Timeline of pterosaur research

Source: Wikipedia, the free encyclopedia.

Life restoration of the first scientifically studied pterosaur, Pterodactylus

This timeline of pterosaur research is a chronologically ordered list of important

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

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.

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

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

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

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
    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

Type specimen of Pterodactylus

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

Portrait of Georges Cuvier, the naturalist who recognized pterosaurs as flying reptiles

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
    waterfowl.[21]

1809

1810s

Samuel Thomas von Soemmerring

1812

  • In contrast to Cuvier and Blumenbach,
    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

Illustration of the holotype skull of Rhamphorhynchus.

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
    Pterodactylus muensteri based on the specimen.[21]

1827

1829

  • Solnhofen lithographic limestone.[25]

1830s

Pterodactylus reconstructed as an aquatic animal
August Goldfuss and William Buckland

1830

  • Pterodactylus banthensis from Franconia.[25]
  • 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

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

The pterosaur Pterodactylus portrayed as the handiwork of Satan in the Book of the Great Sea-Dragons by Thomas Hawkins (1840)

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

  • order Pterosauria.[30]

1843

1847

1850s

1851

1855

  • von Meyer described the new genus and species
    Ctenochasma roemeri.[35]
Illustrated skeletal reconstruction and life restoration of Dimorphodon.

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

  • 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
    unique taxonomic name.[36]

1850s – 1860s

18591860

  • 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 "
    Cerin, France.[30]

1860s

Scaphognathus crassirostris
.

1860

1861

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

1862

  • Oppel described the new species
    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

1870s

Type specimen of Cycnorhamphus
Early restoration of Ornithostoma
Othniel Charles Marsh (left) and his rival Edward Drinker Cope (right)
Type specimen of Coloborhynchus
The short-crested female type specimen of Pteranodon
Life restoration of a male Pteranodon

1870

November, late

November – December 31st

1871

1872

March 7th

  • Marsh described the species
    Pterodactylus velox.[49]

March 12th

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

May – December 31st

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

Illustration of the type specimen and life restoration of Rhamphorhynchus "phyllurus" from an 1882 publication by Othniel Charles Marsh
Ptenodracon" (actually just a juvenile Ctenochasma
)

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

  • Tribelesodon longobardicus from Besano, Italy as the first known Triassic pterosaur.[62]

1887

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

1888

  • 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

Samuel Wendell Williston in 1891

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

The Nyctosaurus specimen FMNH 25026.

1901

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]
Skeletal reconstruction of Scleromochlus

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
    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]
Life restoration of Scleromochlus

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

1910s

1910

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]
Reconstruction of the skull of Lonchodectes
Skull of Parapsicephalus

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
    penguins."[8]

1918

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]
Life restoration of Anurognathus
Life restoration of Campylognathoides

1921

1922

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

1923

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

1929

  • 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

Fossil of a dead horseshoe crab at the end of a type of fossil trackway once attributed to pterosaurs

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

Geosternbergia
) sternbergi

1952

1954

  • Ambroggi and Lapparent described the new
    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

Life restoration of Germanodactylus.
Skull of Dsungaripterus.

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

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

  • Colbert described the new genus and species
    Nesodactylus hesperius.[92]

1970s

Life restoration of Pterodaustro
Sordes pilosus
Fossil skeleton of Eudimorphodon
Skeletal mount of Quetzalcoatlus

1970

  • John Ostrom reported that the type specimen of "Pterodactylus" crassipes actually represented a fourth specimen of Archaeopteryx. This was the fourth specimen referrable to that genus ever discovered.[21]
  • Bonaparte described the new genus and species
    filter-feeding pterosaur known to science.[94]

1971

1973

1974

  • Bramwell and Whitfield re-examined the biomechanics of Pteranodon flight after an extended lull in research on the topic. They estimated that a Pteranodon with a 7 m wingspan would have a mass of about 16 kg. To stay aloft, such a Pteranodon would need to fly at least 6.7 m/s, which is regarded as an "extremely low" minimum speed.[99] Such a load would have allowed it to take off or land "gently". Bramwell and Witfield argued that the biomechanics may have left Pteranodon vulnerable to increasing wind speeds resulting from climate change as the Late Cretaceous proceeded and even tentatively suggested that this may have been the cause of extinction for the genus.[100]
  • Fabre described the new genus
    Gallodactylus.[101]
  • Casamiquela described the new genus and species

1975

  • Stein published an article on the biomechanics of Pteranodon flight.[96] He crafted model Pteranodon wings and tested them in a wind tunnel. He found that Pteranodon was a capable, maneuverable flyer but was best adapted to long distance flights at low velocities.[103] Stein calculated that a large Pteranodon would have to fly at least 10 miles an hour to stay airborne. He concluded that Pteranodon would have had to land on its hind feet because making the front feet ready for landing would collapse the wings, which would no longer be useful for keeping the pterosaur aloft.[78] Stein's conclusions contradicted the previous findings of Bramwell and Whitfield.[96]
  • Lawson described the new genus and species
    Quetzalcoatlus northropi.[104]

