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Old page wikitext, before the edit (old_wikitext) | '{{short description|Genus of pterodactyloid pterosaur from the Late Jurassic}}
{{Other uses|Pterodactyl (disambiguation)}}
{{hatnote|Pterodactyl is also an informal name for members of the order [[Pterosaur]]ia.}}
{{Use mdy dates|date=January 2021}}
{{Automatic taxobox
| fossil_range = Early [[Tithonian]], <br />{{fossil range|150.8|148.5}}
| image = Bsp as i 739 modified.png
| image_caption = Sub-adult [[type specimen]] of ''P. antiquus'', Bavarian State Colection for Palaeontology and Geology
| taxon = Pterodactylus
| authority = [[Constantine Samuel Rafinesque|Rafinesque]], 1815
| type_species = {{extinct}}'''''Ornithocephalus antiquus'''''
| type_species_authority = [[Samuel Thomas von Sömmerring|Sömmerring]], 1812
| subdivision_ranks = [[Species]]
| subdivision =
*{{extinct}}'''''P. antiquus''''' <br/><small>(Sömmerring, 1812)</small>
*{{extinct}}''[[Diopecephalus|P. kochi]]''? <br/><small>([[Johann Andreas Wagner|Wagner]], 1837)</small>
*{{extinct}}''[[Aerodactylus|P. scolopaciceps]]''? <br/><small>([[Hermann von Meyer|Meyer]], 1860)</small>
| synonyms =
{{collapsible list|bullets = true
|title=<small>Genus synonymy</small>
|''Ornithocephalus'' <br/><small>Sömmering, 1812</small>
|''Pterotherium'' <br/><small>Fischer, 1813<ref>{{Cite book|last=Fischer von Waldheim|first=Gotthelf|url=https://www.biodiversitylibrary.org/item/92502|title=Zoognosia tabulis synopticis illustrata : in usum praelectionum mperalis Medico-Chirurgicae Mosquenis edita|date=1813|publisher=Typis Nicolai S. Vsevolozsky|volume=1 |location=Mosquae [Moscow]|pages=466 }}</ref></small>
|''Macrotrachelus'' <br/><small>[[Christoph Gottfried Andreas Giebel|Giebel]], 1852</small>
|''[[Diopecephalus]]''? <br/><small>[[Harry Seeley|Seeley]], 1871</small>
|''[[Aerodactylus]]''? <br/><small>Vidovic & Martill, 2014</small>
}}
{{collapsible list|bullets = false
|title=<small>Species synonymy</small>
|''Ornithocephalus antiquus'' <br/><small>Sömmering, 1812</small>
|''Pterodactylus longirostris'' <br/><small>[[Georges Cuvier|Cuvier]], 1819</small>
|''Ornithocephalus longirostris'' <br/><small>(Cuvier, 1819)</small>
|''Macrotrachelus longirostris'' <br/><small>(Cuvier, 1819)</small>
|''Pterodactylus crocodilocephaloides'' <br/><small>Ritgen, 1826</small>
|''Ornithocephalus kochi''? <br/><small>Wagner, 1837</small>
|''Diopecephalus kochi''? <br/><small>(Wagner, 1837)</small>
|''Pterodactylus spectabilis'' <br/><small>Meyer, 1861</small>
|''Pterodactylus westmani'' <br/><small>Wiman, 1925</small>
|''Pterodactylus cormoranus'' <br/><small>Döderlein, 1929</small>
|''Aerodactylus scolopaciceps'' <br/><small>(Meyer, 1860)</small>
}}
}}
'''''Pterodactylus''''' (from Greek {{lang|grc-Latn|pterodáktylos}} ({{wikt-lang|grc|πτεροδάκτυλος}}) meaning 'winged finger'<ref>{{Cite journal|last=Gudger|first=E.W.|date=1944|title=The Earliest Winged Fish-Catchers|jstor=18398|journal=The Scientific Monthly|volume=59|issue=2|pages=120–129|bibcode=1944SciMo..59..120G}}</ref>) is an extinct [[genus]] of [[pterosaur]]s, whose members are commonly known as '''pterodactyls''' ({{IPAc-en|ˌ|t|ɛr|ə|ˈ|d|æ|k|t|ᵻ|l|z}}). It is thought to contain only a single [[species]], '''''Pterodactylus antiquus''''', which was the first pterosaur to be named and identified as a flying [[reptile]].
[[Fossil]] remains of ''Pterodactylus'' have primarily been found in the [[Solnhofen limestone]] of [[Bavaria]], [[Germany]], which dates back to the [[Late Jurassic]] period (early [[Tithonian]] stage), about 150.8 to 148.5 million years ago. More fragmentary remains of ''Pterodactylus'' have tentatively been identified from elsewhere in [[Europe]], as well as in [[Africa]].<ref name="ammonitedate">{{cite journal|author=Schweigert, G. |year=2007|title=Ammonite biostratigraphy as a tool for dating Upper Jurassic lithographic limestones from South Germany – first results and open questions|journal=Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen|volume=245|issue=1|pages= 117–125|doi=10.1127/0077-7749/2007/0245-0117}}</ref>
''Pterodactylus'' was a generalist [[carnivore]] that probably fed on a variety of invertebrates and vertebrates. Like all pterosaurs, ''Pterodactylus'' had wings formed by a skin and muscle membrane stretching from its elongated fourth finger to its hind limbs. It was supported internally by [[collagen]] fibres and externally by [[keratin]]ous ridges. ''Pterodactylus'' was a small pterosaur compared to other famous genera such as ''[[Pteranodon]]'' and ''[[Quetzalcoatlus]]'', and it also lived earlier, during the Late Jurassic period, while both ''Pteranodon'' and ''Quetzalcoatlus'' lived during the [[Late Cretaceous]]. ''Pterodactylus'' lived alongside other small pterosaurs such as the well-known ''[[Rhamphorhynchus]]'', as well as other genera such as ''[[Scaphognathus]]'', ''[[Anurognathus]]'' and ''[[Ctenochasma]]''. ''Pterodactylus'' is classified as an early-branching member of the [[Euctenochasmatia|ctenochasmatid lineage]], within the pterosaur clade [[Pterodactyloidea]].<ref name="unwin2003" /><ref name=longrichetal2018 />
==Discovery and history==
[[File:Pterodactylus holotype Collini 1784.jpg|thumb|left|Original copper engraving of the ''P. antiquus'' [[holotype]] by Egid Verhelst II and published by Italian scientist {{lang|it|italics=unset|[[Cosimo Alessandro Collini]]}}, 1784]]
The [[type specimen]] of the animal now known as ''Pterodactylus antiquus'' was the first pterosaur fossil ever to be identified. The first ''Pterodactylus'' specimen was described by the Italian scientist {{lang|it|italics=unset|[[Cosimo Alessandro Collini]]}} in 1784, based on a fossil skeleton that had been unearthed from the Solnhofen limestone of Bavaria. Collini was the curator of the {{lang|de|Naturalienkabinett}}, or nature [[cabinet of curiosities]] (a precursor to the modern concept of the [[natural history]] museum), in the palace of [[Charles Theodore, Elector of Bavaria]] at [[Mannheim]].<ref name=pester /><ref name=DU06>{{cite book |last=Unwin |first=David M. |title=The Pterosaurs: From Deep Time |year=2006 |publisher=Pi Press |location=New York |isbn=0-13-146308-X |pages=246}}</ref> The specimen had been given to the collection by Count {{lang|de|italics=unset|[[Pappenheim (state)|Friedrich Ferdinand zu Pappenheim]]}} around 1780, having been recovered from a [[lithographic limestone]] quarry in {{lang|de|italics=unset|[[Eichstätt]]}}.<ref name=brougham1844>{{cite journal | last1 =Brougham|first1= Henry P. |date=1844|title=Dialogues on instinct; with analytical view of the researches on fossil osteology|volume=19 |journal= Knight's Weekly Volume for All Readers|url=https://www.biodiversitylibrary.org/item/229475#page/5/mode/1up}}</ref> The actual date of the specimen's discovery and entry into the collection is unknown however, and it was not mentioned in a catalogue of the collection taken in 1767, so it must have been acquired at some point between that date and its 1784 description by Collini. This makes it potentially the earliest documented pterosaur find; the "Pester Exemplar" of the species ''[[Pterodactylus micronyx]]'' was described in 1779 and possibly discovered earlier than the Mannheim specimen, but it was at first considered to be a fossilized crustacean, and it was not until 1856 that this species was properly described as a pterosaur by German paleontologist {{lang|de|italics=unset|[[Christian Erich Hermann von Meyer|Hermann von Meyer]]}}.<ref name=pester>{{cite journal | last1 = Ősi | first1 = A. | last2 = Prondvai | first2 = E. | last3 = Géczy | first3 = B. | year = 2010 | title = The history of Late Jurassic pterosaurs housed in Hungarian collections and the revision of the holotype of ''Pterodactylus micronyx'' Meyer 1856 (a 'Pester Exemplar') | journal = Geological Society, London, Special Publications | volume = 343 | issue = 1| pages = 277–286 | url = https://www.researchgate.net/publication/258391479 |doi=10.1144/SP343.17 | bibcode = 2010GSLSP.343..277O | s2cid = 129805068 }}</ref>
[[File:Aquatic Pterodactylus.jpg|thumb|left|Wagler's 1830 restoration of an aquatic ''Pterodactylus'']]
In his first description of the Mannheim specimen, Collini did not conclude that it was a flying animal. In fact, Collini could not fathom what kind of animal it might have been, rejecting affinities with the birds or the bats. He speculated that it may have been a sea creature, not for any anatomical reason, but because he thought the ocean depths were more likely to have housed unknown types of animals.<ref name="collini1784">{{Cite journal |last=Collini| first=C A. |date=1784|title=Sur quelques Zoolithes du Cabinet d'Histoire naturelle de S. A. S. E. Palatine & de Bavière, à Mannheim|journal=Acta Theodoro-Palatinae Mannheim |volume=5 Physicum|pages= 58–103 (1 plate)|url=http://www.rhinoresourcecenter.com/index.php?s=1&act=pdfviewer&id=1292315455&folder=129|language=fr}}</ref><ref name="taquet&padian2004">{{Cite journal | doi = 10.1016/j.crpv.2004.02.002 |last1=Taquet|first1= P. |last2=Padian|first2= K. | year = 2004 | title = The earliest known restoration of a pterosaur and the philosophical origins of Cuvier's ''Ossemens Fossiles'' | journal = Comptes Rendus Palevol | volume = 3 | issue = 2| pages = 157–175 }}</ref> The idea that pterosaurs were aquatic animals persisted among a minority of scientists as late as 1830, when the German zoologist [[Johann Georg Wagler]] published a text on "amphibians" which included an illustration of ''Pterodactylus'' using its wings as flippers. Wagler went so far as to classify ''Pterodactylus'', along with other aquatic vertebrates (namely [[plesiosaur]]s, [[ichthyosaur]]s, and [[monotreme]]s), in the class Gryphi, between birds and mammals.<ref>{{Cite book|last=Wagler|first=Johann Georg|url=https://www.biodiversitylibrary.org/item/120243|title=Natürliches System der Amphibien : mit vorangehender Classification der Säugethiere und Vögel : ein Beitrag zur vergleichenden Zoologie|date=1830|location=München|language=de}}</ref>
[[File:Hermann pterodactylus restoration1.png|thumb|Hermann's original life restoration, the first of any pterosaur, 1800]]
The German/French scientist [[Johann Hermann]] was the one who first stated that ''Pterodactylus'' used its long fourth finger to support a wing membrane. Back in March 1800, Hermann alerted the prominent French scientist [[Georges Cuvier]] to the existence of Collini's fossil, believing that it had been captured by the occupying armies of [[Napoleon]] and sent to the French collections in [[Paris]] (and perhaps to Cuvier himself) as war booty; at the time special French [[political commissar]]s systematically seized art treasures and objects of scientific interest. Hermann sent Cuvier a letter containing his own interpretation of the specimen (though he had not examined it personally), which he believed to be a [[mammal]], including the first known life restoration of a pterosaur. Hermann restored the animal with wing membranes extending from the long fourth finger to the ankle and a covering of fur (neither wing membranes nor fur had been preserved in the specimen). Hermann also added a membrane between the neck and wrist, as is the condition in [[bat]]s. Cuvier agreed with this interpretation, and at Hermann's suggestion, Cuvier became the first to publish these ideas in December 1800 in a very short description.<ref name="taquet&padian2004" /> However, contrary to Hermann, Cuvier was convinced the animal was a [[reptile]].<ref name="cuvier1801">{{Cite journal | last= Cuvier|first= G. | year = 1801 | title =Extrait d'un ouvrage sur les espèces de quadrupèdes dont on a trouvé les ossemens dans l'intérieur de la terre |quote= Reptile volant| journal = Journal de Physique, de Chimie et d'Histoire Naturelle | volume = 52 | pages = 253–267 |url=https://gallica.bnf.fr/ark:/12148/bpt6k9604723x/f1.image|language=fr}}</ref> The specimen had not in fact been seized by the French. Rather, in 1802, following the death of Charles Theodore, it was brought to [[Munich]], where Baron [[Johann Paul Carl von Moll]] had obtained a general exemption of confiscation for the Bavarian collections.<ref name=pester /> Cuvier asked von Moll to study the fossil but was informed it could not be found. In 1809 Cuvier published a somewhat longer description, in which he named the animal ''Petro-Dactyle'',<ref name="cuvier1809">{{Cite journal | last= Cuvier|first= G. | year = 1809 | title = Mémoire sur le squelette fossile d'un reptile volant des environs d'Aichstedt, que quelques naturalistes ont pris pour un oiseau, et dont nous formons un genre de Sauriens, sous le nom de Petro-Dactyle | journal = Annales du Muséum national d'Histoire Naturelle, Paris | volume = 13 | pages = 424–437 |url=https://archive.org/details/cbarchive_48783_cuvierg1809mmoiresurlesquelett1809/mode/2up}}</ref> this was a typographical error however, and was later corrected by him to ''Ptéro-Dactyle''.<ref name="taquet&padian2004" /> He also refuted a hypothesis by [[Johann Friedrich Blumenbach]] that it would have been a shore bird.<ref name="cuvier1809" /> Cuvier remarked: "It is not possible to doubt that the long finger served to support a membrane that, by lengthening the anterior extremity of this animal, formed a good wing."<ref>Cuvier, G. (1809), p. 436 : "II n'est guère possible de douter que ce long doigt n'ait servi à supporter une membrane qui formoit ''[sic]'' à l'animal, d'après la longueur de l'extrémité antérieure, une aile bien plus puissante que celle du dragon, et au moins égale en force à celle de la chauve-souris."</ref>
[[File:Pterodactylus antiquus soemmerring.png|thumb|Von Sömmerring's incorrect 1817 restoration of the species ''[[Pterodactylus brevirostris]]'']]
Contrary to von Moll's report, the fossil was not missing; it was being studied by [[Samuel Thomas von Sömmerring]], who gave a public lecture about it on December 27, 1810. In January 1811, von Sömmerring wrote a letter to Cuvier deploring the fact that he had only recently been informed of Cuvier's request for information. His lecture was published in 1812, and in it von Sömmerring named the species '''''Ornithocephalus antiquus'''''.<ref name=sommerring1812>{{cite journal | last1 = von Sömmerring | first1 = S. T. | year = 1812 | title=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: vorgelesen in der mathematisch-physikalischen Classe am 27. Dec. 1810 und Nachtrag vorgelesen am 8. April 1811|journal=Denkschriften der Königlichen Bayerischen Akademie der Wissenschaften | url=http://publikationen.ub.uni-frankfurt.de/frontdoor/index/index/year/2008/docId/15367 | location= München | volume = 3 | pages = 89–158 }}</ref> The animal was described as being both a bat, and a form in between mammals and birds, i.e. not intermediate in descent but in "affinity" or [[archetype]]. Cuvier disagreed, and the same year in his ''Ossemens fossiles'' provided a lengthy description in which he restated that the animal was a reptile.<ref>{{Cite book|last=Cuvier|first=G.|url=https://www.biodiversitylibrary.org/item/123716#page/660/mode/1up|title=Recherches sur les ossemens fossiles de quadrupèdes : où l'on rétablit les caractères de plusieurs espèces d'animaux que les révolutions du globe paroissent avoir détruites|date=1812|publisher=Deterville|volume=Tome 4|location=Paris|language=fr|chapter=Article V – Sur le squelette fossile d’un reptile volant des environs d'Aichstedt, que quelques naturalistes ont pris pour un oiseau et dont nous formons un genre de sauriens, sous le nom de ptéro-dactyle|doi=10.5962/bhl.title.60807}}</ref> It was not until 1817 that a second specimen of ''Pterodactylus'' came to light, again from [[Solnhofen]]. This tiny specimen was that year described by von Sömmerring as ''[[Ornithocephalus brevirostris]]'', named for its short snout, now understood to be a juvenile character (this specimen is now thought to represent a juvenile specimen of a different genus, probably ''[[Ctenochasma]]'').<ref name=BennettPZ /> He provided a restoration of the skeleton, the first one published for any pterosaur.<ref name="taquet&padian2004" /> This restoration was very inaccurate, von Sömmerring mistaking the long [[metacarpal]]s for the bones of the lower arm, the lower arm for the [[humerus]], this upper arm for the [[breast bone]] and this sternum again for the [[shoulder blade]]s.<ref>{{Cite journal|last=von Sömmerring|first=S.T.|date=1817|title=Ueber einen Ornithocephalus brevirostris der Vorwelt|url=http://publikationen.ub.uni-frankfurt.de/frontdoor/index/index/year/2008/docId/15330|journal=Denkschriften der Bayerischen Akademie der Wissenschaften|language=de|volume=6|pages=89–104}}</ref> Sömmerring did not change his opinion that these forms were bats and this "bat model" for interpreting pterosaurs would remain influential long after a consensus had been reached around 1860 that they were reptiles. The standard assumptions were that pterosaurs were quadrupedal, clumsy on the ground, furred, warmblooded and had a wing membrane reaching the ankle. Some of these elements have been confirmed, some refuted by modern research, while others remain disputed.<ref>{{cite book|last=Padian|first= K. |chapter=The case of the bat-winged pterosaur. Typological taxonomy and the influence of pictorial representation on scientific perception | editor-last1 = Czerkas |editor-first1 = S. J.|editor-last2 =Olson | editor-first2 = E. C.| title = Dinosaurs past and present | publisher = Natural History Museum of Los Angeles County in association with University of Washington Press, Seattle and London | location = Los Angeles | year = 1987 | isbn = 978-0-938644-23-1|pages= 65–81|volume=2}}</ref>
In 1815, the generic name ''Ptéro-Dactyle'' was latinized to ''Pterodactylus'' by [[Constantine Samuel Rafinesque]].<ref>{{cite book|last=Rafinesque|first=C.S.|author-link=Constantine Samuel Rafinesque|year=1815|title=Analyse de la nature, ou tableau de l'univers et des corps organisés|edition=L'Imprimerie de Jean Barravecchia|page=224|url=https://www.biodiversitylibrary.org/page/48310144#page/10/mode/1up}}</ref> Unaware of Rafinesque's publication however, Cuvier himself in 1819 latinized the name ''Ptéro-Dactyle'' again to ''Pterodactylus'',<ref name=":0">{{Cite book|last=Cuvier|first=G.|url=http://archive.org/details/isisoderencyclop1819oken|title=Isis (oder Encyclopädische Zeitung) von Oken|date=1819|publisher=Jena : Expedition der Isis|editor-last=Oken|editor-first=Lorenz|pages=[https://archive.org/details/isisoderencyclop1819oken/page/n586/mode/1up 1126] ; [https://archive.org/details/isisoderencyclop1819oken/page/n914/mode/1up 1788]|language=de|chapter=Pterodactylus longirostris}}</ref> but the specific name he then gave, ''longirostris'', has to give precedence to von Sömmerring's ''antiquus''.<ref name=":0" /> In 1888, English [[naturalist]] [[Richard Lydekker]] designated ''Pterodactylus antiquus'' as the [[type species]] of ''Pterodactylus'', and considered ''Ornithocephalus antiquus'' a synonym. He also designated specimen BSP AS.I.739 as the [[holotype]] of the genus.<ref name=Lydekker />
==Description==
[[File:Pterodactylus scale mmartyniuk wiki.png|thumb|upright|left|Size of the sub-adult holotype (blue) and adult (green) specimens in both flying and standing postures, compared with a human]]
''Pterodactylus'' is known from over 30 fossil specimens, and though most belong to juveniles, many preserve complete skeletons.<ref name=BennettPZ /><ref name=bennett1996a>{{Cite journal| doi = 10.1080/02724634.1996.10011332 | author = Bennett, S.C. | year = 1996 | title = Year-classes of pterosaurs from the Solnhofen Limestone of Germany: Taxonomic and Systematic Implications | journal = Journal of Vertebrate Paleontology | volume = 16 | issue = 3| pages = 432–444}}</ref> ''Pterodactylus antiquus'' was a relatively small pterosaur, with an estimated adult wingspan of about {{convert|1.04|m|ftin|sp=us}}, based on the only known adult specimen, which is represented by an isolated skull.<ref name=BennettPZ /> Other "species" were once thought to have been smaller.<ref name=Lydekker>{{cite book|last=Lydekker|first=Richard|year=1888|title=Catalogue of Fossil Reptilia and Amphibia in the British Museum (Natural History). Part I. Containing the Orders Ornithosauria, Crocodilia, Dinosauria, Squamata, Rhynchocephalia and Pterosauria|publisher=Taylor and Francis|pages=2–37|url=https://www.biodiversitylibrary.org/item/125711#page/1/mode/1up}}</ref> However, these smaller specimens have been shown to represent juveniles of ''Pterodactylus'', as well as its contemporary relatives including ''Ctenochasma'', ''[[Germanodactylus]]'', ''[[Aurorazhdarcho]]'', ''[[Gnathosaurus]]'', and hypothetically ''[[Aerodactylus]]'' if this genus is truly valid.<ref name ="bennett2002">{{Cite journal | doi = 10.1671/0272-4634(2002)022[0043:STPOTC]2.0.CO;2 | author = Bennett, S.C. | year = 2002 | title = Soft tissue preservation of the cranial crest of the pterosaur ''Germanodactylus'' from Solnhofen|jstor=4524192 | journal = Journal of Vertebrate Paleontology | volume = 22 | issue = 1| pages = 43–48 }}</ref>
The skulls of adult ''Pterodactylus'' were long and thin, with about 90 narrow and conical teeth. The teeth extended back from the tips of both jaws, and became smaller farther away from the jaw tips, this was unlike the ones seen in most relatives, where teeth were absent in the upper jaw tip, and were relatively uniform in size. The teeth of ''Pterodactylus'' also extended farther back into the jaw compared to close relatives, and some were present below the front of the ''nasoantorbital fenestra'', which is the largest opening in the skull.<ref name=BennettPZ /> Another [[autapomorphy]] that ''Pterodactylus'' has is that the skull and jaws were straight, which are unlike the upwardly curved jaws seen in the related [[ctenochasmatid]]s.<ref name="jouve2004" />
[[File:Pterodactylus BMMS7 life.png|thumb|Life [[Paleoart|restoration]] of BMMS 7, the largest known ''Pterodactylus'' specimen. The shape of the soft tissue crest is based on the specimen BSP 1929 I 18]]
''Pterodactylus'', like related pterosaurs, had a crest on its skull composed mainly of soft tissues. In adult ''Pterodactylus'', this crest extended between the back edge of the [[antorbital fenestra]] and the back of the skull. In at least one specimen, the crest had a short bony base, also seen in related pterosaurs like ''Germanodactylus''. Solid crests have only been found on large, fully adult specimens of ''Pterodactylus'', indicating that this was a display structure that became larger and more well developed as individuals reached maturity.<ref name=BennettPZ>{{cite journal |year=2013 |title=New information on body size and cranial display structures of ''Pterodactylus antiquus'', with a revision of the genus |journal=Paläontologische Zeitschrift |volume=87|issue=2 |doi=10.1007/s12542-012-0159-8 |author=Bennett, S. Christopher |pages=269–289|s2cid=83722829 }}</ref><ref name="frey&martill1998">{{Cite journal |last1=Frey|first1= E. |last2=Martill|first2= D.M. | year = 1998 | title = Soft tissue preservation in a specimen of ''Pterodactylus kochi'' (Wagner) from the Upper Jurassic of Germany | journal = Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen | volume = 210 |issue=3 | pages = 421–441 |doi=10.1127/njgpa/210/1998/421 }}</ref> In 2013, pterosaur researcher [[S. Christopher Bennett]] noted that other authors claimed that the soft tissue crest of ''Pterodactylus'' extended backward behind the skull; Bennett himself, however, didn't find any evidence for the crest extending past the back of the skull.<ref name=BennettPZ /> Two specimens of ''P. antiquus'' (the holotype specimen BSP AS I 739 and the incomplete skull BMMS 7, the largest known skull of ''P. antiquus'') have a low bony crest on their skulls; in BMMS 7 it is 47.5 mm long (1.87 inches, more or less 24% of the estimated total length of its skull) and has a maximum height of 0.9 mm (0.035 inches) above the orbit.<ref name=BennettPZ /> Several specimens previously referred to ''P. antiquus'' preserved evidence of the soft tissue extensions of these crests, including an "occipital lappet", a flexible, tab-like structure extending from the back of the skull. Most of these specimens have been reclassified in the related species ''Aerodactylus scolopaciceps'', which may however be nothing more than a junior synonym. Even if ''Aerodactylus'' were valid, at least one specimen with these features is still considered to belong to ''Pterodactylus'', BSP 1929 I 18, which has an occipital lappet similar to the proposed ''Aerodactylus'' definition, and also possesses a small triangular soft tissue crest with the peak of the crest positioned above the eyes.<ref name=BennettPZ />
==Paleobiology==
===Life history===
[[File:Pterodactylus spectabilis 4.JPG|thumb|left|Holotype specimen of the species ''P. spectabilis'', now considered a juvenile specimen of ''P. antiquus'']]
Like other pterosaurs (most notably ''[[Rhamphorhynchus (pterosaur)|Rhamphorhynchus]]''), ''Pterodactylus'' specimens can vary considerably based on age or level of maturity. Both the proportions of the limb bones, size and shape of the skull, and size and number of teeth changed as the animals grew. Historically, this has led to various growth stages (including growth stages of related pterosaurs) being mistaken for new species of ''Pterodactylus''. Several detailed studies using various methods to measure growth curves among known specimens have suggested that there is actually only one valid species of ''Pterodactylus'', ''P. antiquus''.<ref name="jouve2004" />
The youngest immature specimens of ''Pterodactylus antiquus'' (alternately interpreted as young specimens of the distinct species ''P. kochi'') have a small number of teeth, as few as 15 in some, and the teeth have a relatively broad base.<ref name=bennett1996a /> The teeth of other ''P. antiquus'' specimens are both narrower and more numerous (up to 90 teeth are present in several specimens).<ref name="jouve2004" />
''Pterodactylus'' specimens can be divided into two distinct year classes. In the first year class, the skulls are only {{convert|15|to|45|mm|in|sp=us}} in length. The second year class is characterized by skulls of around {{convert|55|to|95|mm|in|sp=us}} long, but are still immature however. These first two size groups were once classified as juveniles and adults of the species ''P. kochi'', until further study showed that even the supposed "adults" were immature, and possibly belong to a distinct genus. A third year class is represented by specimens of the "traditional" ''P. antiquus'', as well as a few isolated, large specimens once assigned to ''P. kochi'' that overlap ''P. antiquus'' in size. However, all specimens in this third year class also show sign of immaturity. Fully mature ''Pterodactylus'' specimens remain unknown, or may have been mistakenly classified as a different genus.<ref name=bennett1996a />
===Growth and breeding seasons===
[[File:Pterodactylus grandipelvis Teylers 2.JPG|thumb|Fossil pelvis of a large specimen, now referred to the dubious species ''P. grandipelvis'']]
The distinct year classes of ''Pterodactylus antiquus'' specimens show that this species, like the contemporary ''Rhamphorhynchus muensteri'', likely bred seasonally and grew consistently during its lifetime. A new generation of 1st year class ''P. antiquus'' would have been produced seasonally, and reached 2nd-year size by the time the next generation hatched, creating distinct 'clumps' of similarly-sized and aged individuals in the fossil record. The smallest size class probably consisted of individuals that had just begun to fly and were less than one year old.<ref name=bennett1996a /><ref name="Wellnhofer1970">{{cite book|last=Wellnhofer|first=Peter|url=https://www.zobodat.at/pdf/Abhandlungen-Akademie-Bayern_NF_141_0001-0133.pdf|title=Die Pterodactyloidea (Pterosauria) der Oberjura-Plattenkalke Süddeutschlands|publisher=Bayerische Akademie der Wissenschaften, Mathematisch-Wissenschaftlichen Klasse, Abhandlungen|year=1970|volume=141|page=133}}</ref> The second year class represents individuals one to two years old, and the rare third year class is composed of specimens over two years old. This growth pattern is similar to modern [[crocodilia]]ns, rather than the rapid growth of modern [[bird]]s.<ref name=bennett1996a />
===Daily activity patterns===
Comparisons between the [[sclerotic ring|scleral rings]] of ''Pterodactylus antiquus'' and modern birds and reptiles suggest that it may have been [[Diurnality|diurnal]]. This may also indicate [[niche partitioning]] with contemporary pterosaurs inferred to be [[nocturnal]], such as ''Ctenochasma'' and ''Rhamphorhynchus''.<ref>{{cite journal|author1=Schmitz, L. |author2=Motani, R. |year=2011 |title=Nocturnality in Dinosaurs Inferred from Scleral Ring and Orbit Morphology |journal=Science |volume=332 |issue= 6030|pages= 705–8|doi=10.1126/science.1200043 |pmid=21493820 |bibcode=2011Sci...332..705S|s2cid=33253407 |url=http://pdfs.semanticscholar.org/8922/c462d9cd7f64033fe9989f5a1dc0e7e8aac9.pdf |archive-url=https://web.archive.org/web/20190228084617/http://pdfs.semanticscholar.org/8922/c462d9cd7f64033fe9989f5a1dc0e7e8aac9.pdf |url-status=dead |archive-date=2019-02-28 }}</ref>
===Diet===
Based on the shape, size, and arrangement of its teeth, ''Pterodactylus'' has long been recognized as a carnivore specializing in small animals. A 2020 study of pterosaur tooth wear supported the hypothesis that ''Pterodactylus'' preyed mainly on invertebrates and had a generalist feeding strategy, indicated by a relatively high bite force.<ref name=diet2020>Bestwick, J., Unwin, D.M., Butler, R.J. et al. Dietary diversity and evolution of the earliest flying vertebrates revealed by dental microwear texture analysis. Nat Commun 11, 5293 (2020). https://doi.org/10.1038/s41467-020-19022-2</ref>
==Paleoecology==
[[File:Pterodactylus holotype fly mmartyniuk.png|thumb|left|Hypothetical restoration of ''Pterodactylus'' in its environment]]
Specimens of ''Pterodactylus'' have been found mainly in the [[Solnhofen limestone]] (geologically known as the Altmühltal Formation) of [[Bavaria]], [[Germany]]. The main composition of this formation is fine-grained [[limestone]] that originated mainly from the nearby towns [[Solnhofen]] and Eichstätt, which is formed by mud [[silt]] deposits.<ref name="ammonitedate" /> The Solnhofen Limestone is a diverse [[Lagerstätte]] that contains a wide range of different creatures, including highly detailed fossilized imprints of soft bodied organisms such as [[jellyfish]]es. Abundant specimens of pterosaurs similar to ''Pterodactylus'' were also found within the formation, these include the [[rhamphorhynchid]]s ''Rhamphorhynchus'' and ''[[Scaphognathus]]'',<ref>{{Cite journal|last=Bennett|first=S. C.|date=2004|title=New information on the pterosaur ''Scaphognathus crassirostris'' and the pterosaurian cervical series|journal=Journal of Vertebrate Paleontology|language=en|volume=24|issue=Supplement 003|pages=38A|doi=10.1080/02724634.2004.10010643|s2cid=220415208|issn=0272-4634}}</ref> several [[gallodactylid]]s such as ''Aerodactylus'',<ref name=bennett1996a /> ''[[Ardeadactylus]]'', ''Aurorazhdarcho'' and ''[[Cycnorhamphus]]'',<ref name=DU06 /> the [[ctenochasmatid]]s ''Ctenochasma''<ref name=bennett2007>{{cite journal | last1 = Bennett | first1 = S.C. | year = 2007 | title = A review of the pterosaur ''Ctenochasma'': taxonomy and ontogeny | journal = Neues Jahrbuch für Geologie und Paläontologie – Abhandlungen | volume = 245 | issue = 1| pages = 23–31 | doi=10.1127/0077-7749/2007/0245-0023}}</ref> and ''Gnathosaurus'', the [[anurognathid]] ''[[Anurognathus]]'', the [[germanodactylid]] ''Germanodactylus'', as well as the basal [[euctenochasmatia]]n ''[[Diopecephalus]]''.<ref name=W1837 /> Fossil remains of the [[dinosaur]]s ''[[Archaeopteryx]]'' and ''[[Compsognathus]]'' were also found within the limestone, these specimens were related to early evolution of [[feathers]], since they were some of the only ones that had them during the Jurassic period.<ref name="FasWeis04">{{cite book|title=The Evolution and Extinction of the Dinosaurs|year=2005|chapter=Theropoda I: nature red in tooth and claw|editor-last1=Fastovsky| editor-first1=D.E.|editor-last2=Weishampel|editor-first2=D.B. |last1=Fastovsky|first1=D.E.|last2=Weishampel |first2=D.B. |pages=265–299|publisher=Cambridge University Press|isbn=978-0-521-81172-9|edition=2nd|doi=10.1017/9781316471623.009}}</ref> Various [[lizard]] remains were also found alongside those of ''Pterodactylus'', with several specimens assigned to ''[[Ardeosaurus]]'', ''[[Bavarisaurus]]'' and ''[[Eichstaettisaurus]]''.<ref name="hoffstetter1966">{{cite journal |last=Hoffstetter |first=R. |year=1966 |title=A propos des genres ''Ardeosaurus'' et ''Eichstaettisaurus'' (Reptilia, Sauria, Gekkonoidea) du Jurassique Supèrieur de Franconie |trans-title=On the genera ''Ardeosaurus'' and ''Eichstaettisaurus'' (Reptilia, Sauria, Gekkonoidea) from the Upper Jurassic of France |journal=Bulletin de la Société Géologique de France |volume=8 |issue=4 |pages=592–595 |doi=10.2113/gssgfbull.S7-VIII.4.592 |url=https://pubs.geoscienceworld.org/sgf/bsgf/article-abstract/S7-VIII/4/592/518861/A-propos-des-genres-Ardeosaurus-et}}</ref><ref>{{cite journal |last= Evans|first= S.E.|year=1994|title= The Solnhofen (Jurassic: Tithonian) lizard genus ''Bavarisaurus'': new skull material and a reinterpretation |journal=Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen |volume=192 |pages=37–52}}</ref> [[Crocodylomorph]] specimens were widely distributed within the fossil site, most were assigned to the [[metriorhynchid]] genera ''[[Cricosaurus]]'', ''[[Dakosaurus]]'',<ref>{{Cite journal|last1=Brandalise de Andrade|first1=Marco|last2=Young|first2=Mark T.|last3=Desojo|first3=Julia B.|last4=Brusatte|first4=Stephen L.|date=2010|title=The evolution of extreme hypercarnivory in Metriorhynchidae (Mesoeucrocodylia: Thalattosuchia) based on evidence from microscopic denticle morphology|journal=Journal of Vertebrate Paleontology|volume=30|issue=5|pages=1451–1465|doi=10.1080/02724634.2010.501442|s2cid=83985855|issn=0272-4634}}</ref> ''[[Geosaurus]]'' and ''[[Rhacheosaurus]]''.<ref name="Andrade & Young, 2008">{{Cite journal|last1=Brandalise de Andrade|first1=Marco|last2=Young|first2=Mark T.|year=2008|url=http://www.svpca.org/general/pages/abstractPage.php?i=1378&r=talksAndPosters.php&y=2008 |title= High diversity of thalattosuchian crocodylians and the niche partition in the Solnhofen Sea |archive-url=https://web.archive.org/web/20110603232400/http://www.svpca.org/general/pages/abstractPage.php?i=1378&r=talksAndPosters.php&y=2008 |archive-date=June 3, 2011 |journal= The 56th Symposium of Vertebrate Palaeontology and Comparative Anatomy|pages=14–15}}</ref> These genera are colloquially called as marine or sea crocodiles due to their similar built.<ref>{{Cite journal|last1=Young|first1=Mark T.|last2=Brusatte|first2=Stephen L.|last3=de Andrade|first3=Marco Brandalise|last4=Desojo|first4=Julia B.|last5=Beatty|first5=Brian L.|last6=Steel|first6=Lorna|last7=Fernández|first7=Marta S.|last8=Sakamoto|first8=Manabu|last9=Ruiz-Omeñaca|first9=Jose Ignacio|last10=Schoch|first10=Rainer R.|date=September 18, 2012|editor-last=Butler|editor-first=Richard J.|title=The Cranial Osteology and Feeding Ecology of the Metriorhynchid Crocodylomorph Genera Dakosaurus and Plesiosuchus from the Late Jurassic of Europe|journal=PLOS ONE|language=en|volume=7|issue=9|pages=e44985|doi=10.1371/journal.pone.0044985|issn=1932-6203|pmc=3445579|pmid=23028723|bibcode=2012PLoSO...744985Y}}</ref> The turtle genera ''[[Eurysternum]]'' and ''[[Paleomedusa]]'' were also found within the formation.<ref name="WGJ">{{cite journal|first=Walter G.|last= Joyce|year=2003|title=A new Late Jurassic turtle specimen and the taxonomy of ''Palaeomedusa testa'' and ''Eurysternum wagleri''|journal=PaleoBios|volume=23|issue=3|pages=1–8|url=http://www.geo.uni-tuebingen.de/fileadmin/website/arbeitsbereich/palaeo/biogeologie/Images/Joyce_Publications/3__Joyce_2003.pdf|archive-url=https://web.archive.org/web/20151001071809/http://www.geo.uni-tuebingen.de/fileadmin/website/arbeitsbereich/palaeo/biogeologie/Images/Joyce_Publications/3__Joyce_2003.pdf|url-status=dead|archive-date=October 1, 2015}}</ref> Fossils of the [[ichthyosaur]] ''[[Aegirosaurus]]'' also appeared to be present in the site,<ref>{{Cite journal|last1=Bardet|first1=Nathalie|last2=Fernández|first2=Marta S.|date=2000|title=A new ichthyosaur from the Upper Jurassic lithographic limestones of Bavaria|url=https://www.researchgate.net/publication/40662676|journal=Journal of Paleontology|language=en|volume=74|issue=3|pages=503–511|doi=10.1017/S0022336000031760|issn=0022-3360}}</ref> as well as fish remains, with many specimens assigned to [[ray-finned fish]]es such as the [[halecomorph]]s ''[[Lepidotes]]'',<ref name="lamberstable-others"/> ''[[Propterus]]'',<ref name=Marine /> ''[[Gyrodus]]'', ''[[Mesturus]]'', ''[[Proscinetes]]'', ''[[Caturus]]'',<ref name=Marine>{{cite journal|last=Sepkoski |first=Jack |title=A compendium of fossil marine animal genera |journal=Bulletins of American Paleontology |volume=364 |page=560 |year=2002 |url=http://strata.ummp.lsa.umich.edu/jack/showgenera.php?taxon=611&rank=class |access-date=February 27, 2009 |url-status=dead |archive-url=https://web.archive.org/web/20110723131237/http://strata.ummp.lsa.umich.edu/jack/showgenera.php?taxon=611&rank=class |archive-date=July 23, 2011 }}</ref> ''[[Ophiopsis]]''<ref>{{Cite journal|last1=Lane|first1=Jennifer A.|last2=Ebert|first2=Martin|date=2015|title=A taxonomic reassessment of Ophiopsis (Halecomorphi, Ionoscopiformes), with a revision of Upper Jurassic species from the Solnhofen Archipelago, and a new genus of Ophiopsidae|journal=Journal of Vertebrate Paleontology|language=en|volume=35|issue=1|pages=e883238|doi=10.1080/02724634.2014.883238|s2cid=86350086|issn=0272-4634}}</ref> and ''[[Ophiopsiella]]'',<ref name="lamberstable-others">{{Cite journal|last=Lambers|first=Paul H.|date=1999|title=The actinopterygian fish fauna of the Late Kimmeridgian and Early Tithonian 'Plattenkalke' near Solnhofen (Bavaria, Germany): state of the art|journal=Geologie en Mijnbouw |volume=78|issue=2|pages=215–229 |doi=10.1023/A:1003855831015|s2cid=127676896|quote=Table 1: List of actinopterygians from the Solnhofen lithographic limestone – Halecomorphi; 'Amiidae' and 'Ophiopsidae'" (p. 216)}}</ref> the [[Pachycormidae|pachycormids]] ''[[Asthenocormus]]'', ''[[Hypsocormus]]'' and ''[[Orthocormus]]'',<ref name=EoDP>{{cite book |editor=Palmer, D.|year=1999 |title= The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals|publisher= Marshall Editions|location=London|page= 38|isbn= 1-84028-152-9}}</ref> as well as the [[Aspidorhynchidae|aspidorhynchid]] ''[[Aspidorhynchus]]'', and the [[ichthyodectid]] ''[[Thrissops]]''.<ref name=FT12>{{cite journal |last1=Frey |first1=E. |last2=Tischlinger|first2= H. |year=2012 |title=The Late Jurassic pterosaur ''Rhamphorhynchus'', a frequent victim of the ganoid fish ''Aspidorhynchus''? |journal=PLOS ONE |volume=7 |issue=3 |pages=e31945 |doi=10.1371/journal.pone.0031945 |bibcode = 2012PLoSO...7E1945F |pmid=22412850 |pmc=3296705}}</ref><ref>{{Cite book|last=Nybelin|first=Orvar|title=Versuch einer taxonomischen Revision der jurassischen Fischgattung Thrissops Agassiz.|date=1964|publisher=Wettergren and Kerber|location=Göteborg|language=de|oclc=2672427}}</ref>
