Editing Pterodactylus (section)

==Classification==
[[File:Pterodactylus antiquus AMNH.jpg|thumb|A ''P. antiquus'' specimen and counterplate (AMNH 1942) showing muscle impressions in the chest and wing membranes]]
Initial classifications for ''Pterodactylus'' started when paleontologist Hermann von Meyer used the name Pterodactyli to contain ''Pterodactylus'' and other pterosaurs known at the time. This was emended to the [[Family (biology)|family]] [[Pterodactylidae]] by Prince [[Charles Lucien Bonaparte]] in 1838. However, this group has more recently been given several competing definitions.<ref name="unwin2003" /><ref>{{Cite journal |last=Kellner |first=Alexander W. A. |date=2003 |title=Pterosaur phylogeny and comments on the evolutionary history of the group |journal=Geological Society, London, Special Publications |series=Evolution and Palaeobiology of Pterosaurs |language=en |volume=217 |issue=1 |pages=105–137 |doi=10.1144/GSL.SP.2003.217.01.10 |bibcode=2003GSLSP.217..105K |s2cid=128892642 |issn=0305-8719}}</ref>

Beginning in 2014, researchers Steven Vidovic and David Martill constructed an analysis in which several pterosaurs traditionally thought of as [[archaeopterodactyloid]]s closely related to the [[ctenochasmatoid]]s may have been more closely related to the more advanced [[dsungaripteroid]]s, or in some cases, fall outside both groups. Their conclusion was published in 2017, in which they placed ''Pterodactylus'' as a basal member of the suborder [[Pterodactyloidea]].<ref name=W1837/>
{{clade| style=font-size:90%;line-height:90%
       |label1=[[Pterodactyloidea]]
        |1={{clade
         |1=''[[Eosipterus yangi]]''
          |2={{clade
            |1='''''Pterodactylus antiquus'''''[[File:Pterodactylus BMMS7 life.png|70 px]]
       |label2=[[Lophocratia]]
        |2={{clade
           |label1=[[Euctenochasmatia]]
            |1={{clade
              |1=''[[Diopecephalus kochi]]''
              |label2=[[Ctenochasmatoidea]]
               |2={{clade
                |1=[[Gallodactylidae]] [[File:Aerodactylus MCZ 1505.png|70 px]]
                |2=[[Ctenochasmatidae]] [[File:Pterodaustro BW.jpg|80px]] }} }}
             |label2=[[Eupterodactyloidea]]
              |2={{clade
                |1=''[[Altmuehlopterus ramphastinus]]''[[File:Altmuehlopterus DB.jpg|70 px]]
                |2=[[Dsungaripteroidea]] [[File:Pteranodon longiceps mmartyniuk wiki.png|50 px]] [[File:Quetzalcoatlus07.jpg|70px]]
}} }} }} }} }}

As illustrated below, the results of a different [[topology]] are based on a phylogenetic analysis made by Longrich, Martill, and Andres in 2018. Unlike the previous results above, they placed ''Pterodactylus'' within the clade [[Euctenochasmatia]], resulting in a more derived position.<ref name=longrichetal2018>{{cite journal |last1=Longrich |first1=N.R. |last2=Martill |first2=D.M. |last3=Andres |first3=B. |year=2018 |title=Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary |journal=PLOS Biology |volume=16 |issue=3 |page=e2001663 |doi=10.1371/journal.pbio.2001663 |pmid=29534059 |pmc=5849296 |doi-access=free}}</ref>
{{clade| style=font-size:90%;line-height:90%
|label1=[[Archaeopterodactyloidea]]
|1={{clade
  |label1=[[Germanodactylidae]]
