Editing Pterosaur (section)

==History of discovery==
{{see also|Timeline of pterosaur research}}

===First finds===
[[File:Pterodactylus holotype Collini 1784.jpg|thumb|left|Engraving of the original ''[[Pterodactylus antiquus]]'' specimen by [[Egid Verhelst the Younger|Egid Verhelst II]], 1784]]
Pterosaur [[fossil]]s 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]],{{sfn|Witton|2013|p=5}} became much sought after by rich collectors.{{sfn|Wellnhofer|1991|p=22}} In 1784, Italian naturalist [[Cosimo Alessandro Collini]] was the first scientist to describe a pterosaur fossil.{{sfn|Witton|2013|p=6}} 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.{{sfn|Witton|2013|pp=6–7}} The discovery of pterosaurs would thus play an important role in the progress of modern paleontology and geology.{{sfn|Witton|2013|p=7}} Scientific opinion at the time was that if such creatures were still alive, only the sea was a credible habitat; Collini suggested it might be a swimming animal that used its long front limbs as paddles.<ref name="collini1784">Collini, C.A. (1784). "Sur quelques Zoolithes du Cabinet d'Histoire naturelle de S. A. S. E. Palatine & de Bavière, à Mannheim." ''Acta Theodoro-Palatinae Mannheim 5 Pars Physica'', pp. 58–103 (1 plate).</ref> 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]].<ref name="wagler1830">Wagler, J. (1830). ''Natürliches System der Amphibien'' Munich, 1830: 1–354.</ref>
[[File:Pterodactyl reconstruction Newman 1843.jpg|thumb|Newman's marsupial pterosaurs]]
In 1800, [[Johann Hermann]] first suggested that it represented a flying creature in a letter to [[Georges Cuvier]]. Cuvier agreed in 1801, understanding it was an extinct flying reptile.<ref name="cuvier1801">{{cite journal | author = Cuvier G | year = 1801 | title = [Reptile volant]. In: Extrait d'un ouvrage sur les espèces de quadrupèdes dont on a trouvé les ossemens dans l'intérieur de la terre | journal = Journal de Physique, de Chimie et d'Histoire Naturelle | volume = 52 | pages = 253–67}}</ref> In 1809, he coined the name ''Ptéro-Dactyle'', "wing-finger".<ref>Cuvier, G., 1809, "Mémoire sur le squelette fossile d'un Reptil volant des environs d'Aichstedt, que quelques naturalistes ont pris pour un oiseau, et donc nous formons un genre de Sauriens, sous le nom de Ptero-Dactyle", ''Annales du Musée d'Histoire Naturelle'', Paris, '''13''' pp. 424–37</ref> This was in 1815 Latinised to ''[[Pterodactylus]]''.<ref>Rafinesque, C.S., 1815, ''Analyse de la Nature ou tableau de l'univers et des corps organisés'', Palermo</ref> At first most species were assigned to this genus and ultimately "pterodactyl" was popularly and incorrectly applied to all members of Pterosauria.<ref name="myths"/> Today, paleontologists limit the term to the genus ''Pterodactylus'' or members of the [[Pterodactyloidea]].<ref name="alexander"/>

In 1812 and 1817, [[Samuel Thomas von Soemmerring]] redescribed the original specimen and an additional one.<ref>Von Soemmerring, S. T., 1812, "Über einen Ornithocephalus oder über das unbekannten Thier der Vorwelt, dessen Fossiles Gerippe Collini im 5. Bande der Actorum Academiae Theodoro-Palatinae nebst einer Abbildung in natürlicher Grösse im Jahre 1784 beschrieb, und welches Gerippe sich gegenwärtig in der Naturalien-Sammlung der königlichen Akademie der Wissenschaften zu München befindet", ''Denkschriften der königlichen bayerischen Akademie der Wissenschaften'', München: mathematisch-physikalische Classe '''3''': 89–158</ref> 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.{{sfn|Wellnhofer|1991|p=27}} In 1843, [[Edward Newman (entomologist)|Edward Newman]] thought pterosaurs were flying [[marsupials]].<ref>{{cite journal | last1 = Newman | first1 = E | year = 1843 | title = Note on the Pterodactyle Tribe considered as Marsupial Bats | journal = Zoologist | volume = 1 | pages = 129–31 }}</ref> Ironically, as the "bat model" depicted pterosaurs as warm-blooded and furred, it would turn out to be more correct in certain aspects than Cuvier's "reptile model" in the long run. In 1834, [[Johann Jakob Kaup]] coined the term Pterosauria.<ref>{{cite journal | last1 = Kaup | first1 = J. | year = 1834 | title = Versuch einer Eintheilung der Säugethiere in 6 Stämme und der Amphibien in 6 Ordnungen | journal = Isis von Oken | volume = 1834 | pages = 311–315 |url=https://www.biodiversitylibrary.org/page/27509907}}</ref>