==Classification==
[[File:Pterodactylus muscle impressions.jpg|thumb|upright|A ''P. antiquus'' specimen (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 }}</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|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 |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=":1">{{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}}</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 if 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=":1" /> 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, M.P.|year=2010|title=''Pteranodon'' and beyond: The history of giant pterosaurs from 1870 onwards. Geological Society of London Special Publications|volume=343|pages=313–323|url=https://www.researchgate.net/publication/258391482}}</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 }}</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" /> though after additional research Bennett assigned it to the genus ''Aurorazhdarcho''.<ref name=BennettPZ /> 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 }}</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 }}</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}}</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|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 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''
|-
|''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}}
==Cultural significance==
[[File:Pteranodon longiceps mmartyniuk wiki.png|upright|thumb|Life reconstruction of a ''Pteranodon'', popularly called "pterodactyl"]]
''Pterodactylus'' is regarded as one of the most iconic prehistoric creatures, with multiple appearances in books, movies, as well as television series and several videogames. The informal name "'''pterodactyl'''" is sometimes used to refer to any kind of animal belonging to the order [[Pterosauria]], though most of the times to ''Pterodactylus'', as it's the most well-known member of the group.<ref name="myths">{{cite web|url=http://www.pterosaur.net/myths.php|title=Pterosaurs: Myths and Misconceptions|author=Naish, Darren |publisher=Pterosaur.net|access-date=June 18, 2011}}</ref> The popular aspect of ''Pterodactylus'' consists of an elongated head crest, and potentially large wings. Studies of ''Pterodactylus'' however, conclude that it may even lack a bony cranial crest, though several analysis have proven that ''Pterodactylus'' may in fact have a crest made up of soft tissue instead of bone.<ref name=BennettPZ />
''Pterodactylus'' is the star character of the 2005 [[horror film]] ''[[Pterodactyl (film)|Pterodactyl]]'', where it is identified with the informal name "pterodactyl", hence the name of the film. In the film, the "pterodactyls" resemble the aspect of the distantly related genus ''Pteranodon'' due to the elongated bony cranial crest, and their enormous size. One peculiar feature that ''Pterodactylus'' had in the film is the possession of teeth, while this is generally accurate for ''Pterodactylus'', the overall appearance of the creatures in the film is similar to that of ''Pteranodon'', as well as the large size, this makes them resemble some kind of pterosaur identical to ''Pteranodon'', but with the possession of teeth.<ref>{{cite web|url=https://www.imdb.com/title/tt0452668/|title=Pterodactyl (2005)|work=Sci Fi Channel|publisher=IMDb|access-date=August 27, 2005}}</ref>
Another appearance of ''Pterodactylus''-like creatures is in [[J. R. R. Tolkien]]'s [[Middle-earth]] [[Tolkien's legendarium|legendarium]]. In this novel, the [[Nazgûl]], introduced as the Black Riders, are nine characters who rode flying monsters that looked similarly built to ''Pterodactylus''. [[Christopher Tolkien]], the son of the author, described the flying monsters as "Nazgûl-birds"; his father described the appearance of the steeds as somewhat "pterodactylic", and acknowledged that these were obviously "new mythology".<ref name="Letter 211">''[[Letters of J. R. R. Tolkien]]'', #211 to Rhona Beare, October 14, 1958</ref><ref name=T>''[[Letters of J. R. R. Tolkien]]'', #100 to [[Christopher Tolkien]], May 29, 1945, expressing his "loathing" for the [[Royal Air Force]]: "My sentiments are more or less those that [[Frodo]] would have had if he discovered some [[Hobbit]]s learning to ride Nazgûl-birds, 'for the liberation of the Shire'."</ref>
==See also==
* [[List of pterosaur genera]]
* [[Timeline of pterosaur research]]
==References==
{{Reflist|35em}}
==External links==
* {{Commons category-inline|Pterodactylus|''Pterodactylus''}}
* {{Wikispecies-inline|Pterodactylus|''Pterodactylus''}}
{{Pterosauria|Ar.}}
{{fins, limbs and wings}}
{{Portal bar|Paleontology|Germany|Biology}}
{{Taxonbar|from=Q309490}}
{{Authority control}}
{{good article}}
[[Category:Late Jurassic pterosaurs of Europe]]
[[Category:Pterodactyloids]]
[[Category:Solnhofen fauna]]
[[Category:Taxa named by Constantine Samuel Rafinesque]]
[[Category:Fossil taxa described in 1809]]' |
New page wikitext, after the edit (new_wikitext) | '{{short description|Genus of terodactyloid terosaur from the Late Jurassic}}
{{Other uses|terodactyl (disambiguation)}}
{{hatnote|terodactyl is also an informal name for members of the order [[terosaur]]ia.}}
{{Use mdy dates|date=January 2021}}
{{Automatic taxobox
| fossil_range = Early [[Tithonian]], <br />{{fossil range|150.8|148.5}}
| image = Bsp as i 739 modified.png
| image_caption = Sub-adult [[type specimen]] of ''P. antiquus'', Bavarian State Colection for Palaeontology and Geology
| taxon = terodactylus
| authority = [[Constantine Samuel Rafinesque|Rafinesque]], 1815
| type_species = {{extinct}}'''''Ornithocephalus antiquus'''''
| type_species_authority = [[Samuel Thomas von Sömmerring|Sömmerring]], 1812
| subdivision_ranks = [[Species]]
| subdivision =
*{{extinct}}'''''P. antiquus''''' <br/><small>(Sömmerring, 1812)</small>
*{{extinct}}''[[Diopecephalus|P. kochi]]''? <br/><small>([[Johann Andreas Wagner|Wagner]], 1837)</small>
*{{extinct}}''[[Aerodactylus|P. scolopaciceps]]''? <br/><small>([[Hermann von Meyer|Meyer]], 1860)</small>
| synonyms =
{{collapsible list|bullets = true
|title=<small>Genus synonymy</small>
|''Ornithocephalus'' <br/><small>Sömmering, 1812</small>
|''terotherium'' <br/><small>Fischer, 1813<ref>{{Cite book|last=Fischer von Waldheim|first=Gotthelf|url=https://www.biodiversitylibrary.org/item/92502|title=Zoognosia tabulis synopticis illustrata : in usum praelectionum mperalis Medico-Chirurgicae Mosquenis edita|date=1813|publisher=Typis Nicolai S. Vsevolozsky|volume=1 |location=Mosquae [Moscow]|pages=466 }}</ref></small>
|''Macrotrachelus'' <br/><small>[[Christoph Gottfried Andreas Giebel|Giebel]], 1852</small>
|''[[Diopecephalus]]''? <br/><small>[[Harry Seeley|Seeley]], 1871</small>
|''[[Aerodactylus]]''? <br/><small>Vidovic & Martill, 2014</small>
}}
{{collapsible list|bullets = false
|title=<small>Species synonymy</small>
|''Ornithocephalus antiquus'' <br/><small>Sömmering, 1812</small>
|''terodactylus longirostris'' <br/><small>[[Georges Cuvier|Cuvier]], 1819</small>
|''Ornithocephalus longirostris'' <br/><small>(Cuvier, 1819)</small>
|''Macrotrachelus longirostris'' <br/><small>(Cuvier, 1819)</small>
|''terodactylus crocodilocephaloides'' <br/><small>Ritgen, 1826</small>
|''Ornithocephalus kochi''? <br/><small>Wagner, 1837</small>
|''Diopecephalus kochi''? <br/><small>(Wagner, 1837)</small>
|''terodactylus spectabilis'' <br/><small>Meyer, 1861</small>
|''terodactylus westmani'' <br/><small>Wiman, 1925</small>
|''terodactylus cormoranus'' <br/><small>Döderlein, 1929</small>
|''Aerodactylus scolopaciceps'' <br/><small>(Meyer, 1860)</small>
}}
}}
'''''terodactylus''''' (from Greek {{lang|grc-Latn|terodáktylos}} ({{wikt-lang|grc|πτεροδάκτυλος}}) meaning 'winged finger'<ref>{{Cite journal|last=Gudger|first=E.W.|date=1944|title=The Earliest Winged Fish-Catchers|jstor=18398|journal=The Scientific Monthly|volume=59|issue=2|pages=120–129|bibcode=1944SciMo..59..120G}}</ref>) is an extinct [[genus]] of [[terosaur]]s, whose members are commonly known as '''terodactyls''' ({{IPAc-en|ˌ|t|ɛr|ə|ˈ|d|æ|k|t|ᵻ|l|z}}). It is thought to contain only a single [[species]], '''''terodactylus antiquus''''', which was the first terosaur to be named and identified as a flying [[reptile]].
[[Fossil]] remains of ''terodactylus'' have primarily been found in the [[Solnhofen limestone]] of [[Bavaria]], [[Germany]], which dates back to the [[Late Jurassic]] period (early [[Tithonian]] stage), about 150.8 to 148.5 million years ago. More fragmentary remains of ''terodactylus'' have tentatively been identified from elsewhere in [[Europe]], as well as in [[Africa]].<ref name="ammonitedate">{{cite journal|author=Schweigert, G. |year=2007|title=Ammonite biostratigraphy as a tool for dating Upper Jurassic lithographic limestones from South Germany – first results and open questions|journal=Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen|volume=245|issue=1|pages= 117–125|doi=10.1127/0077-7749/2007/0245-0117}}</ref>
''terodactylus'' was a generalist [[carnivore]] that probably fed on a variety of invertebrates and vertebrates. Like all terosaurs, ''terodactylus'' had wings formed by a skin and muscle membrane stretching from its elongated fourth finger to its hind limbs. It was supported internally by [[collagen]] fibres and externally by [[keratin]]ous ridges. ''terodactylus'' was a small terosaur compared to other famous genera such as ''[[teranodon]]'' and ''[[Quetzalcoatlus]]'', and it also lived earlier, during the Late Jurassic period, while both ''teranodon'' and ''Quetzalcoatlus'' lived during the [[Late Cretaceous]]. ''terodactylus'' lived alongside other small terosaurs such as the well-known ''[[Rhamphorhynchus]]'', as well as other genera such as ''[[Scaphognathus]]'', ''[[Anurognathus]]'' and ''[[Ctenochasma]]''. ''terodactylus'' is classified as an early-branching member of the [[Euctenochasmatia|ctenochasmatid lineage]], within the terosaur clade [[terodactyloidea]].<ref name="unwin2003" /><ref name=longrichetal2018 />
==Discovery and history==
[[File:terodactylus holotype Collini 1784.jpg|thumb|left|Original copper engraving of the ''P. antiquus'' [[holotype]] by Egid Verhelst II and published by Italian scientist {{lang|it|italics=unset|[[Cosimo Alessandro Collini]]}}, 1784]]
The [[type specimen]] of the animal now known as ''terodactylus antiquus'' was the first terosaur fossil ever to be identified. The first ''terodactylus'' specimen was described by the Italian scientist {{lang|it|italics=unset|[[Cosimo Alessandro Collini]]}} in 1784, based on a fossil skeleton that had been unearthed from the Solnhofen limestone of Bavaria. Collini was the curator of the {{lang|de|Naturalienkabinett}}, or nature [[cabinet of curiosities]] (a precursor to the modern concept of the [[natural history]] museum), in the palace of [[Charles Theodore, Elector of Bavaria]] at [[Mannheim]].<ref name=pester /><ref name=DU06>{{cite book |last=Unwin |first=David M. |title=The terosaurs: From Deep Time |year=2006 |publisher=Pi Press |location=New York |isbn=0-13-146308-X |pages=246}}</ref> The specimen had been given to the collection by Count {{lang|de|italics=unset|[[Pappenheim (state)|Friedrich Ferdinand zu Pappenheim]]}} around 1780, having been recovered from a [[lithographic limestone]] quarry in {{lang|de|italics=unset|[[Eichstätt]]}}.<ref name=brougham1844>{{cite journal | last1 =Brougham|first1= Henry P. |date=1844|title=Dialogues on instinct; with analytical view of the researches on fossil osteology|volume=19 |journal= Knight's Weekly Volume for All Readers|url=https://www.biodiversitylibrary.org/item/229475#page/5/mode/1up}}</ref> The actual date of the specimen's discovery and entry into the collection is unknown however, and it was not mentioned in a catalogue of the collection taken in 1767, so it must have been acquired at some point between that date and its 1784 description by Collini. This makes it potentially the earliest documented pterosaur find; the "Pester Exemplar" of the species ''[[Pterodactylus micronyx]]'' was described in 1779 and possibly discovered earlier than the Mannheim specimen, but it was at first considered to be a fossilized crustacean, and it was not until 1856 that this species was properly described as a pterosaur by German paleontologist {{lang|de|italics=unset|[[Christian Erich Hermann von Meyer|Hermann von Meyer]]}}.<ref name=pester>{{cite journal | last1 = Ősi | first1 = A. | last2 = Prondvai | first2 = E. | last3 = Géczy | first3 = B. | year = 2010 | title = The history of Late Jurassic pterosaurs housed in Hungarian collections and the revision of the holotype of ''Pterodactylus micronyx'' Meyer 1856 (a 'Pester Exemplar') | journal = Geological Society, London, Special Publications | volume = 343 | issue = 1| pages = 277–286 | url = https://www.researchgate.net/publication/258391479 |doi=10.1144/SP343.17 | bibcode = 2010GSLSP.343..277O | s2cid = 129805068 }}</ref>
[[File:Aquatic Pterodactylus.jpg|thumb|left|Wagler's 1830 restoration of an aquatic ''Pterodactylus'']]
In his first description of the Mannheim specimen, Collini did not conclude that it was a flying animal. In fact, Collini could not fathom what kind of animal it might have been, rejecting affinities with the birds or the bats. He speculated that it may have been a sea creature, not for any anatomical reason, but because he thought the ocean depths were more likely to have housed unknown types of animals.<ref name="collini1784">{{Cite journal |last=Collini| first=C A. |date=1784|title=Sur quelques Zoolithes du Cabinet d'Histoire naturelle de S. A. S. E. Palatine & de Bavière, à Mannheim|journal=Acta Theodoro-Palatinae Mannheim |volume=5 Physicum|pages= 58–103 (1 plate)|url=http://www.rhinoresourcecenter.com/index.php?s=1&act=pdfviewer&id=1292315455&folder=129|language=fr}}</ref><ref name="taquet&padian2004">{{Cite journal | doi = 10.1016/j.crpv.2004.02.002 |last1=Taquet|first1= P. |last2=Padian|first2= K. | year = 2004 | title = The earliest known restoration of a pterosaur and the philosophical origins of Cuvier's ''Ossemens Fossiles'' | journal = Comptes Rendus Palevol | volume = 3 | issue = 2| pages = 157–175 }}</ref> The idea that pterosaurs were aquatic animals persisted among a minority of scientists as late as 1830, when the German zoologist [[Johann Georg Wagler]] published a text on "amphibians" which included an illustration of ''Pterodactylus'' using its wings as flippers. Wagler went so far as to classify ''Pterodactylus'', along with other aquatic vertebrates (namely [[plesiosaur]]s, [[ichthyosaur]]s, and [[monotreme]]s), in the class Gryphi, between birds and mammals.<ref>{{Cite book|last=Wagler|first=Johann Georg|url=https://www.biodiversitylibrary.org/item/120243|title=Natürliches System der Amphibien : mit vorangehender Classification der Säugethiere und Vögel : ein Beitrag zur vergleichenden Zoologie|date=1830|location=München|language=de}}</ref>
[[File:Hermann pterodactylus restoration1.png|thumb|Hermann's original life restoration, the first of any pterosaur, 1800]]
The German/French scientist [[Johann Hermann]] was the one who first stated that ''Pterodactylus'' used its long fourth finger to support a wing membrane. Back in March 1800, Hermann alerted the prominent French scientist [[Georges Cuvier]] to the existence of Collini's fossil, believing that it had been captured by the occupying armies of [[Napoleon]] and sent to the French collections in [[Paris]] (and perhaps to Cuvier himself) as war booty; at the time special French [[political commissar]]s systematically seized art treasures and objects of scientific interest. Hermann sent Cuvier a letter containing his own interpretation of the specimen (though he had not examined it personally), which he believed to be a [[mammal]], including the first known life restoration of a pterosaur. Hermann restored the animal with wing membranes extending from the long fourth finger to the ankle and a covering of fur (neither wing membranes nor fur had been preserved in the specimen). Hermann also added a membrane between the neck and wrist, as is the condition in [[bat]]s. Cuvier agreed with this interpretation, and at Hermann's suggestion, Cuvier became the first to publish these ideas in December 1800 in a very short description.<ref name="taquet&padian2004" /> However, contrary to Hermann, Cuvier was convinced the animal was a [[reptile]].<ref name="cuvier1801">{{Cite journal | last= Cuvier|first= G. | year = 1801 | title =Extrait d'un ouvrage sur les espèces de quadrupèdes dont on a trouvé les ossemens dans l'intérieur de la terre |quote= Reptile volant| journal = Journal de Physique, de Chimie et d'Histoire Naturelle | volume = 52 | pages = 253–267 |url=https://gallica.bnf.fr/ark:/12148/bpt6k9604723x/f1.image|language=fr}}</ref> The specimen had not in fact been seized by the French. Rather, in 1802, following the death of Charles Theodore, it was brought to [[Munich]], where Baron [[Johann Paul Carl von Moll]] had obtained a general exemption of confiscation for the Bavarian collections.<ref name=pester /> Cuvier asked von Moll to study the fossil but was informed it could not be found. In 1809 Cuvier published a somewhat longer description, in which he named the animal ''Petro-Dactyle'',<ref name="cuvier1809">{{Cite journal | last= Cuvier|first= G. | year = 1809 | title = Mémoire sur le squelette fossile d'un reptile volant des environs d'Aichstedt, que quelques naturalistes ont pris pour un oiseau, et dont nous formons un genre de Sauriens, sous le nom de Petro-Dactyle | journal = Annales du Muséum national d'Histoire Naturelle, Paris | volume = 13 | pages = 424–437 |url=https://archive.org/details/cbarchive_48783_cuvierg1809mmoiresurlesquelett1809/mode/2up}}</ref> this was a typographical error however, and was later corrected by him to ''Ptéro-Dactyle''.<ref name="taquet&padian2004" /> He also refuted a hypothesis by [[Johann Friedrich Blumenbach]] that it would have been a shore bird.<ref name="cuvier1809" /> Cuvier remarked: "It is not possible to doubt that the long finger served to support a membrane that, by lengthening the anterior extremity of this animal, formed a good wing."<ref>Cuvier, G. (1809), p. 436 : "II n'est guère possible de douter que ce long doigt n'ait servi à supporter une membrane qui formoit ''[sic]'' à l'animal, d'après la longueur de l'extrémité antérieure, une aile bien plus puissante que celle du dragon, et au moins égale en force à celle de la chauve-souris."</ref>
[[File:Pterodactylus antiquus soemmerring.png|thumb|Von Sömmerring's incorrect 1817 restoration of the species ''[[Pterodactylus brevirostris]]'']]
Contrary to von Moll's report, the fossil was not missing; it was being studied by [[Samuel Thomas von Sömmerring]], who gave a public lecture about it on December 27, 1810. In January 1811, von Sömmerring wrote a letter to Cuvier deploring the fact that he had only recently been informed of Cuvier's request for information. His lecture was published in 1812, and in it von Sömmerring named the species '''''Ornithocephalus antiquus'''''.<ref name=sommerring1812>{{cite journal | last1 = von Sömmerring | first1 = S. T. | year = 1812 | title=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: vorgelesen in der mathematisch-physikalischen Classe am 27. Dec. 1810 und Nachtrag vorgelesen am 8. April 1811|journal=Denkschriften der Königlichen Bayerischen Akademie der Wissenschaften | url=http://publikationen.ub.uni-frankfurt.de/frontdoor/index/index/year/2008/docId/15367 | location= München | volume = 3 | pages = 89–158 }}</ref> The animal was described as being both a bat, and a form in between mammals and birds, i.e. not intermediate in descent but in "affinity" or [[archetype]]. Cuvier disagreed, and the same year in his ''Ossemens fossiles'' provided a lengthy description in which he restated that the animal was a reptile.<ref>{{Cite book|last=Cuvier|first=G.|url=https://www.biodiversitylibrary.org/item/123716#page/660/mode/1up|title=Recherches sur les ossemens fossiles de quadrupèdes : où l'on rétablit les caractères de plusieurs espèces d'animaux que les révolutions du globe paroissent avoir détruites|date=1812|publisher=Deterville|volume=Tome 4|location=Paris|language=fr|chapter=Article V – Sur le squelette fossile d’un reptile volant des environs d'Aichstedt, que quelques naturalistes ont pris pour un oiseau et dont nous formons un genre de sauriens, sous le nom de ptéro-dactyle|doi=10.5962/bhl.title.60807}}</ref> It was not until 1817 that a second specimen of ''Pterodactylus'' came to light, again from [[Solnhofen]]. This tiny specimen was that year described by von Sömmerring as ''[[Ornithocephalus brevirostris]]'', named for its short snout, now understood to be a juvenile character (this specimen is now thought to represent a juvenile specimen of a different genus, probably ''[[Ctenochasma]]'').<ref name=BennettPZ /> He provided a restoration of the skeleton, the first one published for any pterosaur.<ref name="taquet&padian2004" /> This restoration was very inaccurate, von Sömmerring mistaking the long [[metacarpal]]s for the bones of the lower arm, the lower arm for the [[humerus]], this upper arm for the [[breast bone]] and this sternum again for the [[shoulder blade]]s.<ref>{{Cite journal|last=von Sömmerring|first=S.T.|date=1817|title=Ueber einen Ornithocephalus brevirostris der Vorwelt|url=http://publikationen.ub.uni-frankfurt.de/frontdoor/index/index/year/2008/docId/15330|journal=Denkschriften der Bayerischen Akademie der Wissenschaften|language=de|volume=6|pages=89–104}}</ref> Sömmerring did not change his opinion that these forms were bats and this "bat model" for interpreting pterosaurs would remain influential long after a consensus had been reached around 1860 that they were reptiles. The standard assumptions were that pterosaurs were quadrupedal, clumsy on the ground, furred, warmblooded and had a wing membrane reaching the ankle. Some of these elements have been confirmed, some refuted by modern research, while others remain disputed.<ref>{{cite book|last=Padian|first= K. |chapter=The case of the bat-winged pterosaur. Typological taxonomy and the influence of pictorial representation on scientific perception | editor-last1 = Czerkas |editor-first1 = S. J.|editor-last2 =Olson | editor-first2 = E. C.| title = Dinosaurs past and present | publisher = Natural History Museum of Los Angeles County in association with University of Washington Press, Seattle and London | location = Los Angeles | year = 1987 | isbn = 978-0-938644-23-1|pages= 65–81|volume=2}}</ref>
In 1815, the generic name ''Ptéro-Dactyle'' was latinized to ''Pterodactylus'' by [[Constantine Samuel Rafinesque]].<ref>{{cite book|last=Rafinesque|first=C.S.|author-link=Constantine Samuel Rafinesque|year=1815|title=Analyse de la nature, ou tableau de l'univers et des corps organisés|edition=L'Imprimerie de Jean Barravecchia|page=224|url=https://www.biodiversitylibrary.org/page/48310144#page/10/mode/1up}}</ref> Unaware of Rafinesque's publication however, Cuvier himself in 1819 latinized the name ''Ptéro-Dactyle'' again to ''Pterodactylus'',<ref name=":0">{{Cite book|last=Cuvier|first=G.|url=http://archive.org/details/isisoderencyclop1819oken|title=Isis (oder Encyclopädische Zeitung) von Oken|date=1819|publisher=Jena : Expedition der Isis|editor-last=Oken|editor-first=Lorenz|pages=[https://archive.org/details/isisoderencyclop1819oken/page/n586/mode/1up 1126] ; [https://archive.org/details/isisoderencyclop1819oken/page/n914/mode/1up 1788]|language=de|chapter=Pterodactylus longirostris}}</ref> but the specific name he then gave, ''longirostris'', has to give precedence to von Sömmerring's ''antiquus''.<ref name=":0" /> In 1888, English [[naturalist]] [[Richard Lydekker]] designated ''Pterodactylus antiquus'' as the [[type species]] of ''Pterodactylus'', and considered ''Ornithocephalus antiquus'' a synonym. He also designated specimen BSP AS.I.739 as the [[holotype]] of the genus.<ref name=Lydekker />
==Description==
[[File:Pterodactylus scale mmartyniuk wiki.png|thumb|upright|left|Size of the sub-adult holotype (blue) and adult (green) specimens in both flying and standing postures, compared with a human]]
''Pterodactylus'' is known from over 30 fossil specimens, and though most belong to juveniles, many preserve complete skeletons.<ref name=BennettPZ /><ref name=bennett1996a>{{Cite journal| doi = 10.1080/02724634.1996.10011332 | author = Bennett, S.C. | year = 1996 | title = Year-classes of pterosaurs from the Solnhofen Limestone of Germany: Taxonomic and Systematic Implications | journal = Journal of Vertebrate Paleontology | volume = 16 | issue = 3| pages = 432–444}}</ref> ''Pterodactylus antiquus'' was a relatively small pterosaur, with an estimated adult wingspan of about {{convert|1.04|m|ftin|sp=us}}, based on the only known adult specimen, which is represented by an isolated skull.<ref name=BennettPZ /> Other "species" were once thought to have been smaller.<ref name=Lydekker>{{cite book|last=Lydekker|first=Richard|year=1888|title=Catalogue of Fossil Reptilia and Amphibia in the British Museum (Natural History). Part I. Containing the Orders Ornithosauria, Crocodilia, Dinosauria, Squamata, Rhynchocephalia and Pterosauria|publisher=Taylor and Francis|pages=2–37|url=https://www.biodiversitylibrary.org/item/125711#page/1/mode/1up}}</ref> However, these smaller specimens have been shown to represent juveniles of ''Pterodactylus'', as well as its contemporary relatives including ''Ctenochasma'', ''[[Germanodactylus]]'', ''[[Aurorazhdarcho]]'', ''[[Gnathosaurus]]'', and hypothetically ''[[Aerodactylus]]'' if this genus is truly valid.<ref name ="bennett2002">{{Cite journal | doi = 10.1671/0272-4634(2002)022[0043:STPOTC]2.0.CO;2 | author = Bennett, S.C. | year = 2002 | title = Soft tissue preservation of the cranial crest of the pterosaur ''Germanodactylus'' from Solnhofen|jstor=4524192 | journal = Journal of Vertebrate Paleontology | volume = 22 | issue = 1| pages = 43–48 }}</ref>
The skulls of adult ''Pterodactylus'' were long and thin, with about 90 narrow and conical teeth. The teeth extended back from the tips of both jaws, and became smaller farther away from the jaw tips, this was unlike the ones seen in most relatives, where teeth were absent in the upper jaw tip, and were relatively uniform in size. The teeth of ''Pterodactylus'' also extended farther back into the jaw compared to close relatives, and some were present below the front of the ''nasoantorbital fenestra'', which is the largest opening in the skull.<ref name=BennettPZ /> Another [[autapomorphy]] that ''Pterodactylus'' has is that the skull and jaws were straight, which are unlike the upwardly curved jaws seen in the related [[ctenochasmatid]]s.<ref name="jouve2004" />
[[File:Pterodactylus BMMS7 life.png|thumb|Life [[Paleoart|restoration]] of BMMS 7, the largest known ''Pterodactylus'' specimen. The shape of the soft tissue crest is based on the specimen BSP 1929 I 18]]
''Pterodactylus'', like related pterosaurs, had a crest on its skull composed mainly of soft tissues. In adult ''Pterodactylus'', this crest extended between the back edge of the [[antorbital fenestra]] and the back of the skull. In at least one specimen, the crest had a short bony base, also seen in related pterosaurs like ''Germanodactylus''. Solid crests have only been found on large, fully adult specimens of ''Pterodactylus'', indicating that this was a display structure that became larger and more well developed as individuals reached maturity.<ref name=BennettPZ>{{cite journal |year=2013 |title=New information on body size and cranial display structures of ''Pterodactylus antiquus'', with a revision of the genus |journal=Paläontologische Zeitschrift |volume=87|issue=2 |doi=10.1007/s12542-012-0159-8 |author=Bennett, S. Christopher |pages=269–289|s2cid=83722829 }}</ref><ref name="frey&martill1998">{{Cite journal |last1=Frey|first1= E. |last2=Martill|first2= D.M. | year = 1998 | title = Soft tissue preservation in a specimen of ''Pterodactylus kochi'' (Wagner) from the Upper Jurassic of Germany | journal = Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen | volume = 210 |issue=3 | pages = 421–441 |doi=10.1127/njgpa/210/1998/421 }}</ref> In 2013, pterosaur researcher [[S. Christopher Bennett]] noted that other authors claimed that the soft tissue crest of ''Pterodactylus'' extended backward behind the skull; Bennett himself, however, didn't find any evidence for the crest extending past the back of the skull.<ref name=BennettPZ /> Two specimens of ''P. antiquus'' (the holotype specimen BSP AS I 739 and the incomplete skull BMMS 7, the largest known skull of ''P. antiquus'') have a low bony crest on their skulls; in BMMS 7 it is 47.5 mm long (1.87 inches, more or less 24% of the estimated total length of its skull) and has a maximum height of 0.9 mm (0.035 inches) above the orbit.<ref name=BennettPZ /> Several specimens previously referred to ''P. antiquus'' preserved evidence of the soft tissue extensions of these crests, including an "occipital lappet", a flexible, tab-like structure extending from the back of the skull. Most of these specimens have been reclassified in the related species ''Aerodactylus scolopaciceps'', which may however be nothing more than a junior synonym. Even if ''Aerodactylus'' were valid, at least one specimen with these features is still considered to belong to ''Pterodactylus'', BSP 1929 I 18, which has an occipital lappet similar to the proposed ''Aerodactylus'' definition, and also possesses a small triangular soft tissue crest with the peak of the crest positioned above the eyes.<ref name=BennettPZ />
==Paleobiology==
===Life history===
[[File:Pterodactylus spectabilis 4.JPG|thumb|left|Holotype specimen of the species ''P. spectabilis'', now considered a juvenile specimen of ''P. antiquus'']]
Like other pterosaurs (most notably ''[[Rhamphorhynchus (pterosaur)|Rhamphorhynchus]]''), ''Pterodactylus'' specimens can vary considerably based on age or level of maturity. Both the proportions of the limb bones, size and shape of the skull, and size and number of teeth changed as the animals grew. Historically, this has led to various growth stages (including growth stages of related pterosaurs) being mistaken for new species of ''Pterodactylus''. Several detailed studies using various methods to measure growth curves among known specimens have suggested that there is actually only one valid species of ''Pterodactylus'', ''P. antiquus''.<ref name="jouve2004" />