   |1={{clade
     |1=''[[Germanodactylus cristatus]]''
     |2=''[[Germanodactylus rhamphastinus]]''[[File:Altmuehlopterus DB.jpg|70px]] }}
  |label2=[[Euctenochasmatia]]
   |2={{clade
      |1='''''Pterodactylus antiquus'''''[[File:Pterodactylus BMMS7 life.png|70px]]
     |label2=[[Ctenochasmatoidea]]
      |2={{clade
         |label1=[[Gallodactylidae]]
          |1={{clade
           |1=''[[Cycnorhamphus suevicus]]''
           |2=''[[Normannognathus wellnhoferi]]'' }}
         |2=[[Ctenochasmatidae]] [[File:Pterodaustro BW.jpg|70px]]
}} }} }} }}

===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 />

===Controversial species===
[[File:Bsp as xix.png|thumb|Juvenile type specimen of ''Pterodactylus kochi'', now reassigned as ''[[Diopecephalus kochi]]'']]
The only well-known and well-supported species left by the first decades of the 21st century were ''P. antiquus'' and ''P. kochi''. However, most studies between 1995 and 2010 found little reason to separate even these two species, and treated them as synonymous.<ref name="unwin2003">{{cite journal |last1=Unwin |first1=D. M. |year=2003 |title=On the phylogeny and evolutionary history of pterosaurs |journal=Geological Society, London, Special Publications |volume=217 |issue=1 |pages=139–190 |doi=10.1144/GSL.SP.2003.217.01.11 |bibcode=2003GSLSP.217..139U |s2cid=86710955}}</ref> More recent studies of pterosaur relationships have found anurognathids and pterodactyloids to be sister groups, which would limit the more inclusive group [[Caelidracones]] to just two clades.<ref name=LoneStarPterosaurs>{{Cite journal |last1=Andres |first1=B. |last2=Myers |first2=T. S. |doi=10.1017/S1755691013000303 |title=Lone Star Pterosaurs |journal=Earth and Environmental Science Transactions of the Royal Society of Edinburgh |pages=383–398 |year=2013 |volume=103 |issue=3–4 |s2cid=84617119}}</ref><ref name=SCB06 /> In 1996, Bennett suggested that the differences between specimens of ''P. kochi'' and ''P. antiquus'' could be explained by differences in age, with ''P. kochi'' (including specimens alternately classified in the species ''P. scolopaciceps'') representing an immature growth stage of ''P. antiquus''. In a 2004 paper, Jouve used a different method of analysis and recovered the same result, showing that the "distinctive" features of ''P. kochi'' were age-related, and using mathematical comparison to show that the two forms are different growth stages of the same species.<ref name="jouve2004" /> An additional review of the specimens published in 2013 demonstrated that some of the supposed differences between ''P. kochi'' and ''P. antiquus'' were due to measurement errors, further supporting their synonymy.<ref name=BennettPZ />

By the 2010s, a large body of research had been developed based on the idea that ''P. kochi'' and ''P. scolopaciceps'' were early growth stages of ''P. antiquus''. However, in 2014, two scientists began publishing research that challenged this paradigm. Steven Vidovic and David Martill concluded that differences between specimens of ''P. kochi'', ''P. scolopaciceps'', and ''P. antiquus'', such as different lengths of neck vertebrae, thinner or thicker teeth, more rounded skulls, and how