and Watson in the first major study of pterosaur flight biomechanics concluded that on the ground these reptiles were altogether helpless and could only scoot along on their stomachs like penguins.^[9] The debate gained steam in 1957 when William Stokes reported unusual tracks left by a four-footed animal he suspected was a pterosaur walking along the ground.^[10] In 1984, Kevin Padian, who had recently argued that pterosaurs walked on their hind legs, dismissed Stokes's tracks as those of a crocodilian.^[11] However, in the mid-1990s, Jean-Michel Mazin and others reported that fossil footprints in Crayssac, France were similar to those reported by Stokes from the US. Mazin's tracks were more obviously pterosaurian in origin and settled the debate in favor of pterosaurs walking on all fours.^[10]

Pterosaur paleontology continues to progress into the 21st century. In fact, according to David Hone the early 21st century has seen more progress in pterosaur paleontology than in "the preceding two centuries" combined. He compared this transformative period in pterosaur paleontology to the Dinosaur Renaissance of the 1970s.^[12] He also observed that roughly one-third of known pterosaurs were discovered during this brief interval.^[13] One of the most notable of these was Darwinopterus, whose body resembled the more primitive long-tailed "rhamphorynchoids", while its skull resembled those of the more advanced short-tailed pterodactyloids.^[14] These traits establish the species as an important transitional form, documenting one of the most important phases of pterosaur evolution.^[15] Another important new species is Faxinalipterus minima, which might well be the world's oldest pterosaur.^[16] The first confirmed pterosaur eggs were also reported from China during the early 21st century.^[17]

Prescientific

The Cheyenne people of Nebraska believed in mythical thunderbirds and water monsters that were in endless conflict with each other. The thunderbirds were said to resemble giant eagles and killed both people and animals with arrows made of lightning. People occasionally discovered stony arrowheads thought to come from the thunderbirds' arrows. According to folklorist Adrienne Mayor, these supposed arrowheads were likely fossil belemnites, which were compared to missiles by other indigenous American cultures, like the Zuni people.^[18]

The fossils of the Niobrara chalk may have been influential on these stories. The pterosaur Pteranodon and marine reptiles like mosasaurs are preserved in Niobrara Chalk deposits and associated remains may have been interpreted as evidence for antagonism between immense flying animals and serpentine aquatic reptiles. Fossils of the large toothed diving bird Hesperornis are also found in the Niobrara chalk, sometimes preserved inside specimens of large predatory marine reptiles. Observations of similar fossils in the past may have been seen as further evidence for thunderbird-water monster conflict.^[19]

• The Sioux people of South Dakota believed that the first creatures in creation were the insects and reptiles, who were ruled by the Water Monster Unktehi. Reptiles were very diverse and came in all shapes and sizes, but they became violent and bloodthirsty until they were petrified by lightning sent by the Thunder Birds. The physical bodies of the Thunder Beings killed by the lightning, including Unktehi, also ended up being buried. The Sioux believe that earth has a history of four distinct ages. These events occurred during the Age of Rock. This portrayal of the Thunder Birds may also have been influenced by associations of fossils of Pteranodon with marine reptiles of the same age in the western United States.^[20]

18th century

1780s

1784

• Cosimo Alessandro Collini, keeper of the natural history collections of Mannheim, reported the skeleton of an unusual animal to the scientific literature. It had strange arms that could have supported a membrane like that of a bat's wing, yet it was preserved in rocks characterized by fossil of marine life. Based on these associations, he tentatively concluded that the animal was aquatic.^[2]

19th century

1800s

1801

• French anatomist Georges Cuvier restudied Collini's bizarre fossil, based on his published illustration. He reinterpreted its forelimbs as wings and deemed it a flying reptile.^[3]

1802

• The strange fossil described by Collini was moved from Mannheim to Munich.^[21]

1807

• German anatomist Johann Friedrich Blumenbach interpreted Collini's animal as a bird, specifically, as some kind of waterfowl.^[21]