The youngest immature specimens of ''Pterodactylus antiquus'' (alternately interpreted as young specimens of the distinct species ''P. kochi'') have a small number of teeth, as few as 15 in some, and the teeth have a relatively broad base.<ref name=bennett1996a /> The teeth of other ''P. antiquus'' specimens are both narrower and more numerous (up to 90 teeth are present in several specimens).<ref name="jouve2004" />
''Pterodactylus'' specimens can be divided into two distinct year classes. In the first year class, the skulls are only {{convert|15|to|45|mm|in|sp=us}} in length. The second year class is characterized by skulls of around {{convert|55|to|95|mm|in|sp=us}} long, but are still immature however. These first two size groups were once classified as juveniles and adults of the species ''P. kochi'', until further study showed that even the supposed "adults" were immature, and possibly belong to a distinct genus. A third year class is represented by specimens of the "traditional" ''P. antiquus'', as well as a few isolated, large specimens once assigned to ''P. kochi'' that overlap ''P. antiquus'' in size. However, all specimens in this third year class also show sign of immaturity. Fully mature ''Pterodactylus'' specimens remain unknown, or may have been mistakenly classified as a different genus.<ref name=bennett1996a />
===Growth and breeding seasons===
[[File:Pterodactylus grandipelvis Teylers 2.JPG|thumb|Fossil pelvis of a large specimen, now referred to the dubious species ''P. grandipelvis'']]
The distinct year classes of ''Pterodactylus antiquus'' specimens show that this species, like the contemporary ''Rhamphorhynchus muensteri'', likely bred seasonally and grew consistently during its lifetime. A new generation of 1st year class ''P. antiquus'' would have been produced seasonally, and reached 2nd-year size by the time the next generation hatched, creating distinct 'clumps' of similarly-sized and aged individuals in the fossil record. The smallest size class probably consisted of individuals that had just begun to fly and were less than one year old.<ref name=bennett1996a /><ref name="Wellnhofer1970">{{cite book|last=Wellnhofer|first=Peter|url=https://www.zobodat.at/pdf/Abhandlungen-Akademie-Bayern_NF_141_0001-0133.pdf|title=Die Pterodactyloidea (Pterosauria) der Oberjura-Plattenkalke Süddeutschlands|publisher=Bayerische Akademie der Wissenschaften, Mathematisch-Wissenschaftlichen Klasse, Abhandlungen|year=1970|volume=141|page=133}}</ref> The second year class represents individuals one to two years old, and the rare third year class is composed of specimens over two years old. This growth pattern is similar to modern [[crocodilia]]ns, rather than the rapid growth of modern [[bird]]s.<ref name=bennett1996a />
===Daily activity patterns===
Comparisons between the [[sclerotic ring|scleral rings]] of ''Pterodactylus antiquus'' and modern birds and reptiles suggest that it may have been [[Diurnality|diurnal]]. This may also indicate [[niche partitioning]] with contemporary pterosaurs inferred to be [[nocturnal]], such as ''Ctenochasma'' and ''Rhamphorhynchus''.<ref>{{cite journal|author1=Schmitz, L. |author2=Motani, R. |year=2011 |title=Nocturnality in Dinosaurs Inferred from Scleral Ring and Orbit Morphology |journal=Science |volume=332 |issue= 6030|pages= 705–8|doi=10.1126/science.1200043 |pmid=21493820 |bibcode=2011Sci...332..705S|s2cid=33253407 |url=http://pdfs.semanticscholar.org/8922/c462d9cd7f64033fe9989f5a1dc0e7e8aac9.pdf |archive-url=https://web.archive.org/web/20190228084617/http://pdfs.semanticscholar.org/8922/c462d9cd7f64033fe9989f5a1dc0e7e8aac9.pdf |url-status=dead |archive-date=2019-02-28 }}</ref>
===Diet===
Based on the shape, size, and arrangement of its teeth, ''Pterodactylus'' has long been recognized as a carnivore specializing in small animals. A 2020 study of pterosaur tooth wear supported the hypothesis that ''Pterodactylus'' preyed mainly on invertebrates and had a generalist feeding strategy, indicated by a relatively high bite force.<ref name=diet2020>Bestwick, J., Unwin, D.M., Butler, R.J. et al. Dietary diversity and evolution of the earliest flying vertebrates revealed by dental microwear texture analysis. Nat Commun 11, 5293 (2020). https://doi.org/10.1038/s41467-020-19022-2</ref>
==Paleoecology==
[[File:Pterodactylus holotype fly mmartyniuk.png|thumb|left|Hypothetical restoration of ''Pterodactylus'' in its environment]]
Specimens of ''Pterodactylus'' have been found mainly in the [[Solnhofen limestone]] (geologically known as the Altmühltal Formation) of [[Bavaria]], [[Germany]]. The main composition of this formation is fine-grained [[limestone]] that originated mainly from the nearby towns [[Solnhofen]] and Eichstätt, which is formed by mud [[silt]] deposits.<ref name="ammonitedate" /> The Solnhofen Limestone is a diverse [[Lagerstätte]] that contains a wide range of different creatures, including highly detailed fossilized imprints of soft bodied organisms such as [[jellyfish]]es. Abundant specimens of pterosaurs similar to ''Pterodactylus'' were also found within the formation, these include the [[rhamphorhynchid]]s ''Rhamphorhynchus'' and ''[[Scaphognathus]]'',<ref>{{Cite journal|last=Bennett|first=S. C.|date=2004|title=New information on the pterosaur ''Scaphognathus crassirostris'' and the pterosaurian cervical series|journal=Journal of Vertebrate Paleontology|language=en|volume=24|issue=Supplement 003|pages=38A|doi=10.1080/02724634.2004.10010643|s2cid=220415208|issn=0272-4634}}</ref> several [[gallodactylid]]s such as ''Aerodactylus'',<ref name=bennett1996a /> ''[[Ardeadactylus]]'', ''Aurorazhdarcho'' and ''[[Cycnorhamphus]]'',<ref name=DU06 /> the [[ctenochasmatid]]s ''Ctenochasma''<ref name=bennett2007>{{cite journal | last1 = Bennett | first1 = S.C. | year = 2007 | title = A review of the pterosaur ''Ctenochasma'': taxonomy and ontogeny | journal = Neues Jahrbuch für Geologie und Paläontologie – Abhandlungen | volume = 245 | issue = 1| pages = 23–31 | doi=10.1127/0077-7749/2007/0245-0023}}</ref> and ''Gnathosaurus'', the [[anurognathid]] ''[[Anurognathus]]'', the [[germanodactylid]] ''Germanodactylus'', as well as the basal [[euctenochasmatia]]n ''[[Diopecephalus]]''.<ref name=W1837 /> Fossil remains of the [[dinosaur]]s ''[[Archaeopteryx]]'' and ''[[Compsognathus]]'' were also found within the limestone, these specimens were related to early evolution of [[feathers]], since they were some of the only ones that had them during the Jurassic period.<ref name="FasWeis04">{{cite book|title=The Evolution and Extinction of the Dinosaurs|year=2005|chapter=Theropoda I: nature red in tooth and claw|editor-last1=Fastovsky| editor-first1=D.E.|editor-last2=Weishampel|editor-first2=D.B. |last1=Fastovsky|first1=D.E.|last2=Weishampel |first2=D.B. |pages=265–299|publisher=Cambridge University Press|isbn=978-0-521-81172-9|edition=2nd|doi=10.1017/9781316471623.009}}</ref> Various [[lizard]] remains were also found alongside those of ''Pterodactylus'', with several specimens assigned to ''[[Ardeosaurus]]'', ''[[Bavarisaurus]]'' and ''[[Eichstaettisaurus]]''.<ref name="hoffstetter1966">{{cite journal |last=Hoffstetter |first=R. |year=1966 |title=A propos des genres ''Ardeosaurus'' et ''Eichstaettisaurus'' (Reptilia, Sauria, Gekkonoidea) du Jurassique Supèrieur de Franconie |trans-title=On the genera ''Ardeosaurus'' and ''Eichstaettisaurus'' (Reptilia, Sauria, Gekkonoidea) from the Upper Jurassic of France |journal=Bulletin de la Société Géologique de France |volume=8 |issue=4 |pages=592–595 |doi=10.2113/gssgfbull.S7-VIII.4.592 |url=https://pubs.geoscienceworld.org/sgf/bsgf/article-abstract/S7-VIII/4/592/518861/A-propos-des-genres-Ardeosaurus-et}}</ref><ref>{{cite journal |last= Evans|first= S.E.|year=1994|title= The Solnhofen (Jurassic: Tithonian) lizard genus ''Bavarisaurus'': new skull material and a reinterpretation |journal=Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen |volume=192 |pages=37–52}}</ref> [[Crocodylomorph]] specimens were widely distributed within the fossil site, most were assigned to the [[metriorhynchid]] genera ''[[Cricosaurus]]'', ''[[Dakosaurus]]'',<ref>{{Cite journal|last1=Brandalise de Andrade|first1=Marco|last2=Young|first2=Mark T.|last3=Desojo|first3=Julia B.|last4=Brusatte|first4=Stephen L.|date=2010|title=The evolution of extreme hypercarnivory in Metriorhynchidae (Mesoeucrocodylia: Thalattosuchia) based on evidence from microscopic denticle morphology|journal=Journal of Vertebrate Paleontology|volume=30|issue=5|pages=1451–1465|doi=10.1080/02724634.2010.501442|s2cid=83985855|issn=0272-4634}}</ref> ''[[Geosaurus]]'' and ''[[Rhacheosaurus]]''.<ref name="Andrade & Young, 2008">{{Cite journal|last1=Brandalise de Andrade|first1=Marco|last2=Young|first2=Mark T.|year=2008|url=http://www.svpca.org/general/pages/abstractPage.php?i=1378&r=talksAndPosters.php&y=2008 |title= High diversity of thalattosuchian crocodylians and the niche partition in the Solnhofen Sea |archive-url=https://web.archive.org/web/20110603232400/http://www.svpca.org/general/pages/abstractPage.php?i=1378&r=talksAndPosters.php&y=2008 |archive-date=June 3, 2011 |journal= The 56th Symposium of Vertebrate Palaeontology and Comparative Anatomy|pages=14–15}}</ref> These genera are colloquially called as marine or sea crocodiles due to their similar built.<ref>{{Cite journal|last1=Young|first1=Mark T.|last2=Brusatte|first2=Stephen L.|last3=de Andrade|first3=Marco Brandalise|last4=Desojo|first4=Julia B.|last5=Beatty|first5=Brian L.|last6=Steel|first6=Lorna|last7=Fernández|first7=Marta S.|last8=Sakamoto|first8=Manabu|last9=Ruiz-Omeñaca|first9=Jose Ignacio|last10=Schoch|first10=Rainer R.|date=September 18, 2012|editor-last=Butler|editor-first=Richard J.|title=The Cranial Osteology and Feeding Ecology of the Metriorhynchid Crocodylomorph Genera Dakosaurus and Plesiosuchus from the Late Jurassic of Europe|journal=PLOS ONE|language=en|volume=7|issue=9|pages=e44985|doi=10.1371/journal.pone.0044985|issn=1932-6203|pmc=3445579|pmid=23028723|bibcode=2012PLoSO...744985Y}}</ref> The turtle genera ''[[Eurysternum]]'' and ''[[Paleomedusa]]'' were also found within the formation.<ref name="WGJ">{{cite journal|first=Walter G.|last= Joyce|year=2003|title=A new Late Jurassic turtle specimen and the taxonomy of ''Palaeomedusa testa'' and ''Eurysternum wagleri''|journal=PaleoBios|volume=23|issue=3|pages=1–8|url=http://www.geo.uni-tuebingen.de/fileadmin/website/arbeitsbereich/palaeo/biogeologie/Images/Joyce_Publications/3__Joyce_2003.pdf|archive-url=https://web.archive.org/web/20151001071809/http://www.geo.uni-tuebingen.de/fileadmin/website/arbeitsbereich/palaeo/biogeologie/Images/Joyce_Publications/3__Joyce_2003.pdf|url-status=dead|archive-date=October 1, 2015}}</ref> Fossils of the [[ichthyosaur]] ''[[Aegirosaurus]]'' also appeared to be present in the site,<ref>{{Cite journal|last1=Bardet|first1=Nathalie|last2=Fernández|first2=Marta S.|date=2000|title=A new ichthyosaur from the Upper Jurassic lithographic limestones of Bavaria|url=https://www.researchgate.net/publication/40662676|journal=Journal of Paleontology|language=en|volume=74|issue=3|pages=503–511|doi=10.1017/S0022336000031760|issn=0022-3360}}</ref> as well as fish remains, with many specimens assigned to [[ray-finned fish]]es such as the [[halecomorph]]s ''[[Lepidotes]]'',<ref name="lamberstable-others"/> ''[[Propterus]]'',<ref name=Marine /> ''[[Gyrodus]]'', ''[[Mesturus]]'', ''[[Proscinetes]]'', ''[[Caturus]]'',<ref name=Marine>{{cite journal|last=Sepkoski |first=Jack |title=A compendium of fossil marine animal genera |journal=Bulletins of American Paleontology |volume=364 |page=560 |year=2002 |url=http://strata.ummp.lsa.umich.edu/jack/showgenera.php?taxon=611&rank=class |access-date=February 27, 2009 |url-status=dead |archive-url=https://web.archive.org/web/20110723131237/http://strata.ummp.lsa.umich.edu/jack/showgenera.php?taxon=611&rank=class |archive-date=July 23, 2011 }}</ref> ''[[Ophiopsis]]''<ref>{{Cite journal|last1=Lane|first1=Jennifer A.|last2=Ebert|first2=Martin|date=2015|title=A taxonomic reassessment of Ophiopsis (Halecomorphi, Ionoscopiformes), with a revision of Upper Jurassic species from the Solnhofen Archipelago, and a new genus of Ophiopsidae|journal=Journal of Vertebrate Paleontology|language=en|volume=35|issue=1|pages=e883238|doi=10.1080/02724634.2014.883238|s2cid=86350086|issn=0272-4634}}</ref> and ''[[Ophiopsiella]]'',<ref name="lamberstable-others">{{Cite journal|last=Lambers|first=Paul H.|date=1999|title=The actinopterygian fish fauna of the Late Kimmeridgian and Early Tithonian 'Plattenkalke' near Solnhofen (Bavaria, Germany): state of the art|journal=Geologie en Mijnbouw |volume=78|issue=2|pages=215–229 |doi=10.1023/A:1003855831015|s2cid=127676896|quote=Table 1: List of actinopterygians from the Solnhofen lithographic limestone – Halecomorphi; 'Amiidae' and 'Ophiopsidae'" (p. 216)}}</ref> the [[Pachycormidae|pachycormids]] ''[[Asthenocormus]]'', ''[[Hypsocormus]]'' and ''[[Orthocormus]]'',<ref name=EoDP>{{cite book |editor=Palmer, D.|year=1999 |title= The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals|publisher= Marshall Editions|location=London|page= 38|isbn= 1-84028-152-9}}</ref> as well as the [[Aspidorhynchidae|aspidorhynchid]] ''[[Aspidorhynchus]]'', and the [[ichthyodectid]] ''[[Thrissops]]''.<ref name=FT12>{{cite journal |last1=Frey |first1=E. |last2=Tischlinger|first2= H. |year=2012 |title=The Late Jurassic pterosaur ''Rhamphorhynchus'', a frequent victim of the ganoid fish ''Aspidorhynchus''? |journal=PLOS ONE |volume=7 |issue=3 |pages=e31945 |doi=10.1371/journal.pone.0031945 |bibcode = 2012PLoSO...7E1945F |pmid=22412850 |pmc=3296705}}</ref><ref>{{Cite book|last=Nybelin|first=Orvar|title=Versuch einer taxonomischen Revision der jurassischen Fischgattung Thrissops Agassiz.|date=1964|publisher=Wettergren and Kerber|location=Göteborg|language=de|oclc=2672427}}</ref>
==Classification==
[[File:Pterodactylus muscle impressions.jpg|thumb|upright|A ''P. antiquus'' specimen (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 }}</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|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 |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=":1">{{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}}</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 if 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=":1" /> 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, M.P.|year=2010|title=''Pteranodon'' and beyond: The history of giant pterosaurs from 1870 onwards. Geological Society of London Special Publications|volume=343|pages=313–323|url=https://www.researchgate.net/publication/258391482}}</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 }}</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" /> though after additional research Bennett assigned it to the genus ''Aurorazhdarcho''.<ref name=BennettPZ /> 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 }}</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 }}</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}}</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|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 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''
|-
|''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}}
==Cultural significance==
[[File:Pteranodon longiceps mmartyniuk wiki.png|upright|thumb|Life reconstruction of a ''Pteranodon'', popularly called "pterodactyl"]]
''Pterodactylus'' is regarded as one of the most iconic prehistoric creatures, with multiple appearances in books, movies, as well as television series and several videogames. The informal name "'''pterodactyl'''" is sometimes used to refer to any kind of animal belonging to the order [[Pterosauria]], though most of the times to ''Pterodactylus'', as it's the most well-known member of the group.<ref name="myths">{{cite web|url=http://www.pterosaur.net/myths.php|title=Pterosaurs: Myths and Misconceptions|author=Naish, Darren |publisher=Pterosaur.net|access-date=June 18, 2011}}</ref> The popular aspect of ''Pterodactylus'' consists of an elongated head crest, and potentially large wings. Studies of ''Pterodactylus'' however, conclude that it may even lack a bony cranial crest, though several analysis have proven that ''Pterodactylus'' may in fact have a crest made up of soft tissue instead of bone.<ref name=BennettPZ />
''Pterodactylus'' is the star character of the 2005 [[horror film]] ''[[Pterodactyl (film)|Pterodactyl]]'', where it is identified with the informal name "pterodactyl", hence the name of the film. In the film, the "pterodactyls" resemble the aspect of the distantly related genus ''Pteranodon'' due to the elongated bony cranial crest, and their enormous size. One peculiar feature that ''Pterodactylus'' had in the film is the possession of teeth, while this is generally accurate for ''Pterodactylus'', the overall appearance of the creatures in the film is similar to that of ''Pteranodon'', as well as the large size, this makes them resemble some kind of pterosaur identical to ''Pteranodon'', but with the possession of teeth.<ref>{{cite web|url=https://www.imdb.com/title/tt0452668/|title=Pterodactyl (2005)|work=Sci Fi Channel|publisher=IMDb|access-date=August 27, 2005}}</ref>
Another appearance of ''Pterodactylus''-like creatures is in [[J. R. R. Tolkien]]'s [[Middle-earth]] [[Tolkien's legendarium|legendarium]]. In this novel, the [[Nazgûl]], introduced as the Black Riders, are nine characters who rode flying monsters that looked similarly built to ''Pterodactylus''. [[Christopher Tolkien]], the son of the author, described the flying monsters as "Nazgûl-birds"; his father described the appearance of the steeds as somewhat "pterodactylic", and acknowledged that these were obviously "new mythology".<ref name="Letter 211">''[[Letters of J. R. R. Tolkien]]'', #211 to Rhona Beare, October 14, 1958</ref><ref name=T>''[[Letters of J. R. R. Tolkien]]'', #100 to [[Christopher Tolkien]], May 29, 1945, expressing his "loathing" for the [[Royal Air Force]]: "My sentiments are more or less those that [[Frodo]] would have had if he discovered some [[Hobbit]]s learning to ride Nazgûl-birds, 'for the liberation of the Shire'."</ref>
==See also==
* [[List of pterosaur genera]]
* [[Timeline of pterosaur research]]
==References==
{{Reflist|35em}}
==External links==
* {{Commons category-inline|Pterodactylus|''Pterodactylus''}}
* {{Wikispecies-inline|Pterodactylus|''Pterodactylus''}}
{{Pterosauria|Ar.}}
{{fins, limbs and wings}}
{{Portal bar|Paleontology|Germany|Biology}}
{{Taxonbar|from=Q309490}}
{{Authority control}}
{{good article}}
[[Category:Late Jurassic pterosaurs of Europe]]
[[Category:Pterodactyloids]]
[[Category:Solnhofen fauna]]
[[Category:Taxa named by Constantine Samuel Rafinesque]]
[[Category:Fossil taxa described in 1809]]' |
Unified diff of changes made by edit (edit_diff) | '@@ -1,5 +1,5 @@
-{{short description|Genus of pterodactyloid pterosaur from the Late Jurassic}}
-{{Other uses|Pterodactyl (disambiguation)}}
-{{hatnote|Pterodactyl is also an informal name for members of the order [[Pterosaur]]ia.}}
+{{short description|Genus of terodactyloid terosaur from the Late Jurassic}}
+{{Other uses|terodactyl (disambiguation)}}
+{{hatnote|terodactyl is also an informal name for members of the order [[terosaur]]ia.}}
{{Use mdy dates|date=January 2021}}
{{Automatic taxobox
@@ -7,5 +7,5 @@
| image = Bsp as i 739 modified.png
| image_caption = Sub-adult [[type specimen]] of ''P. antiquus'', Bavarian State Colection for Palaeontology and Geology
-| taxon = Pterodactylus
+| taxon = terodactylus
| authority = [[Constantine Samuel Rafinesque|Rafinesque]], 1815
| type_species = {{extinct}}'''''Ornithocephalus antiquus'''''
@@ -20,5 +20,5 @@
|title=<small>Genus synonymy</small>
|''Ornithocephalus'' <br/><small>Sömmering, 1812</small>
-|''Pterotherium'' <br/><small>Fischer, 1813<ref>{{Cite book|last=Fischer von Waldheim|first=Gotthelf|url=https://www.biodiversitylibrary.org/item/92502|title=Zoognosia tabulis synopticis illustrata : in usum praelectionum mperalis Medico-Chirurgicae Mosquenis edita|date=1813|publisher=Typis Nicolai S. Vsevolozsky|volume=1 |location=Mosquae [Moscow]|pages=466 }}</ref></small>
+|''terotherium'' <br/><small>Fischer, 1813<ref>{{Cite book|last=Fischer von Waldheim|first=Gotthelf|url=https://www.biodiversitylibrary.org/item/92502|title=Zoognosia tabulis synopticis illustrata : in usum praelectionum mperalis Medico-Chirurgicae Mosquenis edita|date=1813|publisher=Typis Nicolai S. Vsevolozsky|volume=1 |location=Mosquae [Moscow]|pages=466 }}</ref></small>
|''Macrotrachelus'' <br/><small>[[Christoph Gottfried Andreas Giebel|Giebel]], 1852</small>
|''[[Diopecephalus]]''? <br/><small>[[Harry Seeley|Seeley]], 1871</small>
@@ -28,26 +28,26 @@
|title=<small>Species synonymy</small>
|''Ornithocephalus antiquus'' <br/><small>Sömmering, 1812</small>
-|''Pterodactylus longirostris'' <br/><small>[[Georges Cuvier|Cuvier]], 1819</small>
+|''terodactylus longirostris'' <br/><small>[[Georges Cuvier|Cuvier]], 1819</small>
|''Ornithocephalus longirostris'' <br/><small>(Cuvier, 1819)</small>
|''Macrotrachelus longirostris'' <br/><small>(Cuvier, 1819)</small>
-|''Pterodactylus crocodilocephaloides'' <br/><small>Ritgen, 1826</small>
+|''terodactylus crocodilocephaloides'' <br/><small>Ritgen, 1826</small>
|''Ornithocephalus kochi''? <br/><small>Wagner, 1837</small>
|''Diopecephalus kochi''? <br/><small>(Wagner, 1837)</small>
-|''Pterodactylus spectabilis'' <br/><small>Meyer, 1861</small>
-|''Pterodactylus westmani'' <br/><small>Wiman, 1925</small>
-|''Pterodactylus cormoranus'' <br/><small>Döderlein, 1929</small>
+|''terodactylus spectabilis'' <br/><small>Meyer, 1861</small>
+|''terodactylus westmani'' <br/><small>Wiman, 1925</small>
+|''terodactylus cormoranus'' <br/><small>Döderlein, 1929</small>
|''Aerodactylus scolopaciceps'' <br/><small>(Meyer, 1860)</small>
}}
}}
-'''''Pterodactylus''''' (from Greek {{lang|grc-Latn|pterodáktylos}} ({{wikt-lang|grc|πτεροδάκτυλος}}) meaning 'winged finger'<ref>{{Cite journal|last=Gudger|first=E.W.|date=1944|title=The Earliest Winged Fish-Catchers|jstor=18398|journal=The Scientific Monthly|volume=59|issue=2|pages=120–129|bibcode=1944SciMo..59..120G}}</ref>) is an extinct [[genus]] of [[pterosaur]]s, whose members are commonly known as '''pterodactyls''' ({{IPAc-en|ˌ|t|ɛr|ə|ˈ|d|æ|k|t|ᵻ|l|z}}). It is thought to contain only a single [[species]], '''''Pterodactylus antiquus''''', which was the first pterosaur to be named and identified as a flying [[reptile]].
+'''''terodactylus''''' (from Greek {{lang|grc-Latn|terodáktylos}} ({{wikt-lang|grc|πτεροδάκτυλος}}) meaning 'winged finger'<ref>{{Cite journal|last=Gudger|first=E.W.|date=1944|title=The Earliest Winged Fish-Catchers|jstor=18398|journal=The Scientific Monthly|volume=59|issue=2|pages=120–129|bibcode=1944SciMo..59..120G}}</ref>) is an extinct [[genus]] of [[terosaur]]s, whose members are commonly known as '''terodactyls''' ({{IPAc-en|ˌ|t|ɛr|ə|ˈ|d|æ|k|t|ᵻ|l|z}}). It is thought to contain only a single [[species]], '''''terodactylus antiquus''''', which was the first terosaur to be named and identified as a flying [[reptile]].
-[[Fossil]] remains of ''Pterodactylus'' have primarily been found in the [[Solnhofen limestone]] of [[Bavaria]], [[Germany]], which dates back to the [[Late Jurassic]] period (early [[Tithonian]] stage), about 150.8 to 148.5 million years ago. More fragmentary remains of ''Pterodactylus'' have tentatively been identified from elsewhere in [[Europe]], as well as in [[Africa]].<ref name="ammonitedate">{{cite journal|author=Schweigert, G. |year=2007|title=Ammonite biostratigraphy as a tool for dating Upper Jurassic lithographic limestones from South Germany – first results and open questions|journal=Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen|volume=245|issue=1|pages= 117–125|doi=10.1127/0077-7749/2007/0245-0117}}</ref>
+[[Fossil]] remains of ''terodactylus'' have primarily been found in the [[Solnhofen limestone]] of [[Bavaria]], [[Germany]], which dates back to the [[Late Jurassic]] period (early [[Tithonian]] stage), about 150.8 to 148.5 million years ago. More fragmentary remains of ''terodactylus'' have tentatively been identified from elsewhere in [[Europe]], as well as in [[Africa]].<ref name="ammonitedate">{{cite journal|author=Schweigert, G. |year=2007|title=Ammonite biostratigraphy as a tool for dating Upper Jurassic lithographic limestones from South Germany – first results and open questions|journal=Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen|volume=245|issue=1|pages= 117–125|doi=10.1127/0077-7749/2007/0245-0117}}</ref>
-''Pterodactylus'' was a generalist [[carnivore]] that probably fed on a variety of invertebrates and vertebrates. Like all pterosaurs, ''Pterodactylus'' had wings formed by a skin and muscle membrane stretching from its elongated fourth finger to its hind limbs. It was supported internally by [[collagen]] fibres and externally by [[keratin]]ous ridges. ''Pterodactylus'' was a small pterosaur compared to other famous genera such as ''[[Pteranodon]]'' and ''[[Quetzalcoatlus]]'', and it also lived earlier, during the Late Jurassic period, while both ''Pteranodon'' and ''Quetzalcoatlus'' lived during the [[Late Cretaceous]]. ''Pterodactylus'' lived alongside other small pterosaurs such as the well-known ''[[Rhamphorhynchus]]'', as well as other genera such as ''[[Scaphognathus]]'', ''[[Anurognathus]]'' and ''[[Ctenochasma]]''. ''Pterodactylus'' is classified as an early-branching member of the [[Euctenochasmatia|ctenochasmatid lineage]], within the pterosaur clade [[Pterodactyloidea]].<ref name="unwin2003" /><ref name=longrichetal2018 />
+''terodactylus'' was a generalist [[carnivore]] that probably fed on a variety of invertebrates and vertebrates. Like all terosaurs, ''terodactylus'' had wings formed by a skin and muscle membrane stretching from its elongated fourth finger to its hind limbs. It was supported internally by [[collagen]] fibres and externally by [[keratin]]ous ridges. ''terodactylus'' was a small terosaur compared to other famous genera such as ''[[teranodon]]'' and ''[[Quetzalcoatlus]]'', and it also lived earlier, during the Late Jurassic period, while both ''teranodon'' and ''Quetzalcoatlus'' lived during the [[Late Cretaceous]]. ''terodactylus'' lived alongside other small terosaurs such as the well-known ''[[Rhamphorhynchus]]'', as well as other genera such as ''[[Scaphognathus]]'', ''[[Anurognathus]]'' and ''[[Ctenochasma]]''. ''terodactylus'' is classified as an early-branching member of the [[Euctenochasmatia|ctenochasmatid lineage]], within the terosaur clade [[terodactyloidea]].<ref name="unwin2003" /><ref name=longrichetal2018 />
==Discovery and history==
-[[File:Pterodactylus holotype Collini 1784.jpg|thumb|left|Original copper engraving of the ''P. antiquus'' [[holotype]] by Egid Verhelst II and published by Italian scientist {{lang|it|italics=unset|[[Cosimo Alessandro Collini]]}}, 1784]]
-The [[type specimen]] of the animal now known as ''Pterodactylus antiquus'' was the first pterosaur fossil ever to be identified. The first ''Pterodactylus'' specimen was described by the Italian scientist {{lang|it|italics=unset|[[Cosimo Alessandro Collini]]}} in 1784, based on a fossil skeleton that had been unearthed from the Solnhofen limestone of Bavaria. Collini was the curator of the {{lang|de|Naturalienkabinett}}, or nature [[cabinet of curiosities]] (a precursor to the modern concept of the [[natural history]] museum), in the palace of [[Charles Theodore, Elector of Bavaria]] at [[Mannheim]].<ref name=pester /><ref name=DU06>{{cite book |last=Unwin |first=David M. |title=The Pterosaurs: From Deep Time |year=2006 |publisher=Pi Press |location=New York |isbn=0-13-146308-X |pages=246}}</ref> The specimen had been given to the collection by Count {{lang|de|italics=unset|[[Pappenheim (state)|Friedrich Ferdinand zu Pappenheim]]}} around 1780, having been recovered from a [[lithographic limestone]] quarry in {{lang|de|italics=unset|[[Eichstätt]]}}.<ref name=brougham1844>{{cite journal | last1 =Brougham|first1= Henry P. |date=1844|title=Dialogues on instinct; with analytical view of the researches on fossil osteology|volume=19 |journal= Knight's Weekly Volume for All Readers|url=https://www.biodiversitylibrary.org/item/229475#page/5/mode/1up}}</ref> The actual date of the specimen's discovery and entry into the collection is unknown however, and it was not mentioned in a catalogue of the collection taken in 1767, so it must have been acquired at some point between that date and its 1784 description by Collini. This makes it potentially the earliest documented pterosaur find; the "Pester Exemplar" of the species ''[[Pterodactylus micronyx]]'' was described in 1779 and possibly discovered earlier than the Mannheim specimen, but it was at first considered to be a fossilized crustacean, and it was not until 1856 that this species was properly described as a pterosaur by German paleontologist {{lang|de|italics=unset|[[Christian Erich Hermann von Meyer|Hermann von Meyer]]}}.<ref name=pester>{{cite journal | last1 = Ősi | first1 = A. | last2 = Prondvai | first2 = E. | last3 = Géczy | first3 = B. | year = 2010 | title = The history of Late Jurassic pterosaurs housed in Hungarian collections and the revision of the holotype of ''Pterodactylus micronyx'' Meyer 1856 (a 'Pester Exemplar') | journal = Geological Society, London, Special Publications | volume = 343 | issue = 1| pages = 277–286 | url = https://www.researchgate.net/publication/258391479 |doi=10.1144/SP343.17 | bibcode = 2010GSLSP.343..277O | s2cid = 129805068 }}</ref>
+[[File:terodactylus holotype Collini 1784.jpg|thumb|left|Original copper engraving of the ''P. antiquus'' [[holotype]] by Egid Verhelst II and published by Italian scientist {{lang|it|italics=unset|[[Cosimo Alessandro Collini]]}}, 1784]]
+The [[type specimen]] of the animal now known as ''terodactylus antiquus'' was the first terosaur fossil ever to be identified. The first ''terodactylus'' specimen was described by the Italian scientist {{lang|it|italics=unset|[[Cosimo Alessandro Collini]]}} in 1784, based on a fossil skeleton that had been unearthed from the Solnhofen limestone of Bavaria. Collini was the curator of the {{lang|de|Naturalienkabinett}}, or nature [[cabinet of curiosities]] (a precursor to the modern concept of the [[natural history]] museum), in the palace of [[Charles Theodore, Elector of Bavaria]] at [[Mannheim]].<ref name=pester /><ref name=DU06>{{cite book |last=Unwin |first=David M. |title=The terosaurs: From Deep Time |year=2006 |publisher=Pi Press |location=New York |isbn=0-13-146308-X |pages=246}}</ref> The specimen had been given to the collection by Count {{lang|de|italics=unset|[[Pappenheim (state)|Friedrich Ferdinand zu Pappenheim]]}} around 1780, having been recovered from a [[lithographic limestone]] quarry in {{lang|de|italics=unset|[[Eichstätt]]}}.<ref name=brougham1844>{{cite journal | last1 =Brougham|first1= Henry P. |date=1844|title=Dialogues on instinct; with analytical view of the researches on fossil osteology|volume=19 |journal= Knight's Weekly Volume for All Readers|url=https://www.biodiversitylibrary.org/item/229475#page/5/mode/1up}}</ref> The actual date of the specimen's discovery and entry into the collection is unknown however, and it was not mentioned in a catalogue of the collection taken in 1767, so it must have been acquired at some point between that date and its 1784 description by Collini. This makes it potentially the earliest documented pterosaur find; the "Pester Exemplar" of the species ''[[Pterodactylus micronyx]]'' was described in 1779 and possibly discovered earlier than the Mannheim specimen, but it was at first considered to be a fossilized crustacean, and it was not until 1856 that this species was properly described as a pterosaur by German paleontologist {{lang|de|italics=unset|[[Christian Erich Hermann von Meyer|Hermann von Meyer]]}}.<ref name=pester>{{cite journal | last1 = Ősi | first1 = A. | last2 = Prondvai | first2 = E. | last3 = Géczy | first3 = B. | year = 2010 | title = The history of Late Jurassic pterosaurs housed in Hungarian collections and the revision of the holotype of ''Pterodactylus micronyx'' Meyer 1856 (a 'Pester Exemplar') | journal = Geological Society, London, Special Publications | volume = 343 | issue = 1| pages = 277–286 | url = https://www.researchgate.net/publication/258391479 |doi=10.1144/SP343.17 | bibcode = 2010GSLSP.343..277O | s2cid = 129805068 }}</ref>
[[File:Aquatic Pterodactylus.jpg|thumb|left|Wagler's 1830 restoration of an aquatic ''Pterodactylus'']]
In his first description of the Mannheim specimen, Collini did not conclude that it was a flying animal. In fact, Collini could not fathom what kind of animal it might have been, rejecting affinities with the birds or the bats. He speculated that it may have been a sea creature, not for any anatomical reason, but because he thought the ocean depths were more likely to have housed unknown types of animals.<ref name="collini1784">{{Cite journal |last=Collini| first=C A. |date=1784|title=Sur quelques Zoolithes du Cabinet d'Histoire naturelle de S. A. S. E. Palatine & de Bavière, à Mannheim|journal=Acta Theodoro-Palatinae Mannheim |volume=5 Physicum|pages= 58–103 (1 plate)|url=http://www.rhinoresourcecenter.com/index.php?s=1&act=pdfviewer&id=1292315455&folder=129|language=fr}}</ref><ref name="taquet&padian2004">{{Cite journal | doi = 10.1016/j.crpv.2004.02.002 |last1=Taquet|first1= P. |last2=Padian|first2= K. | year = 2004 | title = The earliest known restoration of a pterosaur and the philosophical origins of Cuvier's ''Ossemens Fossiles'' | journal = Comptes Rendus Palevol | volume = 3 | issue = 2| pages = 157–175 }}</ref> The idea that pterosaurs were aquatic animals persisted among a minority of scientists as late as 1830, when the German zoologist [[Johann Georg Wagler]] published a text on "amphibians" which included an illustration of ''Pterodactylus'' using its wings as flippers. Wagler went so far as to classify ''Pterodactylus'', along with other aquatic vertebrates (namely [[plesiosaur]]s, [[ichthyosaur]]s, and [[monotreme]]s), in the class Gryphi, between birds and mammals.<ref>{{Cite book|last=Wagler|first=Johann Georg|url=https://www.biodiversitylibrary.org/item/120243|title=Natürliches System der Amphibien : mit vorangehender Classification der Säugethiere und Vögel : ein Beitrag zur vergleichenden Zoologie|date=1830|location=München|language=de}}</ref>
' |
New page size (new_size) | 82922 |
Old page size (old_size) | 82959 |
Size change in edit (edit_delta) | -37 |
Lines added in edit (added_lines) | [
0 => '{{short description|Genus of terodactyloid terosaur from the Late Jurassic}}',
1 => '{{Other uses|terodactyl (disambiguation)}}',
2 => '{{hatnote|terodactyl is also an informal name for members of the order [[terosaur]]ia.}}',
3 => '| taxon = terodactylus',