far the teeth extended back in the jaws, were significant enough to separate them into three distinct species. Vidovic and Martill also performed a phylogenetic analysis which treated all relevant specimens as distinct units, and found that the ''P. kochi'' type specimen did not form a natural group with that of ''P. antiquus''. They concluded that the genus ''Diopecephalus'' could be returned to use to distinguish ''"P". kochi'' from ''P. antiquus''. They named the new genus ''[[Aerodactylus]]'' for ''P. scolopaciceps'' as well. So, what Bennett considered early growth stages of one species, Vidovic and Martill considered representatives of new species.<ref name=Aerodactylus>{{Cite journal |doi=10.1371/journal.pone.0110646 |title=''Pterodactylus scolopaciceps'' Meyer, 1860 (Pterosauria, Pterodactyloidea) from the Upper Jurassic of Bavaria, Germany: The Problem of Cryptic Pterosaur Taxa in Early Ontogeny |journal=PLOS ONE |volume=9 |issue=10 |pages=e110646 |year=2014 |last1=Vidovic |first1=S. U. |last2=Martill |first2=D. M. |pmid=25337830 |pmc=4206445 |bibcode=2014PLoSO...9k0646V |doi-access=free}}</ref><ref name=W1837>{{cite journal |last1=Vidovic |first1=Steven U. |last2=Martill |first2=David M. |title=The taxonomy and phylogeny of ''Diopecephalus kochi'' (Wagner, 1837) and ''"Germanodactylus rhamphastinus"'' (Wagner, 1851) |journal=Geological Society, London, Special Publications |volume=455 |date=2017 |issue=1 |pages=125–147 |url=https://eprints.soton.ac.uk/423063/1/Vidovic_Martill_2017_Taxonomy_of_Diopecephalus_and_Germanodactylus_AM_with_Figures.pdf |doi=10.1144/SP455.12 |bibcode=2018GSLSP.455..125V |s2cid=219204038}}</ref>

In 2017, Bennett challenged this hypothesis, he claimed that while Vidovic and Martill had identified real differences between these three groups of specimens, they had not provided any rationale that the differences were enough to distinguish them as species, rather than just individual variation, growth changes, or simply due to crushing and distortion during the fossilization process. Bennett pointed in particular to the data used to distinguish ''Aerodactylus'', which was so different from the data for related species, it might be due to an unnatural assemblage of specimens. As a result, Bennett continued to consider ''Diopecephalus'' and ''Aerodactylus'' simply as year-classes of immature ''Pterodactylus antiquus''.<ref name=bennett_2017_pteranodon_juv>{{cite journal |last1=Bennett |first1=S.C. |year=2017 |title=New smallest specimen of the pterosaur ''Pteranodon'' and ontogenetic niches in pterosaurs |journal=Journal of Paleontology |volume=92 |issue=2 |pages=1–18 |doi=10.1017/jpa.2017.84 |s2cid=90893067}}</ref>

===List of species===
During its over-200-year history, the various species of ''Pterodactylus'' have gone through a number of changes in classification and thus have acquired a large number of synonyms. Additionally, a number of species assigned to ''Pterodactylus'' are based on poor remains that have proven difficult to assign to one species or another and are therefore considered {{lang|la|nomina dubia}} ({{lit|doubtful names}}). The following list includes names that were used to identify new pterosaur species that