Editing Pterodactylus (section)

===Formerly assigned species===
[[File:Rhamphorhynchus Lauer.jpg|thumb|left|Fossil specimen of the species ''[[Rhamphorhynchus muensteri]]'', which was previously assigned as the species ''Pterodactylus münsteri'']]
Numerous species have been assigned to ''Pterodactylus'' in the years since its discovery. In the first half of the 19th century any new pterosaur species would be named ''Pterodactylus'', which thus became a "[[wastebasket taxon]]".<ref name=W1837 /> Even after clearly different forms had later been given their own generic name, new species would be created from the very productive sites, throughout Europe and North America, often based on only slightly different material.<ref name=Aerodactylus />

The earliest reassignments of pterosaur species to ''Pterodactylus'' started in 1825, with the description of ''Rhamphorhynchus''; fossil collector [[Georg Graf zu Münster]] alerted the German paleontologist [[Samuel Thomas von Sömmerring]] about several distinct fossil specimens, Sömmerring thought that they belonged to an ancient bird.<ref name=Pterosauria>{{cite book |last=Witton |first=Mark |title=Pterosaurs: Natural History, Evolution, Anatomy |publisher=Princeton University Press |year=2013 |isbn=978-0-691-15061-1}}</ref> Further fossil preparations had uncovered teeth, to which Graf zu Münster created a skull cast. He later sent the cast to Professor [[Georg August Goldfuss]], who recognized it as a pterosaur, specifically a species of ''Pterodactylus''. At the time however, most paleontologists incorrectly consider the genus ''Ornithocephalus'' ({{lit|bird-head}}) to be the valid name for ''Pterodactylus'', and therefore the specimen found was named as ''Ornithocephalus Münsteri'', which was first mentioned by Graf zu Münster himself.<ref name="Münster1830">{{cite book |last=Münster |first=Georg Graf zu |url=https://books.google.com/books?id=RWuWamFBZaAC&pg=PA2 |title=Nachtrag zu der Abhandlung des professor Goldfuss ueber den Ornithocephalus Münsteri (Goldf.) |publisher=F. C. Birner |year=1830 |location=Bayreuth}}</ref> Another specimen was found and described by Graf zu Münster in 1839, he assigned this specimen to a new separate species called ''Ornithocephalus longicaudus''; the [[specific name (zoology)|specific name]] means 'long tail', in reference to the animal's tail size.<ref>{{Cite book |last=Münster |first=Georg Graf zu |title=Neues Jahrbuch für Mineralogie, Geognosie, Geologie und Petrefaktenkunde |date=1839 |publisher=E. Schweizerbart's Verlagshandlung |location=Stuttgart |pages=676–682 |chapter=Über einige neue Versteinerungen in der lithographischen Schiefer von Baiern |volume=1839 |chapter-url=https://www.biodiversitylibrary.org/item/110811#page/696/mode/1up}}</ref> German paleontologist [[Hermann von Meyer]] in 1845 officially emended that the genus ''Pterodactylus'' had priority over ''Ornithocephalus'', so he reassigned the species ''O. münsteri'' and ''O. longicaudus'' into ''Pterodactylus münsteri'' and ''Pterodactylus longicaudus''.<ref name="vonmeyer1845">{{Cite journal |last1=von Meyer |first1=Hermann |year=1845 |title=System der fossilen Saurier |trans-title=Taxonomy of fossil saurians |url=https://www.biodiversitylibrary.org/item/110636#page/300/mode/1up |journal=Neues Jahrbuch für Mineralogie, Geognosie, Geologie und Petrefaktenkunde |volume=1845 |language=de |location=Stuttgart |publisher=E. Schweizerbart's Verlagshandlung |pages=278–285}}</ref> In 1846, von Meyer created the new species ''Pterodactylus gemmingi'' based on long-tailed remains; the specific name honors the fossil collector [[Carl Eming von Gemming]].