4 => '|''terotherium'' <br/><small>Fischer, 1813<ref>{{Cite book|last=Fischer von Waldheim|first=Gotthelf|url=https://www.biodiversitylibrary.org/item/92502|title=Zoognosia tabulis synopticis illustrata : in usum praelectionum mperalis Medico-Chirurgicae Mosquenis edita|date=1813|publisher=Typis Nicolai S. Vsevolozsky|volume=1 |location=Mosquae [Moscow]|pages=466 }}</ref></small>',
5 => '|''terodactylus longirostris'' <br/><small>[[Georges Cuvier|Cuvier]], 1819</small>',
6 => '|''terodactylus crocodilocephaloides'' <br/><small>Ritgen, 1826</small>',
7 => '|''terodactylus spectabilis'' <br/><small>Meyer, 1861</small>',
8 => '|''terodactylus westmani'' <br/><small>Wiman, 1925</small>',
9 => '|''terodactylus cormoranus'' <br/><small>Döderlein, 1929</small>',
10 => ''''''terodactylus''''' (from Greek {{lang|grc-Latn|terodáktylos}} ({{wikt-lang|grc|πτεροδάκτυλος}}) meaning 'winged finger'<ref>{{Cite journal|last=Gudger|first=E.W.|date=1944|title=The Earliest Winged Fish-Catchers|jstor=18398|journal=The Scientific Monthly|volume=59|issue=2|pages=120–129|bibcode=1944SciMo..59..120G}}</ref>) is an extinct [[genus]] of [[terosaur]]s, whose members are commonly known as '''terodactyls''' ({{IPAc-en|ˌ|t|ɛr|ə|ˈ|d|æ|k|t|ᵻ|l|z}}). It is thought to contain only a single [[species]], '''''terodactylus antiquus''''', which was the first terosaur to be named and identified as a flying [[reptile]].',
11 => '[[Fossil]] remains of ''terodactylus'' have primarily been found in the [[Solnhofen limestone]] of [[Bavaria]], [[Germany]], which dates back to the [[Late Jurassic]] period (early [[Tithonian]] stage), about 150.8 to 148.5 million years ago. More fragmentary remains of ''terodactylus'' have tentatively been identified from elsewhere in [[Europe]], as well as in [[Africa]].<ref name="ammonitedate">{{cite journal|author=Schweigert, G. |year=2007|title=Ammonite biostratigraphy as a tool for dating Upper Jurassic lithographic limestones from South Germany – first results and open questions|journal=Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen|volume=245|issue=1|pages= 117–125|doi=10.1127/0077-7749/2007/0245-0117}}</ref>',
12 => '''terodactylus'' was a generalist [[carnivore]] that probably fed on a variety of invertebrates and vertebrates. Like all terosaurs, ''terodactylus'' had wings formed by a skin and muscle membrane stretching from its elongated fourth finger to its hind limbs. It was supported internally by [[collagen]] fibres and externally by [[keratin]]ous ridges. ''terodactylus'' was a small terosaur compared to other famous genera such as ''[[teranodon]]'' and ''[[Quetzalcoatlus]]'', and it also lived earlier, during the Late Jurassic period, while both ''teranodon'' and ''Quetzalcoatlus'' lived during the [[Late Cretaceous]]. ''terodactylus'' lived alongside other small terosaurs such as the well-known ''[[Rhamphorhynchus]]'', as well as other genera such as ''[[Scaphognathus]]'', ''[[Anurognathus]]'' and ''[[Ctenochasma]]''. ''terodactylus'' is classified as an early-branching member of the [[Euctenochasmatia|ctenochasmatid lineage]], within the terosaur clade [[terodactyloidea]].<ref name="unwin2003" /><ref name=longrichetal2018 />',
13 => '[[File:terodactylus holotype Collini 1784.jpg|thumb|left|Original copper engraving of the ''P. antiquus'' [[holotype]] by Egid Verhelst II and published by Italian scientist {{lang|it|italics=unset|[[Cosimo Alessandro Collini]]}}, 1784]]',
14 => 'The [[type specimen]] of the animal now known as ''terodactylus antiquus'' was the first terosaur fossil ever to be identified. The first ''terodactylus'' specimen was described by the Italian scientist {{lang|it|italics=unset|[[Cosimo Alessandro Collini]]}} in 1784, based on a fossil skeleton that had been unearthed from the Solnhofen limestone of Bavaria. Collini was the curator of the {{lang|de|Naturalienkabinett}}, or nature [[cabinet of curiosities]] (a precursor to the modern concept of the [[natural history]] museum), in the palace of [[Charles Theodore, Elector of Bavaria]] at [[Mannheim]].<ref name=pester /><ref name=DU06>{{cite book |last=Unwin |first=David M. |title=The terosaurs: From Deep Time |year=2006 |publisher=Pi Press |location=New York |isbn=0-13-146308-X |pages=246}}</ref> The specimen had been given to the collection by Count {{lang|de|italics=unset|[[Pappenheim (state)|Friedrich Ferdinand zu Pappenheim]]}} around 1780, having been recovered from a [[lithographic limestone]] quarry in {{lang|de|italics=unset|[[Eichstätt]]}}.<ref name=brougham1844>{{cite journal | last1 =Brougham|first1= Henry P. |date=1844|title=Dialogues on instinct; with analytical view of the researches on fossil osteology|volume=19 |journal= Knight's Weekly Volume for All Readers|url=https://www.biodiversitylibrary.org/item/229475#page/5/mode/1up}}</ref> The actual date of the specimen's discovery and entry into the collection is unknown however, and it was not mentioned in a catalogue of the collection taken in 1767, so it must have been acquired at some point between that date and its 1784 description by Collini. This makes it potentially the earliest documented pterosaur find; the "Pester Exemplar" of the species ''[[Pterodactylus micronyx]]'' was described in 1779 and possibly discovered earlier than the Mannheim specimen, but it was at first considered to be a fossilized crustacean, and it was not until 1856 that this species was properly described as a pterosaur by German paleontologist {{lang|de|italics=unset|[[Christian Erich Hermann von Meyer|Hermann von Meyer]]}}.<ref name=pester>{{cite journal | last1 = Ősi | first1 = A. | last2 = Prondvai | first2 = E. | last3 = Géczy | first3 = B. | year = 2010 | title = The history of Late Jurassic pterosaurs housed in Hungarian collections and the revision of the holotype of ''Pterodactylus micronyx'' Meyer 1856 (a 'Pester Exemplar') | journal = Geological Society, London, Special Publications | volume = 343 | issue = 1| pages = 277–286 | url = https://www.researchgate.net/publication/258391479 |doi=10.1144/SP343.17 | bibcode = 2010GSLSP.343..277O | s2cid = 129805068 }}</ref>'
] |
Lines removed in edit (removed_lines) | [
0 => '{{short description|Genus of pterodactyloid pterosaur from the Late Jurassic}}',
1 => '{{Other uses|Pterodactyl (disambiguation)}}',
2 => '{{hatnote|Pterodactyl is also an informal name for members of the order [[Pterosaur]]ia.}}',
3 => '| taxon = Pterodactylus',
4 => '|''Pterotherium'' <br/><small>Fischer, 1813<ref>{{Cite book|last=Fischer von Waldheim|first=Gotthelf|url=https://www.biodiversitylibrary.org/item/92502|title=Zoognosia tabulis synopticis illustrata : in usum praelectionum mperalis Medico-Chirurgicae Mosquenis edita|date=1813|publisher=Typis Nicolai S. Vsevolozsky|volume=1 |location=Mosquae [Moscow]|pages=466 }}</ref></small>',
5 => '|''Pterodactylus longirostris'' <br/><small>[[Georges Cuvier|Cuvier]], 1819</small>',
6 => '|''Pterodactylus crocodilocephaloides'' <br/><small>Ritgen, 1826</small>',
7 => '|''Pterodactylus spectabilis'' <br/><small>Meyer, 1861</small>',
8 => '|''Pterodactylus westmani'' <br/><small>Wiman, 1925</small>',
9 => '|''Pterodactylus cormoranus'' <br/><small>Döderlein, 1929</small>',
10 => ''''''Pterodactylus''''' (from Greek {{lang|grc-Latn|pterodáktylos}} ({{wikt-lang|grc|πτεροδάκτυλος}}) meaning 'winged finger'<ref>{{Cite journal|last=Gudger|first=E.W.|date=1944|title=The Earliest Winged Fish-Catchers|jstor=18398|journal=The Scientific Monthly|volume=59|issue=2|pages=120–129|bibcode=1944SciMo..59..120G}}</ref>) is an extinct [[genus]] of [[pterosaur]]s, whose members are commonly known as '''pterodactyls''' ({{IPAc-en|ˌ|t|ɛr|ə|ˈ|d|æ|k|t|ᵻ|l|z}}). It is thought to contain only a single [[species]], '''''Pterodactylus antiquus''''', which was the first pterosaur to be named and identified as a flying [[reptile]].',
11 => '[[Fossil]] remains of ''Pterodactylus'' have primarily been found in the [[Solnhofen limestone]] of [[Bavaria]], [[Germany]], which dates back to the [[Late Jurassic]] period (early [[Tithonian]] stage), about 150.8 to 148.5 million years ago. More fragmentary remains of ''Pterodactylus'' have tentatively been identified from elsewhere in [[Europe]], as well as in [[Africa]].<ref name="ammonitedate">{{cite journal|author=Schweigert, G. |year=2007|title=Ammonite biostratigraphy as a tool for dating Upper Jurassic lithographic limestones from South Germany – first results and open questions|journal=Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen|volume=245|issue=1|pages= 117–125|doi=10.1127/0077-7749/2007/0245-0117}}</ref>',
12 => '''Pterodactylus'' was a generalist [[carnivore]] that probably fed on a variety of invertebrates and vertebrates. Like all pterosaurs, ''Pterodactylus'' had wings formed by a skin and muscle membrane stretching from its elongated fourth finger to its hind limbs. It was supported internally by [[collagen]] fibres and externally by [[keratin]]ous ridges. ''Pterodactylus'' was a small pterosaur compared to other famous genera such as ''[[Pteranodon]]'' and ''[[Quetzalcoatlus]]'', and it also lived earlier, during the Late Jurassic period, while both ''Pteranodon'' and ''Quetzalcoatlus'' lived during the [[Late Cretaceous]]. ''Pterodactylus'' lived alongside other small pterosaurs such as the well-known ''[[Rhamphorhynchus]]'', as well as other genera such as ''[[Scaphognathus]]'', ''[[Anurognathus]]'' and ''[[Ctenochasma]]''. ''Pterodactylus'' is classified as an early-branching member of the [[Euctenochasmatia|ctenochasmatid lineage]], within the pterosaur clade [[Pterodactyloidea]].<ref name="unwin2003" /><ref name=longrichetal2018 />',
13 => '[[File:Pterodactylus holotype Collini 1784.jpg|thumb|left|Original copper engraving of the ''P. antiquus'' [[holotype]] by Egid Verhelst II and published by Italian scientist {{lang|it|italics=unset|[[Cosimo Alessandro Collini]]}}, 1784]]',
14 => 'The [[type specimen]] of the animal now known as ''Pterodactylus antiquus'' was the first pterosaur fossil ever to be identified. The first ''Pterodactylus'' specimen was described by the Italian scientist {{lang|it|italics=unset|[[Cosimo Alessandro Collini]]}} in 1784, based on a fossil skeleton that had been unearthed from the Solnhofen limestone of Bavaria. Collini was the curator of the {{lang|de|Naturalienkabinett}}, or nature [[cabinet of curiosities]] (a precursor to the modern concept of the [[natural history]] museum), in the palace of [[Charles Theodore, Elector of Bavaria]] at [[Mannheim]].<ref name=pester /><ref name=DU06>{{cite book |last=Unwin |first=David M. |title=The Pterosaurs: From Deep Time |year=2006 |publisher=Pi Press |location=New York |isbn=0-13-146308-X |pages=246}}</ref> The specimen had been given to the collection by Count {{lang|de|italics=unset|[[Pappenheim (state)|Friedrich Ferdinand zu Pappenheim]]}} around 1780, having been recovered from a [[lithographic limestone]] quarry in {{lang|de|italics=unset|[[Eichstätt]]}}.<ref name=brougham1844>{{cite journal | last1 =Brougham|first1= Henry P. |date=1844|title=Dialogues on instinct; with analytical view of the researches on fossil osteology|volume=19 |journal= Knight's Weekly Volume for All Readers|url=https://www.biodiversitylibrary.org/item/229475#page/5/mode/1up}}</ref> The actual date of the specimen's discovery and entry into the collection is unknown however, and it was not mentioned in a catalogue of the collection taken in 1767, so it must have been acquired at some point between that date and its 1784 description by Collini. This makes it potentially the earliest documented pterosaur find; the "Pester Exemplar" of the species ''[[Pterodactylus micronyx]]'' was described in 1779 and possibly discovered earlier than the Mannheim specimen, but it was at first considered to be a fossilized crustacean, and it was not until 1856 that this species was properly described as a pterosaur by German paleontologist {{lang|de|italics=unset|[[Christian Erich Hermann von Meyer|Hermann von Meyer]]}}.<ref name=pester>{{cite journal | last1 = Ősi | first1 = A. | last2 = Prondvai | first2 = E. | last3 = Géczy | first3 = B. | year = 2010 | title = The history of Late Jurassic pterosaurs housed in Hungarian collections and the revision of the holotype of ''Pterodactylus micronyx'' Meyer 1856 (a 'Pester Exemplar') | journal = Geological Society, London, Special Publications | volume = 343 | issue = 1| pages = 277–286 | url = https://www.researchgate.net/publication/258391479 |doi=10.1144/SP343.17 | bibcode = 2010GSLSP.343..277O | s2cid = 129805068 }}</ref>'
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Parsed HTML source of the new revision (new_html) | '<div class="mw-parser-output"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Genus of terodactyloid terosaur from the Late Jurassic</div>
<div role="note" class="hatnote navigation-not-searchable">For other uses, see <a href="/enwiki/w/index.php?title=Terodactyl_(disambiguation)&action=edit&redlink=1" class="new" title="Terodactyl (disambiguation) (page does not exist)">terodactyl (disambiguation)</a>.</div>
<div role="note" class="hatnote navigation-not-searchable">terodactyl is also an informal name for members of the order <a href="/enwiki/w/index.php?title=Terosaur&action=edit&redlink=1" class="new" title="Terosaur (page does not exist)">terosauria</a>.</div>
<p class="mw-empty-elt">
</p>
<table class="infobox biota" style="text-align: left; width: 200px; font-size: 100%">
<tbody><tr>
<th colspan="2" style="text-align: center; background-color: transparent">Pterodactylus<br /><div style="font-size: 85%;">Temporal range: Early <a href="/enwiki/wiki/Tithonian" title="Tithonian">Tithonian</a>, <br /><span style="display:inline-block;"></span><span style="display:inline-block;">150.8–148.5 <a href="/enwiki/wiki/Megaannum" class="mw-redirect" title="Megaannum">Ma</a></span> <span style="display:inline-block;"></span><div id="Timeline-row" style="margin: 4px auto 0; clear:both; width:220px; padding:0px; height:18px; overflow:visible; border:1px #666; border-style:solid none; position:relative; z-index:0; font-size:13px;">
<div style="position:absolute; height:100%; left:0px; width:208.07692307692px; padding-left:5px; text-align:left; background-color:rgb(254,214,123); background-image: linear-gradient(to right, rgba(255,255,255,1), rgba(254,217,106,1) 15%, rgba(254,217,106,1));"><a href="/enwiki/wiki/Precambrian" title="Precambrian">PreꞒ</a></div>
<div style="position:absolute; height:100%; text-align:center; background-color:rgb(129,170,114); left:36.892307692308px; width:18.818461538462px;"><a href="/enwiki/wiki/Cambrian" title="Cambrian">Ꞓ</a></div>
<div style="position:absolute; height:100%; text-align:center; background-color:rgb(0,169,138); left:55.710769230769px; width:14.08px;"><a href="/enwiki/wiki/Ordovician" title="Ordovician">O</a></div>
<div style="position:absolute; height:100%; text-align:center; background-color:rgb(166,223,197); left:69.790769230769px; width:8.3261538461539px;"><a href="/enwiki/wiki/Silurian" title="Silurian">S</a></div>
<div style="position:absolute; height:100%; text-align:center; background-color:rgb(221,150,81); left:78.116923076923px; width:20.409230769231px;"><a href="/enwiki/wiki/Devonian" title="Devonian">D</a></div>
<div style="position:absolute; height:100%; text-align:center; background-color:rgb(63,174,173); left:98.526153846154px; width:20.307692307692px;"><a href="/enwiki/wiki/Carboniferous" title="Carboniferous">C</a></div>
<div style="position:absolute; height:100%; text-align:center; background-color:rgb(247,88,60); left:118.83384615385px; width:15.907015384615px;"><a href="/enwiki/wiki/Permian" title="Permian">P</a></div>
<div style="position:absolute; height:100%; text-align:center; background-color:rgb(153,78,150); left:134.74086153846px; width:17.126830769231px;"><a href="/enwiki/wiki/Triassic" title="Triassic">T</a></div>
<div style="position:absolute; height:100%; text-align:center; background-color:rgb(0,187,231); left:151.86769230769px; width:19.055384615385px;"><a href="/enwiki/wiki/Jurassic" title="Jurassic">J</a></div>
<div style="position:absolute; height:100%; text-align:center; background-color:rgb(111,200,107); left:170.92307692308px; width:26.738461538462px;"><a href="/enwiki/wiki/Cretaceous" title="Cretaceous">K</a></div>
<div style="position:absolute; height:100%; text-align:center; background-color:rgb(254,161,99); left:197.66153846154px; width:14.543692307692px;"><a href="/enwiki/wiki/Paleogene" title="Paleogene"><small>Pg</small></a></div>
<div style="position:absolute; height:100%; text-align:center; background-color:rgb(254,221,45); left:212.20523076923px; width:6.9215384615385px;"><a href="/enwiki/wiki/Neogene" title="Neogene"><small>N</small></a></div>
<div id="end-border" style="position:absolute; height:100%; background-color:#666; width:1px; left:219px"></div><div style="margin:0 auto; line-height:0; clear:both; width:220px; padding:0px; height:8px; overflow:visible; background-color:transparent; position:relative; top:-4px; z-index:100;">
<div style="position:absolute; left:168.96px; font-size:50%"><div style="position:relative; left:-0.42em">↓</div></div>
</div>
</div></div>
</th></tr>
<tr>
<td colspan="2" style="text-align: center"><a href="/enwiki/wiki/File:Bsp_as_i_739_modified.png" class="image"><img alt="Bsp as i 739 modified.png" src="/upwiki/wikipedia/commons/thumb/f/f9/Bsp_as_i_739_modified.png/220px-Bsp_as_i_739_modified.png" decoding="async" width="220" height="141" srcset="/upwiki/wikipedia/commons/thumb/f/f9/Bsp_as_i_739_modified.png/330px-Bsp_as_i_739_modified.png 1.5x, /upwiki/wikipedia/commons/thumb/f/f9/Bsp_as_i_739_modified.png/440px-Bsp_as_i_739_modified.png 2x" data-file-width="856" data-file-height="549" /></a>
</td></tr>
<tr>
<td colspan="2" style="text-align: center; font-size: 88%">Sub-adult <a href="/enwiki/wiki/Type_specimen" class="mw-redirect" title="Type specimen">type specimen</a> of <i>P. antiquus</i>, Bavarian State Colection for Palaeontology and Geology
</td></tr>
<tr>
<th colspan="2" style="min-width:15em; text-align: center; background-color: transparent"><a href="/enwiki/wiki/Taxonomy_(biology)" title="Taxonomy (biology)">Scientific classification</a> <span class="plainlinks" style="font-size:smaller; float:right; padding-right:0.4em; margin-left:-3em;"><a href="/enwiki/wiki/Template:Taxonomy/terodactylus" title="e"><img alt="e" src="/upwiki/wikipedia/commons/7/74/Red_Pencil_Icon.png" decoding="async" width="16" height="16" data-file-width="16" data-file-height="16" /></a></span>
</th></tr>
<tr>
<td><span style="color: Red;">Missing taxonomy template</span> (<span class="plainlinks"><a class="external text" href="https://en.wikipedia.org/enwiki/w/index.php?action=edit&title=Template:taxonomy/Terodactylus&preload=Template:Taxonomy/preload">fix</a></span>):
</td>
<td><a href="/enwiki/w/index.php?title=Terodactylus&action=edit&redlink=1" class="new" title="Terodactylus (page does not exist)">terodactylus</a>
</td></tr>
<tr>
<th colspan="2" style="text-align: center; background-color: transparent"><a href="/enwiki/wiki/Type_species" title="Type species">Type species</a>
</th></tr>
<tr>
<td colspan="2" style="text-align: center"><abbr title="Extinct" aria-label="Extinct" style="border: none; text-decoration: none; cursor: inherit; font-weight: normal; font-style: normal;">†</abbr><i><b>Ornithocephalus antiquus</b></i><br /><div style="font-size: 85%;"><a href="/enwiki/wiki/Samuel_Thomas_von_S%C3%B6mmerring" title="Samuel Thomas von Sömmerring">Sömmerring</a>, 1812</div>
</td></tr>
<tr>
<th colspan="2" style="text-align: center; background-color: transparent"><a href="/enwiki/wiki/Species" title="Species">Species</a>
</th></tr>
<tr>
<td colspan="2" style="text-align: left">
<ul><li><abbr title="Extinct" aria-label="Extinct" style="border: none; text-decoration: none; cursor: inherit; font-weight: normal; font-style: normal;">†</abbr><i><b>P. antiquus</b></i> <br /><small>(Sömmerring, 1812)</small></li>
<li><abbr title="Extinct" aria-label="Extinct" style="border: none; text-decoration: none; cursor: inherit; font-weight: normal; font-style: normal;">†</abbr><i><a href="/enwiki/wiki/Diopecephalus" title="Diopecephalus">P. kochi</a></i>? <br /><small>(<a href="/enwiki/wiki/Johann_Andreas_Wagner" title="Johann Andreas Wagner">Wagner</a>, 1837)</small></li>
<li><abbr title="Extinct" aria-label="Extinct" style="border: none; text-decoration: none; cursor: inherit; font-weight: normal; font-style: normal;">†</abbr><i><a href="/enwiki/wiki/Aerodactylus" title="Aerodactylus">P. scolopaciceps</a></i>? <br /><small>(<a href="/enwiki/wiki/Hermann_von_Meyer" class="mw-redirect" title="Hermann von Meyer">Meyer</a>, 1860)</small></li></ul>
</td></tr>
<tr>
<th colspan="2" style="text-align: center; background-color: transparent"><a href="/enwiki/wiki/Synonym_(taxonomy)" title="Synonym (taxonomy)">Synonyms</a>
</th></tr>
<tr>
<td colspan="2" style="text-align: left">
<div class="mw-collapsible mw-collapsed" style="text-align: center; font-size: 95%;">
<div style="line-height: 1.6em; font-weight: bold; font-size: 100%; text-align: left;"><div><small>Genus synonymy</small></div></div>
<ul class="mw-collapsible-content" style="font-size: 105%; margin-top: 0; margin-bottom: 0; line-height: inherit; text-align: left;"><li style="line-height: inherit; margin: 0"><i>Ornithocephalus</i> <br /><small>Sömmering, 1812</small>
</li><li style="line-height: inherit; margin: 0"><i>terotherium</i> <br /><small>Fischer, 1813<sup id="cite_ref-1" class="reference"><a href="#cite_note-1">[1]</a></sup></small>
</li><li style="line-height: inherit; margin: 0"><i>Macrotrachelus</i> <br /><small><a href="/enwiki/wiki/Christoph_Gottfried_Andreas_Giebel" title="Christoph Gottfried Andreas Giebel">Giebel</a>, 1852</small>
</li><li style="line-height: inherit; margin: 0"><i><a href="/enwiki/wiki/Diopecephalus" title="Diopecephalus">Diopecephalus</a></i>? <br /><small><a href="/enwiki/wiki/Harry_Seeley" title="Harry Seeley">Seeley</a>, 1871</small>
</li><li style="line-height: inherit; margin: 0"><i><a href="/enwiki/wiki/Aerodactylus" title="Aerodactylus">Aerodactylus</a></i>? <br /><small>Vidovic & Martill, 2014</small>
</li></ul>
</div>
<div class="mw-collapsible mw-collapsed" style="text-align: center; font-size: 95%;">
<div style="line-height: 1.6em; font-weight: bold; font-size: 100%; text-align: left;"><div><small>Species synonymy</small></div></div>
<ul class="mw-collapsible-content" style="font-size: 105%; margin-top: 0; margin-bottom: 0; line-height: inherit; text-align: left;"><li style="line-height: inherit; margin: 0"><i>Ornithocephalus antiquus</i> <br /><small>Sömmering, 1812</small>
</li><li style="line-height: inherit; margin: 0"><i>terodactylus longirostris</i> <br /><small><a href="/enwiki/wiki/Georges_Cuvier" title="Georges Cuvier">Cuvier</a>, 1819</small>
</li><li style="line-height: inherit; margin: 0"><i>Ornithocephalus longirostris</i> <br /><small>(Cuvier, 1819)</small>
</li><li style="line-height: inherit; margin: 0"><i>Macrotrachelus longirostris</i> <br /><small>(Cuvier, 1819)</small>
</li><li style="line-height: inherit; margin: 0"><i>terodactylus crocodilocephaloides</i> <br /><small>Ritgen, 1826</small>
</li><li style="line-height: inherit; margin: 0"><i>Ornithocephalus kochi</i>? <br /><small>Wagner, 1837</small>
</li><li style="line-height: inherit; margin: 0"><i>Diopecephalus kochi</i>? <br /><small>(Wagner, 1837)</small>
</li><li style="line-height: inherit; margin: 0"><i>terodactylus spectabilis</i> <br /><small>Meyer, 1861</small>
</li><li style="line-height: inherit; margin: 0"><i>terodactylus westmani</i> <br /><small>Wiman, 1925</small>
</li><li style="line-height: inherit; margin: 0"><i>terodactylus cormoranus</i> <br /><small>Döderlein, 1929</small>
</li><li style="line-height: inherit; margin: 0"><i>Aerodactylus scolopaciceps</i> <br /><small>(Meyer, 1860)</small>
</li></ul>
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<p><i><b>terodactylus</b></i> (from Greek <i lang="grc-Latn" title="Ancient Greek (to 1453)-language text">terodáktylos</i> (<span lang="grc"><a href="https://en.wiktionary.org/wiki/%CF%80%CF%84%CE%B5%CF%81%CE%BF%CE%B4%CE%AC%CE%BA%CF%84%CF%85%CE%BB%CE%BF%CF%82#Ancient_Greek" class="extiw" title="wikt:πτεροδάκτυλος">πτεροδάκτυλος</a></span>) meaning 'winged finger'<sup id="cite_ref-2" class="reference"><a href="#cite_note-2">[2]</a></sup>) is an extinct <a href="/enwiki/wiki/Genus" title="Genus">genus</a> of <a href="/enwiki/w/index.php?title=Terosaur&action=edit&redlink=1" class="new" title="Terosaur (page does not exist)">terosaurs</a>, whose members are commonly known as <b>terodactyls</b> (<span class="rt-commentedText nowrap"><span class="IPA nopopups noexcerpt"><a href="/enwiki/wiki/Help:IPA/English" title="Help:IPA/English">/<span style="border-bottom:1px dotted"><span title="/ˌ/: secondary stress follows">ˌ</span><span title="'t' in 'tie'">t</span><span title="/ɛr/: 'err' in 'merry'">ɛr</span><span title="/ə/: 'a' in 'about'">ə</span><span title="/ˈ/: primary stress follows">ˈ</span><span title="'d' in 'dye'">d</span><span title="/æ/: 'a' in 'bad'">æ</span><span title="'k' in 'kind'">k</span><span title="'t' in 'tie'">t</span><span title="/ɪ/: 'i' in 'kit'">ɪ</span><span title="'l' in 'lie'">l</span><span title="'z' in 'zoom'">z</span></span>/</a></span></span>). It is thought to contain only a single <a href="/enwiki/wiki/Species" title="Species">species</a>, <i><b>terodactylus antiquus</b></i>, which was the first terosaur to be named and identified as a flying <a href="/enwiki/wiki/Reptile" title="Reptile">reptile</a>.
</p><p><a href="/enwiki/wiki/Fossil" title="Fossil">Fossil</a> remains of <i>terodactylus</i> have primarily been found in the <a href="/enwiki/wiki/Solnhofen_limestone" class="mw-redirect" title="Solnhofen limestone">Solnhofen limestone</a> of <a href="/enwiki/wiki/Bavaria" title="Bavaria">Bavaria</a>, <a href="/enwiki/wiki/Germany" title="Germany">Germany</a>, which dates back to the <a href="/enwiki/wiki/Late_Jurassic" title="Late Jurassic">Late Jurassic</a> period (early <a href="/enwiki/wiki/Tithonian" title="Tithonian">Tithonian</a> stage), about 150.8 to 148.5 million years ago. More fragmentary remains of <i>terodactylus</i> have tentatively been identified from elsewhere in <a href="/enwiki/wiki/Europe" title="Europe">Europe</a>, as well as in <a href="/enwiki/wiki/Africa" title="Africa">Africa</a>.<sup id="cite_ref-ammonitedate_3-0" class="reference"><a href="#cite_note-ammonitedate-3">[3]</a></sup>
</p><p><i>terodactylus</i> was a generalist <a href="/enwiki/wiki/Carnivore" title="Carnivore">carnivore</a> that probably fed on a variety of invertebrates and vertebrates. Like all terosaurs, <i>terodactylus</i> had wings formed by a skin and muscle membrane stretching from its elongated fourth finger to its hind limbs. It was supported internally by <a href="/enwiki/wiki/Collagen" title="Collagen">collagen</a> fibres and externally by <a href="/enwiki/wiki/Keratin" title="Keratin">keratinous</a> ridges. <i>terodactylus</i> was a small terosaur compared to other famous genera such as <i><a href="/enwiki/w/index.php?title=Teranodon&action=edit&redlink=1" class="new" title="Teranodon (page does not exist)">teranodon</a></i> and <i><a href="/enwiki/wiki/Quetzalcoatlus" title="Quetzalcoatlus">Quetzalcoatlus</a></i>, and it also lived earlier, during the Late Jurassic period, while both <i>teranodon</i> and <i>Quetzalcoatlus</i> lived during the <a href="/enwiki/wiki/Late_Cretaceous" title="Late Cretaceous">Late Cretaceous</a>. <i>terodactylus</i> lived alongside other small terosaurs such as the well-known <i><a href="/enwiki/wiki/Rhamphorhynchus" title="Rhamphorhynchus">Rhamphorhynchus</a></i>, as well as other genera such as <i><a href="/enwiki/wiki/Scaphognathus" title="Scaphognathus">Scaphognathus</a></i>, <i><a href="/enwiki/wiki/Anurognathus" title="Anurognathus">Anurognathus</a></i> and <i><a href="/enwiki/wiki/Ctenochasma" title="Ctenochasma">Ctenochasma</a></i>. <i>terodactylus</i> is classified as an early-branching member of the <a href="/enwiki/wiki/Euctenochasmatia" title="Euctenochasmatia">ctenochasmatid lineage</a>, within the terosaur clade <a href="/enwiki/w/index.php?title=Terodactyloidea&action=edit&redlink=1" class="new" title="Terodactyloidea (page does not exist)">terodactyloidea</a>.<sup id="cite_ref-unwin2003_4-0" class="reference"><a href="#cite_note-unwin2003-4">[4]</a></sup><sup id="cite_ref-longrichetal2018_5-0" class="reference"><a href="#cite_note-longrichetal2018-5">[5]</a></sup>
</p>
<div id="toc" class="toc" role="navigation" aria-labelledby="mw-toc-heading"><input type="checkbox" role="button" id="toctogglecheckbox" class="toctogglecheckbox" style="display:none" /><div class="toctitle" lang="en" dir="ltr"><h2 id="mw-toc-heading">Contents</h2><span class="toctogglespan"><label class="toctogglelabel" for="toctogglecheckbox"></label></span></div>
<ul>
<li class="toclevel-1 tocsection-1"><a href="#Discovery_and_history"><span class="tocnumber">1</span> <span class="toctext">Discovery and history</span></a></li>
<li class="toclevel-1 tocsection-2"><a href="#Description"><span class="tocnumber">2</span> <span class="toctext">Description</span></a></li>
<li class="toclevel-1 tocsection-3"><a href="#Paleobiology"><span class="tocnumber">3</span> <span class="toctext">Paleobiology</span></a>
<ul>
<li class="toclevel-2 tocsection-4"><a href="#Life_history"><span class="tocnumber">3.1</span> <span class="toctext">Life history</span></a></li>
<li class="toclevel-2 tocsection-5"><a href="#Growth_and_breeding_seasons"><span class="tocnumber">3.2</span> <span class="toctext">Growth and breeding seasons</span></a></li>
<li class="toclevel-2 tocsection-6"><a href="#Daily_activity_patterns"><span class="tocnumber">3.3</span> <span class="toctext">Daily activity patterns</span></a></li>
<li class="toclevel-2 tocsection-7"><a href="#Diet"><span class="tocnumber">3.4</span> <span class="toctext">Diet</span></a></li>
</ul>
</li>
<li class="toclevel-1 tocsection-8"><a href="#Paleoecology"><span class="tocnumber">4</span> <span class="toctext">Paleoecology</span></a></li>
<li class="toclevel-1 tocsection-9"><a href="#Classification"><span class="tocnumber">5</span> <span class="toctext">Classification</span></a>
<ul>
<li class="toclevel-2 tocsection-10"><a href="#Formerly_assigned_species"><span class="tocnumber">5.1</span> <span class="toctext">Formerly assigned species</span></a></li>
<li class="toclevel-2 tocsection-11"><a href="#Controversial_species"><span class="tocnumber">5.2</span> <span class="toctext">Controversial species</span></a></li>
<li class="toclevel-2 tocsection-12"><a href="#List_of_species"><span class="tocnumber">5.3</span> <span class="toctext">List of species</span></a></li>
</ul>
</li>
<li class="toclevel-1 tocsection-13"><a href="#Cultural_significance"><span class="tocnumber">6</span> <span class="toctext">Cultural significance</span></a></li>
<li class="toclevel-1 tocsection-14"><a href="#See_also"><span class="tocnumber">7</span> <span class="toctext">See also</span></a></li>
<li class="toclevel-1 tocsection-15"><a href="#References"><span class="tocnumber">8</span> <span class="toctext">References</span></a></li>
<li class="toclevel-1 tocsection-16"><a href="#External_links"><span class="tocnumber">9</span> <span class="toctext">External links</span></a></li>
</ul>
</div>
<h2><span class="mw-headline" id="Discovery_and_history">Discovery and history</span><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Pterodactylus&action=edit&section=1" title="Edit section: Discovery and history">edit</a><span class="mw-editsection-bracket">]</span></span></h2>
<div class="thumb tleft"><div class="thumbinner" style="width:182px;"><a href="/enwiki//en.wikipedia.org/wiki/Special:Upload?wpDestFile=Terodactylus_holotype_Collini_1784.jpg" class="new" title="File:Terodactylus holotype Collini 1784.jpg">File:Terodactylus holotype Collini 1784.jpg</a> <div class="thumbcaption">Original copper engraving of the <i>P. antiquus</i> <a href="/enwiki/wiki/Holotype" title="Holotype">holotype</a> by Egid Verhelst II and published by Italian scientist <span lang="it" title="Italian-language text"><a href="/enwiki/wiki/Cosimo_Alessandro_Collini" title="Cosimo Alessandro Collini">Cosimo Alessandro Collini</a></span>, 1784</div></div></div>
<p>The <a href="/enwiki/wiki/Type_specimen" class="mw-redirect" title="Type specimen">type specimen</a> of the animal now known as <i>terodactylus antiquus</i> was the first terosaur fossil ever to be identified. The first <i>terodactylus</i> specimen was described by the Italian scientist <span lang="it" title="Italian-language text"><a href="/enwiki/wiki/Cosimo_Alessandro_Collini" title="Cosimo Alessandro Collini">Cosimo Alessandro Collini</a></span> in 1784, based on a fossil skeleton that had been unearthed from the Solnhofen limestone of Bavaria. Collini was the curator of the <i lang="de" title="German-language text">Naturalienkabinett</i>, or nature <a href="/enwiki/wiki/Cabinet_of_curiosities" title="Cabinet of curiosities">cabinet of curiosities</a> (a precursor to the modern concept of the <a href="/enwiki/wiki/Natural_history" title="Natural history">natural history</a> museum), in the palace of <a href="/enwiki/wiki/Charles_Theodore,_Elector_of_Bavaria" title="Charles Theodore, Elector of Bavaria">Charles Theodore, Elector of Bavaria</a> at <a href="/enwiki/wiki/Mannheim" title="Mannheim">Mannheim</a>.<sup id="cite_ref-pester_6-0" class="reference"><a href="#cite_note-pester-6">[6]</a></sup><sup id="cite_ref-DU06_7-0" class="reference"><a href="#cite_note-DU06-7">[7]</a></sup> The specimen had been given to the collection by Count <span lang="de" title="German-language text"><a href="/enwiki/wiki/Pappenheim_(state)" title="Pappenheim (state)">Friedrich Ferdinand zu Pappenheim</a></span> around 1780, having been recovered from a <a href="/enwiki/wiki/Lithographic_limestone" title="Lithographic limestone">lithographic limestone</a> quarry in <span lang="de" title="German-language text"><a href="/enwiki/wiki/Eichst%C3%A4tt" title="Eichstätt">Eichstätt</a></span>.<sup id="cite_ref-brougham1844_8-0" class="reference"><a href="#cite_note-brougham1844-8">[8]</a></sup> The actual date of the specimen's discovery and entry into the collection is unknown however, and it was not mentioned in a catalogue of the collection taken in 1767, so it must have been acquired at some point between that date and its 1784 description by Collini. This makes it potentially the earliest documented pterosaur find; the "Pester Exemplar" of the species <i><a href="/enwiki/wiki/Pterodactylus_micronyx" class="mw-redirect" title="Pterodactylus micronyx">Pterodactylus micronyx</a></i> was described in 1779 and possibly discovered earlier than the Mannheim specimen, but it was at first considered to be a fossilized crustacean, and it was not until 1856 that this species was properly described as a pterosaur by German paleontologist <span lang="de" title="German-language text"><a href="/enwiki/wiki/Christian_Erich_Hermann_von_Meyer" title="Christian Erich Hermann von Meyer">Hermann von Meyer</a></span>.<sup id="cite_ref-pester_6-1" class="reference"><a href="#cite_note-pester-6">[6]</a></sup>