now have been reclassified, or until recently thought to be pertaining to ''Pterodactylus'' proper, and names based on other material that has as yet not been assigned to other genera. This list also includes species that are {{lang|la|[[nomina nuda]]}} ('naked names'), which are species that were not published formally. Species that are {{lang|la|[[nomina oblita]]}} ('forgotten names') are the ones that have been disused, and species that are {{lang|la|[[nomina rejecta]]}} ('rejected names') are the ones that have been rejected because a more preferable name had been accepted instead.<ref name=Lydekker /><ref>{{cite book |first=Caroline |last=Arnold |title=Pterosaurs: Rulers of the Skies in the Dinosaur Age |year=2014 |publisher=StarWalk Kids Media |isbn=978-1-63083-412-8}}</ref>
{{collapse top|List of species}}
{| class="wikitable sortable"
|-
! Name
! Author
! Year
! Status
! class="unsortable" | Notes
|-
|''Pterodactylus antiquus''
|([[Samuel Thomas von Sömmerring|Sömmerring]])
|(1812)
|Valid
|Designated as the [[type species]] of ''Pterodactylus'', and replacing ''Ornithocephalus antiquus'' due to being an obsolete name
|-
|''Pterodactylus "suevicus"''
|Oken
|1816<ref>{{Cite book |last=Oken |first=Lorenz |url=https://www.biodiversitylibrary.org/item/276843 |title=Okens Lehrbuch der Naturgeschichte |date=1816 |publisher=August Schmid und Comp. |volume=3 |location=Leipzig |pages=312–314}}</ref>
|''[[Nomen nudum]]''
|
|-
|''Pterodactylus brevirostris''
|(Sömmerring)
|(1817)
|Synonym of ''[[Ctenochasma elegans]]''
|Reclassified from ''Ornithocephalus brevirostris''
|-
|''Pterodactylus giganteus''
|(Oken)
|(1819)
|''[[Nomen dubium]]''
|Synonym of ''[[Rhamphorhynchus muensteri]]''; reclassified from ''Ornithocephalus giganteus''
|-
|''Pterodactylus longirostris''
|[[Georges Cuvier|Cuvier]]
|1819
|Synonym of ''Pterodactylus antiquus''
|
|-
|''Pterodactylus grandis''
|Cuvier
|1824
|''Nomen dubium''
|Synonym of ''Rhamphorhynchus muensteri''
|-
|''Pterodactylus crocodilocephaloides''
|Ritgen
|1826
|Synonym of ''Pterodactylus antiquus''
|
|-
|''Pterodactylus nettecephaloides''
|Ritgen
|1826
|''[[Nomen oblitum]]''
|Synonym of ''[[Aurorazhdarcho micronyx]]''
|-
|''Pterodactylus macronyx''
|[[William Buckland|Buckland]]
|1829
|Reclassified as ''[[Dimorphodon macronyx]]''
|
|-
|''Pterodactylus (Rhamphorhynchus) macronyx''
|(Buckland)
|(1829)
|Synonym of ''Dimorphodon macronyx''
|Reclassified from ''Dimorphodon macronyx''
|-
|''Pterodactylus banthensis''
|(Theodori)
|(1830)
|Reclassified as ''[[Dorygnathus banthensis]]''
|Reclassified from ''Ornithocephalus banthensis''
|-
|''Pterodactylus (Rhamphorhynchus) (ensirostris) banthensis''
|(Theodori)
|(1830)
|''Nomen oblitum''
|Synonym of ''Dorygnathus banthensis''
|-
|''Pterodactylus goldfussi''
|Theodori
|1830
|Synonym of ''Dorygnathus banthensis''
|
|-
|''Pterodactylus crassirostris''
|[[Georg August Goldfuss|Goldfuss]]
|1831
|Reclassified as ''[[Scaphognathus crassirostris]]''
|
|-
|''Pterodactylus macronyx''
|[[Christian Erich Hermann von Meyer|Meyer]] ''non'' Buckland
|1831 ''non'' 1829
|Synonym of ''Dorygnathus banthensis''
|
|-
|''Pterodactylus medius''
|[[Georg zu Münster|Münster]]
|1831
|''Nomen dubium''