1977

1978

1979

  • Kevin Padian published a study on the biomechanics of pterosaur flight.[96]
  • Stokes and Madsen reported additional potential pterosaur tracks from the Navajo Formation.[106]

1980s

1980

  • Brower published an article on the biomechanics of Pteranodon flight. His conclusions contradicted the findings of Bramwell and Whitfield, however.[96]
  • de Buisonjé described the new genus and species
    Santanadactylus brasilensis.[107]

1981

  • Frey and Riess published a study on the biomechanics of pterosaur flight.[96]
  • Galton described the new genus and species
    Comodactylus ostromi.[35]

1982

  • Dong described the new genus and species
    Huanhepterus quingyangensis.[108]
biped
Fossil wing bones of Azhdarcho

1983

  • Frey and Riess published a study on the biomechanics of pterosaur flight.[96]
  • Padian argued that the wing membrane of pterosaurus probably did not attach to the hindlimb and that pterosaurs had narrow wings comparable in proportion to those of modern soaring sea birds.[70] He also argued based on the skeletal anatomy of Dimorphodon that it and other pterosaurs probably walked on their hind legs when not airborne.[8]
  • Padian published an additional paper arguing for bipedal pterosaurs.[38]
  • Brower studied the aerodynamics of Nyctosaurus and Pteranodon by comparing them to hang gliders. He concluded that both were unable to ascend or descend at high speed. He thought that they spent most of their time soaring rather than actively flapping. Brower thought Pteranodon itself to be entirely incapable of flapping flight.[78] His conclusions contradicted the previous findings of Bramwell and Whitfield.[96]
  • He, Yan and Su described the new genus and species
    Angustinaripterus longicephalus.[109]
  • Wild described the new genus and species
  • Leonardi and Borgomanero described the new genus and species

1984

1985

  • Schultz and others argued that the type specimen of Apatomerus mirus was not actually a pterosaur fossils.[75]
  • Aeronautical engineer Alan McCready designed a flying model of Quetzalcoatlus in an experimental attempt to ascertain how a creature of its size could even be capable of flight. However, his results were inconclusive.[77]
  • Campos and Kellner described the new genus and species
    Anhanguera blittersdorffi.[82]

1986

1987

  • Bennett observed that the tail of Pteranodon was longer than generally thought, being at least 19 cm on a Pteranodon with a 7.5 m wingspan. He hypothesized that this lengthier tail could have supported an additional membrane that would have assisted the animal's pitch during flight. However, Bennett has subsequently disavowed the idea that Pteranodon supported a membrane with its tail.[78]
  • Nessov and Borkin described the new genus Arambourgiania.[97]
  • Padian published another paper arguing for bipedal pterosaurs.[38]
  • Conrad and others "assumed" that the ichnogenus Purbeckopus was produced by a crocodylian.[38]
  • Unwin published a paper regarding the debate over the identity of the Pteraichnus trackmaker.[38]
  • Leonardi suggested that the supposed pterosaur footprints Stokes reported from the Navajo Formation in the 1970s were actually produced by the same kind of non-pterosaurian animal that made Batrachopus.[106]
Skull of Tupuxuara

1988

  • Russell estimated that by this time 878 Pteranodon specimens were known.[44]
  • Pennycuick published a study on the biomechanics of pterosaur flight.[96]
  • Wellnhofer studied the range of motion in the hip and hind limbs of three-dimensionally preserved pterosaur fossils from the Crato Formation of Brazil for insight into their terrestrial gait.[8] He concluded that they walked on all fours.[105]
  • Kellner and Campos described the new genus and species
    Tupuxuara longicristatus.[113]

1989

1990s

P. sternbergi
. The grey areas were not preserved in their respective specimens.

1990

  • Michael Everhart discovered his first Pteranodon specimen in Kansas. In life it would have had a roughly 14 foot wingspan.[44]
  • Stewart reported that Nyctosaurus and Pteranodon made their first appearances in the stratigraphic column in the middle of the Smoky Hill Chalk, which dates back to the Santonian.[115]
  • Bennett published a study of the biostratigraphy of Pteranodon. He found that the type specimen of P. sternbergi was discovered in the lower chalk, while the type specimen of P. longiceps was discovered in the upper portion of the chalk.[115]

1991

  • Peter Wellnhofer published The Illustrated Encyclopedia of Pterosaurs. Wellnhofer's book was only the second serious book about pterosaurs ever published.[69] In it, he argued that Othniel Charles Marsh was correct to reconstruct the tail vane of Rhamphorhynchus with a vertical orientation based on its asymmetry and also provided additional evidence for this orientation based on the shape of its tail vertebrae.[61] Wellnhofer also observed that pterosaurs had large, birdlike brains.[47] Wellnhofer also argued in favor of quadrupedal pterosaurs.[105]
  • Chatterjee described the new genus and species
    Protoavis texensis.[116]
Life restoration of Tupuxuara
Life restoration of Zhejiangopterus