1809

• Cuvier named the flying reptile reported by Collini Ptéro-Dactyle.^[21]

1810s

1812

• In contrast to Cuvier and Blumenbach, Samuel Thomas von Soemmering interpreted Collini's fossil as a mammal.^[22] Specifically, he interpreted it as an unusual bat, which morphologically linked mammals with birds. He named this strange creature Ornithocephalus. Soemmering may have interpreted this series of forms in an evolutionary sense, following the early evolutionary ideas of Jean-Baptiste de Lamarck. Soemmering's analysis of the specimen was blemished by anatomical errors, like the misidentifications of bones. Soemmering agreed with Cuvier that the creature was a flying insectivore, however.^[21] Soemmering argued that pterosaurs walked on all fours like bats when on the ground. His advocacy for this interpretation of pterosaur terrestrial gait has been regarded as the beginning of a multi-century debate on the subject.^[8]

1817

• Soemmering reported the discovery of a second pterosaur specimen. This second specimen was smaller than the first, with a 25 cm wingspan, and possessed a shorter snout. These traits mislead Soemmering into greater confidence in his interpretation of pterosaurs as bats. This specimen reminded him of the parti-colored bat.^[21]

1819

• Cuvier renames Ptéro-Dactyle into the current generic name Pterodactylus, but assigning P. longirostris (now considered a synonym of the type species P. antiquus) as the type species of the genus.^[23]

1820s

1824

• Cuvier reiterated his previous conclusions that the Ptero-Dactyle was a reptile that flew with membranous wings. He also advanced novel speculations about its paleobiology, like that it used the claws on its wings to climb trees and "crawled" quadrupedally when not in flight rather than walking on its hind limbs.^[21]

c. 1825

• Paleontologist Georg Graf Munster discovered an unusual skull. He sent the fossil to Soemmering, who thought it belonged to an ancient sea bird. He also sent a cast of the skull to August Georg Goldfuss, who recognized it as a pterosaur. Goldfuss described the new species Pterodactylus muensteri based on the specimen.^[21]

1827

• Gideon Mantell discovered some fossil bones in the Wealden beds (Wessex Formation) of Sussex, England that he believed were the remains of ancient birds.^[24]
• Mantell recognized that his "bird" bones were actually pterosaur fossils and reported his findings to the scientific literature. These were the first Cretaceous pterosaur fossils ever described.^[24]

1829

• William Buckland described the new species Pterodactylus macronyx based on fossil remains discovered by Mary Anning in December the previous year from the Lias Group of the Dorset Coast of England. These were the first scientifically documented pterosaurs to be discovered outside of the Solnhofen lithographic limestone.^[25]

1830s

1830

• Carl von Theodori described the new species Pterodactylus banthensis from Franconia.^[25]
• Georg Wagler argued that pterosaurs represented a distinct class of aquatic vertebrates that he called Gryphi. Like Collini, Wagler thought that pterosaurs swam underwater using their forelimbs as flippers.^[26]

1831

• August Goldfuss depicted pterosaurs as flying reptiles that used their wing claws to climb cliffs.^[26] He hypothesized that on land, they would have had to travel on all fours.^[8] He also suggested that they may have been covered in hair.^[27]

1832

• von Meyer described the new genus Rhamphocephalus.^[28]

1833

• von Meyer described the new genus and species Gnathosaurus subulatus.^[29]

1834

• Johann Jacob Kaup first referred to pterosaurs by that name.^[30]

1836

• William Buckland depicted pterosaurs as cliff-climbing winged reptiles in a manner heavily influenced by Goldfuss.^[26]

1839

• Graf Munster received a complete skeleton of "Pterodactylus" munsteri which revealed the presence of a long bony tail in this species.^[31]

1840s

1840

• Thomas Hawkins published The Book of the Great Sea-Dragons, wherein he suggested that the great reptiles of the Mesozoic were created by the devil.^[32] He described pterosaurs as "an engrafted-by-Evil stock" and depicted them as bat-like scavengers that combed the ancient seashore.^[33]