===Expanding research===
[[File:Dimorphodon_reconstruction_Seeley_1901.jpg|thumb|left|Historical reconstruction of ''[[Dimorphodon]]'' as a biped by Seeley]]
In 1828, [[Mary Anning]] found in England the first pterosaur genus outside Germany,{{sfn|Wellnhofer|1991|p=28}} named as ''[[Dimorphodon]]'' by [[Richard Owen]], also the first non-pterodactyloid pterosaur known.{{sfn|Wellnhofer|1991|p=29}} Later in the century, the [[Early Cretaceous]] [[Cambridge Greensand]] produced thousands of pterosaur fossils, that however, were of poor quality, consisting mostly of strongly eroded fragments.{{sfn|Wellnhofer|1991|p=33}} Nevertheless, based on these, numerous genera and species would be named.{{sfn|Witton|2013|p=7}} Many were described by [[Harry Govier Seeley]], at the time the main English expert on the subject, who also wrote the first pterosaur book, ''Ornithosauria'',<ref>Seeley, H.G., 1870, ''Ornithosauria – an elementary study of the bones of Pterodactyles'', Cambridge University Press</ref> and in 1901 the first popular book,{{sfn|Witton|2013|p=7}} ''Dragons of the Air''. Seeley thought that pterosaurs were warm-blooded and dynamic creatures, closely related to birds.<ref>Seeley, H.G., 1901, ''Dragons of the Air: An account of extinct flying reptiles'', Londen: Methuen</ref> Earlier, the evolutionist [[St. George Jackson Mivart]] had suggested pterosaurs were the direct ancestors of birds.<ref>{{cite journal | last1 = Mivart | first1 = G | year = 1881 | title = A popular account of chamaeleons | journal = Nature | volume = 24 | issue = 615 | pages = 309–38 | doi = 10.1038/024335f0 | bibcode = 1881Natur..24..335. | s2cid = 30819954 }}</ref> Owen opposed the views of both men, seeing pterosaurs as cold-blooded "true" reptiles.{{sfn|Wellnhofer|1991|p=35}}