|Synonym of ''[[Altmuehlopterus rhamphastinus]]''
|-
|''Pterodactylus muensteri''
|(Goldfuss)
|(1831)
|Synonym of ''Rhamphorhynchus muensteri''
|Reclassified from ''Ornithocephalus muensteri''
|-
|''Pterodactylus (Rhamphorhynchus) muensteri''
|(Goldfuss)
|(1831)
|Synonym of ''Rhamphorhynchus muensteri''
|Reclassified from ''Pterodactylus muensteri''
|-
|''Pterodactylus bucklandi''
|Meyer
|1832
|Indeterminate [[Rhamphorhynchidae|rhamphorhynchine]]
|Once assigned to the genus ''[[Rhamphocephalus]]''
|-
|''Pterodactylus longipes''
|Münster
|1836
|Synonym of ''[[Ardeadactylus longicollum]]''
|
|-
|''Pterodactylus kochi''
|Wagner
|1837
|Reclassified as ''[[Diopecephalus kochi]]''
|Reclassified from ''Ornithocephalus kochi''; possible synonym of ''Pterodactylus antiquus'' (Bennett, 2013 <ref>{{Cite journal|last1=Vidovic|first1=Steven U. |last2=Martill|first2=David M.  |date=22 October 2014
|title=Pterodactylus scolopaciceps Meyer, 1860 (Pterosauria, Pterodactyloidea) from the Upper Jurassic of Bavaria, Germany: The Problem of Cryptic Pterosaur Taxa in Early Ontogeny|journal=PLOS ONE|volume=9|issue=10|pages=e110646|doi=10.1371/journal.pone.0110646|doi-access=free |pmid=25337830 |pmc=4206445 |bibcode=2014PLoSO...9k0646V }}</ref>) 
|-
|''Pterodactylus lavateri''
|Meyer
|1838
|Synonym of ''Rhamphorhynchus muensteri''
|
|-
|''Pterodactylus (Ornithopterus) lavateri''
|(Meyer)
|(1838)
|Synonym of ''Rhamphorhynchus muensteri''
|Reclassified from ''Pterodactylus lavateri''
|-
|''Pterodactylus longicaudus''
|Münster
|1839
|Reclassified as ''Rhamphorhynchus longicaudus''
|
|-
|''Pterodactylus meyeri''
|Münster
|1842
|Synonym of ''Diopecephalus kochi''
|
|-
|''Pterodactylus secundarius''
|Meyer
|1843
|''Nomen dubium''
|Synonym of ''Rhamphorhynchus muensteri''
|-
|''Pterodactylus cliftii''
|Mantell
|1844
|Synonym of ''[[Palaeornis cliftii]]''
|Reclassified from ''Palaeornis cliftii''
|-
|''Pterodactylus diomedeus''
|([[Richard Owen|Owen]])
|(1846)
|Synonym of ''[[Cimoliornis diomedeus]]''
|Reclassified from ''Cimoliornis diomedeus''
|-
|''Pterodactylus gemmingi''
|Meyer
|1846
|Synonym of ''Rhamphorhynchus muensteri''
|
|-
|''Pterodactylus (Rhamphorhynchus) gemmingi''
|(Meyer)
|(1846)
|Synonym of ''Rhamphorhynchus muensteri''
|Reclassified from ''Pterodactylus gemmingi''
|-
|''Pterodactylus giganteus''
|[[James Scott Bowerbank|Bowerbank]] ''non'' Oken
|1846 ''non'' 1819
|Reclassified as ''[[Lonchodraco giganteus]]''
|
|-
|''Pterodactylus ornis''
|[[Christoph Gottfried Andreas Giebel|Giebel]]
|1847
|Synonym of ''Palaeornis cliftii''
|
|-
|''Pterodactylus conirostris''
|Owen
|1850
|Synonym of ''Lonchodraco giganteus''
|
|-
|''Pterodactylus cuvieri''
|Bowerbank
|1851
|Reclassified as ''[[Cimoliopterus cuvieri]]''
|
|-
|''Pterodactylus compressirostris''
|Owen
|1851
|Reclassified as ''[[Lonchodectes compressirostris]]''
|
|-
|''Pterodactylus redenbacheri''
|(Wagner)
|(1851)
|Synonym of ''Aurorazhdarcho micronyx''
|Reclassified from ''Ornithocephalus redenbacheri''
|-
|''Pterodactylus rhamphastinus''
|(Wagner)
|(1851)
|Reclassified as ''Altmuehlopterus rhamphastinus''
|Reclassified from ''Ornithocephalus rhamphastinus''
|-
|''Pterodactylus giganteus''
|Morris ''non'' Oken ''non'' Bowerbank