</p>
<div class="thumb tleft"><div class="thumbinner" style="width:222px;"><a href="/enwiki/wiki/File:Aquatic_Pterodactylus.jpg" class="image"><img alt="" src="/upwiki/wikipedia/commons/thumb/7/7b/Aquatic_Pterodactylus.jpg/220px-Aquatic_Pterodactylus.jpg" decoding="async" width="220" height="69" class="thumbimage" srcset="/upwiki/wikipedia/commons/thumb/7/7b/Aquatic_Pterodactylus.jpg/330px-Aquatic_Pterodactylus.jpg 1.5x, /upwiki/wikipedia/commons/thumb/7/7b/Aquatic_Pterodactylus.jpg/440px-Aquatic_Pterodactylus.jpg 2x" data-file-width="1499" data-file-height="467" /></a> <div class="thumbcaption"><div class="magnify"><a href="/enwiki/wiki/File:Aquatic_Pterodactylus.jpg" class="internal" title="Enlarge"></a></div>Wagler's 1830 restoration of an aquatic <i>Pterodactylus</i></div></div></div>
<p>In his first description of the Mannheim specimen, Collini did not conclude that it was a flying animal. In fact, Collini could not fathom what kind of animal it might have been, rejecting affinities with the birds or the bats. He speculated that it may have been a sea creature, not for any anatomical reason, but because he thought the ocean depths were more likely to have housed unknown types of animals.<sup id="cite_ref-collini1784_9-0" class="reference"><a href="#cite_note-collini1784-9">[9]</a></sup><sup id="cite_ref-taquet&padian2004_10-0" class="reference"><a href="#cite_note-taquet&padian2004-10">[10]</a></sup> The idea that pterosaurs were aquatic animals persisted among a minority of scientists as late as 1830, when the German zoologist <a href="/enwiki/wiki/Johann_Georg_Wagler" title="Johann Georg Wagler">Johann Georg Wagler</a> published a text on "amphibians" which included an illustration of <i>Pterodactylus</i> using its wings as flippers. Wagler went so far as to classify <i>Pterodactylus</i>, along with other aquatic vertebrates (namely <a href="/enwiki/wiki/Plesiosaur" class="mw-redirect" title="Plesiosaur">plesiosaurs</a>, <a href="/enwiki/wiki/Ichthyosaur" title="Ichthyosaur">ichthyosaurs</a>, and <a href="/enwiki/wiki/Monotreme" title="Monotreme">monotremes</a>), in the class Gryphi, between birds and mammals.<sup id="cite_ref-11" class="reference"><a href="#cite_note-11">[11]</a></sup>
</p>
<div class="thumb tright"><div class="thumbinner" style="width:222px;"><a href="/enwiki/wiki/File:Hermann_pterodactylus_restoration1.png" class="image"><img alt="" src="/upwiki/wikipedia/commons/thumb/f/f3/Hermann_pterodactylus_restoration1.png/220px-Hermann_pterodactylus_restoration1.png" decoding="async" width="220" height="147" class="thumbimage" srcset="/upwiki/wikipedia/commons/thumb/f/f3/Hermann_pterodactylus_restoration1.png/330px-Hermann_pterodactylus_restoration1.png 1.5x, /upwiki/wikipedia/commons/thumb/f/f3/Hermann_pterodactylus_restoration1.png/440px-Hermann_pterodactylus_restoration1.png 2x" data-file-width="719" data-file-height="480" /></a> <div class="thumbcaption"><div class="magnify"><a href="/enwiki/wiki/File:Hermann_pterodactylus_restoration1.png" class="internal" title="Enlarge"></a></div>Hermann's original life restoration, the first of any pterosaur, 1800</div></div></div>
<p>The German/French scientist <a href="/enwiki/wiki/Johann_Hermann" title="Johann Hermann">Johann Hermann</a> was the one who first stated that <i>Pterodactylus</i> used its long fourth finger to support a wing membrane. Back in March 1800, Hermann alerted the prominent French scientist <a href="/enwiki/wiki/Georges_Cuvier" title="Georges Cuvier">Georges Cuvier</a> to the existence of Collini's fossil, believing that it had been captured by the occupying armies of <a href="/enwiki/wiki/Napoleon" title="Napoleon">Napoleon</a> and sent to the French collections in <a href="/enwiki/wiki/Paris" title="Paris">Paris</a> (and perhaps to Cuvier himself) as war booty; at the time special French <a href="/enwiki/wiki/Political_commissar" title="Political commissar">political commissars</a> systematically seized art treasures and objects of scientific interest. Hermann sent Cuvier a letter containing his own interpretation of the specimen (though he had not examined it personally), which he believed to be a <a href="/enwiki/wiki/Mammal" title="Mammal">mammal</a>, including the first known life restoration of a pterosaur. Hermann restored the animal with wing membranes extending from the long fourth finger to the ankle and a covering of fur (neither wing membranes nor fur had been preserved in the specimen). Hermann also added a membrane between the neck and wrist, as is the condition in <a href="/enwiki/wiki/Bat" title="Bat">bats</a>. Cuvier agreed with this interpretation, and at Hermann's suggestion, Cuvier became the first to publish these ideas in December 1800 in a very short description.<sup id="cite_ref-taquet&padian2004_10-1" class="reference"><a href="#cite_note-taquet&padian2004-10">[10]</a></sup> However, contrary to Hermann, Cuvier was convinced the animal was a <a href="/enwiki/wiki/Reptile" title="Reptile">reptile</a>.<sup id="cite_ref-cuvier1801_12-0" class="reference"><a href="#cite_note-cuvier1801-12">[12]</a></sup> The specimen had not in fact been seized by the French. Rather, in 1802, following the death of Charles Theodore, it was brought to <a href="/enwiki/wiki/Munich" title="Munich">Munich</a>, where Baron <a href="/enwiki/w/index.php?title=Johann_Paul_Carl_von_Moll&action=edit&redlink=1" class="new" title="Johann Paul Carl von Moll (page does not exist)">Johann Paul Carl von Moll</a> had obtained a general exemption of confiscation for the Bavarian collections.<sup id="cite_ref-pester_6-2" class="reference"><a href="#cite_note-pester-6">[6]</a></sup> Cuvier asked von Moll to study the fossil but was informed it could not be found. In 1809 Cuvier published a somewhat longer description, in which he named the animal <i>Petro-Dactyle</i>,<sup id="cite_ref-cuvier1809_13-0" class="reference"><a href="#cite_note-cuvier1809-13">[13]</a></sup> this was a typographical error however, and was later corrected by him to <i>Ptéro-Dactyle</i>.<sup id="cite_ref-taquet&padian2004_10-2" class="reference"><a href="#cite_note-taquet&padian2004-10">[10]</a></sup> He also refuted a hypothesis by <a href="/enwiki/wiki/Johann_Friedrich_Blumenbach" title="Johann Friedrich Blumenbach">Johann Friedrich Blumenbach</a> that it would have been a shore bird.<sup id="cite_ref-cuvier1809_13-1" class="reference"><a href="#cite_note-cuvier1809-13">[13]</a></sup> Cuvier remarked: "It is not possible to doubt that the long finger served to support a membrane that, by lengthening the anterior extremity of this animal, formed a good wing."<sup id="cite_ref-14" class="reference"><a href="#cite_note-14">[14]</a></sup>
</p>
<div class="thumb tright"><div class="thumbinner" style="width:222px;"><a href="/enwiki/wiki/File:Pterodactylus_antiquus_soemmerring.png" class="image"><img alt="" src="/upwiki/wikipedia/commons/thumb/6/6d/Pterodactylus_antiquus_soemmerring.png/220px-Pterodactylus_antiquus_soemmerring.png" decoding="async" width="220" height="106" class="thumbimage" srcset="/upwiki/wikipedia/commons/thumb/6/6d/Pterodactylus_antiquus_soemmerring.png/330px-Pterodactylus_antiquus_soemmerring.png 1.5x, /upwiki/wikipedia/commons/thumb/6/6d/Pterodactylus_antiquus_soemmerring.png/440px-Pterodactylus_antiquus_soemmerring.png 2x" data-file-width="744" data-file-height="358" /></a> <div class="thumbcaption"><div class="magnify"><a href="/enwiki/wiki/File:Pterodactylus_antiquus_soemmerring.png" class="internal" title="Enlarge"></a></div>Von Sömmerring's incorrect 1817 restoration of the species <i><a href="/enwiki/wiki/Pterodactylus_brevirostris" class="mw-redirect" title="Pterodactylus brevirostris">Pterodactylus brevirostris</a></i></div></div></div>
<p>Contrary to von Moll's report, the fossil was not missing; it was being studied by <a href="/enwiki/wiki/Samuel_Thomas_von_S%C3%B6mmerring" title="Samuel Thomas von Sömmerring">Samuel Thomas von Sömmerring</a>, who gave a public lecture about it on December 27, 1810. In January 1811, von Sömmerring wrote a letter to Cuvier deploring the fact that he had only recently been informed of Cuvier's request for information. His lecture was published in 1812, and in it von Sömmerring named the species <i><b>Ornithocephalus antiquus</b></i>.<sup id="cite_ref-sommerring1812_15-0" class="reference"><a href="#cite_note-sommerring1812-15">[15]</a></sup> The animal was described as being both a bat, and a form in between mammals and birds, i.e. not intermediate in descent but in "affinity" or <a href="/enwiki/wiki/Archetype" title="Archetype">archetype</a>. Cuvier disagreed, and the same year in his <i>Ossemens fossiles</i> provided a lengthy description in which he restated that the animal was a reptile.<sup id="cite_ref-16" class="reference"><a href="#cite_note-16">[16]</a></sup> It was not until 1817 that a second specimen of <i>Pterodactylus</i> came to light, again from <a href="/enwiki/wiki/Solnhofen" title="Solnhofen">Solnhofen</a>. This tiny specimen was that year described by von Sömmerring as <i><a href="/enwiki/wiki/Ornithocephalus_brevirostris" class="mw-redirect" title="Ornithocephalus brevirostris">Ornithocephalus brevirostris</a></i>, named for its short snout, now understood to be a juvenile character (this specimen is now thought to represent a juvenile specimen of a different genus, probably <i><a href="/enwiki/wiki/Ctenochasma" title="Ctenochasma">Ctenochasma</a></i>).<sup id="cite_ref-BennettPZ_17-0" class="reference"><a href="#cite_note-BennettPZ-17">[17]</a></sup> He provided a restoration of the skeleton, the first one published for any pterosaur.<sup id="cite_ref-taquet&padian2004_10-3" class="reference"><a href="#cite_note-taquet&padian2004-10">[10]</a></sup> This restoration was very inaccurate, von Sömmerring mistaking the long <a href="/enwiki/wiki/Metacarpal" class="mw-redirect" title="Metacarpal">metacarpals</a> for the bones of the lower arm, the lower arm for the <a href="/enwiki/wiki/Humerus" title="Humerus">humerus</a>, this upper arm for the <a href="/enwiki/wiki/Breast_bone" class="mw-redirect" title="Breast bone">breast bone</a> and this sternum again for the <a href="/enwiki/wiki/Shoulder_blade" class="mw-redirect" title="Shoulder blade">shoulder blades</a>.<sup id="cite_ref-18" class="reference"><a href="#cite_note-18">[18]</a></sup> Sömmerring did not change his opinion that these forms were bats and this "bat model" for interpreting pterosaurs would remain influential long after a consensus had been reached around 1860 that they were reptiles. The standard assumptions were that pterosaurs were quadrupedal, clumsy on the ground, furred, warmblooded and had a wing membrane reaching the ankle. Some of these elements have been confirmed, some refuted by modern research, while others remain disputed.<sup id="cite_ref-19" class="reference"><a href="#cite_note-19">[19]</a></sup>
</p><p>In 1815, the generic name <i>Ptéro-Dactyle</i> was latinized to <i>Pterodactylus</i> by <a href="/enwiki/wiki/Constantine_Samuel_Rafinesque" title="Constantine Samuel Rafinesque">Constantine Samuel Rafinesque</a>.<sup id="cite_ref-20" class="reference"><a href="#cite_note-20">[20]</a></sup> Unaware of Rafinesque's publication however, Cuvier himself in 1819 latinized the name <i>Ptéro-Dactyle</i> again to <i>Pterodactylus</i>,<sup id="cite_ref-:0_21-0" class="reference"><a href="#cite_note-:0-21">[21]</a></sup> but the specific name he then gave, <i>longirostris</i>, has to give precedence to von Sömmerring's <i>antiquus</i>.<sup id="cite_ref-:0_21-1" class="reference"><a href="#cite_note-:0-21">[21]</a></sup> In 1888, English <a href="/enwiki/wiki/Naturalist" class="mw-redirect" title="Naturalist">naturalist</a> <a href="/enwiki/wiki/Richard_Lydekker" title="Richard Lydekker">Richard Lydekker</a> designated <i>Pterodactylus antiquus</i> as the <a href="/enwiki/wiki/Type_species" title="Type species">type species</a> of <i>Pterodactylus</i>, and considered <i>Ornithocephalus antiquus</i> a synonym. He also designated specimen BSP AS.I.739 as the <a href="/enwiki/wiki/Holotype" title="Holotype">holotype</a> of the genus.<sup id="cite_ref-Lydekker_22-0" class="reference"><a href="#cite_note-Lydekker-22">[22]</a></sup>
</p>
<h2><span class="mw-headline" id="Description">Description</span><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Pterodactylus&action=edit&section=2" title="Edit section: Description">edit</a><span class="mw-editsection-bracket">]</span></span></h2>
<div class="thumb tleft"><div class="thumbinner" style="width:172px;"><a href="/enwiki/wiki/File:Pterodactylus_scale_mmartyniuk_wiki.png" class="image"><img alt="" src="/upwiki/wikipedia/commons/thumb/e/e3/Pterodactylus_scale_mmartyniuk_wiki.png/170px-Pterodactylus_scale_mmartyniuk_wiki.png" decoding="async" width="170" height="173" class="thumbimage" srcset="/upwiki/wikipedia/commons/thumb/e/e3/Pterodactylus_scale_mmartyniuk_wiki.png/255px-Pterodactylus_scale_mmartyniuk_wiki.png 1.5x, /upwiki/wikipedia/commons/thumb/e/e3/Pterodactylus_scale_mmartyniuk_wiki.png/340px-Pterodactylus_scale_mmartyniuk_wiki.png 2x" data-file-width="1117" data-file-height="1138" /></a> <div class="thumbcaption"><div class="magnify"><a href="/enwiki/wiki/File:Pterodactylus_scale_mmartyniuk_wiki.png" class="internal" title="Enlarge"></a></div>Size of the sub-adult holotype (blue) and adult (green) specimens in both flying and standing postures, compared with a human</div></div></div>
<p><i>Pterodactylus</i> is known from over 30 fossil specimens, and though most belong to juveniles, many preserve complete skeletons.<sup id="cite_ref-BennettPZ_17-1" class="reference"><a href="#cite_note-BennettPZ-17">[17]</a></sup><sup id="cite_ref-bennett1996a_23-0" class="reference"><a href="#cite_note-bennett1996a-23">[23]</a></sup> <i>Pterodactylus antiquus</i> was a relatively small pterosaur, with an estimated adult wingspan of about 1.04 meters (3 ft 5 in), based on the only known adult specimen, which is represented by an isolated skull.<sup id="cite_ref-BennettPZ_17-2" class="reference"><a href="#cite_note-BennettPZ-17">[17]</a></sup> Other "species" were once thought to have been smaller.<sup id="cite_ref-Lydekker_22-1" class="reference"><a href="#cite_note-Lydekker-22">[22]</a></sup> However, these smaller specimens have been shown to represent juveniles of <i>Pterodactylus</i>, as well as its contemporary relatives including <i>Ctenochasma</i>, <i><a href="/enwiki/wiki/Germanodactylus" title="Germanodactylus">Germanodactylus</a></i>, <i><a href="/enwiki/wiki/Aurorazhdarcho" title="Aurorazhdarcho">Aurorazhdarcho</a></i>, <i><a href="/enwiki/wiki/Gnathosaurus" title="Gnathosaurus">Gnathosaurus</a></i>, and hypothetically <i><a href="/enwiki/wiki/Aerodactylus" title="Aerodactylus">Aerodactylus</a></i> if this genus is truly valid.<sup id="cite_ref-bennett2002_24-0" class="reference"><a href="#cite_note-bennett2002-24">[24]</a></sup>
</p><p>The skulls of adult <i>Pterodactylus</i> were long and thin, with about 90 narrow and conical teeth. The teeth extended back from the tips of both jaws, and became smaller farther away from the jaw tips, this was unlike the ones seen in most relatives, where teeth were absent in the upper jaw tip, and were relatively uniform in size. The teeth of <i>Pterodactylus</i> also extended farther back into the jaw compared to close relatives, and some were present below the front of the <i>nasoantorbital fenestra</i>, which is the largest opening in the skull.<sup id="cite_ref-BennettPZ_17-3" class="reference"><a href="#cite_note-BennettPZ-17">[17]</a></sup> Another <a href="/enwiki/wiki/Autapomorphy" title="Autapomorphy">autapomorphy</a> that <i>Pterodactylus</i> has is that the skull and jaws were straight, which are unlike the upwardly curved jaws seen in the related <a href="/enwiki/wiki/Ctenochasmatid" class="mw-redirect" title="Ctenochasmatid">ctenochasmatids</a>.<sup id="cite_ref-jouve2004_25-0" class="reference"><a href="#cite_note-jouve2004-25">[25]</a></sup>
</p>
<div class="thumb tright"><div class="thumbinner" style="width:222px;"><a href="/enwiki/wiki/File:Pterodactylus_BMMS7_life.png" class="image"><img alt="" src="/upwiki/wikipedia/commons/thumb/a/a0/Pterodactylus_BMMS7_life.png/220px-Pterodactylus_BMMS7_life.png" decoding="async" width="220" height="104" class="thumbimage" srcset="/upwiki/wikipedia/commons/thumb/a/a0/Pterodactylus_BMMS7_life.png/330px-Pterodactylus_BMMS7_life.png 1.5x, /upwiki/wikipedia/commons/thumb/a/a0/Pterodactylus_BMMS7_life.png/440px-Pterodactylus_BMMS7_life.png 2x" data-file-width="931" data-file-height="438" /></a> <div class="thumbcaption"><div class="magnify"><a href="/enwiki/wiki/File:Pterodactylus_BMMS7_life.png" class="internal" title="Enlarge"></a></div>Life <a href="/enwiki/wiki/Paleoart" title="Paleoart">restoration</a> of BMMS 7, the largest known <i>Pterodactylus</i> specimen. The shape of the soft tissue crest is based on the specimen BSP 1929 I 18</div></div></div>
<p><i>Pterodactylus</i>, like related pterosaurs, had a crest on its skull composed mainly of soft tissues. In adult <i>Pterodactylus</i>, this crest extended between the back edge of the <a href="/enwiki/wiki/Antorbital_fenestra" title="Antorbital fenestra">antorbital fenestra</a> and the back of the skull. In at least one specimen, the crest had a short bony base, also seen in related pterosaurs like <i>Germanodactylus</i>. Solid crests have only been found on large, fully adult specimens of <i>Pterodactylus</i>, indicating that this was a display structure that became larger and more well developed as individuals reached maturity.<sup id="cite_ref-BennettPZ_17-4" class="reference"><a href="#cite_note-BennettPZ-17">[17]</a></sup><sup id="cite_ref-frey&martill1998_26-0" class="reference"><a href="#cite_note-frey&martill1998-26">[26]</a></sup> In 2013, pterosaur researcher <a href="/enwiki/w/index.php?title=S._Christopher_Bennett&action=edit&redlink=1" class="new" title="S. Christopher Bennett (page does not exist)">S. Christopher Bennett</a> noted that other authors claimed that the soft tissue crest of <i>Pterodactylus</i> extended backward behind the skull; Bennett himself, however, didn't find any evidence for the crest extending past the back of the skull.<sup id="cite_ref-BennettPZ_17-5" class="reference"><a href="#cite_note-BennettPZ-17">[17]</a></sup> Two specimens of <i>P. antiquus</i> (the holotype specimen BSP AS I 739 and the incomplete skull BMMS 7, the largest known skull of <i>P. antiquus</i>) have a low bony crest on their skulls; in BMMS 7 it is 47.5 mm long (1.87 inches, more or less 24% of the estimated total length of its skull) and has a maximum height of 0.9 mm (0.035 inches) above the orbit.<sup id="cite_ref-BennettPZ_17-6" class="reference"><a href="#cite_note-BennettPZ-17">[17]</a></sup> Several specimens previously referred to <i>P. antiquus</i> preserved evidence of the soft tissue extensions of these crests, including an "occipital lappet", a flexible, tab-like structure extending from the back of the skull. Most of these specimens have been reclassified in the related species <i>Aerodactylus scolopaciceps</i>, which may however be nothing more than a junior synonym. Even if <i>Aerodactylus</i> were valid, at least one specimen with these features is still considered to belong to <i>Pterodactylus</i>, BSP 1929 I 18, which has an occipital lappet similar to the proposed <i>Aerodactylus</i> definition, and also possesses a small triangular soft tissue crest with the peak of the crest positioned above the eyes.<sup id="cite_ref-BennettPZ_17-7" class="reference"><a href="#cite_note-BennettPZ-17">[17]</a></sup>
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<h2><span class="mw-headline" id="Paleobiology">Paleobiology</span><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Pterodactylus&action=edit&section=3" title="Edit section: Paleobiology">edit</a><span class="mw-editsection-bracket">]</span></span></h2>
<h3><span class="mw-headline" id="Life_history">Life history</span><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Pterodactylus&action=edit&section=4" title="Edit section: Life history">edit</a><span class="mw-editsection-bracket">]</span></span></h3>
<div class="thumb tleft"><div class="thumbinner" style="width:222px;"><a href="/enwiki/wiki/File:Pterodactylus_spectabilis_4.JPG" class="image"><img alt="" src="/upwiki/wikipedia/commons/thumb/c/cc/Pterodactylus_spectabilis_4.JPG/220px-Pterodactylus_spectabilis_4.JPG" decoding="async" width="220" height="136" class="thumbimage" srcset="/upwiki/wikipedia/commons/thumb/c/cc/Pterodactylus_spectabilis_4.JPG/330px-Pterodactylus_spectabilis_4.JPG 1.5x, /upwiki/wikipedia/commons/thumb/c/cc/Pterodactylus_spectabilis_4.JPG/440px-Pterodactylus_spectabilis_4.JPG 2x" data-file-width="2764" data-file-height="1703" /></a> <div class="thumbcaption"><div class="magnify"><a href="/enwiki/wiki/File:Pterodactylus_spectabilis_4.JPG" class="internal" title="Enlarge"></a></div>Holotype specimen of the species <i>P. spectabilis</i>, now considered a juvenile specimen of <i>P. antiquus</i></div></div></div>
<p>Like other pterosaurs (most notably <i><a href="/enwiki/wiki/Rhamphorhynchus_(pterosaur)" class="mw-redirect" title="Rhamphorhynchus (pterosaur)">Rhamphorhynchus</a></i>), <i>Pterodactylus</i> specimens can vary considerably based on age or level of maturity. Both the proportions of the limb bones, size and shape of the skull, and size and number of teeth changed as the animals grew. Historically, this has led to various growth stages (including growth stages of related pterosaurs) being mistaken for new species of <i>Pterodactylus</i>. Several detailed studies using various methods to measure growth curves among known specimens have suggested that there is actually only one valid species of <i>Pterodactylus</i>, <i>P. antiquus</i>.<sup id="cite_ref-jouve2004_25-1" class="reference"><a href="#cite_note-jouve2004-25">[25]</a></sup>
</p><p>The youngest immature specimens of <i>Pterodactylus antiquus</i> (alternately interpreted as young specimens of the distinct species <i>P. kochi</i>) have a small number of teeth, as few as 15 in some, and the teeth have a relatively broad base.<sup id="cite_ref-bennett1996a_23-1" class="reference"><a href="#cite_note-bennett1996a-23">[23]</a></sup> The teeth of other <i>P. antiquus</i> specimens are both narrower and more numerous (up to 90 teeth are present in several specimens).<sup id="cite_ref-jouve2004_25-2" class="reference"><a href="#cite_note-jouve2004-25">[25]</a></sup>
</p><p><i>Pterodactylus</i> specimens can be divided into two distinct year classes. In the first year class, the skulls are only 15 to 45 millimeters (0.59 to 1.77 in) in length. The second year class is characterized by skulls of around 55 to 95 millimeters (2.2 to 3.7 in) long, but are still immature however. These first two size groups were once classified as juveniles and adults of the species <i>P. kochi</i>, until further study showed that even the supposed "adults" were immature, and possibly belong to a distinct genus. A third year class is represented by specimens of the "traditional" <i>P. antiquus</i>, as well as a few isolated, large specimens once assigned to <i>P. kochi</i> that overlap <i>P. antiquus</i> in size. However, all specimens in this third year class also show sign of immaturity. Fully mature <i>Pterodactylus</i> specimens remain unknown, or may have been mistakenly classified as a different genus.<sup id="cite_ref-bennett1996a_23-2" class="reference"><a href="#cite_note-bennett1996a-23">[23]</a></sup>
</p>
<h3><span class="mw-headline" id="Growth_and_breeding_seasons">Growth and breeding seasons</span><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Pterodactylus&action=edit&section=5" title="Edit section: Growth and breeding seasons">edit</a><span class="mw-editsection-bracket">]</span></span></h3>
<div class="thumb tright"><div class="thumbinner" style="width:222px;"><a href="/enwiki/wiki/File:Pterodactylus_grandipelvis_Teylers_2.JPG" class="image"><img alt="" src="/upwiki/wikipedia/commons/thumb/e/ec/Pterodactylus_grandipelvis_Teylers_2.JPG/220px-Pterodactylus_grandipelvis_Teylers_2.JPG" decoding="async" width="220" height="105" class="thumbimage" srcset="/upwiki/wikipedia/commons/thumb/e/ec/Pterodactylus_grandipelvis_Teylers_2.JPG/330px-Pterodactylus_grandipelvis_Teylers_2.JPG 1.5x, /upwiki/wikipedia/commons/thumb/e/ec/Pterodactylus_grandipelvis_Teylers_2.JPG/440px-Pterodactylus_grandipelvis_Teylers_2.JPG 2x" data-file-width="3489" data-file-height="1669" /></a> <div class="thumbcaption"><div class="magnify"><a href="/enwiki/wiki/File:Pterodactylus_grandipelvis_Teylers_2.JPG" class="internal" title="Enlarge"></a></div>Fossil pelvis of a large specimen, now referred to the dubious species <i>P. grandipelvis</i></div></div></div>
<p>The distinct year classes of <i>Pterodactylus antiquus</i> specimens show that this species, like the contemporary <i>Rhamphorhynchus muensteri</i>, likely bred seasonally and grew consistently during its lifetime. A new generation of 1st year class <i>P. antiquus</i> would have been produced seasonally, and reached 2nd-year size by the time the next generation hatched, creating distinct 'clumps' of similarly-sized and aged individuals in the fossil record. The smallest size class probably consisted of individuals that had just begun to fly and were less than one year old.<sup id="cite_ref-bennett1996a_23-3" class="reference"><a href="#cite_note-bennett1996a-23">[23]</a></sup><sup id="cite_ref-Wellnhofer1970_27-0" class="reference"><a href="#cite_note-Wellnhofer1970-27">[27]</a></sup> The second year class represents individuals one to two years old, and the rare third year class is composed of specimens over two years old. This growth pattern is similar to modern <a href="/enwiki/wiki/Crocodilia" title="Crocodilia">crocodilians</a>, rather than the rapid growth of modern <a href="/enwiki/wiki/Bird" title="Bird">birds</a>.<sup id="cite_ref-bennett1996a_23-4" class="reference"><a href="#cite_note-bennett1996a-23">[23]</a></sup>
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<h3><span class="mw-headline" id="Daily_activity_patterns">Daily activity patterns</span><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Pterodactylus&action=edit&section=6" title="Edit section: Daily activity patterns">edit</a><span class="mw-editsection-bracket">]</span></span></h3>
<p>Comparisons between the <a href="/enwiki/wiki/Sclerotic_ring" title="Sclerotic ring">scleral rings</a> of <i>Pterodactylus antiquus</i> and modern birds and reptiles suggest that it may have been <a href="/enwiki/wiki/Diurnality" title="Diurnality">diurnal</a>. This may also indicate <a href="/enwiki/wiki/Niche_partitioning" class="mw-redirect" title="Niche partitioning">niche partitioning</a> with contemporary pterosaurs inferred to be <a href="/enwiki/wiki/Nocturnal" class="mw-redirect" title="Nocturnal">nocturnal</a>, such as <i>Ctenochasma</i> and <i>Rhamphorhynchus</i>.<sup id="cite_ref-28" class="reference"><a href="#cite_note-28">[28]</a></sup>
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<h3><span class="mw-headline" id="Diet">Diet</span><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Pterodactylus&action=edit&section=7" title="Edit section: Diet">edit</a><span class="mw-editsection-bracket">]</span></span></h3>
<p>Based on the shape, size, and arrangement of its teeth, <i>Pterodactylus</i> has long been recognized as a carnivore specializing in small animals. A 2020 study of pterosaur tooth wear supported the hypothesis that <i>Pterodactylus</i> preyed mainly on invertebrates and had a generalist feeding strategy, indicated by a relatively high bite force.<sup id="cite_ref-diet2020_29-0" class="reference"><a href="#cite_note-diet2020-29">[29]</a></sup>
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<h2><span class="mw-headline" id="Paleoecology">Paleoecology</span><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Pterodactylus&action=edit&section=8" title="Edit section: Paleoecology">edit</a><span class="mw-editsection-bracket">]</span></span></h2>
<div class="thumb tleft"><div class="thumbinner" style="width:222px;"><a href="/enwiki/wiki/File:Pterodactylus_holotype_fly_mmartyniuk.png" class="image"><img alt="" src="/upwiki/wikipedia/commons/thumb/a/ae/Pterodactylus_holotype_fly_mmartyniuk.png/220px-Pterodactylus_holotype_fly_mmartyniuk.png" decoding="async" width="220" height="123" class="thumbimage" srcset="/upwiki/wikipedia/commons/thumb/a/ae/Pterodactylus_holotype_fly_mmartyniuk.png/330px-Pterodactylus_holotype_fly_mmartyniuk.png 1.5x, /upwiki/wikipedia/commons/thumb/a/ae/Pterodactylus_holotype_fly_mmartyniuk.png/440px-Pterodactylus_holotype_fly_mmartyniuk.png 2x" data-file-width="500" data-file-height="280" /></a> <div class="thumbcaption"><div class="magnify"><a href="/enwiki/wiki/File:Pterodactylus_holotype_fly_mmartyniuk.png" class="internal" title="Enlarge"></a></div>Hypothetical restoration of <i>Pterodactylus</i> in its environment</div></div></div>
<p>Specimens of <i>Pterodactylus</i> have been found mainly in the <a href="/enwiki/wiki/Solnhofen_limestone" class="mw-redirect" title="Solnhofen limestone">Solnhofen limestone</a> (geologically known as the Altmühltal Formation) of <a href="/enwiki/wiki/Bavaria" title="Bavaria">Bavaria</a>, <a href="/enwiki/wiki/Germany" title="Germany">Germany</a>. The main composition of this formation is fine-grained <a href="/enwiki/wiki/Limestone" title="Limestone">limestone</a> that originated mainly from the nearby towns <a href="/enwiki/wiki/Solnhofen" title="Solnhofen">Solnhofen</a> and Eichstätt, which is formed by mud <a href="/enwiki/wiki/Silt" title="Silt">silt</a> deposits.<sup id="cite_ref-ammonitedate_3-1" class="reference"><a href="#cite_note-ammonitedate-3">[3]</a></sup> The Solnhofen Limestone is a diverse <a href="/enwiki/wiki/Lagerst%C3%A4tte" title="Lagerstätte">Lagerstätte</a> that contains a wide range of different creatures, including highly detailed fossilized imprints of soft bodied organisms such as <a href="/enwiki/wiki/Jellyfish" title="Jellyfish">jellyfishes</a>. Abundant specimens of pterosaurs similar to <i>Pterodactylus</i> were also found within the formation, these include the <a href="/enwiki/wiki/Rhamphorhynchid" class="mw-redirect" title="Rhamphorhynchid">rhamphorhynchids</a> <i>Rhamphorhynchus</i> and <i><a href="/enwiki/wiki/Scaphognathus" title="Scaphognathus">Scaphognathus</a></i>,<sup id="cite_ref-30" class="reference"><a href="#cite_note-30">[30]</a></sup> several <a href="/enwiki/wiki/Gallodactylid" class="mw-redirect" title="Gallodactylid">gallodactylids</a> such as <i>Aerodactylus</i>,<sup id="cite_ref-bennett1996a_23-5" class="reference"><a href="#cite_note-bennett1996a-23">[23]</a></sup> <i><a href="/enwiki/wiki/Ardeadactylus" title="Ardeadactylus">Ardeadactylus</a></i>, <i>Aurorazhdarcho</i> and <i><a href="/enwiki/wiki/Cycnorhamphus" title="Cycnorhamphus">Cycnorhamphus</a></i>,<sup id="cite_ref-DU06_7-1" class="reference"><a href="#cite_note-DU06-7">[7]</a></sup> the <a href="/enwiki/wiki/Ctenochasmatid" class="mw-redirect" title="Ctenochasmatid">ctenochasmatids</a> <i>Ctenochasma</i><sup id="cite_ref-bennett2007_31-0" class="reference"><a href="#cite_note-bennett2007-31">[31]</a></sup> and <i>Gnathosaurus</i>, the <a href="/enwiki/wiki/Anurognathid" class="mw-redirect" title="Anurognathid">anurognathid</a> <i><a href="/enwiki/wiki/Anurognathus" title="Anurognathus">Anurognathus</a></i>, the <a href="/enwiki/wiki/Germanodactylid" class="mw-redirect" title="Germanodactylid">germanodactylid</a> <i>Germanodactylus</i>, as well as the basal <a href="/enwiki/wiki/Euctenochasmatia" title="Euctenochasmatia">euctenochasmatian</a> <i><a href="/enwiki/wiki/Diopecephalus" title="Diopecephalus">Diopecephalus</a></i>.<sup id="cite_ref-W1837_32-0" class="reference"><a href="#cite_note-W1837-32">[32]</a></sup> Fossil remains of the <a href="/enwiki/wiki/Dinosaur" title="Dinosaur">dinosaurs</a> <i><a href="/enwiki/wiki/Archaeopteryx" title="Archaeopteryx">Archaeopteryx</a></i> and <i><a href="/enwiki/wiki/Compsognathus" title="Compsognathus">Compsognathus</a></i> were also found within the limestone, these specimens were related to early evolution of <a href="/enwiki/wiki/Feathers" class="mw-redirect" title="Feathers">feathers</a>, since they were some of the only ones that had them during the Jurassic period.<sup id="cite_ref-FasWeis04_33-0" class="reference"><a href="#cite_note-FasWeis04-33">[33]</a></sup> Various <a href="/enwiki/wiki/Lizard" title="Lizard">lizard</a> remains were also found alongside those of <i>Pterodactylus</i>, with several specimens assigned to <i><a href="/enwiki/wiki/Ardeosaurus" title="Ardeosaurus">Ardeosaurus</a></i>, <i><a href="/enwiki/wiki/Bavarisaurus" title="Bavarisaurus">Bavarisaurus</a></i> and <i><a href="/enwiki/wiki/Eichstaettisaurus" title="Eichstaettisaurus">Eichstaettisaurus</a></i>.<sup id="cite_ref-hoffstetter1966_34-0" class="reference"><a href="#cite_note-hoffstetter1966-34">[34]</a></sup><sup id="cite_ref-35" class="reference"><a href="#cite_note-35">[35]</a></sup> <a href="/enwiki/wiki/Crocodylomorph" class="mw-redirect" title="Crocodylomorph">Crocodylomorph</a> specimens were widely distributed within the fossil site, most were assigned to the <a href="/enwiki/wiki/Metriorhynchid" class="mw-redirect" title="Metriorhynchid">metriorhynchid</a> genera <i><a href="/enwiki/wiki/Cricosaurus" title="Cricosaurus">Cricosaurus</a></i>, <i><a href="/enwiki/wiki/Dakosaurus" title="Dakosaurus">Dakosaurus</a></i>,<sup id="cite_ref-36" class="reference"><a href="#cite_note-36">[36]</a></sup> <i><a href="/enwiki/wiki/Geosaurus" title="Geosaurus">Geosaurus</a></i> and <i><a href="/enwiki/wiki/Rhacheosaurus" title="Rhacheosaurus">Rhacheosaurus</a></i>.<sup id="cite_ref-Andrade_&_Young,_2008_37-0" class="reference"><a href="#cite_note-Andrade_&_Young,_2008-37">[37]</a></sup> These genera are colloquially called as marine or sea crocodiles due to their similar built.<sup id="cite_ref-38" class="reference"><a href="#cite_note-38">[38]</a></sup> The turtle genera <i><a href="/enwiki/wiki/Eurysternum" title="Eurysternum">Eurysternum</a></i> and <i><a href="/enwiki/wiki/Paleomedusa" class="mw-redirect" title="Paleomedusa">Paleomedusa</a></i> were also found within the formation.<sup id="cite_ref-WGJ_39-0" class="reference"><a href="#cite_note-WGJ-39">[39]</a></sup> Fossils of the <a href="/enwiki/wiki/Ichthyosaur" title="Ichthyosaur">ichthyosaur</a> <i><a href="/enwiki/wiki/Aegirosaurus" title="Aegirosaurus">Aegirosaurus</a></i> also appeared to be present in the site,<sup id="cite_ref-40" class="reference"><a href="#cite_note-40">[40]</a></sup> as well as fish remains, with many specimens assigned to <a href="/enwiki/wiki/Ray-finned_fish" class="mw-redirect" title="Ray-finned fish">ray-finned fishes</a> such as the <a href="/enwiki/wiki/Halecomorph" class="mw-redirect" title="Halecomorph">halecomorphs</a> <i><a href="/enwiki/wiki/Lepidotes" title="Lepidotes">Lepidotes</a></i>,<sup id="cite_ref-lamberstable-others_41-0" class="reference"><a href="#cite_note-lamberstable-others-41">[41]</a></sup> <i><a href="/enwiki/wiki/Propterus" title="Propterus">Propterus</a></i>,<sup id="cite_ref-Marine_42-0" class="reference"><a href="#cite_note-Marine-42">[42]</a></sup> <i><a href="/enwiki/wiki/Gyrodus" title="Gyrodus">Gyrodus</a></i>, <i><a href="/enwiki/wiki/Mesturus" title="Mesturus">Mesturus</a></i>, <i><a href="/enwiki/wiki/Proscinetes" title="Proscinetes">Proscinetes</a></i>, <i><a href="/enwiki/wiki/Caturus" title="Caturus">Caturus</a></i>,<sup id="cite_ref-Marine_42-1" class="reference"><a href="#cite_note-Marine-42">[42]</a></sup> <i><a href="/enwiki/wiki/Ophiopsis" title="Ophiopsis">Ophiopsis</a></i><sup id="cite_ref-43" class="reference"><a href="#cite_note-43">[43]</a></sup> and <i><a href="/enwiki/wiki/Ophiopsiella" title="Ophiopsiella">Ophiopsiella</a></i>,<sup id="cite_ref-lamberstable-others_41-1" class="reference"><a href="#cite_note-lamberstable-others-41">[41]</a></sup> the <a href="/enwiki/wiki/Pachycormidae" class="mw-redirect" title="Pachycormidae">pachycormids</a> <i><a href="/enwiki/wiki/Asthenocormus" title="Asthenocormus">Asthenocormus</a></i>, <i><a href="/enwiki/wiki/Hypsocormus" title="Hypsocormus">Hypsocormus</a></i> and <i><a href="/enwiki/wiki/Orthocormus" title="Orthocormus">Orthocormus</a></i>,<sup id="cite_ref-EoDP_44-0" class="reference"><a href="#cite_note-EoDP-44">[44]</a></sup> as well as the <a href="/enwiki/wiki/Aspidorhynchidae" class="mw-redirect" title="Aspidorhynchidae">aspidorhynchid</a> <i><a href="/enwiki/wiki/Aspidorhynchus" title="Aspidorhynchus">Aspidorhynchus</a></i>, and the <a href="/enwiki/wiki/Ichthyodectid" class="mw-redirect" title="Ichthyodectid">ichthyodectid</a> <i><a href="/enwiki/wiki/Thrissops" title="Thrissops">Thrissops</a></i>.<sup id="cite_ref-FT12_45-0" class="reference"><a href="#cite_note-FT12-45">[45]</a></sup><sup id="cite_ref-46" class="reference"><a href="#cite_note-46">[46]</a></sup>