<ref name="vonmeyer1846">{{Cite journal |last1=von Meyer |first1=Hermann |date=1846 |title=Pterodactylus (Rhamphorhynchus) gemmingi aus dem Kalkschiefer von Solenhofen |url=https://www.biodiversitylibrary.org/item/43708#page/17/mode/1up |journal=Palaeontographica |location=Cassel |publication-date=1851 |volume=1 |pages=1–20}}</ref> Later, in 1847, von Meyer finally erected the generic name ''Rhamphorhynchus'' ({{lit|beak snout}}) due to the distinctively long tails seen in the specimens found, which are much longer than those seen in ''Pterodactylus''. He assigned the species ''P. longicaudus'' as the type species of ''Rhamphorhynchus'', which resulted in a new combination called ''Rhamphorhynchus longicaudus''.<ref name="vonmeyer1847">{{cite book |last=von Meyer |first=Hermann |title=Homoeosaurus maximiliani und Rhamphorhynchus (Pterodactylus) longicaudus: Zwei fossile Reptilien aus dem Kalkschiefer von Solenhofen |url=http://data.onb.ac.at/rep/10B61475 |year=1847 |publisher=S. Schmerber'schen buchhandlung |location=Frankfurt |language=de}}</ref> The species ''R. münsteri'' was later changed to ''R. muensteri'' by Lydekker in 1888, due to the [[International Commission on Zoological Nomenclature|ICZN]] rule that prohibits non-standard Latin characters, such as ''ü'', in scientific names.<ref name=Lydekker />

[[File:PZSL1851PlateReptilia04.png|thumb|Holotype jaw fragments and teeth of ''[[Cimoliopterus]]'', which was previously known as ''Pterodactylus cuvieri'']]
Beginning in 1846, many pterosaur specimens were found near the village of [[Burham]] in [[Kent]], [[England]] by British paleontologists [[James Scott Bowerbank]] and Sir [[Richard Owen]]. Bowerbank had assigned fossil remains to two new species; the first was named in 1846 as ''Pterodactylus giganteus'';<ref name=Bowerbank1846>{{cite journal |last1=Bowerbank |first1=J.S. |year=1846 |title=On a new species of pterodactyl found in the Upper Chalk of Kent (''Pterodactylus giganteus'') |url=https://zenodo.org/record/1448505 |journal=Quarterly Journal of the Geological Society of London |volume=2 |issue=1–2 |pages=7–9 |doi=10.1144/gsl.jgs.1846.002.01-02.05 |s2cid=129389179}}</ref> the specific name means 'the gigantic one' in Latin, in reference to the large size of the remains, and the second species was named in 1851 as ''Pterodactylus cuvieri'', in honor of the French scientist Georges Cuvier.<ref name="Bowerbank1851">{{cite journal |last1=Bowerbank |first1=J.S. |year=1851 |title=On the pterodactyles of the Chalk Formation |url=https://zenodo.org/record/1447536 |journal=Proceedings of the Zoological Society of London |volume=19 |pages=14–20 |doi=10.1111/j.1096-3642.1851.tb01125.x}}</ref> Later in 1851, Owen named and described new pterosaur specimens that have been found yet again in England. He assigned these specimens to a new species called ''Pterodactylus compressirostris''.<ref name=Owen1851>Owen, R. (1851). Monograph on the fossil Reptilia of the Cretaceous Formations. ''The Palaeontographical Society'' '''5'''(11):1–118.</ref> In 1914 however, paleontologist [[Reginald Hooley]] redescribed ''P. compressirostris'', to which he erected the genus ''[[Lonchodectes]]'' ({{lit|[[lance]] biter}}), and therefore made ''P. compressirostris'' the type species, and created the new combination ''L. compressirostris''.