1842

• Sir Richard Owen formally named the order Pterosauria.^[30]

1843

• Edward Newman interpreted pterosaurs as mammals in a similar fashion to Soemmering. However, Newman specifically considered pterosaurs to be carnivorous flying marsupials.^[26]

1847

• von Meyer erected the genus Rhamphorhynchus.^[34]

1850s

1851

• Two pterosaurs sculpted by Benjamin Waterhouse Hawkins were put on display in England's Crystal Palace. These were the first three-dimensional life-size restorations of pterosaurs.^[26]

1855

• von Meyer described the new genus and species Ctenochasma roemeri.^[35]

1855

• August Quenstedt described the species Pterodactylus suevicus from the Nusplingen lithographic limestone.^[25] In this publication he argued that pterosaurs would have walked bipedally on the ground.^[8]

1856

• Albert Oppel reported the discovery of a pterosaur lower jaw from the Posidonia shales of Holzmaden. This was the first pterosaur specimen to be reported from these deposits, which would go on to produce many pterosaur fossils of exceptional quality.^[25]

1859

• Sir Richard Owen erected the new genus Dimorphodon for the species "Pterodactylus" macronyx.^[25]
• Buckman described a clutch of 4.5 cm long oval-shaped fossil eggs from Middle Jurassic marine rock in the United Kingdom. He erected the new oogenus and oospecies Oolithes bathonicae for them, the first time fossil eggs had been given their own unique taxonomic name.^[36]

1850s – 1860s

1859 – 1860

• Meyer described 40 specimens of Pterodactylus. Among these specimens he reported more than 20 species. Most of these species are not recognized as distinct today and generally represent the misguided application of new names to members of known species at different ages. One species was not even a pterosaur; the "Pterodactylus" crassipes type specimen would later be recognized as the "Haarlem" specimen of Archaeopteryx.^[21] He also reported the presence of pterosaurs in the lithographic limestone of Cerin, France.^[30]

1860s

1860

• Andreas Wagner erected the new genus Dorygnathus for the species Pterodactylus banthensis.^[25]

1861

• Wagner described the new genus Scaphognathus for the species Pterodactylus crassirostris.^[21]

1862

• Oppel described the new species Ctenochasma gracile.^[35] He also interpreted some tracks from the Solnhofen lithographic limestone as pterosaur tracks.^[37] This was the first report of potential pterosaur tracks to the scientific literature.^[38]

1863

• The book "La Terre avant Le Deluge" by Louis Figuier was published. It included an early restoration of a Rhamphorhynchus walking across the ground on all fours. This depiction was based on fossil footprints from the Solnhofen limestone attributed to the taxon.^[26]

1869

• Harry Govier Seeley described the new genus Ornithocheirus.^[39]

1870s

1870

• Seeley described the new genus Cycnorhamphus and the new species Ornithocheirus huxleyi.^[40]
• Seeley argued that pterosaurs represented the evolutionary transitional form between reptiles and birds, distinguished from the traditional reptiles by a warm-blooded metabolism as well as bird like anatomy, physiology and terrestrial gait. The claim ignited a "violent controversy" with the Owen due to his more traditional perspectives on pterosaurs and his hostility to evolutionary theory.^[41]
• Othniel Charles Marsh described the new genus and species Laornis edvardsianus.^[42]

November, late

• Marsh's Yale Peabody Museum crews visited Kansas.^[43] Marsh discovered the first Pteranodon wing bones.^[44] These were the first scientifically described pterosaur fossils from North America.^[7]

November – December 31st

• Sir Richard Owen expressed astonishment at the North American discovery of pterosaurs exceeding the size of warm-blooded birds and mammals, given his interpretation of the group as typical cold blooded reptiles.^[41]