|1854 ''non'' 1819 ''non'' 1846
|Synonym of ''Cimoliornis diomedeus''
|
|-
|''Pterodactylus longicollum''
|Meyer
|1854
|Reclassified as ''Ardeadactylus longicollum''
|
|-
|''Pterodactylus wuerttembergicus''
|[[Friedrich August Quenstedt|Quenstedt]]
|1854
|''Nomen oblitum''
|Synonym of ''[[Cycnorhamphus suevicus]]''
|-
|''Pterodactylus suevicus''
|Quenstedt
|1855
|Reclassified as ''Cycnorhamphus suevicus''
|
|-
|''Pterodactylus micronyx''
|Meyer
|1856
|Reclassified as ''Aurorazhdarcho micronyx''
|
|-
|''Pterodactylus crassipes''
|Meyer
|1857
|''[[Nomen rejectum]]''
|Reclassified as ''[[Ostromia crassipes]]''
|-
|''Pterodactylus (Rhamphorhynchus) crassipes''
|(Meyer)
|(1857)
|''Nomen rejectum''
|Synonym of ''Ostromia crassipes''; reclassified from ''Pterodactylus crassipes''
|-
|''Pterodactylus eurychirus''
|Wagner
|1857
|Synonym of ''Cycnorhamphus suevicus''
|
|-
|''Pterodactylus (Ornithocephalus) eurychirus''
|(Wagner)
|(1857)
|Synonym of ''Cycnorhamphus suevicus''
|Reclassified from ''Pterodactylus eurychirus''
|-
|''Pterodactylus suevicus eurychirus''
|(Wagner)
|(1857)
|Synonym of ''Cycnorhamphus suevicus''
|
|-
|''Pterodactylus hirundinaceus''
|Wagner
|1857
|Synonym of ''Rhamphorhynchus muensteri''
|
|-
|''Pterodactylus (Rhamphorhynchus) hirundinaceus''
|Wagner
|1857
|Synonym of ''Rhamphorhynchus muensteri''
|
|-
|''Pterodactylus (Ornithocephalus) propinquus''
|Wagner
|1857
|''Nomen dubium''
|Possible synonym of ''Altmuehlopterus rhamphastinus''
|-
|''Pterodactylus propinquus''
|(Wagner)
|(1857)
|''Nomen dubium''
|Reclassified from ''Pterodactylus (Ornithocephalus) propinquus''; possible synonym of ''Altmuehlopterus rhamphastinus''
|-
|''Pterodactylus propinquus medius''
|Wagner
|1857
|''Nomen dubium''
|Synonym of ''Altmuehlopterus rhamphastinus''
|-
|''Pterodactylus (Ornithocephalus) vulturinus''
|Wagner
|1857
|Synonym of ''Ardeadactylus longicollum''
|
|-
|''Pterodactylus vulturinus''
|(Wagner)
|(1857)
|Synonym of ''Ardeadactylus longicollum''
|Reclassified from ''Pterodactylus (Ornithocephalus) vulturinus''
|-
|''Pterodactylus liasicus''
|Quenstedt
|1858
|Reclassified as ''[[Campylognathoides liasicus]]''
|
|-
|''Pterodactylus fittoni''
|Owen
|1859
|''Nomen dubium''
|Reclassified as either ''[[Ornithocheirus]] fittoni'' or ''[[Anhanguera (pterosaur)|Anhanguera]] fittoni''; possible synonym of ''Cimoliopterus cuvieri''
|-
|''Pterodactylus sedgwickii''
|Owen
|1859
|Reclassified as ''[[Camposipterus sedgwickii]]''
|
|-
|''Pterodactylus silvestris''
|Owen
|1859
|Synonym of ''Palaeornis cliftii''
|
|-
|''Pterodactylus cerinensis''
|Meyer
|1860
|''Nomen dubium''
|
|-
|''Pterodactylus grandipelvis''
|Meyer
|1860
|''Nomen dubium''
|
|-
|''Pterodactylus scolopaciceps''
|Meyer
|1860
|Reclassified as ''[[Aerodactylus scolopaciceps]]''
|
|-
|''Pterodactylus elegans''
|Wagner
|1861
|Reclassified as ''Ctenochasma elegans''
|
|-
|''Pterodactylus pulchellus''
|Meyer
|1861
|Synonym of ''Aurorazhdarcho micronyx''
|
|-
|''Pterodactylus simus''
|Owen
|1861
|Reclassified as ''Ornithocheirus simus''
|
|-
|''Pterodactylus spectabilis''
|Meyer
|1861
|Synonym of ''Pterodactylus antiquus''
|
|-
|''Pterodactylus woodwardi''
|Owen
|1861
|''Nomen dubium''
|Reclassified as ''Ornithocheirus woodwardi''