<ref name="Hooley1914">{{Cite journal |last=Hooley |first=Reginald Walter |date=1914 |title=On the Ornithosaurian genus ''Ornithocheirus'', with a review of the specimens from the Cambridge Greensand in the Sedgwick Museum, Cambridge |url=https://zenodo.org/record/2207691 |journal=Annals and Magazine of Natural History |language=en |volume=13 |issue=78 |pages=529–557 |doi=10.1080/00222931408693521 |issn=0374-5481}}</ref> In a 2013 review, ''P. giganteus'' and ''P. cuvieri'' were reassigned to new genera; ''P. giganteus'' was reassigned to a genus called  ''[[Lonchodraco]]'' ('lance dragon'), which resulted in a new combination called ''L. giganteus'', and ''P. cuvieri'' was reassigned to the new genus ''[[Cimoliopterus]]'' ('chalk wing'), creating ''C. cuvieri''.<ref name="Rodrigues & Kellner 2013">{{Cite journal |last1=Rodrigues |first1=T. |last2=Kellner |first2=A. |doi=10.3897/zookeys.308.5559 |title=Taxonomic review of the ''Ornithocheirus'' complex (Pterosauria) from the Cretaceous of England |journal=ZooKeys |pages=1–112 |year=2013 |issue=308 |pmid=23794925 |pmc=3689139 |doi-access=free |bibcode=2013ZooK..308....1R}}</ref> Back in 1859, Owen had found remains the front part of a snout in the [[Cambridge Greensand]], and assigned it into the species ''Pterodactylus segwickii''; in honor of [[Adam Sedgwick]], a British geologist.<ref>Owen, R. (1859). ''Monograph on the fossil Reptilia of the Cretaceous formations. Supplement no. I''. Palaeontographical Society, London, p. 19</ref> This species however, was reassigned to the genus ''[[Camposipterus]]'' in 2013, therefore creating the new combination ''Camposipterus segwickii''.<ref name="Rodrigues & Kellner 2013"/> Later, in 1861, Owen had uncovered multiple distinctively looking fossil remains yet again in the Cambridge Greensand, these were assigned to a new species named ''Pterodactylus simus'',<ref>Martill, David. (2010). The early history of pterosaur discovery in Great Britain. Geological Society of London, Special Publications. 343. 287–311. {{doi|10.1144/SP343.18}}</ref> though the British paleontologist [[Harry Govier Seeley]] had created a separate generic name called ''[[Ornithocheirus]]'', and reassigned ''P. simus'' as the type species, which created the combination ''Ornithocheirus simus''.<ref name=S1869>{{Cite journal |last=Seeley |first=Harry Govier |date=1869 |title=Index to the fossil remains of Aves, Ornithosauria, and Reptilia, from the Secondary System of Strata, arranged in the Woodwardian Museum of the University of Cambridge |url=https://www.biodiversitylibrary.org/part/59487 |journal=Annals and Magazine of Natural History |language=en |volume=5 |issue=27 |pages=225–226 |doi=10.1080/00222937008696143 |issn=0374-5481}}</ref> Between the years 1869 and 1870, Seeley had reassigned many pterosaur species into ''Ornithocheirus'', while also creating several new species.<ref name=S1869/><ref name=HGS70>{{cite journal |author=Seeley, H.G. |year=1870 |title=The Ornithosauria: an Elementary Study of the Bones of Pterodactyles |journal=Cambridge |pages=112–128 |url=https://books.google.com/books?id=0M8yAQAAMAAJ}}</ref> Many of these species however, are now reclassified to other genera, or considered {{lang|la|[[nomina dubia]]}}.<ref name="Rodrigues & Kellner 2013"/> In 1874, further specimens were found in England, again by Owen, these ones were assigned to a new species called ''Pterodactylus sagittirostris'',<ref>Owen, R. 1874. "A Monograph on the Fossil Reptilia of the Mesozoic Formations. 1. Pterosauria." ''The Palaeontographical Society'' Monograph '''27''': 1–14</ref> this species however, was reassigned to the genus ''Lonchodectes'' in 1914 by Hooley, which resulted in an ''L. sagittirostris''.<ref name="Hooley1914" /> This conclusion was revised by Rigal ''et al.'' in 2017, who disagreed with Hooley's reassignment, and therefore created the genus ''[[Serradraco]]'', which afterwards resulted in a new combination called ''S. sagittirostris''.