1871

• Seeley described the new genus Ornithostoma for a toothless pterosaur from the Cambridge Greensand.^[45] He also described the new genus Diopecephalus.^[46] He observed that pterosaurs had large, birdlike brains.^[47]
• Marsh discovered further Pteranodon remains.^[48]
• Marsh discovered the type specimen of Pterodactylus velox.^[49]
• Marsh described the new species Pterodactlus oweni based on some Pteranodon wing bones. He attributed teeth to the species.^[48]
• Edward Drinker Cope discovered at least two Pteranodon specimens.^[49]
• William Carruthers described two new oospecies of Oolithes and suggested that these fossil eggs were laid by pterosaurs.^[50]

1872

March 7th

• Marsh described the species Pterodactylus ingens. He continued to attribute teeth to Pteranodon.^[48] Marsh realized that his species name Pterodactylus oweni had already been used, so renamed it Pterodactylus occidentalis.^[49] Marsh also named Pterodactylus velox.^[49]

March 12th

• Cope also attributed teeth to Pteranodon.^[48]
• Cope described the new species Ornithochirus umbrosus and O. harpyia.^[49]
• Cope published an additional paper on his "Ornithochirus" species. This paper was read before the American Philosophical Society.^[51]

March 12th – December 31st

• An anonymous review synonymized Cope's Ornithochirus umbrosus and O. harpyia with Marsh's Pterodactylus ingens and Pterodactylus occidentalis, respectively.^[51]

1874

• Cope acknowledged the validity of Marsh's Pterodactylus ingens and P. occidentalis, but continued to insist that his O. umbrosus was a valid species, although he came to refer it to Pterodactylus. This paper included the first illustrations of Pteranodon wing bones.^[52]
• A pterosaur fossil bearing an impression of the wing membrane was discovered. This was the first physical evidence of the structure which had previously been inferred purely from skeletal anatomy.^[21]
• Owen described the new genus Coloborhynchus.^[53]

1875

• Cope argued that his Ornithochirus species supposedly synonymous with Marsh's Pterodactylus species actually had priority because while Marsh's description was published first, Cope's would have been if not for delays caused by a fire at the publisher.^[52]
• Mudge discovered the type specimen of Pteranodon comptus.^[54]
• Seeley described the new genus Doratorhynchus.^[46]

1876

May

• Samuel Wendell Williston discovered the type specimen of Pteranodon.^[45]

May – December 31st

• Marsh described the new genus and species Pteranodon longiceps and referred the toothless American pterosaurs to the new order Pteranodontia.^[45] He also described the new species Pteranodon comptus and P. gracilis.^[55]
• Marsh published additional research on American pterosaurs. He acknowledged that his referral of teeth to the genus was an error.^[45] Marsh described the new genus Nyctosaurus for "Pteranodon" gracilis.^[54]
• Williston helped excavate the Pteranodon specimen YPM 1177.^[56]
• Brous and Williston helped excavate the Pteranodon specimen YPM 2473.^[57]

1877

• Charles H. Sternberg discovered AMNH 5098, a Pteranodon specimen with fossil fish and crustacean remains preserved in its throat pouch in Kansas.^[56]

1880s

1881

• Marsh described the new genus Dermodactylus as well as the new genus and species Laopteryx priscus.^[58]
• Marsh renamed Nyctosaurus Nyctodactylus under the mistaken belief that the latter genus was preoccupied.^[54]

1882

• Marsh described the new species Rhamphorhynchus phyllurus from the Solnhofen lithographic limestone. The type specimen displayed exquisitely preserved impressions of the animal's wing membranes as well as a diamond shaped fin at the end of the tail.^[59] Marsh thought that this fin was oriented vertically because it was slightly asymmetrical and could be used help the pterosaur steer as it flew.^[60] However, Marsh's hypothesis regarding the orientation of the tail vane would later become controversial.^[61]
• Karl Alfred von Zittel described a fossil Rhamphorhynchus wing from the Solnhofen lithographic limestone that also preserved lifelike impressions of the wing membrane. He observed that the wing of Rhamphorhynchus was strengthened by fibrous tissue.^[59] Based on this specimen, Zittel concluded that in life Rhamphorhynchus had relatively narrow wings, whereas Marsh thought the wings were much wider. The wing Zittel studied has been known as the "Zittel wing" in his honor ever since.^[60]