1992

  • Bennett published a study of sexual dimorphism in Pteranodon using 400 specimens. He found there to be two size based morphs, a larger form with a larger crest and narrow pelvis and a small form with a small crest and wide pelvis. He concluded that the larger form was male and the smaller form was female. He found that female Pteranodon outnumbered male Pteranodon by 2:1.[115] Bennett argued that since large crests were only associated with one size morph, that it functioned purely as display.[117] He also hypothesized that Pteranodon sternbergi was the direct ancestor of Pteranodon longiceps.[117]
  • Bennett argued that the purported pterosaur footprints reported by Gillette and Thomas were actually produced by crocodylians.[105]
  • Lockley and others reported the presence of Early Cretaceous pterosaur tracks in Spain.[118]

1993

  • Moratalla reported the presence of Early Cretaceous pterosaur tracks in Spain.[118]

1994

Life restoration of Plataleorhynchus

1995

1996

June 1st

  • Pamela Everhart discovered a Pteranodon specimen in Kansas.[44] Pam and her husband Michael partially excavated the specimen and covered the rest of the fossils until a more thorough excavation was possible.[128]

June 29–30th

  • Michael and Pamela Everhart returned to finish excavating the Pteranodon, which had a roughly 17 foot wingspan.[129]
Tapejara imperator

1997

1998

May

  • Chris Bennet referred the 1996 Everhart Pteranodon specimen to the species P. sternbergi.[47]

June

  • Michael and Pamela Everhart returned to the site of their 1996 Pteranodon discovery to search for additional remains of the animal. However, extensive digging only produced one additional bone from the specimen.[47]

1999

21st century

2000s

Life restoration of Nyctosaurus

2000

2001

  • Jenkens and others described the new species
    Eudimorphodon cromptonellus.[130]
  • Chris Bennett published the first monograph-length discussion of Pteranodon in more than 90 years.[77] One of his more notable conclusions was that the reconstructions used by previous researchers to study Pteranodon flight biomechanics were so inaccurate that any conclusions drawn from them were completely invalid.[96]
  • Wang and
    Haopterus gracilis.[135]
  • Howse, Milner, and Martill described the new genus Istiodactylus.[136]
  • Dalla Vecchia and others reported the discovery of pterosaur fossils in Lebanon.[17]
  • Paleontologists gathered at Toulouse for a scientific conference dedicated solely to pterosaurs.[69]
  • Fuentes Vidarte published research on Early Cretaceous pterosaur tracks in Spain.[137]
  • Mazin and others reported the existence of fossil pterosaur tracks preserving the impression of a fifth toe. This suggests that the trackmaker was a primitive long-tailed pterosaur.[138]
  • Garcia Ramos and others published research on exceptionally well-preserved Late Jurassic pterosaur tracks in Asturias, Spain.[134]
  • Lockley and others argued that there were two different types of fossil footprint assemblages that include Pteraichnus that differed by the type of a rock they were preserved in. One type of Pteraichnus-bearing track assemblage is associated with carbonate rocks, and the other with clastic rocks.[139]
3-dimensional restoration of Hatzegopteryx

2002

2003

Life restoration of Ludodactylus.
Fossil skeleton of Sinopterus.
Skeletal reconstruction of a pterosaur being bitten by the spinosaur Irritator

2004

  • Chiappe and others reported the first confirmed pterosaur eggs to the scientific literature.[17]
  • Ji and others, in the same issue of Nature as Chiappe and his colleagues, reported additional pterosaur egg fossils.[17]
  • Wang and Zhou reported the discovery of an Early Cretaceous fossilized pterosaur embryo still preserved inside the egg.[117]
  • Peters described the new genus and species Avgodectes pseudembryon.[159]
  • Gasparini, Fernández, and de la Fuente described the new genus and species
    Cacibupteryx caribensis.[160]

May

  • Michael Everhart examined the Apatomerus type specimen and deterimened that it was not a pterosaur fossil.[75]

July

  • Everhart discovered a bone similar in the collections of Kansas University that was similar to the Apatomerus type specimen. This bone was associated with plesiosaur vertebrae, thus revealing the true identity of Apatomerus.[75]

2005

Life restoration of Nurhachius.
  • Unwin published the book The Pterosaurs from Deep Time. This was only the third "serious boo[k]" about pterosaurs ever published.[69] In it, Unwin argued that young pterosaurs were born well-developed and requiring little investment of parental care.[17]
  • A pterosaur-focused exhibit went on tour in Japan.[69]
  • Peinkowski and Niedzwiedzki published a study on pterosaur tracks from Poland.[137]
  • Meyer and others reported the presence of pterosaur footprints in the Upper Cretaceous Cerro del Pueblo Formation of Mexico. However, they are now actually thought to be poorly preserved dinosaur footprints.[118]
  • Kleeman reported the presence of pterosaur footprints in the Upper Cretaceous Cerro del Pueblo Formation of Mexico. However, they are now actually thought to be poorly preserved dinosaur footprints.[118]
Life restoration of Muzquizopteryx