<ref name="Martill2017">{{cite journal |last1=Rigal |first1=S. |last2=Martill |first2=D. M. |last3=Sweetman |first3=S. C. |title=A new pterosaur specimen from the Upper Tunbridge Wells Sand Formation (Cretaceous, Valanginian) of southern England and a review of ''Lonchodectes sagittirostris'' (Owen 1874) |journal=Geological Society, London, Special Publications |date=2017 |volume=455 |pages=221–232 |doi=10.1144/SP455.5 |s2cid=133080548 |url=https://www.researchgate.net/publication/313966182}}</ref>

[[File:Pteranodon longiceps YPM1177.jpg|thumb|left|Specimen YPM1177, the first uncovered skull of ''[[Pteranodon]]'', which was back then assigned as a species of ''Pterodactylus'']]
Assigning new pterosaur species to ''Pterodactylus'' was not only common in Europe, but also in North America; paleontologists such as [[Othniel Charles Marsh]] in 1871 for example, described several toothless pterosaur specimens, which were accompanied by teeth that belonged to the fish ''[[Xiphactinus]]'', which Marsh assumed that these teeth belonged to the pterosaur specimens he found, since all pterosaurs discovered at the time had teeth. He then assigned these specimens to a new species called ''"Pterodactylus oweni"'', but this was changed to ''Pterodactylus occidentalis'' because ''"P. oweni"'' was found to have been [[preoccupied]] by a pterosaur species described with the same name back in 1864 by Seeley.<ref>{{cite journal |author=Marsh, O.C. |year=1871 |title=Note on a new and gigantic species of Pterodactyle |journal=American Journal of Science |series=3 |volume=1 |issue=6 |page=472 |url=http://oceansofkansas.com/Marsh71.htm}}</ref><ref name="History">{{cite journal |author=Witton |first=Mark P. |year=2010 |title=''Pteranodon'' and beyond: The history of giant pterosaurs from 1870 onwards. Geological Society of London Special Publications |url=https://www.researchgate.net/publication/258391482 |journal=Geological Society, London, Special Publications |volume=343 |pages=313–323|doi=10.1144/SP343.19 }}</ref> In 1872, American paleontologist [[Edward Drinker Cope]] also found various pterosaur specimens in North America, he assigned these to two new species known as ''Ornithochirus umbrosus'' and ''Ornithochirus harpyia'', Cope attempted to assign the specimens he found to the genus ''Ornithocheirus'', but misspelled forgetting the 'e'.<ref>{{cite journal |last1=Cope |first1=E. D. |year=1872 |title=On two new Ornithosaurians from Kansas |journal=Proceedings of the American Philosophical Society |volume=12 |issue=88 |pages=420–422 |jstor=981730}}</ref> In 1875 however, Cope reassigned the species ''O. umbrosus'' and ''O. harpyia'' into ''Pterodactylus umbrosus'' and ''Pterodactylus harpyia'', though these species had been considered {{lang|la|nomina dubia}} ever since.<ref name=cope1875>{{cite journal |last1=Cope |first1=E. D. |year=1875 |title=The Vertebrata of the Cretaceous formations of the West. |journal=Report, U. S. Geological Survey of the Territories (Hayden) |volume=2 |pages=302 pp., 57 pls |url=https://www.biodiversitylibrary.org/item/125656#page/11/mode/1up}}</ref><ref name=History /> Paleontologist [[Samuel Wendell Williston]] unearthed the first skull of the pterosaur, and found that the animal was toothless,<ref name=History /> this made Marsh create the genus ''[[Pteranodon]]'' ({{lit|toothless wing}}), and therefore reassigned all the American pterosaur species, including the ones that he named, from ''Pterodactylus'' to ''Pteranodon''.<ref name=marsh1876a>{{Cite journal |author=Marsh, O.C. |year=1876a |title=Notice of a new sub-order of Pterosauria |url=https://zenodo.org/record/1450032 |journal=American Journal of Science |series=Series 3 |volume=11 |issue=65 |pages=507–509 |doi=10.2475/ajs.s3-11.66.507 |bibcode=1876AmJS...11..507M |s2cid=130203580}}</ref>