In the US, [[Othniel Charles Marsh]] in 1870 discovered ''Pteranodon'' in the [[Niobrara Chalk]], then the largest known pterosaur,{{sfn|Wellnhofer|1991|p=35}} the first toothless one and the first from America.{{sfn|Wellnhofer|1991|p=36}} These layers too rendered thousands of fossils,{{sfn|Wellnhofer|1991|p=36}} also including relatively complete skeletons that were three-dimensionally preserved instead of being strongly compressed as with the Solnhofen specimens. This led to a much better understanding of many anatomical details,{{sfn|Wellnhofer|1991|p=36}} such as the hollow nature of the bones.
[[File:Rhamphorhynchus_reconstruction_Riou_1863.jpg|thumb|Early reconstruction of ''[[Rhamphorhynchus]]'']]
Meanwhile, finds from the Solnhofen had continued, accounting for the majority of complete high-quality specimens discovered.{{sfn|Wellnhofer|1991|p=31}} They allowed to identify most new basal taxa, such as ''[[Rhamphorhynchus]]'', ''[[Scaphognathus]]'' and ''[[Dorygnathus]]''.{{sfn|Wellnhofer|1991|p=31}} This material gave birth to a German school of pterosaur research, which saw flying reptiles as the warm-blooded, furry and active Mesozoic counterparts of modern bats and birds.{{sfn|Wellnhofer|1991|pp=37–38}} In 1882, Marsh and [[Karl Alfred Zittel]] published studies about the wing membranes of specimens of ''Rhamphorhynchus''.<ref>{{cite journal | last1 = Marsh | first1 = O.C. | year = 1882 | title = The wings of Pterodactyles | journal = American Journal of Science | volume = 3 | issue = 16| page = 223 }}</ref><ref>{{cite journal | last1 = Zittel | first1 = K.A. | year = 1882 | title = Über Flugsaurier aus dem lithografischen Schiefer Bayerns | journal = Palaeontographica | volume = 29 | pages = 47–80 }}</ref> German studies continued well into the 1930s, describing new species such as ''[[Anurognathus]]''. In 1927, [[Ferdinand Broili]] discovered hair follicles in pterosaur skin,<ref>Broili, F., 1927, "Ein Ramphorhynchus mit Spuren von Haarbedeckung", ''Sitzungsberichte der Bayerischen Akademie der Wissenschaften'' p. 49-67</ref> and [[paleoneurology|paleoneurologist]] [[Tilly Edinger]] determined that the brains of pterosaurs more resembled those of birds than modern cold-blooded reptiles.<ref>{{cite journal | last1 = Edinger | first1 = T | year = 1927 | title = Das Gehirn der Pterosaurier | url = http://bigcat.fhsu.edu/biology/cbennett/bib-arch-pter/Edinger-1927.pdf | journal = [[Zeitschrift für Anatomie und Entwicklungsgeschichte]] | volume = 83 | issue = 1/3 | pages = 105–12 | doi = 10.1007/bf02117933 | s2cid = 19084773 | access-date = 2019-10-27 | archive-date = 2020-07-28 | archive-url = https://web.archive.org/web/20200728093840/http://bigcat.fhsu.edu/biology/cbennett/bib-arch-pter/Edinger-1927.pdf | url-status = dead }}</ref>

In contrast, English and American paleontologists by the middle of the twentieth century largely lost interest in pterosaurs. They saw them as failed evolutionary experiments, cold-blooded and scaly, that hardly could fly, the larger species only able to glide, being forced to climb trees or throw themselves from cliffs to achieve a take-off. In 1914, for the first-time pterosaur aerodynamics were quantitatively analysed, by [[Ernest Hanbury Hankin]] and [[David Meredith Seares Watson]], but they interpreted ''Pteranodon'' as a pure glider.<ref>Hankin E.H. & Watson D.S.M.; "On the Flight of Pterodactyls", ''The Aeronautical Journal'', October 1914, pp. 324–35</ref> Little research was done on the group during the 1940s and 1950s.{{sfn|Witton|2013|p=7}}

===Pterosaur renaissance===
[[File:Zhenjiangopterus jconway.jpg|thumb|left|This drawing of ''[[Zhejiangopterus]]'' by John Conway exemplifies the "new look" of pterosaurs]]
The situation for dinosaurs was comparable. From the 1960s onwards, a [[dinosaur renaissance]] took place, a quick increase in the number of studies and critical ideas, influenced by the discovery of additional fossils of ''[[Deinonychus]]'', whose spectacular traits refuted what had become entrenched orthodoxy. In 1970, likewise the description of the furry pterosaur ''[[Sordes]]'' began what [[Robert Bakker]] named a renaissance of pterosaurs.<ref>Bakker, Robert, 1986, ''The Dinosaur Heresies'', Londen: Penguin Books, 1988, p. 283</ref> [[Kevin Padian]] especially propagated the new views, publishing a series of studies depicting pterosaurs as warm-blooded, active and running animals.<ref>{{cite journal | last1 = Padian | first1 = K | year = 1979 | title = The wings of pterosaurs: A new look | journal = Discovery | volume = 14 | pages = 20–29 }}</ref><ref>Padian, K., 1980, ''Studies of the structure, evolution, and flight of pterosaurs (reptilia: Pterosauria)'', Ph.D. diss., Department of Biology, Yale University</ref><ref name="Padian1983"/> This coincided with a revival of the German school through the work of [[Peter Wellnhofer]], who in 1970s laid the foundations of modern pterosaur science.{{sfn|Witton|2013|p=5}} In 1978, he published the first pterosaur textbook,{{sfn|Witton|2013|p=9}} the ''Handbuch der Paläoherptologie, Teil 19: Pterosauria'',<ref>Wellnhofer, P., 1978, ''Handbuch der Paläoherpetologie XIX. Pterosauria'', Urban & Fischer, München</ref> and in 1991 the second ever popular science pterosaur book,{{sfn|Witton|2013|p=9}} the ''Encyclopedia of Pterosaurs''.{{sfn|Wellnhofer|1991|pp=1–192}}