2006

Humerus fragment of Aralazhdarcho

2007

  • Averianov described the new genus and species
    Aralazhdarcho bostobensis.[176]
  • Wang and others described the new genus and species
  • Kellner and Campos described the new genus Tupandactylus.[178]
  • Humphries and others debunked the hypothesis that many pterosaurs fed by skim feeding.[162]
  • Veldmeijer and others published a study on pterosaur skull biomechanics.[162]
  • Following the success of the 2001 pterosaur symposium in Toulouse, a regular gathering of pterosaur paleontologists was established and titled "Flugsaurier" after the German word for pterosaur. This debut meeting was held in Munich and dedicated to long-time pterosaur paleontologist Peter Wellnhofer.[69]
  • Harris and others reported the existence of fossil pterosaur tracks preserving the impression of a fifth toe. This suggests that the trackmaker was a primitive long-tailed pterosaur.[138]
  • Garcia Ramos and others published research on exceptionally well-preserved Late Jurassic pterosaur tracks in Asturias, Spain.[134]
  • Pinuela and others published research on exceptionally well-preserved Late Jurassic pterosaur tracks in Asturias, Spain.[118]
  • Lockley and others also reported the existence of fossil pterosaur tracks preserving the impression of a fifth toe. This suggests that the trackmaker was a primitive long-tailed pterosaur.[138]
  • Lockley and others reported the presence of pterosaur tracks in the Dakota Group of Colorado.[118]

April

  • The University of Munich awarded Helmut Tischlinger an honorary doctorate degree for his work studying pterosaur fossil under ultraviolet light to better understand their soft tissues.[61]

2008

Skeletal reconstruction of Shenzhoupterus
Life restoration of Nemicolopterus.
Life restoration of Ningchengopterus.

2009

2010s

The skull of Alanqa
Life restoration of Darwinopterus
Life restoration of Dawndraco
Life restoration of Sericipterus

2010

Type skeleton of Aurorazhdarcho

2011

Life restoration of Bellubrunnus
Skull of Guidraco

2012

Skull of the type specimen of Ardeadactylus
Life restoration of Cuspicephalus

2013

2014

Type skeleton of Aerodactylus

2015

Illustration of the skull of Banguela with jaws both open and closed

2016

2017

  • Zhou and others described the new genus and species
    Liaodactylus primus
    .
  • Wang and others described the new genus and species
    Douzhanopterus zhengi
    .
  • Vidovic and Martill erected the new genus Altmuehlopterus for the species "Ornithocephalus" ramphastinus.
  • Kellner and Calvo described the new genus and species
    Argentinadraco barrealensis
    .

2018

  • Britt and others described the new genus and species
    Caelestiventus hanseni
    .
  • Longrich, Martill and Andres described the new genera and species
    Tethydraco regalis
    .
  • Lü and others described the new genus and species
    Vesperopterylus lamadongensis
    .
  • Martill and others described the new genus and species
    Xericeps curvirostris
    .
  • Rigal, Martill and Sweetman erected the new genus Serradraco for the species "Pterodactylus" sagittirostris.
  • Vullo and others described the new genus and species
    Mistralazhdarcho maggii
    .