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<h2><span class="mw-headline" id="Classification">Classification</span><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Pterodactylus&action=edit&section=9" title="Edit section: Classification">edit</a><span class="mw-editsection-bracket">]</span></span></h2>
<div class="thumb tright"><div class="thumbinner" style="width:172px;"><a href="/enwiki/wiki/File:Pterodactylus_muscle_impressions.jpg" class="image"><img alt="" src="/upwiki/wikipedia/commons/thumb/1/1f/Pterodactylus_muscle_impressions.jpg/170px-Pterodactylus_muscle_impressions.jpg" decoding="async" width="170" height="218" class="thumbimage" srcset="/upwiki/wikipedia/commons/thumb/1/1f/Pterodactylus_muscle_impressions.jpg/255px-Pterodactylus_muscle_impressions.jpg 1.5x, /upwiki/wikipedia/commons/thumb/1/1f/Pterodactylus_muscle_impressions.jpg/340px-Pterodactylus_muscle_impressions.jpg 2x" data-file-width="910" data-file-height="1169" /></a> <div class="thumbcaption"><div class="magnify"><a href="/enwiki/wiki/File:Pterodactylus_muscle_impressions.jpg" class="internal" title="Enlarge"></a></div>A <i>P. antiquus</i> specimen (AMNH 1942) showing muscle impressions in the chest and wing membranes</div></div></div>
<p>Initial classifications for <i>Pterodactylus</i> started when paleontologist Hermann von Meyer used the name Pterodactyli to contain <i>Pterodactylus</i> and other pterosaurs known at the time. This was emended to the <a href="/enwiki/wiki/Family_(biology)" title="Family (biology)">family</a> <a href="/enwiki/wiki/Pterodactylidae" title="Pterodactylidae">Pterodactylidae</a> by Prince <a href="/enwiki/wiki/Charles_Lucien_Bonaparte" title="Charles Lucien Bonaparte">Charles Lucien Bonaparte</a> in 1838. However, this group has more recently been given several competing definitions.<sup id="cite_ref-unwin2003_4-1" class="reference"><a href="#cite_note-unwin2003-4">[4]</a></sup><sup id="cite_ref-47" class="reference"><a href="#cite_note-47">[47]</a></sup>
</p><p>Beginning in 2014, researchers Steven Vidovic and David Martill constructed an analysis in which several pterosaurs traditionally thought of as <a href="/enwiki/wiki/Archaeopterodactyloid" class="mw-redirect" title="Archaeopterodactyloid">archaeopterodactyloids</a> closely related to the <a href="/enwiki/wiki/Ctenochasmatoid" class="mw-redirect" title="Ctenochasmatoid">ctenochasmatoids</a> may have been more closely related to the more advanced <a href="/enwiki/wiki/Dsungaripteroid" class="mw-redirect" title="Dsungaripteroid">dsungaripteroids</a>, or in some cases, fall outside both groups. Their conclusion was published in 2017, in which they placed <i>Pterodactylus</i> as a basal member of the suborder <a href="/enwiki/wiki/Pterodactyloidea" title="Pterodactyloidea">Pterodactyloidea</a>.<sup id="cite_ref-W1837_32-1" class="reference"><a href="#cite_note-W1837-32">[32]</a></sup>
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<p><i><b>Pterodactylus antiquus</b></i><a href="/enwiki/wiki/File:Pterodactylus_BMMS7_life.png" class="image"><img alt="Pterodactylus BMMS7 life.png" src="/upwiki/wikipedia/commons/thumb/a/a0/Pterodactylus_BMMS7_life.png/70px-Pterodactylus_BMMS7_life.png" decoding="async" width="70" height="33" srcset="/upwiki/wikipedia/commons/thumb/a/a0/Pterodactylus_BMMS7_life.png/105px-Pterodactylus_BMMS7_life.png 1.5x, /upwiki/wikipedia/commons/thumb/a/a0/Pterodactylus_BMMS7_life.png/140px-Pterodactylus_BMMS7_life.png 2x" data-file-width="931" data-file-height="438" /></a>
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<p>As illustrated below, the results of a different <a href="/enwiki/wiki/Topology" title="Topology">topology</a> are based on a phylogenetic analysis made by Longrich, Martill, and Andres in 2018. Unlike the previous results above, they placed <i>Pterodactylus</i> within the clade <a href="/enwiki/wiki/Euctenochasmatia" title="Euctenochasmatia">Euctenochasmatia</a>, resulting in a more derived position.<sup id="cite_ref-longrichetal2018_5-1" class="reference"><a href="#cite_note-longrichetal2018-5">[5]</a></sup>
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<p><i><a href="/enwiki/wiki/Germanodactylus_cristatus" class="mw-redirect" title="Germanodactylus cristatus">Germanodactylus cristatus</a></i>
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<p><i><a href="/enwiki/wiki/Germanodactylus_rhamphastinus" class="mw-redirect" title="Germanodactylus rhamphastinus">Germanodactylus rhamphastinus</a></i><a href="/enwiki/wiki/File:Altmuehlopterus_DB.jpg" class="image"><img alt="Altmuehlopterus DB.jpg" src="/upwiki/wikipedia/commons/thumb/0/09/Altmuehlopterus_DB.jpg/70px-Altmuehlopterus_DB.jpg" decoding="async" width="70" height="58" srcset="/upwiki/wikipedia/commons/thumb/0/09/Altmuehlopterus_DB.jpg/105px-Altmuehlopterus_DB.jpg 1.5x, /upwiki/wikipedia/commons/thumb/0/09/Altmuehlopterus_DB.jpg/140px-Altmuehlopterus_DB.jpg 2x" data-file-width="1440" data-file-height="1198" /></a>
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<p><i><b>Pterodactylus antiquus</b></i><a href="/enwiki/wiki/File:Pterodactylus_BMMS7_life.png" class="image"><img alt="Pterodactylus BMMS7 life.png" src="/upwiki/wikipedia/commons/thumb/a/a0/Pterodactylus_BMMS7_life.png/70px-Pterodactylus_BMMS7_life.png" decoding="async" width="70" height="33" srcset="/upwiki/wikipedia/commons/thumb/a/a0/Pterodactylus_BMMS7_life.png/105px-Pterodactylus_BMMS7_life.png 1.5x, /upwiki/wikipedia/commons/thumb/a/a0/Pterodactylus_BMMS7_life.png/140px-Pterodactylus_BMMS7_life.png 2x" data-file-width="931" data-file-height="438" /></a>
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<p><i><a href="/enwiki/wiki/Cycnorhamphus_suevicus" class="mw-redirect" title="Cycnorhamphus suevicus">Cycnorhamphus suevicus</a></i>
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<p><i><a href="/enwiki/wiki/Normannognathus_wellnhoferi" class="mw-redirect" title="Normannognathus wellnhoferi">Normannognathus wellnhoferi</a></i>
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<p><a href="/enwiki/wiki/Ctenochasmatidae" title="Ctenochasmatidae">Ctenochasmatidae</a><a href="/enwiki/wiki/File:Pterodaustro_BW.jpg" class="image"><img alt="Pterodaustro BW.jpg" src="/upwiki/wikipedia/commons/thumb/2/29/Pterodaustro_BW.jpg/70px-Pterodaustro_BW.jpg" decoding="async" width="70" height="50" srcset="/upwiki/wikipedia/commons/thumb/2/29/Pterodaustro_BW.jpg/105px-Pterodaustro_BW.jpg 1.5x, /upwiki/wikipedia/commons/thumb/2/29/Pterodaustro_BW.jpg/140px-Pterodaustro_BW.jpg 2x" data-file-width="633" data-file-height="448" /></a>
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<h3><span class="mw-headline" id="Formerly_assigned_species">Formerly assigned species</span><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Pterodactylus&action=edit&section=10" title="Edit section: Formerly assigned species">edit</a><span class="mw-editsection-bracket">]</span></span></h3>
<div class="thumb tleft"><div class="thumbinner" style="width:222px;"><a href="/enwiki/wiki/File:Rhamphorhynchus_Lauer.jpg" class="image"><img alt="" src="/upwiki/wikipedia/commons/thumb/4/48/Rhamphorhynchus_Lauer.jpg/220px-Rhamphorhynchus_Lauer.jpg" decoding="async" width="220" height="165" class="thumbimage" srcset="/upwiki/wikipedia/commons/thumb/4/48/Rhamphorhynchus_Lauer.jpg/330px-Rhamphorhynchus_Lauer.jpg 1.5x, /upwiki/wikipedia/commons/thumb/4/48/Rhamphorhynchus_Lauer.jpg/440px-Rhamphorhynchus_Lauer.jpg 2x" data-file-width="4032" data-file-height="3021" /></a> <div class="thumbcaption"><div class="magnify"><a href="/enwiki/wiki/File:Rhamphorhynchus_Lauer.jpg" class="internal" title="Enlarge"></a></div>Fossil specimen of the species <i><a href="/enwiki/wiki/Rhamphorhynchus_muensteri" class="mw-redirect" title="Rhamphorhynchus muensteri">Rhamphorhynchus muensteri</a></i>, which was previously assigned as the species <i>Pterodactylus münsteri</i></div></div></div>
<p>Numerous species have been assigned to <i>Pterodactylus</i> in the years since its discovery. In the first half of the 19th century any new pterosaur species would be named <i>Pterodactylus</i>, which thus became a "<a href="/enwiki/wiki/Wastebasket_taxon" title="Wastebasket taxon">wastebasket taxon</a>".<sup id="cite_ref-W1837_32-2" class="reference"><a href="#cite_note-W1837-32">[32]</a></sup> 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.<sup id="cite_ref-Aerodactylus_48-0" class="reference"><a href="#cite_note-Aerodactylus-48">[48]</a></sup>
</p><p>The earliest reassignments of pterosaur species to <i>Pterodactylus</i> started in 1825, with the description of <i>Rhamphorhynchus</i>; fossil collector <a href="/enwiki/wiki/Georg_Graf_zu_M%C3%BCnster" class="mw-redirect" title="Georg Graf zu Münster">Georg Graf zu Münster</a> alerted the German paleontologist <a href="/enwiki/wiki/Samuel_Thomas_von_S%C3%B6mmerring" title="Samuel Thomas von Sömmerring">Samuel Thomas von Sömmerring</a> about several distinct fossil specimens, Sömmerring thought that they belonged to an ancient bird.<sup id="cite_ref-Pterosauria_49-0" class="reference"><a href="#cite_note-Pterosauria-49">[49]</a></sup> Further fossil preparations had uncovered teeth, to which Graf zu Münster created a skull cast. He later sent the cast to Professor <a href="/enwiki/wiki/Georg_August_Goldfuss" title="Georg August Goldfuss">Georg August Goldfuss</a>, who recognized it as a pterosaur, specifically a species of <i>Pterodactylus</i>. At the time however, most paleontologists incorrectly consider the genus <i>Ornithocephalus</i> (<abbr title="literal translation">lit.</abbr><span style="white-space:nowrap"> </span>'bird-head') to be the valid name for <i>Pterodactylus</i>, and therefore the specimen found was named as <i>Ornithocephalus Münsteri</i>, which was first mentioned by Graf zu Münster himself.<sup id="cite_ref-Münster1830_50-0" class="reference"><a href="#cite_note-Münster1830-50">[50]</a></sup> Another specimen was found and described by Graf zu Münster in 1839, he assigned this specimen to a new separate species called <i>Ornithocephalus longicaudus</i>; the <a href="/enwiki/wiki/Specific_name_(zoology)" title="Specific name (zoology)">specific name</a> means 'long tail', in reference to the animal's tail size.<sup id="cite_ref-51" class="reference"><a href="#cite_note-51">[51]</a></sup> German paleontologist <a href="/enwiki/wiki/Hermann_von_Meyer" class="mw-redirect" title="Hermann von Meyer">Hermann von Meyer</a> in 1845 officially emended that the genus <i>Pterodactylus</i> had priority over <i>Ornithocephalus</i>, so he reassigned the species <i>O. münsteri</i> and <i>O. longicaudus</i> into <i>Pterodactylus münsteri</i> and <i>Pterodactylus longicaudus</i>.<sup id="cite_ref-vonmeyer1845_52-0" class="reference"><a href="#cite_note-vonmeyer1845-52">[52]</a></sup> In 1846, von Meyer created the new species <i>Pterodactylus gemmingi</i> based on long-tailed remains; the specific name honors the fossil collector <a href="/enwiki/w/index.php?title=Carl_Eming_von_Gemming&action=edit&redlink=1" class="new" title="Carl Eming von Gemming (page does not exist)">Carl Eming von Gemming</a>.<sup id="cite_ref-vonmeyer1846_53-0" class="reference"><a href="#cite_note-vonmeyer1846-53">[53]</a></sup> Later, in 1847, von Meyer finally erected the generic name <i>Rhamphorhynchus</i> (<abbr title="literal translation">lit.</abbr><span style="white-space:nowrap"> </span>'beak snout') due to the distinctively long tails seen in the specimens found, which are much longer than those seen in <i>Pterodactylus</i>. He assigned the species <i>P. longicaudus</i> as the type species of <i>Rhamphorhynchus</i>, which resulted in a new combination called <i>Rhamphorhynchus longicaudus</i>.<sup id="cite_ref-vonmeyer1847_54-0" class="reference"><a href="#cite_note-vonmeyer1847-54">[54]</a></sup> The species <i>R. münsteri</i> was later changed to <i>R. muensteri</i> by Lydekker in 1888, due to the <a href="/enwiki/wiki/International_Commission_on_Zoological_Nomenclature" title="International Commission on Zoological Nomenclature">ICZN</a> rule that prohibits non-standard Latin characters, such as <i>ü</i>, in scientific names.<sup id="cite_ref-Lydekker_22-2" class="reference"><a href="#cite_note-Lydekker-22">[22]</a></sup>
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<div class="thumb tright"><div class="thumbinner" style="width:222px;"><a href="/enwiki/wiki/File:PZSL1851PlateReptilia04.png" class="image"><img alt="" src="/upwiki/wikipedia/commons/thumb/4/4c/PZSL1851PlateReptilia04.png/220px-PZSL1851PlateReptilia04.png" decoding="async" width="220" height="106" class="thumbimage" srcset="/upwiki/wikipedia/commons/thumb/4/4c/PZSL1851PlateReptilia04.png/330px-PZSL1851PlateReptilia04.png 1.5x, /upwiki/wikipedia/commons/thumb/4/4c/PZSL1851PlateReptilia04.png/440px-PZSL1851PlateReptilia04.png 2x" data-file-width="2636" data-file-height="1273" /></a> <div class="thumbcaption"><div class="magnify"><a href="/enwiki/wiki/File:PZSL1851PlateReptilia04.png" class="internal" title="Enlarge"></a></div>Holotype jaw fragments and teeth of <i><a href="/enwiki/wiki/Cimoliopterus" title="Cimoliopterus">Cimoliopterus</a></i>, which was previously known as <i>Pterodactylus cuvieri</i></div></div></div>
<p>Beginning in 1846, many pterosaur specimens were found near the village of <a href="/enwiki/wiki/Burham" title="Burham">Burham</a> in <a href="/enwiki/wiki/Kent" title="Kent">Kent</a>, <a href="/enwiki/wiki/England" title="England">England</a> by British paleontologists <a href="/enwiki/wiki/James_Scott_Bowerbank" title="James Scott Bowerbank">James Scott Bowerbank</a> and Sir <a href="/enwiki/wiki/Richard_Owen" title="Richard Owen">Richard Owen</a>. Bowerbank had assigned fossil remains to two new species; the first was named in 1846 as <i>Pterodactylus giganteus</i>;<sup id="cite_ref-Bowerbank1846_55-0" class="reference"><a href="#cite_note-Bowerbank1846-55">[55]</a></sup> 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 <i>Pterodactylus cuvieri</i>, in honor of the French scientist Georges Cuvier.<sup id="cite_ref-Bowerbank1851_56-0" class="reference"><a href="#cite_note-Bowerbank1851-56">[56]</a></sup> 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 <i>Pterodactylus compressirostris</i>.<sup id="cite_ref-Owen1851_57-0" class="reference"><a href="#cite_note-Owen1851-57">[57]</a></sup> In 1914 however, paleontologist <a href="/enwiki/wiki/Reginald_Hooley" title="Reginald Hooley">Reginald Hooley</a> redescribed <i>P. compressirostris</i>, to which he erected the genus <i><a href="/enwiki/wiki/Lonchodectes" title="Lonchodectes">Lonchodectes</a></i> (<abbr title="literal translation">lit.</abbr><span style="white-space:nowrap"> </span>'<a href="/enwiki/wiki/Lance" title="Lance">lance</a> biter'), and therefore made <i>P. compressirostris</i> the type species, and created the new combination <i>L. compressirostris</i>.<sup id="cite_ref-:1_58-0" class="reference"><a href="#cite_note-:1-58">[58]</a></sup> In a 2013 review, <i>P. giganteus</i> and <i>P. cuvieri</i> were reassigned to new genera; <i>P. giganteus</i> was reassigned to a genus called <i><a href="/enwiki/wiki/Lonchodraco" title="Lonchodraco">Lonchodraco</a></i> ('lance dragon'), which resulted in a new combination called <i>L. giganteus</i>, and <i>P. cuvieri</i> was reassigned to the new genus <i><a href="/enwiki/wiki/Cimoliopterus" title="Cimoliopterus">Cimoliopterus</a></i> ('chalk wing'), creating <i>C. cuvieri</i>.<sup id="cite_ref-Rodrigues_&_Kellner_2013_59-0" class="reference"><a href="#cite_note-Rodrigues_&_Kellner_2013-59">[59]</a></sup> Back in 1859, Owen had found remains the front part of a snout in the <a href="/enwiki/wiki/Cambridge_Greensand" title="Cambridge Greensand">Cambridge Greensand</a>, and assigned it into the species <i>Pterodactylus segwickii</i>; in honor of <a href="/enwiki/wiki/Adam_Sedgwick" title="Adam Sedgwick">Adam Sedgwick</a>, a British geologist.<sup id="cite_ref-60" class="reference"><a href="#cite_note-60">[60]</a></sup> This species however, was reassigned to the genus <i><a href="/enwiki/wiki/Camposipterus" title="Camposipterus">Camposipterus</a></i> in 2013, therefore creating the new combination <i>Camposipterus segwickii</i>.<sup id="cite_ref-Rodrigues_&_Kellner_2013_59-1" class="reference"><a href="#cite_note-Rodrigues_&_Kellner_2013-59">[59]</a></sup> 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 <i>Pterodactylus simus</i>,<sup id="cite_ref-61" class="reference"><a href="#cite_note-61">[61]</a></sup> though the British paleontologist <a href="/enwiki/wiki/Harry_Govier_Seeley" class="mw-redirect" title="Harry Govier Seeley">Harry Govier Seeley</a> had created a separate generic name called <i><a href="/enwiki/wiki/Ornithocheirus" title="Ornithocheirus">Ornithocheirus</a></i>, and reassigned <i>P. simus</i> as the type species, which created the combination <i>Ornithocheirus simus</i>.<sup id="cite_ref-S1869_62-0" class="reference"><a href="#cite_note-S1869-62">[62]</a></sup> Between the years 1869 and 1870, Seeley had reassigned many pterosaur species into <i>Ornithocheirus</i>, while also creating several new species.<sup id="cite_ref-S1869_62-1" class="reference"><a href="#cite_note-S1869-62">[62]</a></sup><sup id="cite_ref-HGS70_63-0" class="reference"><a href="#cite_note-HGS70-63">[63]</a></sup> Many if these species however, are now reclassified to other genera, or considered <i lang="la" title="Latin-language text"><a href="/enwiki/wiki/Nomina_dubia" class="mw-redirect" title="Nomina dubia">nomina dubia</a></i>.<sup id="cite_ref-Rodrigues_&_Kellner_2013_59-2" class="reference"><a href="#cite_note-Rodrigues_&_Kellner_2013-59">[59]</a></sup> In 1874, further specimens were found in England, again by Owen, these ones were assigned to a new species called <i>Pterodactylus sagittirostris</i>,<sup id="cite_ref-64" class="reference"><a href="#cite_note-64">[64]</a></sup> this species however, was reassigned to the genus <i>Lonchodectes</i> in 1914 by Hooley, which resulted in an <i>L. sagittirostris</i>.<sup id="cite_ref-:1_58-1" class="reference"><a href="#cite_note-:1-58">[58]</a></sup> This conclusion was revised by Rigal <i>et al.</i> in 2017, who disagreed with Hooley's reassignment, and therefore created the genus <i><a href="/enwiki/wiki/Serradraco" title="Serradraco">Serradraco</a></i>, which afterwards resulted in a new combination called <i>S. sagittirostris</i>.<sup id="cite_ref-Martill2017_65-0" class="reference"><a href="#cite_note-Martill2017-65">[65]</a></sup>
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<div class="thumb tleft"><div class="thumbinner" style="width:222px;"><a href="/enwiki/wiki/File:Pteranodon_longiceps_YPM1177.jpg" class="image"><img alt="" src="/upwiki/wikipedia/commons/thumb/f/f8/Pteranodon_longiceps_YPM1177.jpg/220px-Pteranodon_longiceps_YPM1177.jpg" decoding="async" width="220" height="63" class="thumbimage" srcset="/upwiki/wikipedia/commons/thumb/f/f8/Pteranodon_longiceps_YPM1177.jpg/330px-Pteranodon_longiceps_YPM1177.jpg 1.5x, /upwiki/wikipedia/commons/thumb/f/f8/Pteranodon_longiceps_YPM1177.jpg/440px-Pteranodon_longiceps_YPM1177.jpg 2x" data-file-width="1323" data-file-height="381" /></a> <div class="thumbcaption"><div class="magnify"><a href="/enwiki/wiki/File:Pteranodon_longiceps_YPM1177.jpg" class="internal" title="Enlarge"></a></div>Specimen YPM1177, the first uncovered skull of <i><a href="/enwiki/wiki/Pteranodon" title="Pteranodon">Pteranodon</a></i>, which was back then assigned as a species of <i>Pterodactylus</i></div></div></div>
<p>Assigning new pterosaur species to <i>Pterodactylus</i> was not only common in Europe, but also in North America; paleontologists such as <a href="/enwiki/wiki/Othniel_Charles_Marsh" title="Othniel Charles Marsh">Othniel Charles Marsh</a> in 1871 for example, described several toothless pterosaur specimens, which were accompanied by teeth that belonged to the fish <i><a href="/enwiki/wiki/Xiphactinus" title="Xiphactinus">Xiphactinus</a></i>, 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 <i>"Pterodactylus oweni"</i>, but this was changed to <i>Pterodactylus occidentalis</i> because <i>"P. oweni"</i> was found to have been <a href="/enwiki/wiki/Preoccupied" class="mw-redirect" title="Preoccupied">preoccupied</a> by a pterosaur species described with the same name back in 1864 by Seeley.<sup id="cite_ref-66" class="reference"><a href="#cite_note-66">[66]</a></sup><sup id="cite_ref-History_67-0" class="reference"><a href="#cite_note-History-67">[67]</a></sup> In 1872, American paleontologist <a href="/enwiki/wiki/Edward_Drinker_Cope" title="Edward Drinker Cope">Edward Drinker Cope</a> also found various pterosaur specimens in North America, he assigned these to two new species known as <i>Ornithochirus umbrosus</i> and <i>Ornithochirus harpyia</i>, Cope attempted to assign the specimens he found to the genus <i>Ornithocheirus</i>, but misspelled forgetting the 'e'.<sup id="cite_ref-68" class="reference"><a href="#cite_note-68">[68]</a></sup> In 1875 however, Cope reassigned the species <i>O. umbrosus</i> and <i>O. harpyia</i> into <i>Pterodactylus umbrosus</i> and <i>Pterodactylus harpyia</i>, though these species had been considered <i lang="la" title="Latin-language text">nomina dubia</i> ever since.<sup id="cite_ref-cope1875_69-0" class="reference"><a href="#cite_note-cope1875-69">[69]</a></sup><sup id="cite_ref-History_67-1" class="reference"><a href="#cite_note-History-67">[67]</a></sup> Paleontologist <a href="/enwiki/wiki/Samuel_Wendell_Williston" title="Samuel Wendell Williston">Samuel Wendell Williston</a> unearthed the first skull of the pterosaur, and found that the animal was toothless,<sup id="cite_ref-History_67-2" class="reference"><a href="#cite_note-History-67">[67]</a></sup> this made Marsh create the genus <i><a href="/enwiki/wiki/Pteranodon" title="Pteranodon">Pteranodon</a></i> (lit. 'toothless wing'), and therefore reassigned all the American pterosaur species, including the ones that he named, from <i>Pterodactylus</i> to <i>Pteranodon</i>.<sup id="cite_ref-marsh1876a_70-0" class="reference"><a href="#cite_note-marsh1876a-70">[70]</a></sup>
</p><p>Later, in the 1980s, subsequent revisions by <a href="/enwiki/wiki/Peter_Wellnhofer" title="Peter Wellnhofer">Peter Wellnhofer</a> had reduced the number of recognized species to about half a dozen. Many species assigned to <i>Pterodactylus</i> 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. <i>P. elegans</i>, for example, was found by numerous studies to be an immature <i>Ctenochasma</i>.<sup id="cite_ref-jouve2004_25-3" class="reference"><a href="#cite_note-jouve2004-25">[25]</a></sup> Another species of <i>Pterodactylus</i> originally based on small, immature specimens was <i>P. micronyx</i>. However, it has been difficult to determine exactly of what genus and species <i>P. micronyx</i> might be the juvenile form. Stéphane Jouve, Christopher Bennett and others had once suggested that it probably belonged either to <i><a href="/enwiki/wiki/Gnathosaurus" title="Gnathosaurus">Gnathosaurus subulatus</a></i> or one of the species belonging to <i>Ctenochasma</i>,<sup id="cite_ref-bennett2002_24-1" class="reference"><a href="#cite_note-bennett2002-24">[24]</a></sup><sup id="cite_ref-jouve2004_25-4" class="reference"><a href="#cite_note-jouve2004-25">[25]</a></sup> though after additional research Bennett assigned it to the genus <i>Aurorazhdarcho</i>.<sup id="cite_ref-BennettPZ_17-8" class="reference"><a href="#cite_note-BennettPZ-17">[17]</a></sup> Another species with a complex history is <i>P. longicollum</i>, named by von Meyer in 1854, based on a large specimen with a long neck and fewer teeth. Many researchers, including <a href="/enwiki/w/index.php?title=David_Unwin&action=edit&redlink=1" class="new" title="David Unwin (page does not exist)">David Unwin</a>, have found <i>P. longicollum</i> to be distinct from <i>P. kochi</i> and <i>P. antiquus</i>. Unwin found <i>P. longicollum</i> to be closer to <i>Germanodactylus</i> and therefore requiring a new genus name.<sup id="cite_ref-unwin2003_4-2" class="reference"><a href="#cite_note-unwin2003-4">[4]</a></sup> It has sometimes been placed in the genus <i>Diopecephalus</i> because <a href="/enwiki/wiki/Harry_Govier_Seeley" class="mw-redirect" title="Harry Govier Seeley">Harry Govier Seeley</a> based this genus partly on the <i>P. longicollum</i> material. However, it was shown by Bennett that the <a href="/enwiki/wiki/Type_specimen" class="mw-redirect" title="Type specimen">type specimen</a> later designated for <i>Diopecephalus</i> was a fossil belonging to <i>P. kochi</i>, and no longer thought to be separate from <i>Pterodactylus</i>. <i>Diopecephalus</i> is therefore a synonym of <i>Pterodactylus</i>, and as such is unavailable for use as a new genus for <i>"P." longicollum</i>.<sup id="cite_ref-SCB06_71-0" class="reference"><a href="#cite_note-SCB06-71">[71]</a></sup> <i>"P." longicollum</i> was eventually made the type species of a separate genus <i>Ardeadactylus</i>.<sup id="cite_ref-BennettPZ_17-9" class="reference"><a href="#cite_note-BennettPZ-17">[17]</a></sup>
</p>
<h3><span class="mw-headline" id="Controversial_species">Controversial species</span><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Pterodactylus&action=edit&section=11" title="Edit section: Controversial species">edit</a><span class="mw-editsection-bracket">]</span></span></h3>
<div class="thumb tright"><div class="thumbinner" style="width:222px;"><a href="/enwiki/wiki/File:Bsp_as_xix.png" class="image"><img alt="" src="/upwiki/wikipedia/commons/thumb/6/69/Bsp_as_xix.png/220px-Bsp_as_xix.png" decoding="async" width="220" height="142" class="thumbimage" srcset="/upwiki/wikipedia/commons/thumb/6/69/Bsp_as_xix.png/330px-Bsp_as_xix.png 1.5x, /upwiki/wikipedia/commons/thumb/6/69/Bsp_as_xix.png/440px-Bsp_as_xix.png 2x" data-file-width="889" data-file-height="572" /></a> <div class="thumbcaption"><div class="magnify"><a href="/enwiki/wiki/File:Bsp_as_xix.png" class="internal" title="Enlarge"></a></div>Juvenile type specimen of <i>Pterodactylus kochi</i>, now reassigned as <i><a href="/enwiki/wiki/Diopecephalus_kochi" class="mw-redirect" title="Diopecephalus kochi">Diopecephalus kochi</a></i></div></div></div>
<p>The only well-known and well-supported species left by the first decades of the 21st century were <i>P. antiquus</i> and <i>P. kochi</i>. However, most studies between 1995 and 2010 found little reason to separate even these two species, and treated them as synonymous.<sup id="cite_ref-unwin2003_4-3" class="reference"><a href="#cite_note-unwin2003-4">[4]</a></sup> More recent studies of pterosaur relationships have found anurognathids and pterodactyloids to be sister groups, which would limit the more inclusive group <a href="/enwiki/wiki/Caelidracones" title="Caelidracones">Caelidracones</a> to just two clades.<sup id="cite_ref-LoneStarPterosaurs_72-0" class="reference"><a href="#cite_note-LoneStarPterosaurs-72">[72]</a></sup><sup id="cite_ref-SCB06_71-1" class="reference"><a href="#cite_note-SCB06-71">[71]</a></sup> In 1996, Bennett suggested that the differences between specimens of <i>P. kochi</i> and <i>P. antiquus</i> could be explained by differences in age, with <i>P. kochi</i> (including specimens alternately classified in the species <i>P. scolopaciceps</i>) representing an immature growth stage of <i>P. antiquus</i>. In a 2004 paper, Jouve used a different method of analysis and recovered the same result, showing that the "distinctive" features of <i>P. kochi</i> were age-related, and using mathematical comparison to show that the two forms are different growth stages of the same species.<sup id="cite_ref-jouve2004_25-5" class="reference"><a href="#cite_note-jouve2004-25">[25]</a></sup> An additional review of the specimens published in 2013 demonstrated that some of the supposed differences between <i>P. kochi</i> and <i>P. antiquus</i> were due to measurement errors, further supporting their synonymy.<sup id="cite_ref-BennettPZ_17-10" class="reference"><a href="#cite_note-BennettPZ-17">[17]</a></sup>
</p><p>By the 2010s, a large body of research had been developed based on the idea that <i>P. kochi</i> and <i>P. scolopaciceps</i> were early growth stages of <i>P. antiquus</i>. However, in 2014, two scientists began publishing research that challenged this paradigm. Steven Vidovic and David Martill concluded that differences between specimens of <i>P. kochi</i>, <i>P. scolopaciceps</i>, and <i>P. antiquus</i>, 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 <i>P. kochi</i> type specimen did not form a natural group with that of <i>P. antiquus</i>. They concluded that the genus <i>Diopecephalus</i> could be returned to use to distinguish <i>"P". kochi</i> from <i>P. antiquus</i>. They named the new genus <i><a href="/enwiki/wiki/Aerodactylus" title="Aerodactylus">Aerodactylus</a></i> for <i>P. scolopaciceps</i> as well. So, what Bennett considered early growth stages of one species, Vidovic and Martill considered representatives of new species.<sup id="cite_ref-Aerodactylus_48-1" class="reference"><a href="#cite_note-Aerodactylus-48">[48]</a></sup><sup id="cite_ref-W1837_32-3" class="reference"><a href="#cite_note-W1837-32">[32]</a></sup>
</p><p>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 <i>Aerodactylus</i>, 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 <i>Diopecephalus</i> and <i>Aerodactylus</i> simply as year-classes of immature <i>Pterodactylus antiquus</i>.<sup id="cite_ref-bennett_2017_pteranodon_juv_73-0" class="reference"><a href="#cite_note-bennett_2017_pteranodon_juv-73">[73]</a></sup>
</p>
<h3><span class="mw-headline" id="List_of_species">List of species</span><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Pterodactylus&action=edit&section=12" title="Edit section: List of species">edit</a><span class="mw-editsection-bracket">]</span></span></h3>
<p>During its over-200-year history, the various species of <i>Pterodactylus</i> 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 <i>Pterodactylus</i> are based on poor remains that have proven difficult to assign to one species or another and are therefore considered <i lang="la" title="Latin-language text">nomina dubia</i> (<abbr title="literal translation">lit.</abbr><span style="white-space:nowrap"> </span>'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 <i>Pterodactylus</i> proper, and names based on other material that has as yet not been assigned to other genera. This list also includes species that are <i lang="la" title="Latin-language text"><a href="/enwiki/wiki/Nomina_nuda" class="mw-redirect" title="Nomina nuda">nomina nuda</a></i> ('naked names'), which are species that were not published formally. Species that are <i lang="la" title="Latin-language text"><a href="/enwiki/wiki/Nomina_oblita" class="mw-redirect" title="Nomina oblita">nomina oblita</a></i> ('forgotten names') are the ones that have been disused, and species that are <i lang="la" title="Latin-language text"><a href="/enwiki/wiki/Nomina_rejecta" class="mw-redirect" title="Nomina rejecta">nomina rejecta</a></i> ('rejected names') are the ones that have been rejected because a more preferable name had been accepted instead.<sup id="cite_ref-Lydekker_22-3" class="reference"><a href="#cite_note-Lydekker-22">[22]</a></sup><sup id="cite_ref-74" class="reference"><a href="#cite_note-74">[74]</a></sup>
</p>
<div style="margin-left:0">
<table class="mw-collapsible mw-collapsed" style="background: transparent; text-align: left; border: 1px solid Silver; margin: 0.2em auto auto; width:100%; clear: both; padding: 1px;">
<tbody><tr>
<th style="background: #F0F2F5; font-size:87%; padding:0.2em 0.3em; text-align:center;"><div style="display:inline;font-size:115%">List of species</div>
</th></tr>
<tr>
<td style="border: solid 1px Silver; padding: 0.6em; background: White;">
<table class="wikitable sortable">
<tbody><tr>
<th>Name
</th>
<th>Author
</th>
<th>Year
</th>
<th>Status
</th>
<th class="unsortable">Notes
</th></tr>
<tr>
<td><i>Pterodactylus antiquus</i>
</td>
<td>(<a href="/enwiki/wiki/Samuel_Thomas_von_S%C3%B6mmerring" title="Samuel Thomas von Sömmerring">Sömmerring</a>)
</td>
<td>(1812)
</td>
<td>Valid
</td>
<td>Designated as the <a href="/enwiki/wiki/Type_species" title="Type species">type species</a> of <i>Pterodactylus</i>, and replacing <i>Ornithocephalus antiquus</i> due to being an obsolete name
</td></tr>
<tr>
<td><i>Pterodactylus brevirostris</i>
</td>
<td>(Sömmerring)
</td>
<td>(1817)
</td>
<td>Synonym of <i><a href="/enwiki/wiki/Ctenochasma_elegans" class="mw-redirect" title="Ctenochasma elegans">Ctenochasma elegans</a></i>
</td>
<td>Reclassified from <i>Ornithocephalus brevirostris</i>
</td></tr>
<tr>
<td><i>Pterodactylus giganteus</i>
</td>
<td>(Oken)
</td>
<td>(1819)
</td>
<td><i><a href="/enwiki/wiki/Nomen_dubium" title="Nomen dubium">Nomen dubium</a></i>