Later, in the 1980s, subsequent revisions by [[Peter Wellnhofer]] had reduced the number of recognized species to about half a dozen. Many species assigned to ''Pterodactylus'' had been based on juvenile specimens, and subsequently been recognized as immature individuals of other species or genera. By the 1990s it was understood that this was even true for part of the remaining species. ''P. elegans'', for example, was found by numerous studies to be an immature ''Ctenochasma''.<ref name="jouve2004">{{Cite journal |doi=10.1671/0272-4634(2004)024[0542:DOTSOA]2.0.CO;2 |author=Jouve, S. |year=2004 |title=Description of the skull of a ''Ctenochasma'' (Pterosauria) from the latest Jurassic of eastern France, with a taxonomic revision of European Tithonian Pterodactyloidea |journal=Journal of Vertebrate Paleontology |volume=24 |issue=3 |pages=542–554 |s2cid=86019483}}</ref> Another species of ''Pterodactylus'' originally based on small, immature specimens was ''P. micronyx''. However, it has been difficult to determine exactly of what genus and species ''P. micronyx'' might be the juvenile form. Stéphane Jouve, Christopher Bennett and others had once suggested that it probably belonged either to ''[[Gnathosaurus|Gnathosaurus subulatus]]'' or one of the species belonging to ''Ctenochasma''.<ref name ="bennett2002" /><ref name="jouve2004" /> After additional research in 2013, Bennett assigned it to the genus ''Aurorazhdarcho'',<ref name=BennettPZ /> though a subsequent review by this researcher again proposed synonymy of ''P. micronyx'' with ''Gnathosaurus''.<ref name=Bennett2025>{{Cite journal |last1=Bennett |first1=S. Christopher |date=2025-03-11 |title=A review of the pterosaur ''Gnathosaurus subulatus'' from the Tithonian Solnhofen Lithographic Limestones of Germany: taxonomy and ontogeny |url=http://www.schweizerbart.de/papers/njgpa/detail/prepub/106471/A_review_of_the_pterosaur_Gnathosaurus_subulatus_f?af=crossref |journal=Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen |language=en |doi=10.1127/njgpa/2025/1245 |issn=0077-7749}}</ref> Another species with a complex history is ''P. longicollum'', named by von Meyer in 1854, based on a large specimen with a long neck and fewer teeth. Many researchers, including [[David Unwin]], have found ''P. longicollum'' to be distinct from ''P. kochi'' and ''P. antiquus''. Unwin found ''P. longicollum'' to be closer to ''Germanodactylus'' and therefore requiring a new genus name.<ref name="unwin2003" /> It has sometimes been placed in the genus ''Diopecephalus'' because [[Harry Govier Seeley]] based this genus partly on the ''P. longicollum'' material. However, it was shown by Bennett that the [[type specimen]] later designated for ''Diopecephalus'' was a fossil belonging to ''P. kochi'', and no longer thought to be separate from ''Pterodactylus''. ''Diopecephalus'' is therefore a synonym of ''Pterodactylus'', and as such is unavailable for use as a new genus for ''"P." longicollum''.<ref name=SCB06>{{cite journal |last=Bennett |first=S.C. |year=2006 |title=Juvenile specimens of the pterosaur ''Germanodactylus cristatus'', with a review of the genus |journal=Journal of Vertebrate Paleontology |volume=26 |issue=4 |pages=872–878 |doi=10.1671/0272-4634(2006)26[872:JSOTPG]2.0.CO;2 |s2cid=86460861}}</ref> ''"P." longicollum'' was eventually made the type species of a separate genus ''Ardeadactylus''.<ref name=BennettPZ />