|-
|''Pterodactylus hopkinsi''
|[[Harry Govier Seeley|Seeley]]
|1864
|''[[Nomen nudum]]''
|
|-
|''Pterodactylus oweni''
|Seeley
|1864
|Synonym of ''[[Ikrandraco machaerorhynchus]]''
|Not to be confused with the preoccupied species ''"Pterodactylus oweni"
|-
|''Pterodactylus longispinis''
|[[Edward Drinker Cope|Cope]]
|1866
|Reclassified as ''[[Rhabdopelix longispinis]]''
|
|-
|''Pterodactylus macrurus''
|Seeley
|1869
|Reclassified as ''[[Gnathosaurus macrurus]]''
|
|-
|''Pterodactylus nobilis''
|Owen
|1869
|''Nomen dubium''
|Reclassified as ''Ornithocheirus nobilis''
|-
|''Pterodactylus curtus''
|Owen
|1870
|''Nomen dubium''
|Reclassified as ''Ornithocheirus curtus''
|-
|''"Pterodactylus oweni"''
|[[Othniel Marsh|Marsh]]
|1871
|Preoccupied
|Reassigned as ''Pterodactylus occidentalis'' prior to being preoccupied
|-
|''Pterodactylus ingens''
|Marsh
|1872
|Synonym of ''[[Pteranodon longiceps]]''
|Reclassified as ''Pteranodon ingens''
|-
|''Pterodactylus occidentalis''
|Marsh
|1872
|''Nomen dubium''
|Reclassified from the preoccupied species ''"Pterodactylus oweni"''; reclassified as ''Pteranodon occidentalis''
|-
|''Pterodactylus velox''
|Marsh
|1872
|''Nomen dubium''
|Reclassified as ''Pteranodon velox''
|-
|''Pterodactylus suprajurensis''
|Sauvage
|1873
|''Nomen dubium''
|
|-
|''Pterodactylus aclandi''
|Owen
|1874
|''Nomen dubium''
|Possible synonym of ''Rhamphocephalus depressirostris''
|-
|''Pterodactylus daviesii''
|Owen
|1874
|''Nomen dubium''
|
|-
|''Pterodactylus duncani''
|Owen
|1874
|''Nomen dubium''
|Possible synonym of ''Rhamphocephalus bucklandi''
|-
|''Pterodactylus kiddii''
|Owen
|1874
|''Nomen dubium''
|Possible synonym of ''Rhamphocephalus bucklandi''
|-
|''Pterodactylus manseli''
|Owen
|1874
|''Nomen dubium''
|
|-
|''Pterodactylus marderi''
|Owen
|1874
|Synonym of ''Dimorphodon macronyx''
|
|-
|''Pterodactylus pleydelli''
|Owen
|1874
|''Nomen dubium''
|
|-
|''Pterodactylus sagittirostris''
|Owen
|1874
|Reclassified as ''[[Serradraco sagittirostris]]''
|
|-
|''Pterodactylus umbrosus''
|Cope
|1874
|''Nomen dubium''
|Reclassified from ''[[Ornithochirus umbrosus]]''
|-
|''Pterodactylus validus''
|Owen
|1875
|Reclassified as ''[[Doratorhynchus validus]]''
|
|-
|''Pterodactylus montanus''
|Marsh
|1878
|Reclassified as ''[[Dermodactylus montanus]]''
|
|-
|''Pterodactylus suevicus''
|[[Oscar Fraas|Fraas]] ''non'' Quenstedt
|1878 ''non'' 1855
|Synonym of ''Ardeadactylus longicollum''
|
|-
|''Pterodactylus fraasi''
|(Seeley)
|(1901)
|Synonym of ''Ardeadactylus longicollum''
|Reclassified from. ''Cycnorhamphus fraasi''
|-
|''Pterodactylus cristatus''
|[[Carl Wiman|Wiman]]
|1925
|Reclassified as ''[[Germanodactylus cristatus]]''
|
|-
|''Pterodactylus westmani''
|Wiman
|1925
|Synonym of ''Pterodactylus antiquus''
|
|-
|''Pterodactylus cormoranus''
|Döderlein
|1929
|Synonym of ''Pterodactylus antiquus''
|
|-
|''Pterodactylus arningi''
|[[Hans Reck|Reck]]
|1931
|''Nomen dubium''
|
|-
|''Pterodactylus brancai''
|Reck
|1931
|''Nomen dubium''
|Reclassified as ''[[Dsungaripterus brancai]]''
|-
|''Pterodactylus maximus''
|Reck
|1931
|''Nomen dubium''
|
|-
|''Pterodactylus raptor''
|Owen ''vide'' Ingles & Sawyer
|1979
|''Nomen nudum''
|
|-
|}
{{collapse bottom}}