2019

2020s

2020

See also

Footnotes

  1. ^ Mayor (2005); "Cheyenne Fossil Knowledge", pages 209–211 and "The High Plains: Thunder Birds, Water Monsters, and Buffalo-Calling Stones", page 221.
  2. ^ a b Wellnhofer (2008); "2. Early discoveries", page 8.
  3. ^ a b Wellnhofer (2008); "2. Early discoveries", pages 8–9.
  4. ^ For pterosaurs interpreted as birds or bats, see Wellnhofer (2008); "2. Early discoveries", page 9. For an attribution of pterosaurs to the infernal, see O'Connor (2012); page 499 and Hawkins (1840); "Addenda", page 7.
  5. ^ For the description of Rhamphorhynchus, see Hanson (2008); "R", pages 19–20. For the description of Gnathosaurus, see "G", page 9.
  6. ^ For the description of Pterosauria, see Wellnhofer (2008); "2. Early discoveries", page 10. For Dimorphodon, see "3. First pterosaurs from the Lias".
  7. ^ a b Wellnhofer (2008); "5. American discoveries", page 11.
  8. ^ a b c d e f g h Wellnhofer (2008); "9. The problem of terrestrial locomotion", page 14.
  9. ^ For this paper as the first major study of pterosaur biomechanics, see Wellnhofer (2008); "8. Flight biomechanics", page 13. For its conclusion regarding pterosaurian helplessness on the ground, see "9. The problem of terrestrial locomotion", page 14.
  10. ^ a b c d e Wellnhofer (2008); "9. The problem of terrestrial locomotion", page 15.
  11. ^ For Padian's 1983 peper on pterosaur gait, see Wellnhofer (2008); "9. The problem of terrestrial locomotion", page 14. For his criticism of Stokes's pterosaur track claims, see Lockley and Hunt (1995); "What's in a Name?", page 145.
  12. ^ Hone (2012); "Abstract", page 1366.
  13. ^ a b c Hone (2012); "2 What is Out There?", page 1367.
  14. ^ a b Hone (2012); "3 New and Important Finds", page 1367.
  15. ^ a b Hone (2012); "3 New and Important Finds", pages 1367–1368.
  16. ^ a b c Hone (2012); "4 Pterosaur Origins", page 1369.
  17. ^ a b c d e f g h i j k l m n o p q r s t u Hone (2012); "3 New and Important Finds", page 1368.
  18. ^ Mayor (2005); "Cheyenne Fossil Knowledge", pages 209–210.
  19. ^ Mayor (2005); "Cheyenne Fossil Knowledge", page 211.
  20. ^ Mayor (2005); "The High Plains: Thunder Birds, Water Monsters, and Buffalo-Calling Stones", page 221.
  21. ^ a b c d e f g h i j k l m Wellnhofer (2008); "2. Early discoveries", page 9.
  22. ^ For von Soemmerring's full name, see Wellnhofer (2008); "1. Personal Remarks", page 8. For his involvement in early pterosaur research, see "2. Early discoveries", page 9.
  23. ^ Cuvier, G. (1819). "Pterodactylus longirostris". In Oken, Lorenz (ed.). Isis (oder Encyclopädische Zeitung) von Oken (in German). Jena : Expedition der Isis. pp. 1126, 1788.
  24. ^ a b Wellnhofer (2008); "4. Discoveries of Cretaceous pterosaurs", pages 10–11.
  25. ^ a b c d e f g Wellnhofer (2008); "3. First pterosaurs from the Lias", page 10.
  26. ^ a b c d e f Wellnhofer (2008); "7. Early life restorations", page 12.
  27. ^ a b c d Wellnhofer (2008); "10. The evidence of soft parts", page 17.
  28. ^ a b c Hanson (2008); "R", page 19.
  29. ^ Hanson (2008); "G", page 9.
  30. ^ a b c d Wellnhofer (2008); "2. Early discoveries", page 10.
  31. ^ Wellnhofer (2008); "2. Early discoveries", pages 9–10.
  32. ^ O'Connor (2012); page 499.
  33. ^ For Hawkins's interpretation of pterosaurs as "engrafted-by-Evil", see Hawkins (1840); "Addenda", page 7. For his portrayal of pterosaurs as shoreline scavengers, see Wellnhofer (2008); "7. Early life restorations", page 12.
  34. ^ a b Hanson (2008); "R", pages 19–20.
  35. ^ a b c Hanson (2008); "C", page 6.
  36. ^ Carpenter (1999); "England", page 13.
  37. ^ a b Lockley and Meyer (2000); "Turtles and Hopping Dinosaurs", page 178.
  38. ^ a b c d e f g h i j k l m n o p q r s t u Lockley, Harris, and Mitchell (2008); "2. History of discovery and debate", page 186.
  39. ^ Hanson (2008); "O", pages 13–14.
  40. ^ For the description of Cycnorhamphus, see Hanson (2008); "C", pages 6–7. For the description of Ornithocheirus huxleyi, see "O", pages 13–14.
  41. ^ a b Wellnhofer (2008); "4. Discoveries of Cretaceous pterosaurs", page 11.