</td>
<td>Synonym of <i><a href="/enwiki/wiki/Rhamphorhynchus_muensteri" class="mw-redirect" title="Rhamphorhynchus muensteri">Rhamphorhynchus muensteri</a></i>; reclassified from <i>Ornithocephalus giganteus</i>
</td></tr>
<tr>
<td><i>Pterodactylus longirostris</i>
</td>
<td><a href="/enwiki/wiki/Georges_Cuvier" title="Georges Cuvier">Cuvier</a>
</td>
<td>1819
</td>
<td>Synonym of <i>Pterodactylus antiquus</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus grandis</i>
</td>
<td>Cuvier
</td>
<td>1824
</td>
<td><i>Nomen dubium</i>
</td>
<td>Synonym of <i>Rhamphorhynchus muensteri</i>
</td></tr>
<tr>
<td><i>Pterodactylus crocodilocephaloides</i>
</td>
<td>Ritgen
</td>
<td>1826
</td>
<td>Synonym of <i>Pterodactylus antiquus</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus nettecephaloides</i>
</td>
<td>Ritgen
</td>
<td>1826
</td>
<td><i><a href="/enwiki/wiki/Nomen_oblitum" title="Nomen oblitum">Nomen oblitum</a></i>
</td>
<td>Synonym of <i><a href="/enwiki/wiki/Aurorazhdarcho_micronyx" class="mw-redirect" title="Aurorazhdarcho micronyx">Aurorazhdarcho micronyx</a></i>
</td></tr>
<tr>
<td><i>Pterodactylus macronyx</i>
</td>
<td><a href="/enwiki/wiki/William_Buckland" title="William Buckland">Buckland</a>
</td>
<td>1829
</td>
<td>Reclassified as <i><a href="/enwiki/wiki/Dimorphodon_macronyx" class="mw-redirect" title="Dimorphodon macronyx">Dimorphodon macronyx</a></i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus (Rhamphorhynchus) macronyx</i>
</td>
<td>(Buckland)
</td>
<td>(1829)
</td>
<td>Synonym of <i>Dimorphodon macronyx</i>
</td>
<td>Reclassified from <i>Dimorphodon macronyx</i>
</td></tr>
<tr>
<td><i>Pterodactylus banthensis</i>
</td>
<td>(Theodori)
</td>
<td>(1830)
</td>
<td>Reclassified as <i><a href="/enwiki/wiki/Dorygnathus_banthensis" class="mw-redirect" title="Dorygnathus banthensis">Dorygnathus banthensis</a></i>
</td>
<td>Reclassified from <i>Ornithocephalus banthensis</i>
</td></tr>
<tr>
<td><i>Pterodactylus (Rhamphorhynchus) (ensirostris) banthensis</i>
</td>
<td>(Theodori)
</td>
<td>(1830)
</td>
<td><i>Nomen oblitum</i>
</td>
<td>Synonym of <i>Dorygnathus banthensis</i>
</td></tr>
<tr>
<td><i>Pterodactylus goldfussi</i>
</td>
<td>Theodori
</td>
<td>1830
</td>
<td>Synonym of <i>Dorygnathus banthensis</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus crassirostris</i>
</td>
<td><a href="/enwiki/wiki/Georg_August_Goldfuss" title="Georg August Goldfuss">Goldfuss</a>
</td>
<td>1831
</td>
<td>Reclassified as <i><a href="/enwiki/wiki/Scaphognathus_crassirostris" class="mw-redirect" title="Scaphognathus crassirostris">Scaphognathus crassirostris</a></i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus macronyx</i>
</td>
<td><a href="/enwiki/wiki/Christian_Erich_Hermann_von_Meyer" title="Christian Erich Hermann von Meyer">Meyer</a> <i>non</i> Buckland
</td>
<td>1831 <i>non</i> 1829
</td>
<td>Synonym of <i>Dorygnathus banthensis</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus medius</i>
</td>
<td><a href="/enwiki/wiki/Georg_zu_M%C3%BCnster" title="Georg zu Münster">Münster</a>
</td>
<td>1831
</td>
<td><i>Nomen dubium</i>
</td>
<td>Synonym of <i><a href="/enwiki/wiki/Altmuehlopterus_rhamphastinus" class="mw-redirect" title="Altmuehlopterus rhamphastinus">Altmuehlopterus rhamphastinus</a></i>
</td></tr>
<tr>
<td><i>Pterodactylus muensteri</i>
</td>
<td>(Goldfuss)
</td>
<td>(1831)
</td>
<td>Synonym of <i>Rhamphorhynchus muensteri</i>
</td>
<td>Reclassified from <i>Ornithocephalus muensteri</i>
</td></tr>
<tr>
<td><i>Pterodactylus (Rhamphorhynchus) muensteri</i>
</td>
<td>(Goldfuss)
</td>
<td>(1831)
</td>
<td>Synonym of <i>Rhamphorhynchus muensteri</i>
</td>
<td>Reclassified from <i>Pterodactylus muensteri</i>
</td></tr>
<tr>
<td><i>Pterodactylus bucklandi</i>
</td>
<td>Meyer
</td>
<td>1832
</td>
<td>Indeterminate <a href="/enwiki/wiki/Rhamphorhynchidae" title="Rhamphorhynchidae">rhamphorhynchine</a>
</td>
<td>Once assigned to the genus <i><a href="/enwiki/wiki/Rhamphocephalus" title="Rhamphocephalus">Rhamphocephalus</a></i>
</td></tr>
<tr>
<td><i>Pterodactylus longipes</i>
</td>
<td>Münster
</td>
<td>1836
</td>
<td>Synonym of <i><a href="/enwiki/wiki/Ardeadactylus_longicollum" class="mw-redirect" title="Ardeadactylus longicollum">Ardeadactylus longicollum</a></i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus kochi</i>
</td>
<td>Wagner
</td>
<td>1837
</td>
<td>Reclassified as <i><a href="/enwiki/wiki/Diopecephalus_kochi" class="mw-redirect" title="Diopecephalus kochi">Diopecephalus kochi</a></i>
</td>
<td>Reclassified from <i>Ornithocephalus kochi</i>; possible synonym of <i>Pterodactylus antiquus</i>
</td></tr>
<tr>
<td><i>Pterodactylus lavateri</i>
</td>
<td>Meyer
</td>
<td>1838
</td>
<td>Synonym of <i>Rhamphorhynchus muensteri</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus (Ornithopterus) lavateri</i>
</td>
<td>(Meyer)
</td>
<td>(1838)
</td>
<td>Synonym of <i>Rhamphorhynchus muensteri</i>
</td>
<td>Reclassified from <i>Pterodactylus lavateri</i>
</td></tr>
<tr>
<td><i>Pterodactylus longicaudus</i>
</td>
<td>Münster
</td>
<td>1839
</td>
<td>Reclassified as <i>Rhamphorhynchus longicaudus</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus meyeri</i>
</td>
<td>Münster
</td>
<td>1842
</td>
<td>Synonym of <i>Diopecephalus kochi</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus secundarius</i>
</td>
<td>Meyer
</td>
<td>1843
</td>
<td><i>Nomen dubium</i>
</td>
<td>Synonym of <i>Rhamphorhynchus muensteri</i>
</td></tr>
<tr>
<td><i>Pterodactylus cliftii</i>
</td>
<td>Mantell
</td>
<td>1844
</td>
<td>Synonym of <i><a href="/enwiki/wiki/Palaeornis_cliftii" title="Palaeornis cliftii">Palaeornis cliftii</a></i>
</td>
<td>Reclassified from <i>Palaeornis cliftii</i>
</td></tr>
<tr>
<td><i>Pterodactylus diomedeus</i>
</td>
<td>(<a href="/enwiki/wiki/Richard_Owen" title="Richard Owen">Owen</a>)
</td>
<td>(1846)
</td>
<td>Synonym of <i><a href="/enwiki/wiki/Cimoliornis_diomedeus" class="mw-redirect" title="Cimoliornis diomedeus">Cimoliornis diomedeus</a></i>
</td>
<td>Reclassified from <i>Cimoliornis diomedeus</i>
</td></tr>
<tr>
<td><i>Pterodactylus gemmingi</i>
</td>
<td>Meyer
</td>
<td>1846
</td>
<td>Synonym of <i>Rhamphorhynchus muensteri</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus (Rhamphorhynchus) gemmingi</i>
</td>
<td>(Meyer)
</td>
<td>(1846)
</td>
<td>Synonym of <i>Rhamphorhynchus muensteri</i>
</td>
<td>Reclassified from <i>Pterodactylus gemmingi</i>
</td></tr>
<tr>
<td><i>Pterodactylus giganteus</i>
</td>
<td><a href="/enwiki/wiki/James_Scott_Bowerbank" title="James Scott Bowerbank">Bowerbank</a> <i>non</i> Oken
</td>
<td>1846 <i>non</i> 1819
</td>
<td>Reclassified as <i><a href="/enwiki/wiki/Lonchodraco_giganteus" class="mw-redirect" title="Lonchodraco giganteus">Lonchodraco giganteus</a></i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus ornis</i>
</td>
<td><a href="/enwiki/wiki/Christoph_Gottfried_Andreas_Giebel" title="Christoph Gottfried Andreas Giebel">Giebel</a>
</td>
<td>1847
</td>
<td>Synonym of <i>Palaeornis cliftii</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus conirostris</i>
</td>
<td>Owen
</td>
<td>1850
</td>
<td>Synonym of <i>Lonchodraco giganteus</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus cuvieri</i>
</td>
<td>Bowerbank
</td>
<td>1851
</td>
<td>Reclassified as <i><a href="/enwiki/wiki/Cimoliopterus_cuvieri" class="mw-redirect" title="Cimoliopterus cuvieri">Cimoliopterus cuvieri</a></i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus compressirostris</i>
</td>
<td>Owen
</td>
<td>1851
</td>
<td>Reclassified as <i><a href="/enwiki/wiki/Lonchodectes_compressirostris" class="mw-redirect" title="Lonchodectes compressirostris">Lonchodectes compressirostris</a></i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus redenbacheri</i>
</td>
<td>(Wagner)
</td>
<td>(1851)
</td>
<td>Synonym of <i>Aurorazhdarcho micronyx</i>
</td>
<td>Reclassified from <i>Ornithocephalus redenbacheri</i>
</td></tr>
<tr>
<td><i>Pterodactylus rhamphastinus</i>
</td>
<td>(Wagner)
</td>
<td>(1851)
</td>
<td>Reclassified as <i>Altmuehlopterus rhamphastinus</i>
</td>
<td>Reclassified from <i>Ornithocephalus rhamphastinus</i>
</td></tr>
<tr>
<td><i>Pterodactylus giganteus</i>
</td>
<td>Morris <i>non</i> Oken <i>non</i> Bowerbank
</td>
<td>1854 <i>non</i> 1819 <i>non</i> 1846
</td>
<td>Synonym of <i>Cimoliornis diomedeus</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus longicollum</i>
</td>
<td>Meyer
</td>
<td>1854
</td>
<td>Reclassified as <i>Ardeadactylus longicollum</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus wuerttembergicus</i>
</td>
<td><a href="/enwiki/wiki/Friedrich_August_Quenstedt" class="mw-redirect" title="Friedrich August Quenstedt">Quenstedt</a>
</td>
<td>1854
</td>
<td><i>Nomen oblitum</i>
</td>
<td>Synonym of <i><a href="/enwiki/wiki/Cycnorhamphus_suevicus" class="mw-redirect" title="Cycnorhamphus suevicus">Cycnorhamphus suevicus</a></i>
</td></tr>
<tr>
<td><i>Pterodactylus suevicus</i>
</td>
<td>Quenstedt
</td>
<td>1855
</td>
<td>Reclassified as <i>Cycnorhamphus suevicus</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus micronyx</i>
</td>
<td>Meyer
</td>
<td>1856
</td>
<td>Reclassified as <i>Aurorazhdarcho micronyx</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus crassipes</i>
</td>
<td>Meyer
</td>
<td>1857
</td>
<td><i><a href="/enwiki/wiki/Nomen_rejectum" class="mw-redirect" title="Nomen rejectum">Nomen rejectum</a></i>
</td>
<td>Reclassified as <i><a href="/enwiki/wiki/Ostromia_crassipes" class="mw-redirect" title="Ostromia crassipes">Ostromia crassipes</a></i>
</td></tr>
<tr>
<td><i>Pterodactylus (Rhamphorhynchus) crassipes</i>
</td>
<td>(Meyer)
</td>
<td>(1857)
</td>
<td><i>Nomen rejectum</i>
</td>
<td>Synonym of <i>Ostromia crassipes</i>; reclassified from <i>Pterodactylus crassipes</i>
</td></tr>
<tr>
<td><i>Pterodactylus eurychirus</i>
</td>
<td>Wagner
</td>
<td>1857
</td>
<td>Synonym of <i>Cycnorhamphus suevicus</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus (Ornithocephalus) eurychirus</i>
</td>
<td>(Wagner)
</td>
<td>(1857)
</td>
<td>Synonym of <i>Cycnorhamphus suevicus</i>
</td>
<td>Reclassified from <i>Pterodactylus eurychirus</i>
</td></tr>
<tr>
<td><i>Pterodactylus suevicus eurychirus</i>
</td>
<td>(Wagner)
</td>
<td>(1857)
</td>
<td>Synonym of <i>Cycnorhamphus suevicus</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus hirundinaceus</i>
</td>
<td>Wagner
</td>
<td>1857
</td>
<td>Synonym of <i>Rhamphorhynchus muensteri</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus (Rhamphorhynchus) hirundinaceus</i>
</td>
<td>Wagner
</td>
<td>1857
</td>
<td>Synonym of <i>Rhamphorhynchus muensteri</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus (Ornithocephalus) propinquus</i>
</td>
<td>Wagner
</td>
<td>1857
</td>
<td><i>Nomen dubium</i>
</td>
<td>Possible synonym of <i>Altmuehlopterus rhamphastinus</i>
</td></tr>
<tr>
<td><i>Pterodactylus propinquus</i>
</td>
<td>(Wagner)
</td>
<td>(1857)
</td>
<td><i>Nomen dubium</i>
</td>
<td>Reclassified from <i>Pterodactylus (Ornithocephalus) propinquus</i>; possible synonym of <i>Altmuehlopterus rhamphastinus</i>
</td></tr>
<tr>
<td><i>Pterodactylus propinquus medius</i>
</td>
<td>Wagner
</td>
<td>1857
</td>
<td><i>Nomen dubium</i>
</td>
<td>Synonym of <i>Altmuehlopterus rhamphastinus</i>
</td></tr>
<tr>
<td><i>Pterodactylus (Ornithocephalus) vulturinus</i>
</td>
<td>Wagner
</td>
<td>1857
</td>
<td>Synonym of <i>Ardeadactylus longicollum</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus vulturinus</i>
</td>
<td>(Wagner)
</td>
<td>(1857)
</td>
<td>Synonym of <i>Ardeadactylus longicollum</i>
</td>
<td>Reclassified from <i>Pterodactylus (Ornithocephalus) vulturinus</i>
</td></tr>
<tr>
<td><i>Pterodactylus liasicus</i>
</td>
<td>Quenstedt
</td>
<td>1858
</td>
<td>Reclassified as <i><a href="/enwiki/wiki/Campylognathoides_liasicus" class="mw-redirect" title="Campylognathoides liasicus">Campylognathoides liasicus</a></i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus fittoni</i>
</td>
<td>Owen
</td>
<td>1859
</td>
<td><i>Nomen dubium</i>
</td>
<td>Reclassified as either <i><a href="/enwiki/wiki/Ornithocheirus" title="Ornithocheirus">Ornithocheirus</a> fittoni</i> or <i><a href="/enwiki/wiki/Anhanguera_(pterosaur)" title="Anhanguera (pterosaur)">Anhanguera</a> fittoni</i>; possible synonym of <i>Cimoliopterus cuvieri</i>
</td></tr>
<tr>
<td><i>Pterodactylus sedgwickii</i>
</td>
<td>Owen
</td>
<td>1859
</td>
<td>Reclassified as <i><a href="/enwiki/wiki/Camposipterus_sedgwickii" class="mw-redirect" title="Camposipterus sedgwickii">Camposipterus sedgwickii</a></i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus silvestris</i>
</td>
<td>Owen
</td>
<td>1859
</td>
<td>Synonym of <i>Palaeornis cliftii</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus cerinensis</i>
</td>
<td>Meyer
</td>
<td>1860
</td>
<td><i>Nomen dubium</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus grandipelvis</i>
</td>
<td>Meyer
</td>
<td>1860
</td>
<td><i>Nomen dubium</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus scolopaciceps</i>
</td>
<td>Meyer
</td>
<td>1860
</td>
<td>Reclassified as <i><a href="/enwiki/wiki/Aerodactylus_scolopaciceps" class="mw-redirect" title="Aerodactylus scolopaciceps">Aerodactylus scolopaciceps</a></i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus elegans</i>
</td>
<td>Wagner
</td>
<td>1861
</td>
<td>Reclassified as <i>Ctenochasma elegans</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus pulchellus</i>
</td>
<td>Meyer
</td>
<td>1861
</td>
<td>Synonym of <i>Aurorazhdarcho micronyx</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus simus</i>
</td>
<td>Owen
</td>
<td>1861
</td>
<td>Reclassified as <i>Ornithocheirus simus</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus spectabilis</i>
</td>
<td>Meyer
</td>
<td>1861
</td>
<td>Synonym of <i>Pterodactylus antiquus</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus woodwardi</i>
</td>
<td>Owen
</td>
<td>1861
</td>
<td><i>Nomen dubium</i>
</td>
<td>Reclassified as <i>Ornithocheirus woodwardi</i>
</td></tr>
<tr>
<td><i>Pterodactylus hopkinsi</i>
</td>
<td><a href="/enwiki/wiki/Harry_Govier_Seeley" class="mw-redirect" title="Harry Govier Seeley">Seeley</a>
</td>
<td>1864
</td>
<td><i><a href="/enwiki/wiki/Nomen_nudum" title="Nomen nudum">Nomen nudum</a></i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus oweni</i>
</td>
<td>Seeley
</td>
<td>1864
</td>
<td>Synonym of <i><a href="/enwiki/wiki/Ikrandraco_machaerorhynchus" class="mw-redirect" title="Ikrandraco machaerorhynchus">Ikrandraco machaerorhynchus</a></i>
</td>
<td>Not to be confused with the preoccupied species <i>"Pterodactylus oweni"</i>
</td></tr>
<tr>
<td><i>Pterodactylus longispinis</i>
</td>
<td><a href="/enwiki/wiki/Edward_Drinker_Cope" title="Edward Drinker Cope">Cope</a>
</td>
<td>1866
</td>
<td>Reclassified as <i><a href="/enwiki/wiki/Rhabdopelix_longispinis" class="mw-redirect" title="Rhabdopelix longispinis">Rhabdopelix longispinis</a></i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus macrurus</i>
</td>
<td>Seeley
</td>
<td>1869
</td>
<td>Reclassified as <i><a href="/enwiki/wiki/Gnathosaurus_macrurus" class="mw-redirect" title="Gnathosaurus macrurus">Gnathosaurus macrurus</a></i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus nobilis</i>
</td>
<td>Owen
</td>
<td>1869
</td>
<td><i>Nomen dubium</i>
</td>
<td>Reclassified as <i>Ornithocheirus nobilis</i>
</td></tr>
<tr>
<td><i>Pterodactylus curtus</i>
</td>
<td>Owen
</td>
<td>1870
</td>
<td><i>Nomen dubium</i>
</td>
<td>Reclassified as <i>Ornithocheirus curtus</i>
</td></tr>
<tr>
<td><i>"Pterodactylus oweni"</i>
</td>
<td><a href="/enwiki/wiki/Othniel_Marsh" class="mw-redirect" title="Othniel Marsh">Marsh</a>
</td>
<td>1871
</td>
<td>Preoccupied
</td>
<td>Reassigned as <i>Pterodactylus occidentalis</i> prior to being preoccupied
</td></tr>
<tr>
<td><i>Pterodactylus ingens</i>
</td>
<td>Marsh
</td>
<td>1872
</td>
<td>Synonym of <i><a href="/enwiki/wiki/Pteranodon_longiceps" class="mw-redirect" title="Pteranodon longiceps">Pteranodon longiceps</a></i>
</td>
<td>Reclassified as <i>Pteranodon ingens</i>
</td></tr>
<tr>
<td><i>Pterodactylus occidentalis</i>
</td>
<td>Marsh
</td>
<td>1872
</td>
<td><i>Nomen dubium</i>
</td>
<td>Reclassified from the preoccupied species <i>"Pterodactylus oweni"</i>; reclassified as <i>Pteranodon occidentalis</i>
</td></tr>
<tr>
<td><i>Pterodactylus velox</i>
</td>
<td>Marsh
</td>
<td>1872
</td>
<td><i>Nomen dubium</i>
</td>
<td>Reclassified as <i>Pteranodon velox</i>
</td></tr>
<tr>
<td><i>Pterodactylus suprajurensis</i>
</td>
<td>Sauvage
</td>
<td>1873
</td>
<td><i>Nomen dubium</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus aclandi</i>
</td>
<td>Owen
</td>
<td>1874
</td>
<td><i>Nomen dubium</i>
</td>
<td>Possible synonym of <i>Rhamphocephalus depressirostris</i>
</td></tr>
<tr>
<td><i>Pterodactylus daviesii</i>
</td>
<td>Owen
</td>
<td>1874
</td>
<td><i>Nomen dubium</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus duncani</i>
</td>
<td>Owen
</td>
<td>1874
</td>
<td><i>Nomen dubium</i>
</td>
<td>Possible synonym of <i>Rhamphocephalus bucklandi</i>
</td></tr>
<tr>
<td><i>Pterodactylus kiddii</i>
</td>
<td>Owen
</td>
<td>1874
</td>
<td><i>Nomen dubium</i>
</td>
<td>Possible synonym of <i>Rhamphocephalus bucklandi</i>
</td></tr>
<tr>
<td><i>Pterodactylus manseli</i>
</td>
<td>Owen
</td>
<td>1874
</td>
<td><i>Nomen dubium</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus marderi</i>
</td>
<td>Owen
</td>
<td>1874
</td>
<td>Synonym of <i>Dimorphodon macronyx</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus pleydelli</i>
</td>
<td>Owen
</td>
<td>1874
</td>
<td><i>Nomen dubium</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus sagittirostris</i>
</td>
<td>Owen
</td>
<td>1874
</td>
<td>Reclassified as <i><a href="/enwiki/wiki/Serradraco_sagittirostris" class="mw-redirect" title="Serradraco sagittirostris">Serradraco sagittirostris</a></i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus umbrosus</i>
</td>
<td>Cope
</td>
<td>1874
</td>
<td><i>Nomen dubium</i>
</td>
<td>Reclassified from <i><a href="/enwiki/wiki/Ornithochirus_umbrosus" class="mw-redirect" title="Ornithochirus umbrosus">Ornithochirus umbrosus</a></i>
</td></tr>
<tr>
<td><i>Pterodactylus validus</i>
</td>
<td>Owen
</td>
<td>1875
</td>
<td>Reclassified as <i><a href="/enwiki/wiki/Doratorhynchus_validus" class="mw-redirect" title="Doratorhynchus validus">Doratorhynchus validus</a></i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus montanus</i>
</td>
<td>Marsh
</td>
<td>1878
</td>
<td>Reclassified as <i><a href="/enwiki/wiki/Dermodactylus_montanus" class="mw-redirect" title="Dermodactylus montanus">Dermodactylus montanus</a></i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus suevicus</i>
</td>
<td><a href="/enwiki/wiki/Oscar_Fraas" title="Oscar Fraas">Fraas</a> <i>non</i> Quenstedt
</td>
<td>1878 <i>non</i> 1855
</td>
<td>Synonym of <i>Ardeadactylus longicollum</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus fraasi</i>
</td>
<td>(Seeley)
</td>
<td>(1901)
</td>
<td>Synonym of <i>Ardeadactylus longicollum</i>
</td>
<td>Reclassified from. <i>Cycnorhamphus fraasi</i>
</td></tr>
<tr>
<td><i>Pterodactylus cristatus</i>
</td>
<td><a href="/enwiki/wiki/Carl_Wiman" title="Carl Wiman">Wiman</a>
</td>
<td>1925
</td>
<td>Reclassified as <i><a href="/enwiki/wiki/Germanodactylus_cristatus" class="mw-redirect" title="Germanodactylus cristatus">Germanodactylus cristatus</a></i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus westmani</i>
</td>
<td>Wiman
</td>
<td>1925
</td>
<td>Synonym of <i>Pterodactylus antiquus</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus cormoranus</i>
</td>
<td>Döderlein
</td>
<td>1929
</td>
<td>Synonym of <i>Pterodactylus antiquus</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus arningi</i>
</td>
<td><a href="/enwiki/wiki/Hans_Reck" title="Hans Reck">Reck</a>
</td>
<td>1931
</td>
<td><i>Nomen dubium</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus brancai</i>
</td>
<td>Reck
</td>
<td>1931
</td>
<td><i>Nomen dubium</i>
</td>
<td>Reclassified as <i><a href="/enwiki/wiki/Dsungaripterus_brancai" class="mw-redirect" title="Dsungaripterus brancai">Dsungaripterus brancai</a></i>
</td></tr>
<tr>
<td><i>Pterodactylus maximus</i>
</td>
<td>Reck
</td>
<td>1931
</td>
<td><i>Nomen dubium</i>
</td>
<td>
</td></tr>
<tr>
<td><i>Pterodactylus raptor</i>
</td>
<td>Owen <i>vide</i> Ingles & Sawyer
</td>
<td>1979
</td>
<td><i>Nomen nudum</i>
</td>
<td>
</td></tr>
</tbody></table>
</td></tr></tbody></table></div>
<h2><span class="mw-headline" id="Cultural_significance">Cultural significance</span><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Pterodactylus&action=edit&section=13" title="Edit section: Cultural significance">edit</a><span class="mw-editsection-bracket">]</span></span></h2>
<div class="thumb tright"><div class="thumbinner" style="width:172px;"><a href="/enwiki/wiki/File:Pteranodon_longiceps_mmartyniuk_wiki.png" class="image"><img alt="" src="/upwiki/wikipedia/commons/thumb/1/1f/Pteranodon_longiceps_mmartyniuk_wiki.png/170px-Pteranodon_longiceps_mmartyniuk_wiki.png" decoding="async" width="170" height="165" class="thumbimage" srcset="/upwiki/wikipedia/commons/thumb/1/1f/Pteranodon_longiceps_mmartyniuk_wiki.png/255px-Pteranodon_longiceps_mmartyniuk_wiki.png 1.5x, /upwiki/wikipedia/commons/thumb/1/1f/Pteranodon_longiceps_mmartyniuk_wiki.png/340px-Pteranodon_longiceps_mmartyniuk_wiki.png 2x" data-file-width="500" data-file-height="484" /></a> <div class="thumbcaption"><div class="magnify"><a href="/enwiki/wiki/File:Pteranodon_longiceps_mmartyniuk_wiki.png" class="internal" title="Enlarge"></a></div>Life reconstruction of a <i>Pteranodon</i>, popularly called "pterodactyl"</div></div></div>
<p><i>Pterodactylus</i> is regarded as one of the most iconic prehistoric creatures, with multiple appearances in books, movies, as well as television series and several videogames. The informal name "<b>pterodactyl</b>" is sometimes used to refer to any kind of animal belonging to the order <a href="/enwiki/wiki/Pterosauria" class="mw-redirect" title="Pterosauria">Pterosauria</a>, though most of the times to <i>Pterodactylus</i>, as it's the most well-known member of the group.<sup id="cite_ref-myths_75-0" class="reference"><a href="#cite_note-myths-75">[75]</a></sup> The popular aspect of <i>Pterodactylus</i> consists of an elongated head crest, and potentially large wings. Studies of <i>Pterodactylus</i> however, conclude that it may even lack a bony cranial crest, though several analysis have proven that <i>Pterodactylus</i> may in fact have a crest made up of soft tissue instead of bone.<sup id="cite_ref-BennettPZ_17-11" class="reference"><a href="#cite_note-BennettPZ-17">[17]</a></sup>
</p><p><i>Pterodactylus</i> is the star character of the 2005 <a href="/enwiki/wiki/Horror_film" title="Horror film">horror film</a> <i><a href="/enwiki/wiki/Pterodactyl_(film)" title="Pterodactyl (film)">Pterodactyl</a></i>, where it is identified with the informal name "pterodactyl", hence the name of the film. In the film, the "pterodactyls" resemble the aspect of the distantly related genus <i>Pteranodon</i> due to the elongated bony cranial crest, and their enormous size. One peculiar feature that <i>Pterodactylus</i> had in the film is the possession of teeth, while this is generally accurate for <i>Pterodactylus</i>, the overall appearance of the creatures in the film is similar to that of <i>Pteranodon</i>, as well as the large size, this makes them resemble some kind of pterosaur identical to <i>Pteranodon</i>, but with the possession of teeth.<sup id="cite_ref-76" class="reference"><a href="#cite_note-76">[76]</a></sup>
</p><p>Another appearance of <i>Pterodactylus</i>-like creatures is in <a href="/enwiki/wiki/J._R._R._Tolkien" title="J. R. R. Tolkien">J. R. R. Tolkien</a>'s <a href="/enwiki/wiki/Middle-earth" title="Middle-earth">Middle-earth</a> <a href="/enwiki/wiki/Tolkien%27s_legendarium" title="Tolkien's legendarium">legendarium</a>. In this novel, the <a href="/enwiki/wiki/Nazg%C3%BBl" title="Nazgûl">Nazgûl</a>, introduced as the Black Riders, are nine characters who rode flying monsters that looked similarly built to <i>Pterodactylus</i>. <a href="/enwiki/wiki/Christopher_Tolkien" title="Christopher Tolkien">Christopher Tolkien</a>, the son of the author, described the flying monsters as "Nazgûl-birds"; his father described the appearance of the steeds as somewhat "pterodactylic", and acknowledged that these were obviously "new mythology".<sup id="cite_ref-Letter_211_77-0" class="reference"><a href="#cite_note-Letter_211-77">[77]</a></sup><sup id="cite_ref-T_78-0" class="reference"><a href="#cite_note-T-78">[78]</a></sup>
</p>
<h2><span class="mw-headline" id="See_also">See also</span><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Pterodactylus&action=edit&section=14" title="Edit section: See also">edit</a><span class="mw-editsection-bracket">]</span></span></h2>
<ul><li><a href="/enwiki/wiki/List_of_pterosaur_genera" title="List of pterosaur genera">List of pterosaur genera</a></li>
<li><a href="/enwiki/wiki/Timeline_of_pterosaur_research" title="Timeline of pterosaur research">Timeline of pterosaur research</a></li></ul>
<h2><span class="mw-headline" id="References">References</span><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Pterodactylus&action=edit&section=15" title="Edit section: References">edit</a><span class="mw-editsection-bracket">]</span></span></h2>
<style data-mw-deduplicate="TemplateStyles:r1011085734">.mw-parser-output .reflist{font-size:90%;margin-bottom:0.5em;list-style-type:decimal}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width" style="column-width: 35em;">
<ol class="references">
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<li id="cite_note-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-2">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFGudger1944" class="citation journal cs1">Gudger, E.W. (1944). "The Earliest Winged Fish-Catchers". <i>The Scientific Monthly</i>. <b>59</b> (2): 120–129. <a href="/enwiki/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1944SciMo..59..120G">1944SciMo..59..120G</a>. <a href="/enwiki/wiki/JSTOR_(identifier)" class="mw-redirect" title="JSTOR (identifier)">JSTOR</a> <a rel="nofollow" class="external text" href="/enwiki//www.jstor.org/stable/18398">18398</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+Scientific+Monthly&rft.atitle=The+Earliest+Winged+Fish-Catchers&rft.volume=59&rft.issue=2&rft.pages=120-129&rft.date=1944&rft_id=%2F%2Fwww.jstor.org%2Fstable%2F18398%23id-name%3DJSTOR&rft_id=info%3Abibcode%2F1944SciMo..59..120G&rft.aulast=Gudger&rft.aufirst=E.W.&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-ammonitedate-3"><span class="mw-cite-backlink">^ <a href="#cite_ref-ammonitedate_3-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-ammonitedate_3-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFSchweigert,_G.2007" class="citation journal cs1">Schweigert, G. (2007). "Ammonite biostratigraphy as a tool for dating Upper Jurassic lithographic limestones from South Germany – first results and open questions". <i>Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen</i>. <b>245</b> (1): 117–125. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1127%2F0077-7749%2F2007%2F0245-0117">10.1127/0077-7749/2007/0245-0117</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Neues+Jahrbuch+f%C3%BCr+Geologie+und+Pal%C3%A4ontologie%2C+Abhandlungen&rft.atitle=Ammonite+biostratigraphy+as+a+tool+for+dating+Upper+Jurassic+lithographic+limestones+from+South+Germany+%E2%80%93+first+results+and+open+questions&rft.volume=245&rft.issue=1&rft.pages=117-125&rft.date=2007&rft_id=info%3Adoi%2F10.1127%2F0077-7749%2F2007%2F0245-0117&rft.au=Schweigert%2C+G.&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-unwin2003-4"><span class="mw-cite-backlink">^ <a href="#cite_ref-unwin2003_4-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-unwin2003_4-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-unwin2003_4-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-unwin2003_4-3"><sup><i><b>d</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFUnwin2003" class="citation journal cs1">Unwin, D. M. (2003). "On the phylogeny and evolutionary history of pterosaurs". <i>Geological Society, London, Special Publications</i>. <b>217</b> (1): 139–190. <a href="/enwiki/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2003GSLSP.217..139U">2003GSLSP.217..139U</a>. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1144%2FGSL.SP.2003.217.01.11">10.1144/GSL.SP.2003.217.01.11</a>. <a href="/enwiki/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:86710955">86710955</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Geological+Society%2C+London%2C+Special+Publications&rft.atitle=On+the+phylogeny+and+evolutionary+history+of+pterosaurs&rft.volume=217&rft.issue=1&rft.pages=139-190&rft.date=2003&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A86710955%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.1144%2FGSL.SP.2003.217.01.11&rft_id=info%3Abibcode%2F2003GSLSP.217..139U&rft.aulast=Unwin&rft.aufirst=D.+M.&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-longrichetal2018-5"><span class="mw-cite-backlink">^ <a href="#cite_ref-longrichetal2018_5-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-longrichetal2018_5-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFLongrichMartillAndres2018" class="citation journal cs1">Longrich, N.R.; Martill, D.M.; Andres, B. (2018). <a rel="nofollow" class="external text" href="/enwiki//www.ncbi.nlm.nih.gov/pmc/articles/PMC5849296">"Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary"</a>. <i>PLOS Biology</i>. <b>16</b> (3): e2001663. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1371%2Fjournal.pbio.2001663">10.1371/journal.pbio.2001663</a>. <a href="/enwiki/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="cs1-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="/enwiki//www.ncbi.nlm.nih.gov/pmc/articles/PMC5849296">5849296</a></span>. <a href="/enwiki/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="/enwiki//pubmed.ncbi.nlm.nih.gov/29534059">29534059</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=PLOS+Biology&rft.atitle=Late+Maastrichtian+pterosaurs+from+North+Africa+and+mass+extinction+of+Pterosauria+at+the+Cretaceous-Paleogene+boundary&rft.volume=16&rft.issue=3&rft.pages=e2001663&rft.date=2018&rft_id=%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC5849296%23id-name%3DPMC&rft_id=info%3Apmid%2F29534059&rft_id=info%3Adoi%2F10.1371%2Fjournal.pbio.2001663&rft.aulast=Longrich&rft.aufirst=N.R.&rft.au=Martill%2C+D.M.&rft.au=Andres%2C+B.&rft_id=%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC5849296&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
</li>
<li id="cite_note-pester-6"><span class="mw-cite-backlink">^ <a href="#cite_ref-pester_6-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-pester_6-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-pester_6-2"><sup><i><b>c</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFŐsiProndvaiGéczy2010" class="citation journal cs1">Ősi, A.; Prondvai, E.; Géczy, B. (2010). <a rel="nofollow" class="external text" href="https://www.researchgate.net/publication/258391479">"The history of Late Jurassic pterosaurs housed in Hungarian collections and the revision of the holotype of <i>Pterodactylus micronyx</i> Meyer 1856 (a 'Pester Exemplar')"</a>. <i>Geological Society, London, Special Publications</i>. <b>343</b> (1): 277–286. <a href="/enwiki/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2010GSLSP.343..277O">2010GSLSP.343..277O</a>. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1144%2FSP343.17">10.1144/SP343.17</a>. <a href="/enwiki/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:129805068">129805068</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Geological+Society%2C+London%2C+Special+Publications&rft.atitle=The+history+of+Late+Jurassic+pterosaurs+housed+in+Hungarian+collections+and+the+revision+of+the+holotype+of+Pterodactylus+micronyx+Meyer+1856+%28a+%27Pester+Exemplar%27%29&rft.volume=343&rft.issue=1&rft.pages=277-286&rft.date=2010&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A129805068%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.1144%2FSP343.17&rft_id=info%3Abibcode%2F2010GSLSP.343..277O&rft.aulast=%C5%90si&rft.aufirst=A.&rft.au=Prondvai%2C+E.&rft.au=G%C3%A9czy%2C+B.&rft_id=https%3A%2F%2Fwww.researchgate.net%2Fpublication%2F258391479&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-DU06-7"><span class="mw-cite-backlink">^ <a href="#cite_ref-DU06_7-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-DU06_7-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFUnwin2006" class="citation book cs1">Unwin, David M. (2006). <i>The terosaurs: From Deep Time</i>. New York: Pi Press. p. 246. <a href="/enwiki/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/enwiki/wiki/Special:BookSources/0-13-146308-X" title="Special:BookSources/0-13-146308-X"><bdi>0-13-146308-X</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+terosaurs%3A+From+Deep+Time&rft.place=New+York&rft.pages=246&rft.pub=Pi+Press&rft.date=2006&rft.isbn=0-13-146308-X&rft.aulast=Unwin&rft.aufirst=David+M.&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-brougham1844-8"><span class="mw-cite-backlink"><b><a href="#cite_ref-brougham1844_8-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFBrougham1844" class="citation journal cs1">Brougham, Henry P. (1844). <a rel="nofollow" class="external text" href="https://www.biodiversitylibrary.org/item/229475#page/5/mode/1up">"Dialogues on instinct; with analytical view of the researches on fossil osteology"</a>. <i>Knight's Weekly Volume for All Readers</i>. <b>19</b>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Knight%27s+Weekly+Volume+for+All+Readers&rft.atitle=Dialogues+on+instinct%3B+with+analytical+view+of+the+researches+on+fossil+osteology&rft.volume=19&rft.date=1844&rft.aulast=Brougham&rft.aufirst=Henry+P.&rft_id=https%3A%2F%2Fwww.biodiversitylibrary.org%2Fitem%2F229475%23page%2F5%2Fmode%2F1up&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
</li>