1884

• Marsh reported that by this time the Yale Peabody Museum curated over 600 Pteranodon specimens. He also published more information about the skull of the Pteranodon type specimen and illustrated it. Marsh suspected that Pteranodon lacked a sclerotic ring, since one was absent in even well-preserved specimens.^[54]

1886

• Francesco Bassani described the new genus and species Tribelesodon longobardicus from Besano, Italy as the first known Triassic pterosaur.^[62]

1887

• Seeley described the new genus and species Ornithodesmus cluniculus.^[63]

1888

• Richard Lydekker described the new genus Ptenodracon for the second pterosaur specimen to be discovered. This genus is now regarded as a junior synonym of Pterodactylus because the traits supposedly "Ptenodracon" instead indicate that the specimen was a juvenile.^[21]
• Newton reported the discovery of an endocast of a pterosaur brain in the Lias of Whitby, England.^[61] The find revealed that pterosaur brains were more like modern birds than reptiles.^[64]

1890s

1891

• Wiliston published what paleontologist Michael Everhart called the first complete description of Pteranodon this year.^[65] Notable observations in this publication include the discovery of a sclerotic ring in this taxon.^[54] Williston also found a coprolite containing tiny, indeterminate bone fragments preserved in one Pteranodon's pelvic area.^[56] Williston also argued that previous estimates of Pteranodon's wingspan were exaggerated and that the maximum wingspan of the genus was just short of 20 feet.^[66]

1892

• Williston disputed the length of Pteranodon's crest in Marsh's 1884 reconstruction.^[54]
• Williston published a redescription of the skull of Pteranodon based on a more recently discovered and better preserved specimen, KUVP 2212.^[56] He also criticized the length of Pteranodon's crest in Marsh's 1884 reconstruction of the specimen YPM 1177 as being too speculative given the quality of its preservation.^[67] Williston speculated that Pteranodon-like fossils would be one day discovered in Europe, and that in this case Pteranodon was probably a junior synonym of Ornithostoma.^[68]

In this paper Williston also described a new, relatively complete Nyctosaurus specimen. He noted that the only published trait distinguishing the genus from Pterodactylus was an absence of teeth and recommended synonymizing these two genera if "Nyctosaurus" teeth were ever found.^[57]

1893

• Williston argued that Pteranodon was a junior synonym of Ornithostoma. He praised Cope for recognizing these affinities, while lambasting Marsh for being unable to do so despite having a larger number of specimens. According to Everhart, Williston's criticism of Marsh may have been motivated by mistreatment at his hands while he worked for him.^[68]

1895

• Williston published a description of the lower jaw of Pteranodon. This was the first such specimen not to be "crushed from side to side".^[68]

1896

• Williston published a description of another Pteranodon skull and synonymized the genus with Ornithostoma.^[68]

1897

• Williston reported that Seeley was also unable to find any trait in Pteranodon distinguishing it from Ornithostoma.^[68]

20th century

1900s

1901

• Felix Plieninger formally divided the pterosaurs into two suborders, the long-tailed Rhamphorhynchoidea and the short-tailed Pterodactyloids.^[30]
• Seeley published Dragons of the Air. This was the first "serious boo" about pterosaurs.^[69] In it he restored pterosaurs with the wing membrane attached to the hindlimb.^[70]