  42. ^ Hanson (2008); "L", page 10.
  43. ^ Everhart (2005); "Pteranodons: Rulers of the Air", page 195.
  44. ^ a b c d Everhart (2005); "Pteranodons: Rulers of the Air", page 191.
  45. ^ a b c d e Everhart (2005); "Pteranodons: Rulers of the Air", page 199.
  46. ^ a b c d Hanson (2008); "D", page 7.
  47. ^ a b c d e Everhart (2005); "Pteranodons: Rulers of the Air", page 194.
  48. ^ a b c d Everhart (2005); "Pteranodons: Rulers of the Air", page 196.
  49. ^ a b c d e f g Everhart (2005); "Pteranodons: Rulers of the Air", page 197.
  50. ^ For Carruthers's first name and his description of new oospecies, see Carpenter (1999); "England", page 13. For his attribution of Oolithes to pterosaurs, see Carpenter, Hirsch, and Horner (1996); "The discovery of dinosaur eggs", page 1.
  51. ^ a b Everhart (2005); "Pteranodons: Rulers of the Air", pages 197–198.
  52. ^ a b Everhart (2005); "Pteranodons: Rulers of the Air", page 198.
  53. ^ a b c Hanson (2008); "C", pages 5–6.
  54. ^ a b c d e f g h i Everhart (2005); "Pteranodons: Rulers of the Air", page 200.
  55. ^ Everhart (2005); "Pteranodons: Rulers of the Air", pages 199–200.
  56. ^ a b c d e f Everhart (2005); "Pteranodons: Rulers of the Air", page 202.
  57. ^ a b Everhart (2005); "Pteranodons: Rulers of the Air", page 203.
  58. ^ For the description of Dermodactylus in 1881, see Hanson (2008); "D", page 7. For the description of Laopteryx priscus see "L", page 10.
  59. ^ a b Wellnhofer (2008); "10. The evidence of soft parts", page 15.
  60. ^ a b Wellnhofer (2008); "10. The evidence of soft parts", pages 15–16.
  61. ^ a b c d e f Wellnhofer (2008); "10. The evidence of soft parts", page 16.
  62. ^ a b c d e f Wellnhofer (2008); "6. Triassic Pterosaurs", page 12.
  63. ^ Hanson (2008); "O", page 14.
  64. ^ Wellnhofer (2008); "10. The evidence of soft parts", pages 16–17.
  65. ^ Everhart (2005); "Pteranodons: Rulers of the Air", page 201.
  66. ^ Everhart (2005); "Pteranodons: Rulers of the Air", pages 201–202.
  67. ^ For the subject of Williston's critique being the length of Pteranodon's crest in Marsh's 1884 reconstruction, see Everhart (2005); "Pteranodons: Rulers of the Air", page 200. For Williston's characterization of the reconstruction as too speculative for its quality of preservation, see page 202.
  68. ^ a b c d e f Everhart (2005); "Pteranodons: Rulers of the Air", page 204.
  69. ^ a b c d e f g h i Hone (2012); "8 Increased Research and Outreach", page 1372.
  70. ^ a b c d e f Everhart (2005); "Pteranodons: Rulers of the Air", page 209.
  71. ^ a b c d Everhart (2005); "Pteranodons: Rulers of the Air", page 205.
  72. ^ Everhart (2005); "Pteranodons: Rulers of the Air", pages 205–206.
  73. ^ a b c d e f g Everhart (2005); "Pteranodons: Rulers of the Air", page 207.
  74. ^ Everhart (2005); "Pteranodons: Rulers of the Air", pages 200–201.
  75. ^ a b c d e f g h Everhart (2005); "Pteranodons: Rulers of the Air", page 206.
  76. ^ Hanson (2008); "S", page 21.
  77. ^ a b c Wellnhofer (2008); "5. American discoveries", page 12.
  78. ^ a b c d e Everhart (2005); "Pteranodons: Rulers of the Air", page 210.
  79. ^ Hanson (2008); "L", pages 10–11.
  80. ^ Everhart (2005); "Pteranodons: Rulers of the Air", page 209. Wellnhofer (2008); "8. Flight biomechanics", page 13.
  81. ^ a b c d e Hanson (2008); "P", page 15.
  82. ^ a b Hanson (2008); "A", pages 2–3.
  83. ^ a b Hanson (2008); "C", page 5.
  84. ^ a b c d Hanson (2008); "B", page 4.
  85. ^ Hanson (2008); "A", page 24.
  86. ^ a b Lockley, Harris, and Mitchell (2008); "2. History of discovery and debate", page 188.
  87. ^ a b c d e f Everhart (2005); "Pteranodons: Rulers of the Air", page 208.
  88. ^ Hanson (2008); "P", page 25.
  89. ^ Wellnhofer (2008); "11. Recent discoveries", page 18.
  90. ^ Hanson (2008); "G", pages 8–9.
  91. ^ Wellnhofer (2008); "1. Personal remarks", page 7.
  92. ^ a b c Hanson (2008); "N", page 12.
  93. ^ Hanson (2008); "P", page 18.
  94. ^ a b Wellnhofer (2008); "11. Recent discoveries", page 17.