This development accelerated through the exploitation of two new ''Lagerstätten''.{{sfn|Witton|2013|p=9}} During the 1970s, the Early Cretaceous [[Santana Formation]] in Brazil began to produce chalk nodules that, though often limited in size and the completeness of the fossils they contained, perfectly preserved three-dimensional pterosaur skeletal parts.{{sfn|Witton|2013|p=9}} German and Dutch institutes bought such nodules from fossil poachers and prepared them in Europe, allowing their scientists to describe many new species and revealing a whole new fauna. Soon, Brazilian researchers, among them [[Alexander Kellner]], intercepted the trade and named even more species.
[[File:Sinopterus_dongi_NMNS.jpg|thumb|Specimen of ''[[Sinopterus]]'', one of many excellent pterosaurs fossils from [[Liaoning]], [[China]]]]
Even more productive was the Early Cretaceous Chinese [[Jehol Biota]] of [[Liaoning]] that since the 1990s has brought forth hundreds of exquisitely preserved two-dimensional fossils, often showing soft tissue remains. Chinese researchers such as [[Lü Junchang]] have again named many new taxa. As discoveries also increased in other parts of the world, a sudden surge in the total of named genera took place. By 2009, when they had increased to about ninety, this growth showed no sign of levelling-off.<ref>{{cite journal | last1 = Dyke | first1 = G.J. McGowan | last2 = Nudds | first2 = R.L. | last3 = Smith | first3 = D. | year = 2009 | title = The shape of pterosaur evolution: evidence from the fossil record | journal = Journal of Evolutionary Biology | volume = 22 | issue = 4| pages = 890–98 | doi = 10.1111/j.1420-9101.2008.01682.x | pmid = 19210587 | s2cid = 32518380 | doi-access = free }}</ref> In 2013, M.P. Witton indicated that the number of discovered pterosaur species had risen to 130.<ref name=WittonPycnofibres>{{harvnb|Witton|2013}}</ref> Over ninety percent of known taxa has been named during the "renaissance". Many of these were from groups the existence of which had been unknown.{{sfn|Witton|2013|p=9}} Advances in computing power enabled researchers to determine their complex relationships through the [[quantitative method]] of [[cladistics]]. New and old fossils yielded much more information when subjected to modern ultraviolet light or roentgen photography, or [[CAT-scan]]s.{{sfn|Witton|2013|p=10}} Insights from other fields of biology were applied to the data obtained.{{sfn|Witton|2013|p=10}} All this resulted in a substantial progress in pterosaur research, rendering older accounts in popular science books completely outdated.

In 2017 a fossil from a 170-million-year-old pterosaur, later named as the species ''[[Dearc sgiathanach]]'' in 2022, was discovered on the [[Isle of Skye]] in [[Scotland]]. The [[National Museum of Scotland]] claims that it is the largest of its kind ever discovered from the [[Jurassic period]], and it has been described as the world's best-preserved skeleton of a pterosaur.<ref>{{cite web |title='Superbly preserved' pterosaur fossil unearthed in Scotland |date=22 Feb 2022 |website=Associated Press (AP) |url=https://phys.org/news/2022-02-superbly-pterosaur-fossil-unearthed-scotland.html}}</ref>