1902

• Williston published further anatomical description of Nyctosaurus based on a recently discovered well-preserved specimen now catalogued as FMNH 25026.^[68] He estimated its live weight as less than five pounds. He interpreted the skull as completely lacking a crest.^[71]
• Williston published another paper about FMNH 25026 in which he described the skull in detail and photographed it.^[71]
• Williston published a popular article about pterosaurs for Popular Science Monthly.^[71] In this article, Williston restored pterosaurs with the wing membrane attached to the hindlimb.^[70] According to Everhart, by this point Williston had "largely given up" in his attempts to synonymize Pteranodon with Ornithostoma.^[71]

1903

• Williston observed that the generic name Nyctosaurus was not actually preoccupied. He speculated that Marsh probably came to believe that it was preoccupied because of the existence of a higher order taxon called Nyctisauria. Since Nyctosaurus was not preoccupied, Williston reclassified "Nyctodactylus" back to the original genus.^[54] Williston also described the new genus and species Apatomerus mirus for a partial pterosaur femur from the Kiowa Shale of Kansas. This specimen is now catalogued as KUVP 1198.^[72] This paper contained a notable error wherein Williston claimed that Pteranodon lacked a fibula.^[73]
• George Francis Eaton published a paper defending Marsh's research on Pteranodon against Williston.^[54]

1904

• Eaton published a paper defending Marsh's research on Pteranodon against Williston.^[54] Some of Eaton's criticisms have since come under fire. For instance, Everhart has noted that Eaton's criticism of Williston for reporting a sclerotic ring in Pteranodon rather than Nyctosaurus ignored the fact that Eaton had found sclerotic rings in both genera.^[74]
• Williston published a paper on pterosaur fingers.^[75]

1907

• Plieninger described the species Rhamphorhynchus kokeni from the Nusplingen lithographic limestone.^[25]
• Woodward described the new genus and species Scleromochlus taylori.^[76]

1910s

1910

• Eaton published his doctoral dissertation on the osteology of Pteranodon.^[75] This publication was the most significant work about Pteranodon as well as large pterosaurs generally for many decades afterward.^[77] In this monograph, he restored pterosaurs with the wing membrane attached to the hindlimb.^[70] He concurred with earlier work by Marsh and Williston that Pteranodon had a short tail.^[78] According to Everhart, Eaton toned down his former stridently defensive attitude toward Marsh and warmed up somewhat to Williston's work.^[75] He noted that the supposed wing bones of Pteranodon comptus were actually Nyctosaurus tibiae and that P. ingens and P. occidentalis were only distinguishable by their sizes.^[75] Everhart also noted that Eaton actually followed some of Williston's work "too closeley" and repeated Williston's erroneous claim that Pteranodon lacked a fibula.^[73]
• Williston finally confirmed the presence of a fibula in Pteranodon, correcting his previous error that mislead Eaton.^[73]

1911

• Williston published a favorable review of Eaton's dissertation and conceded that his earlier criticism of Marsh's Pteranodon skull reconstruction was baseless.^[73]
• Williston published a restoration of Nyctosaurus. This was his last paper on pterosaurs.^[75]

1913

• Hooley described the new subfamily Ptenodraconinae based on Lydekker's misguided genus Ptenodracon.^[21]

1914

• Hooley described the new genus Lonchodectes.^[79]
• Hankin and Watson published the first study of the aerodynamics of pterosaur flight. They concluded that Pteranodon spent much more time soaring than actively flapping.^[80] On the ground, however, Hankin and Watson argued that pterosaurs would have been "completely helpless" and could only move about by "pushing themselves along, after the manner of penguins."^[8]

1918

• Arthaber described the new genus and species Parapsicephalus.^[81]

1920s

1920

• Wiman published a description of the fossils purchased by the Paleontological Museum in Uppsala, Sweden from C. H. Sternberg, which included Pteranodon fossils. He confirmed the presence of a fibula in some of the specimens.^[73]

1921

• Friedrich von Huene described the new genus Rhabdopelix.^[28]

1922

• Nopcsa described in detail and reconstructed Bassani's purported Triassic pterosaur from Italy, Tribelesodon.^[62]

1923

• Döderlein described the new genus Anurognathus.^[82]