  95. ^ Everhart (2005); "Pteranodons: Rulers of the Air", pages 208–209.
  96. ^ a b c d e f g h i j Wellnhofer (2008); "8. Flight biomechanics", page 14.
  97. ^ a b c Hanson (2008); "A", page 3.
  98. ^ Hanson (2008); "D", pages 7–8.
  99. ^ Wellnhofer (2008); "8. Flight biomechanics", page 13.
  100. ^ Wellnhofer (2008); "8. Flight biomechanics", pages 13–14.
  101. ^ Hanson (2008); "P", pages 17–18.
  102. ^ Hanson (2008); "H", page 9.
  103. ^ Everhart (2005); "Pteranodons: Rulers of the Air", pages 209–210.
  104. ^ Hanson (2008); "Q", page 19.
  105. ^ a b c d e f g h i j k l m Lockley, Harris, and Mitchell (2008); "2. History of discovery and debate", page 187.
  106. ^ a b c d e Lockley, Harris, and Mitchell (2008); "4.2 Lack of convincing, pre-Late Jurassic reports of pterosaurian tracks", page 189.
  107. ^ Hanson (2008); "S", page 20.
  108. ^ Hanson (2008); "H", page 10.
  109. ^ Hanson (2008); "A", page 2.
  110. ^ a b Hanson (2008); "A", page 4.
  111. ^ Lockley and Hunt (1995); "What's in a Name?", page 145.
  112. ^ Hanson (2008); "B", page 4-5.
  113. ^ a b c d e f Hanson (2008); "T", page 22.
  114. ^ a b Hanson (2008); "M", page 12.
  115. ^ a b c Everhart (2005); "Pteranodons: Rulers of the Air", page 211.
  116. ^ Hanson (2008); "P", pages 15–16.
  117. ^ a b c d e f g Everhart (2005); "Pteranodons: Rulers of the Air", page 212.
  118. ^ a b c d e f g h Lockley, Harris, and Mitchell (2008); "4.3 Important new finds that have yet to be studied", page 190.
  119. ^ Everhart (2005); "Pteranodons: Rulers of the Air", pages 197 and 199.
  120. ^ Everhart (2005); "Pteranodons: Rulers of the Air", pages 207–209.
  121. ^ For discussion of Bennett's critique of Miller's Pteranodon taxonomy, see Everhart (2005); "Pteranodons: Rulers of the Air", pages 208–209. For his critique of Harksen's P. sternbergi reconstruction, see pages 207–208.
  122. ^ Hanson (2008); "Z", page 23.
  123. ^ Hanson (2008); "P", pages 24–25.
  124. ^ a b c Hone (2012); "4 Pterosaur Origins", page 1368.
  125. ^ Hanson (2008); "K", page 10.
  126. ^ Carpenter (1999); "England", pages 13–14.
  127. ^ a b Lockley, Harris, and Mitchell (2008); "2. History of discovery and debate", pages 186-187.
  128. ^ Everhart (2005); "Pteranodons: Rulers of the Air", pages 191–192.
  129. ^ Everhart (2005); "Pteranodons: Rulers of the Air", pages 192–193.
  130. ^ a b Hanson (2008); "E", page 8.
  131. ^ Hanson (2008); "D", page 7. For the original description, see Martill et al. (2000).
  132. ^ Kellner and Tomida (2000); in passim.
  133. ^ a b c d e f g h i j k l Hone (2012); "6 Anatomy", page 1370.
  134. ^ a b c d Lockley, Harris, and Mitchell (2008); "4.4.1 Exceptional preservation", page 190.
  135. ^ Wang and Lu (2001); in passim.
  136. ^ Howse, Milner, and Martill (2001); in passim.
  137. ^ a b c d Lockley, Harris, and Mitchell (2008); "3. Current state of knowledge", page 188.
  138. ^ a b c Lockley, Harris, and Mitchell (2008); "4.1 Identifying the track makers", page 189.
  139. ^ Lockley, Harris, and Mitchell (2008); "4.5 Tracks and indicators of pterosaur behavior and paleoecology", page 191.
  140. ^ Hanson (2008); "H", page 24.
  141. ^ Dalla Vecchia et al. (2002); in passim.
  142. ^ Buffetaut, Grigorescu, and Csiki (2002); in passim.
  143. ^ Wang et al. (2002); in passim.
  144. ^ Varricchio (2002); in passim.
  145. ^ Czerkas and Ji (2002); in passim.
  146. ^ Kellner and Campos (2002); in passim.
  147. ^ Czerkas and Mickelson (2002); in passim.
  148. ^ a b c d e f g Hone (2012); "7 Functional Morphology and Ecology", page 1370.
  149. ^ Everhart (2005); "Pteranodons: Rulers of the Air", pages 212–213.
  150. ^ Lü (2003); in passim.
  151. ^ a b Wang and Zhou (2003a); in passim.
  152. ^ Carpenter et al. (2003); in passim.
  153. ^ Dong, Sun, and Wu (2003); in passim.
  154. ^ Frey, Martill, and Buchy (2003); in passim.
  155. ^ Pereda-Suberbiola et al. (2003); in passim.
  156. ^ Wang and Zhou (2003b); in passim.
  157. ^ a b c d Hone (2012); "5 Phylogeny and Taxonomy", page 1370.
  158. ^ Lockley, Harris, and Mitchell (2008); "1. Introduction", page 186.
  159. ^ Peters (2004); in passim.
  160. ^ Gasparini, Fernández, and de la Fuente (2004); in passim.
  161. ^ Maisch, Matzke, and Sun (2004); in passim.
  162. ^ a b c d e f g h Hone (2012); "7 Functional Morphology and Ecology", page 1371.
  163. ^ Ösi, Weishampel, and Jianu (2005); in passim.
  164. ^ Lü and Ji (2005a); in passim.
  165. ^ Steel et al. (2005); in passim.
  166. ^ Lü and Ji (2005b); in passim.
  167. ^ Lü and Zhang (2005); in passim.
  168. ^ Wang et al. (2005); in passim.
  169. ^ Lu and Yuan (2005); in passim.
  170. ^ Dong and Lu (2005); in passim.
  171. ^ Wang and Zhou (2006); in passim.
  172. ^ Fröbisch and Fröbisch (2006); in passim.
  173. ^ Wang et al. (2006); in passim.
  174. ^ Frey et al. (2006); in passim.
  175. ^ Lü et al. (2006); in passim.
  176. ^ Averianov (2007); in passim.
  177. ^ Wang et al. (2007); in passim.
  178. ^ Kellner and Campos (2007); in passim.
  179. ^ Andres and Ji (2008); in passim.
  180. ^ Wang et al. (2008a); in passim.
  181. ^ Molnar and Thulborn (2008); in passim.
  182. ^ Wang et al. (2008b); in passim.
  183. ^ Stecher (2008); in passim.
  184. ^ Lü et al. (2008); in passim.
  185. ^ Averianov, Arkhangelsky, and Pervushov (2008); in passim.
  186. ^ a b Hone (2012); "7 Functional Morphology and Ecology", pages 1370–1371.
  187. ^ Dalla Vecchia (2009); in passim.
  188. ^ Lü (2009a); in passim.
  189. ^ Lü (2009b); in passim.
  190. ^ Wang et al. (2009); in passim.
  191. ^ Myers (2010); in passim.
  192. ^ Ibrahim et al. (2010); in passim.
  193. ^ Lü and Fucha (2010); in passim.
  194. ^ Lü et al. (2010); in passim.
  195. ^ a b Wang et al. (2010); in passim.
  196. ^ a b Kellner (2010); in passim.
  197. ^ Bonaparte, Schultz, and Soares (2010); in passim.
  198. ^ Lü, Fucha, and Chen (2010); in passim.
  199. ^ Fuentes Vidarte and Meijide Calvo (2010); in passim.
  200. ^ Andres, Clark, and Xu (2010); in passim.
  201. ^ Lü (2010); in passim.
  202. ^ Hone (2012); "4 Pterosaur Origins", pages 1368–1369.
  203. ^ Frey, Meyer, and Tischlinger (2011); in passim.
  204. ^ Kellner, Rodrigues, and Costa (2011); in passim.
  205. ^ Elgin and Frey (2011); in passim.
  206. ^ Lü et al. (2011); in passim.
  207. ^ Lü and Bo (2011); in passim.
  208. ^ Sullivan and Fowler (2011); in passim.
  209. ^ Jiang and Wang (2011); in passim.
  210. ^ Martill (2011); in passim.
  211. ^ Novas et al. (2012); in passim.
  212. ^ Hone et al. (2012); in passim.
  213. ^ Lü and Hone (2012); in passim.
  214. ^ Vullo et al. (2012); in passim.
  215. ^ Lü et al. (2012a); in passim.
  216. ^ Wang et al. (2012); in passim.
  217. ^ Cheng et al. (2012); in passim.
  218. ^ Lü et al. (2012b); in passim.
  219. ^ Lü et al. (2012c); in passim.
  220. ^ Hone (2012); "7 Functional Morphology and Ecology", page 1372.
  221. ^ a b Andres and Myers (2013); in passim.
  222. ^ Bennett (2013); in passim.
  223. ^ a b c Rodrigues and Kellner (2013); in passim.
  224. ^ Kellner (2013); in passim.
  225. ^ Martill and Etches (2013); in passim.
  226. ^ Vremir et al. (2013); in passim.
  227. ^ Naish, Simpson and Dyke (2013); in passim.
  228. ^ Codorniú and Gasparini (2013); in passim.
  229. ^ Vidovic and Martill (2014); in passim.
  230. ^ Jiang et al. (2014); in passim.
  231. ^ Manzig et al. (2014); in passim.
  232. ^ Wang et al. (2014a); in passim.
  233. ^ Wang et al. (2014b); in passim.
  234. ^ Andres, Clark and Xu (2014); in passim.
  235. ^ Bantim et al. (2014); in passim.
  236. ^ a b c Kellner (2015); in passim.
  237. ^ Headden and Campos (2015); in passim.
  238. ^ Myers (2015); in passim.
  239. ^ Cheng et al. (2015); in passim.
  240. ^ Rodrigues et al. (2015); in passim.
  241. ^ Lü et al. (2015); in passim.
  242. ^ O'Sullivan and Martill (2015); in passim.
  243. ^ Lü et al. (2016); in passim.
  244. ^ Codorniú et al. (2016); in passim.
  245. S2CID 134439172
    .
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  247. .
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  261. .

References

A-F

G-L

M-R

S-Z

External links