1925

• Abel argued that pterosaurs would have to walk on all fours when not in the air, like a modern bat.^[8]

1927

• Broili described possible fossil evidence for a hair like body covering in pterosaurs from Germany.^[27]

1928

• Strand described the new genus Campylognathoides.^[83]
• Nessov described the new genus Bennettazhia.^[84]

1929

• Bernhard Peyer discovered that the purported Triassic pterosaur Tribelesodon was actually a juvenile Tanystrophaeus, whose long neck vertebrae were mistaken for a wing-finger.^[62]

1930s

1937

• Koh described the new species Rhamphorhynchus intermedius.^[34]

1938

• Kenneth Caster conclusively demonstrated that unusual fossil tracks from the Solnhofen lithographic limestone variously attributed to creatures like Archaeopteryx, little dinosaurs, or pterosaurs were actually made by horseshoe crabs, as specimens had been found literally "dead in their tracks".^[37]

1939

• Broili described the new genus and species Belonochasma aenigmaticum.^[84]

1940s

1940

• Caster reported finding a dead horseshoe crab at the end of a type of fossil trackway once attributed to long-tailed pterosaurs.^[62]

1943

• Brown reported a Pteranodon specimen with the remains of two fish species and a crustacean preserved where its throat pouch would have been in life.^[56]

1948

• Riabinin described the new genus and species Batrachognathus volans.^[84]

1950s

1952

• George F. Sternberg discovered an unusual Pteranodon skull in Kansas, FHSM VP-339 that would later serve as the type specimen of the species Pteranodon sternbergi.^[73]

1954

• Ambroggi and Lapparent described the new ichnogenus and ichnospecies Agadirichnus elegans.^[85] They attributed these tracks to a lizard.^[86]

1956

• Sternberg discovered another specimen of P. sternbergi near WaKeeney, Kansas which is now catalogued as FHSM VP-184. This specimen lacked a skull and was smaller than the type. In life it would have had a roughly 12.5 foot wingspan.^[87]

1957

• Stokes described the new ichnogenus and species Pteraichnus saltwashensis from the Late Jurassic Morrison Formation of Utah, USA, interpreting them as pterosaur tracks.^[10] Stokes reported the presence of an impression left by the putative pterosaurian trackmaker's wing finger, although this claim is probably mistaken.^[38] If his identification of these tracks was correct, it would mean that pterosaurs walked on all fours.^[10]
• Eric von Holst published an experimental study of Rhamphorhynchus flight biomechanics that utilized a flapping scale model. Because the model could only fly when its tail vane was oriented horizontally rather than vertically, von Holst concluded that Marsh's original reconstruction was erroneous.^[61]

1958

• Sternberg and Walker reported the second P. sternbergi specimen to the scientific literature.^[87]
• Kuhn accepted the pterosaurian interpretation of Pteraichnus.^[38]

1960s

1962

• George Sternberg discovered a nearly complete Nyctosaurus specimen near Elkader, Kansas which is now catalogued as FHSM VP-2148.^[87]

1963

• Delair described the new ichnogenus and ichnospecies Purbeckopus pentadactylus.^[88] Delair did not recognize the tracks as pterosaurian.^[38]

1964

• Bonner reported the Elkader Nyctosaurus discovered by Sternberg to the scientific literature and described the new species N. sternbergi based on it.^[87]
• C. C. Young described the new genus Dsungaripterus from Early Cretaceous rocks in China.^[89]
• Young described the new genus Germanodactylus.^[90]
• Kuhn continued to accept the pterosaurian interpretation of Pteraichnus.^[38]

1966

• Harksen described the new species Pteranodon sternbergi. Unlike P. longiceps, this species had a short, wide crest.^[73]

1968

• Peter Wellnhofer published a revision of the taxonomy of Pterodactylus. It was his first publication about pterosaurs.^[91]

1969 Zdroj:https://en.wikipedia.org?pojem=Timeline_of_pterosaur_research
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