<li id="cite_note-collini1784-9"><span class="mw-cite-backlink"><b><a href="#cite_ref-collini1784_9-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFCollini1784" class="citation journal cs1">Collini, C A. (1784). <a rel="nofollow" class="external text" href="http://www.rhinoresourcecenter.com/index.php?s=1&act=pdfviewer&id=1292315455&folder=129">"Sur quelques Zoolithes du Cabinet d'Histoire naturelle de S. A. S. E. Palatine & de Bavière, à Mannheim"</a>. <i>Acta Theodoro-Palatinae Mannheim</i> (in French). 5 Physicum: 58–103 (1 plate).</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Acta+Theodoro-Palatinae+Mannheim&rft.atitle=Sur+quelques+Zoolithes+du+Cabinet+d%27Histoire+naturelle+de+S.+A.+S.+E.+Palatine+%26+de+Bavi%C3%A8re%2C+%C3%A0+Mannheim&rft.volume=5+Physicum&rft.pages=58-103+%281+plate%29&rft.date=1784&rft.aulast=Collini&rft.aufirst=C+A.&rft_id=http%3A%2F%2Fwww.rhinoresourcecenter.com%2Findex.php%3Fs%3D1%26act%3Dpdfviewer%26id%3D1292315455%26folder%3D129&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-taquet&padian2004-10"><span class="mw-cite-backlink">^ <a href="#cite_ref-taquet&padian2004_10-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-taquet&padian2004_10-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-taquet&padian2004_10-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-taquet&padian2004_10-3"><sup><i><b>d</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFTaquetPadian2004" class="citation journal cs1">Taquet, P.; Padian, K. (2004). "The earliest known restoration of a pterosaur and the philosophical origins of Cuvier's <i>Ossemens Fossiles</i>". <i>Comptes Rendus Palevol</i>. <b>3</b> (2): 157–175. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.crpv.2004.02.002">10.1016/j.crpv.2004.02.002</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Comptes+Rendus+Palevol&rft.atitle=The+earliest+known+restoration+of+a+pterosaur+and+the+philosophical+origins+of+Cuvier%27s+Ossemens+Fossiles&rft.volume=3&rft.issue=2&rft.pages=157-175&rft.date=2004&rft_id=info%3Adoi%2F10.1016%2Fj.crpv.2004.02.002&rft.aulast=Taquet&rft.aufirst=P.&rft.au=Padian%2C+K.&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
</li>
<li id="cite_note-11"><span class="mw-cite-backlink"><b><a href="#cite_ref-11">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFWagler1830" class="citation book cs1">Wagler, Johann Georg (1830). <a rel="nofollow" class="external text" href="https://www.biodiversitylibrary.org/item/120243"><i>Natürliches System der Amphibien : mit vorangehender Classification der Säugethiere und Vögel : ein Beitrag zur vergleichenden Zoologie</i></a> (in German). München.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Nat%C3%BCrliches+System+der+Amphibien+%3A+mit+vorangehender+Classification+der+S%C3%A4ugethiere+und+V%C3%B6gel+%3A+ein+Beitrag+zur+vergleichenden+Zoologie&rft.place=M%C3%BCnchen&rft.date=1830&rft.aulast=Wagler&rft.aufirst=Johann+Georg&rft_id=https%3A%2F%2Fwww.biodiversitylibrary.org%2Fitem%2F120243&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
</li>
<li id="cite_note-cuvier1801-12"><span class="mw-cite-backlink"><b><a href="#cite_ref-cuvier1801_12-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFCuvier1801" class="citation journal cs1">Cuvier, G. (1801). <a rel="nofollow" class="external text" href="https://gallica.bnf.fr/ark:/12148/bpt6k9604723x/f1.image">"Extrait d'un ouvrage sur les espèces de quadrupèdes dont on a trouvé les ossemens dans l'intérieur de la terre"</a>. <i>Journal de Physique, de Chimie et d'Histoire Naturelle</i> (in French). <b>52</b>: 253–267. <q>Reptile volant</q></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+de+Physique%2C+de+Chimie+et+d%27Histoire+Naturelle&rft.atitle=Extrait+d%27un+ouvrage+sur+les+esp%C3%A8ces+de+quadrup%C3%A8des+dont+on+a+trouv%C3%A9+les+ossemens+dans+l%27int%C3%A9rieur+de+la+terre&rft.volume=52&rft.pages=253-267&rft.date=1801&rft.aulast=Cuvier&rft.aufirst=G.&rft_id=https%3A%2F%2Fgallica.bnf.fr%2Fark%3A%2F12148%2Fbpt6k9604723x%2Ff1.image&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-cuvier1809-13"><span class="mw-cite-backlink">^ <a href="#cite_ref-cuvier1809_13-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-cuvier1809_13-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFCuvier1809" class="citation journal cs1">Cuvier, G. (1809). <a rel="nofollow" class="external text" href="https://archive.org/details/cbarchive_48783_cuvierg1809mmoiresurlesquelett1809/mode/2up">"Mémoire sur le squelette fossile d'un reptile volant des environs d'Aichstedt, que quelques naturalistes ont pris pour un oiseau, et dont nous formons un genre de Sauriens, sous le nom de Petro-Dactyle"</a>. <i>Annales du Muséum national d'Histoire Naturelle, Paris</i>. <b>13</b>: 424–437.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Annales+du+Mus%C3%A9um+national+d%27Histoire+Naturelle%2C+Paris&rft.atitle=M%C3%A9moire+sur+le+squelette+fossile+d%27un+reptile+volant+des+environs+d%27Aichstedt%2C+que+quelques+naturalistes+ont+pris+pour+un+oiseau%2C+et+dont+nous+formons+un+genre+de+Sauriens%2C+sous+le+nom+de+Petro-Dactyle&rft.volume=13&rft.pages=424-437&rft.date=1809&rft.aulast=Cuvier&rft.aufirst=G.&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fcbarchive_48783_cuvierg1809mmoiresurlesquelett1809%2Fmode%2F2up&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
</li>
<li id="cite_note-14"><span class="mw-cite-backlink"><b><a href="#cite_ref-14">^</a></b></span> <span class="reference-text">Cuvier, G. (1809), p. 436 : "II n'est guère possible de douter que ce long doigt n'ait servi à supporter une membrane qui formoit <i>[sic]</i> à l'animal, d'après la longueur de l'extrémité antérieure, une aile bien plus puissante que celle du dragon, et au moins égale en force à celle de la chauve-souris."</span>
</li>
<li id="cite_note-sommerring1812-15"><span class="mw-cite-backlink"><b><a href="#cite_ref-sommerring1812_15-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFvon_Sömmerring1812" class="citation journal cs1">von Sömmerring, S. T. (1812). <a rel="nofollow" class="external text" href="http://publikationen.ub.uni-frankfurt.de/frontdoor/index/index/year/2008/docId/15367">"Über einen <i>Ornithocephalus</i> 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: vorgelesen in der mathematisch-physikalischen Classe am 27. Dec. 1810 und Nachtrag vorgelesen am 8. April 1811"</a>. <i>Denkschriften der Königlichen Bayerischen Akademie der Wissenschaften</i>. München. <b>3</b>: 89–158.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Denkschriften+der+K%C3%B6niglichen+Bayerischen+Akademie+der+Wissenschaften&rft.atitle=%C3%9Cber+einen+Ornithocephalus+oder+%C3%BCber+das+unbekannten+Thier+der+Vorwelt%2C+dessen+Fossiles+Gerippe+Collini+im+5.+Bande+der+Actorum+Academiae+Theodoro-Palatinae+nebst+einer+Abbildung+in+nat%C3%BCrlicher+Gr%C3%B6sse+im+Jahre+1784+beschrieb%2C+und+welches+Gerippe+sich+gegenw%C3%A4rtig+in+der+Naturalien-Sammlung+der+k%C3%B6niglichen+Akademie+der+Wissenschaften+zu+M%C3%BCnchen+befindet%3A+vorgelesen+in+der+mathematisch-physikalischen+Classe+am+27.+Dec.+1810+und+Nachtrag+vorgelesen+am+8.+April+1811&rft.volume=3&rft.pages=89-158&rft.date=1812&rft.aulast=von+S%C3%B6mmerring&rft.aufirst=S.+T.&rft_id=http%3A%2F%2Fpublikationen.ub.uni-frankfurt.de%2Ffrontdoor%2Findex%2Findex%2Fyear%2F2008%2FdocId%2F15367&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
</li>
<li id="cite_note-16"><span class="mw-cite-backlink"><b><a href="#cite_ref-16">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFCuvier1812" class="citation book cs1">Cuvier, G. (1812). "Article V – Sur le squelette fossile d'un reptile volant des environs d'Aichstedt, que quelques naturalistes ont pris pour un oiseau et dont nous formons un genre de sauriens, sous le nom de ptéro-dactyle". <a rel="nofollow" class="external text" href="https://www.biodiversitylibrary.org/item/123716#page/660/mode/1up"><i>Recherches sur les ossemens fossiles de quadrupèdes : où l'on rétablit les caractères de plusieurs espèces d'animaux que les révolutions du globe paroissent avoir détruites</i></a> (in French). Tome 4. Paris: Deterville. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.5962%2Fbhl.title.60807">10.5962/bhl.title.60807</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Article+V+%E2%80%93+Sur+le+squelette+fossile+d%E2%80%99un+reptile+volant+des+environs+d%27Aichstedt%2C+que+quelques+naturalistes+ont+pris+pour+un+oiseau+et+dont+nous+formons+un+genre+de+sauriens%2C+sous+le+nom+de+pt%C3%A9ro-dactyle&rft.btitle=Recherches+sur+les+ossemens+fossiles+de+quadrup%C3%A8des+%3A+o%C3%B9+l%27on+r%C3%A9tablit+les+caract%C3%A8res+de+plusieurs+esp%C3%A8ces+d%27animaux+que+les+r%C3%A9volutions+du+globe+paroissent+avoir+d%C3%A9truites&rft.place=Paris&rft.pub=Deterville&rft.date=1812&rft_id=info%3Adoi%2F10.5962%2Fbhl.title.60807&rft.aulast=Cuvier&rft.aufirst=G.&rft_id=https%3A%2F%2Fwww.biodiversitylibrary.org%2Fitem%2F123716%23page%2F660%2Fmode%2F1up&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-BennettPZ-17"><span class="mw-cite-backlink">^ <a href="#cite_ref-BennettPZ_17-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-BennettPZ_17-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-BennettPZ_17-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-BennettPZ_17-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-BennettPZ_17-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-BennettPZ_17-5"><sup><i><b>f</b></i></sup></a> <a href="#cite_ref-BennettPZ_17-6"><sup><i><b>g</b></i></sup></a> <a href="#cite_ref-BennettPZ_17-7"><sup><i><b>h</b></i></sup></a> <a href="#cite_ref-BennettPZ_17-8"><sup><i><b>i</b></i></sup></a> <a href="#cite_ref-BennettPZ_17-9"><sup><i><b>j</b></i></sup></a> <a href="#cite_ref-BennettPZ_17-10"><sup><i><b>k</b></i></sup></a> <a href="#cite_ref-BennettPZ_17-11"><sup><i><b>l</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFBennett,_S._Christopher2013" class="citation journal cs1">Bennett, S. Christopher (2013). "New information on body size and cranial display structures of <i>Pterodactylus antiquus</i>, with a revision of the genus". <i>Paläontologische Zeitschrift</i>. <b>87</b> (2): 269–289. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2Fs12542-012-0159-8">10.1007/s12542-012-0159-8</a>. <a href="/enwiki/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:83722829">83722829</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Pal%C3%A4ontologische+Zeitschrift&rft.atitle=New+information+on+body+size+and+cranial+display+structures+of+Pterodactylus+antiquus%2C+with+a+revision+of+the+genus&rft.volume=87&rft.issue=2&rft.pages=269-289&rft.date=2013&rft_id=info%3Adoi%2F10.1007%2Fs12542-012-0159-8&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A83722829%23id-name%3DS2CID&rft.au=Bennett%2C+S.+Christopher&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-18"><span class="mw-cite-backlink"><b><a href="#cite_ref-18">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFvon_Sömmerring1817" class="citation journal cs1">von Sömmerring, S.T. (1817). <a rel="nofollow" class="external text" href="http://publikationen.ub.uni-frankfurt.de/frontdoor/index/index/year/2008/docId/15330">"Ueber einen Ornithocephalus brevirostris der Vorwelt"</a>. <i>Denkschriften der Bayerischen Akademie der Wissenschaften</i> (in German). <b>6</b>: 89–104.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Denkschriften+der+Bayerischen+Akademie+der+Wissenschaften&rft.atitle=Ueber+einen+Ornithocephalus+brevirostris+der+Vorwelt&rft.volume=6&rft.pages=89-104&rft.date=1817&rft.aulast=von+S%C3%B6mmerring&rft.aufirst=S.T.&rft_id=http%3A%2F%2Fpublikationen.ub.uni-frankfurt.de%2Ffrontdoor%2Findex%2Findex%2Fyear%2F2008%2FdocId%2F15330&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-19"><span class="mw-cite-backlink"><b><a href="#cite_ref-19">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFPadian1987" class="citation book cs1">Padian, K. (1987). "The case of the bat-winged pterosaur. Typological taxonomy and the influence of pictorial representation on scientific perception". In Czerkas, S. J.; Olson, E. C. (eds.). <i>Dinosaurs past and present</i>. <b>2</b>. Los Angeles: Natural History Museum of Los Angeles County in association with University of Washington Press, Seattle and London. pp. 65–81. <a href="/enwiki/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/enwiki/wiki/Special:BookSources/978-0-938644-23-1" title="Special:BookSources/978-0-938644-23-1"><bdi>978-0-938644-23-1</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=The+case+of+the+bat-winged+pterosaur.+Typological+taxonomy+and+the+influence+of+pictorial+representation+on+scientific+perception&rft.btitle=Dinosaurs+past+and+present&rft.place=Los+Angeles&rft.pages=65-81&rft.pub=Natural+History+Museum+of+Los+Angeles+County+in+association+with+University+of+Washington+Press%2C+Seattle+and+London&rft.date=1987&rft.isbn=978-0-938644-23-1&rft.aulast=Padian&rft.aufirst=K.&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-20"><span class="mw-cite-backlink"><b><a href="#cite_ref-20">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFRafinesque1815" class="citation book cs1"><a href="/enwiki/wiki/Constantine_Samuel_Rafinesque" title="Constantine Samuel Rafinesque">Rafinesque, C.S.</a> (1815). <a rel="nofollow" class="external text" href="https://www.biodiversitylibrary.org/page/48310144#page/10/mode/1up"><i>Analyse de la nature, ou tableau de l'univers et des corps organisés</i></a> (L'Imprimerie de Jean Barravecchia ed.). p. 224.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Analyse+de+la+nature%2C+ou+tableau+de+l%27univers+et+des+corps+organis%C3%A9s&rft.pages=224&rft.edition=L%27Imprimerie+de+Jean+Barravecchia&rft.date=1815&rft.aulast=Rafinesque&rft.aufirst=C.S.&rft_id=https%3A%2F%2Fwww.biodiversitylibrary.org%2Fpage%2F48310144%23page%2F10%2Fmode%2F1up&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-:0-21"><span class="mw-cite-backlink">^ <a href="#cite_ref-:0_21-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:0_21-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFCuvier1819" class="citation book cs1">Cuvier, G. (1819). "Pterodactylus longirostris". In Oken, Lorenz (ed.). <a rel="nofollow" class="external text" href="http://archive.org/details/isisoderencyclop1819oken"><i>Isis (oder Encyclopädische Zeitung) von Oken</i></a> (in German). Jena : Expedition der Isis. pp. <a rel="nofollow" class="external text" href="https://archive.org/details/isisoderencyclop1819oken/page/n586/mode/1up">1126</a>, <a rel="nofollow" class="external text" href="https://archive.org/details/isisoderencyclop1819oken/page/n914/mode/1up">1788</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Pterodactylus+longirostris&rft.btitle=Isis+%28oder+Encyclop%C3%A4dische+Zeitung%29+von+Oken&rft.pages=1126%2C+1788&rft.pub=Jena+%3A+Expedition+der+Isis&rft.date=1819&rft.aulast=Cuvier&rft.aufirst=G.&rft_id=http%3A%2F%2Farchive.org%2Fdetails%2Fisisoderencyclop1819oken&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-Lydekker-22"><span class="mw-cite-backlink">^ <a href="#cite_ref-Lydekker_22-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Lydekker_22-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Lydekker_22-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Lydekker_22-3"><sup><i><b>d</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFLydekker1888" class="citation book cs1">Lydekker, Richard (1888). <a rel="nofollow" class="external text" href="https://www.biodiversitylibrary.org/item/125711#page/1/mode/1up"><i>Catalogue of Fossil Reptilia and Amphibia in the British Museum (Natural History). Part I. Containing the Orders Ornithosauria, Crocodilia, Dinosauria, Squamata, Rhynchocephalia and Pterosauria</i></a>. Taylor and Francis. pp. 2–37.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Catalogue+of+Fossil+Reptilia+and+Amphibia+in+the+British+Museum+%28Natural+History%29.+Part+I.+Containing+the+Orders+Ornithosauria%2C+Crocodilia%2C+Dinosauria%2C+Squamata%2C+Rhynchocephalia+and+Pterosauria&rft.pages=2-37&rft.pub=Taylor+and+Francis&rft.date=1888&rft.aulast=Lydekker&rft.aufirst=Richard&rft_id=https%3A%2F%2Fwww.biodiversitylibrary.org%2Fitem%2F125711%23page%2F1%2Fmode%2F1up&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-bennett1996a-23"><span class="mw-cite-backlink">^ <a href="#cite_ref-bennett1996a_23-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-bennett1996a_23-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-bennett1996a_23-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-bennett1996a_23-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-bennett1996a_23-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-bennett1996a_23-5"><sup><i><b>f</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFBennett,_S.C.1996" class="citation journal cs1">Bennett, S.C. (1996). "Year-classes of pterosaurs from the Solnhofen Limestone of Germany: Taxonomic and Systematic Implications". <i>Journal of Vertebrate Paleontology</i>. <b>16</b> (3): 432–444. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1080%2F02724634.1996.10011332">10.1080/02724634.1996.10011332</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Vertebrate+Paleontology&rft.atitle=Year-classes+of+pterosaurs+from+the+Solnhofen+Limestone+of+Germany%3A+Taxonomic+and+Systematic+Implications&rft.volume=16&rft.issue=3&rft.pages=432-444&rft.date=1996&rft_id=info%3Adoi%2F10.1080%2F02724634.1996.10011332&rft.au=Bennett%2C+S.C.&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-bennett2002-24"><span class="mw-cite-backlink">^ <a href="#cite_ref-bennett2002_24-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-bennett2002_24-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFBennett,_S.C.2002" class="citation journal cs1">Bennett, S.C. (2002). "Soft tissue preservation of the cranial crest of the pterosaur <i>Germanodactylus</i> from Solnhofen". <i>Journal of Vertebrate Paleontology</i>. <b>22</b> (1): 43–48. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1671%2F0272-4634%282002%29022%5B0043%3ASTPOTC%5D2.0.CO%3B2">10.1671/0272-4634(2002)022[0043:STPOTC]2.0.CO;2</a>. <a href="/enwiki/wiki/JSTOR_(identifier)" class="mw-redirect" title="JSTOR (identifier)">JSTOR</a> <a rel="nofollow" class="external text" href="/enwiki//www.jstor.org/stable/4524192">4524192</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Vertebrate+Paleontology&rft.atitle=Soft+tissue+preservation+of+the+cranial+crest+of+the+pterosaur+Germanodactylus+from+Solnhofen&rft.volume=22&rft.issue=1&rft.pages=43-48&rft.date=2002&rft_id=info%3Adoi%2F10.1671%2F0272-4634%282002%29022%5B0043%3ASTPOTC%5D2.0.CO%3B2&rft_id=%2F%2Fwww.jstor.org%2Fstable%2F4524192%23id-name%3DJSTOR&rft.au=Bennett%2C+S.C.&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-Wellnhofer1970-27"><span class="mw-cite-backlink"><b><a href="#cite_ref-Wellnhofer1970_27-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFWellnhofer1970" class="citation book cs1">Wellnhofer, Peter (1970). <a rel="nofollow" class="external text" href="https://www.zobodat.at/pdf/Abhandlungen-Akademie-Bayern_NF_141_0001-0133.pdf"><i>Die Pterodactyloidea (Pterosauria) der Oberjura-Plattenkalke Süddeutschlands</i></a> <span class="cs1-format">(PDF)</span>. <b>141</b>. Bayerische Akademie der Wissenschaften, Mathematisch-Wissenschaftlichen Klasse, Abhandlungen. p. 133.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Die+Pterodactyloidea+%28Pterosauria%29+der+Oberjura-Plattenkalke+S%C3%BCddeutschlands&rft.pages=133&rft.pub=Bayerische+Akademie+der+Wissenschaften%2C+Mathematisch-Wissenschaftlichen+Klasse%2C+Abhandlungen&rft.date=1970&rft.aulast=Wellnhofer&rft.aufirst=Peter&rft_id=https%3A%2F%2Fwww.zobodat.at%2Fpdf%2FAbhandlungen-Akademie-Bayern_NF_141_0001-0133.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-W1837-32"><span class="mw-cite-backlink">^ <a href="#cite_ref-W1837_32-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-W1837_32-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-W1837_32-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-W1837_32-3"><sup><i><b>d</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFVidovicMartill2017" class="citation journal cs1">Vidovic, Steven U.; Martill, David M. (2017). <a rel="nofollow" class="external text" href="https://eprints.soton.ac.uk/423063/1/Vidovic_Martill_2017_Taxonomy_of_Diopecephalus_and_Germanodactylus_AM_with_Figures.pdf">"The taxonomy and phylogeny of <i>Diopecephalus kochi</i> (Wagner, 1837) and <i>"Germanodactylus rhamphastinus"</i> (Wagner, 1851)"</a> <span class="cs1-format">(PDF)</span>. <i>Geological Society, London, Special Publications</i>. <b>455</b> (1): 125–147. <a href="/enwiki/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2018GSLSP.455..125V">2018GSLSP.455..125V</a>. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1144%2FSP455.12">10.1144/SP455.12</a>. <a href="/enwiki/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:219204038">219204038</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Geological+Society%2C+London%2C+Special+Publications&rft.atitle=The+taxonomy+and+phylogeny+of+Diopecephalus+kochi+%28Wagner%2C+1837%29+and+%22Germanodactylus+rhamphastinus%22+%28Wagner%2C+1851%29&rft.volume=455&rft.issue=1&rft.pages=125-147&rft.date=2017&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A219204038%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.1144%2FSP455.12&rft_id=info%3Abibcode%2F2018GSLSP.455..125V&rft.aulast=Vidovic&rft.aufirst=Steven+U.&rft.au=Martill%2C+David+M.&rft_id=https%3A%2F%2Feprints.soton.ac.uk%2F423063%2F1%2FVidovic_Martill_2017_Taxonomy_of_Diopecephalus_and_Germanodactylus_AM_with_Figures.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-38"><span class="mw-cite-backlink"><b><a href="#cite_ref-38">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFYoungBrusattede_AndradeDesojo2012" class="citation journal cs1">Young, Mark T.; Brusatte, Stephen L.; de Andrade, Marco Brandalise; Desojo, Julia B.; Beatty, Brian L.; Steel, Lorna; Fernández, Marta S.; Sakamoto, Manabu; Ruiz-Omeñaca, Jose Ignacio; Schoch, Rainer R. (September 18, 2012). Butler, Richard J. (ed.). <a rel="nofollow" class="external text" href="/enwiki//www.ncbi.nlm.nih.gov/pmc/articles/PMC3445579">"The Cranial Osteology and Feeding Ecology of the Metriorhynchid Crocodylomorph Genera Dakosaurus and Plesiosuchus from the Late Jurassic of Europe"</a>. <i>PLOS ONE</i>. <b>7</b> (9): e44985. <a href="/enwiki/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2012PLoSO...744985Y">2012PLoSO...744985Y</a>. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1371%2Fjournal.pone.0044985">10.1371/journal.pone.0044985</a>. <a href="/enwiki/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="/enwiki//www.worldcat.org/issn/1932-6203">1932-6203</a>. <a href="/enwiki/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="cs1-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="/enwiki//www.ncbi.nlm.nih.gov/pmc/articles/PMC3445579">3445579</a></span>. <a href="/enwiki/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="/enwiki//pubmed.ncbi.nlm.nih.gov/23028723">23028723</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=PLOS+ONE&rft.atitle=The+Cranial+Osteology+and+Feeding+Ecology+of+the+Metriorhynchid+Crocodylomorph+Genera+Dakosaurus+and+Plesiosuchus+from+the+Late+Jurassic+of+Europe&rft.volume=7&rft.issue=9&rft.pages=e44985&rft.date=2012-09-18&rft_id=%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3445579%23id-name%3DPMC&rft_id=info%3Abibcode%2F2012PLoSO...744985Y&rft_id=info%3Apmid%2F23028723&rft_id=info%3Adoi%2F10.1371%2Fjournal.pone.0044985&rft.issn=1932-6203&rft.aulast=Young&rft.aufirst=Mark+T.&rft.au=Brusatte%2C+Stephen+L.&rft.au=de+Andrade%2C+Marco+Brandalise&rft.au=Desojo%2C+Julia+B.&rft.au=Beatty%2C+Brian+L.&rft.au=Steel%2C+Lorna&rft.au=Fern%C3%A1ndez%2C+Marta+S.&rft.au=Sakamoto%2C+Manabu&rft.au=Ruiz-Ome%C3%B1aca%2C+Jose+Ignacio&rft.au=Schoch%2C+Rainer+R.&rft_id=%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3445579&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-lamberstable-others-41"><span class="mw-cite-backlink">^ <a href="#cite_ref-lamberstable-others_41-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-lamberstable-others_41-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFLambers1999" class="citation journal cs1">Lambers, Paul H. (1999). "The actinopterygian fish fauna of the Late Kimmeridgian and Early Tithonian 'Plattenkalke' near Solnhofen (Bavaria, Germany): state of the art". <i>Geologie en Mijnbouw</i>. <b>78</b> (2): 215–229. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1023%2FA%3A1003855831015">10.1023/A:1003855831015</a>. <a href="/enwiki/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:127676896">127676896</a>. <q>Table 1: List of actinopterygians from the Solnhofen lithographic limestone – Halecomorphi; 'Amiidae' and 'Ophiopsidae'" (p. 216)</q></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Geologie+en+Mijnbouw&rft.atitle=The+actinopterygian+fish+fauna+of+the+Late+Kimmeridgian+and+Early+Tithonian+%27Plattenkalke%27+near+Solnhofen+%28Bavaria%2C+Germany%29%3A+state+of+the+art&rft.volume=78&rft.issue=2&rft.pages=215-229&rft.date=1999&rft_id=info%3Adoi%2F10.1023%2FA%3A1003855831015&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A127676896%23id-name%3DS2CID&rft.aulast=Lambers&rft.aufirst=Paul+H.&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-Marine-42"><span class="mw-cite-backlink">^ <a href="#cite_ref-Marine_42-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Marine_42-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFSepkoski2002" class="citation journal cs1">Sepkoski, Jack (2002). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20110723131237/http://strata.ummp.lsa.umich.edu/jack/showgenera.php?taxon=611&rank=class">"A compendium of fossil marine animal genera"</a>. <i>Bulletins of American Paleontology</i>. <b>364</b>: 560. Archived from <a rel="nofollow" class="external text" href="http://strata.ummp.lsa.umich.edu/jack/showgenera.php?taxon=611&rank=class">the original</a> on July 23, 2011<span class="reference-accessdate">. Retrieved <span class="nowrap">February 27,</span> 2009</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Bulletins+of+American+Paleontology&rft.atitle=A+compendium+of+fossil+marine+animal+genera&rft.volume=364&rft.pages=560&rft.date=2002&rft.aulast=Sepkoski&rft.aufirst=Jack&rft_id=http%3A%2F%2Fstrata.ummp.lsa.umich.edu%2Fjack%2Fshowgenera.php%3Ftaxon%3D611%26rank%3Dclass&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-43"><span class="mw-cite-backlink"><b><a href="#cite_ref-43">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFLaneEbert2015" class="citation journal cs1">Lane, Jennifer A.; Ebert, Martin (2015). "A taxonomic reassessment of Ophiopsis (Halecomorphi, Ionoscopiformes), with a revision of Upper Jurassic species from the Solnhofen Archipelago, and a new genus of Ophiopsidae". <i>Journal of Vertebrate Paleontology</i>. <b>35</b> (1): e883238. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1080%2F02724634.2014.883238">10.1080/02724634.2014.883238</a>. <a href="/enwiki/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="/enwiki//www.worldcat.org/issn/0272-4634">0272-4634</a>. <a href="/enwiki/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:86350086">86350086</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Vertebrate+Paleontology&rft.atitle=A+taxonomic+reassessment+of+Ophiopsis+%28Halecomorphi%2C+Ionoscopiformes%29%2C+with+a+revision+of+Upper+Jurassic+species+from+the+Solnhofen+Archipelago%2C+and+a+new+genus+of+Ophiopsidae&rft.volume=35&rft.issue=1&rft.pages=e883238&rft.date=2015&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A86350086%23id-name%3DS2CID&rft.issn=0272-4634&rft_id=info%3Adoi%2F10.1080%2F02724634.2014.883238&rft.aulast=Lane&rft.aufirst=Jennifer+A.&rft.au=Ebert%2C+Martin&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-EoDP-44"><span class="mw-cite-backlink"><b><a href="#cite_ref-EoDP_44-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFPalmer,_D.1999" class="citation book cs1">Palmer, D., ed. (1999). <i>The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals</i>. London: Marshall Editions. p. 38. <a href="/enwiki/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/enwiki/wiki/Special:BookSources/1-84028-152-9" title="Special:BookSources/1-84028-152-9"><bdi>1-84028-152-9</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Marshall+Illustrated+Encyclopedia+of+Dinosaurs+and+Prehistoric+Animals&rft.place=London&rft.pages=38&rft.pub=Marshall+Editions&rft.date=1999&rft.isbn=1-84028-152-9&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-FT12-45"><span class="mw-cite-backlink"><b><a href="#cite_ref-FT12_45-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFFreyTischlinger2012" class="citation journal cs1">Frey, E.; Tischlinger, H. (2012). <a rel="nofollow" class="external text" href="/enwiki//www.ncbi.nlm.nih.gov/pmc/articles/PMC3296705">"The Late Jurassic pterosaur <i>Rhamphorhynchus</i>, a frequent victim of the ganoid fish <i>Aspidorhynchus</i>?"</a>. <i>PLOS ONE</i>. <b>7</b> (3): e31945. <a href="/enwiki/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2012PLoSO...7E1945F">2012PLoSO...7E1945F</a>. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1371%2Fjournal.pone.0031945">10.1371/journal.pone.0031945</a>. <a href="/enwiki/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="cs1-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="/enwiki//www.ncbi.nlm.nih.gov/pmc/articles/PMC3296705">3296705</a></span>. <a href="/enwiki/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="/enwiki//pubmed.ncbi.nlm.nih.gov/22412850">22412850</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=PLOS+ONE&rft.atitle=The+Late+Jurassic+pterosaur+Rhamphorhynchus%2C+a+frequent+victim+of+the+ganoid+fish+Aspidorhynchus%3F&rft.volume=7&rft.issue=3&rft.pages=e31945&rft.date=2012&rft_id=%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3296705%23id-name%3DPMC&rft_id=info%3Apmid%2F22412850&rft_id=info%3Adoi%2F10.1371%2Fjournal.pone.0031945&rft_id=info%3Abibcode%2F2012PLoSO...7E1945F&rft.aulast=Frey&rft.aufirst=E.&rft.au=Tischlinger%2C+H.&rft_id=%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3296705&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-46"><span class="mw-cite-backlink"><b><a href="#cite_ref-46">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFNybelin1964" class="citation book cs1">Nybelin, Orvar (1964). <i>Versuch einer taxonomischen Revision der jurassischen Fischgattung Thrissops Agassiz</i> (in German). Göteborg: Wettergren and Kerber. <a href="/enwiki/wiki/OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="/enwiki//www.worldcat.org/oclc/2672427">2672427</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Versuch+einer+taxonomischen+Revision+der+jurassischen+Fischgattung+Thrissops+Agassiz.&rft.place=G%C3%B6teborg&rft.pub=Wettergren+and+Kerber&rft.date=1964&rft_id=info%3Aoclcnum%2F2672427&rft.aulast=Nybelin&rft.aufirst=Orvar&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-47"><span class="mw-cite-backlink"><b><a href="#cite_ref-47">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFKellner2003" class="citation journal cs1">Kellner, Alexander W. A. (2003). "Pterosaur phylogeny and comments on the evolutionary history of the group". <i>Geological Society, London, Special Publications</i>. Evolution and Palaeobiology of Pterosaurs. <b>217</b> (1): 105–137. <a href="/enwiki/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2003GSLSP.217..105K">2003GSLSP.217..105K</a>. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1144%2FGSL.SP.2003.217.01.10">10.1144/GSL.SP.2003.217.01.10</a>. <a href="/enwiki/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="/enwiki//www.worldcat.org/issn/0305-8719">0305-8719</a>. <a href="/enwiki/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:128892642">128892642</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Geological+Society%2C+London%2C+Special+Publications&rft.atitle=Pterosaur+phylogeny+and+comments+on+the+evolutionary+history+of+the+group&rft.volume=217&rft.issue=1&rft.pages=105-137&rft.date=2003&rft_id=info%3Adoi%2F10.1144%2FGSL.SP.2003.217.01.10&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A128892642%23id-name%3DS2CID&rft.issn=0305-8719&rft_id=info%3Abibcode%2F2003GSLSP.217..105K&rft.aulast=Kellner&rft.aufirst=Alexander+W.+A.&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-Aerodactylus-48"><span class="mw-cite-backlink">^ <a href="#cite_ref-Aerodactylus_48-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Aerodactylus_48-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFVidovicMartill2014" class="citation journal cs1">Vidovic, S. U.; Martill, D. M. (2014). <a rel="nofollow" class="external text" href="/enwiki//www.ncbi.nlm.nih.gov/pmc/articles/PMC4206445">"<i>Pterodactylus scolopaciceps</i> Meyer, 1860 (Pterosauria, Pterodactyloidea) from the Upper Jurassic of Bavaria, Germany: The Problem of Cryptic Pterosaur Taxa in Early Ontogeny"</a>. <i>PLOS ONE</i>. <b>9</b> (10): e110646. <a href="/enwiki/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2014PLoSO...9k0646V">2014PLoSO...9k0646V</a>. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1371%2Fjournal.pone.0110646">10.1371/journal.pone.0110646</a>. <a href="/enwiki/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="cs1-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="/enwiki//www.ncbi.nlm.nih.gov/pmc/articles/PMC4206445">4206445</a></span>. <a href="/enwiki/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="/enwiki//pubmed.ncbi.nlm.nih.gov/25337830">25337830</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=PLOS+ONE&rft.atitle=Pterodactylus+scolopaciceps+Meyer%2C+1860+%28Pterosauria%2C+Pterodactyloidea%29+from+the+Upper+Jurassic+of+Bavaria%2C+Germany%3A+The+Problem+of+Cryptic+Pterosaur+Taxa+in+Early+Ontogeny&rft.volume=9&rft.issue=10&rft.pages=e110646&rft.date=2014&rft_id=%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4206445%23id-name%3DPMC&rft_id=info%3Apmid%2F25337830&rft_id=info%3Adoi%2F10.1371%2Fjournal.pone.0110646&rft_id=info%3Abibcode%2F2014PLoSO...9k0646V&rft.aulast=Vidovic&rft.aufirst=S.+U.&rft.au=Martill%2C+D.+M.&rft_id=%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4206445&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-Pterosauria-49"><span class="mw-cite-backlink"><b><a href="#cite_ref-Pterosauria_49-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFWitton2013" class="citation book cs1">Witton, Mark (2013). <i>Pterosaurs: Natural History, Evolution, Anatomy</i>. Princeton University Press. <a href="/enwiki/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/enwiki/wiki/Special:BookSources/978-0-691-15061-1" title="Special:BookSources/978-0-691-15061-1"><bdi>978-0-691-15061-1</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Pterosaurs%3A+Natural+History%2C+Evolution%2C+Anatomy&rft.pub=Princeton+University+Press&rft.date=2013&rft.isbn=978-0-691-15061-1&rft.aulast=Witton&rft.aufirst=Mark&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-Münster1830-50"><span class="mw-cite-backlink"><b><a href="#cite_ref-Münster1830_50-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFMünster1830" class="citation book cs1">Münster, Georg Graf zu (1830). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=RWuWamFBZaAC&pg=PA2"><i>Nachtrag zu der Abhandlung des professor Goldfuss ueber den Ornithocephalus Münsteri (Goldf.)</i></a>. Bayreuth: F. C. Birner.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Nachtrag+zu+der+Abhandlung+des+professor+Goldfuss+ueber+den+Ornithocephalus+M%C3%BCnsteri+%28Goldf.%29&rft.place=Bayreuth&rft.pub=F.+C.+Birner&rft.date=1830&rft.aulast=M%C3%BCnster&rft.aufirst=Georg+Graf+zu&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DRWuWamFBZaAC%26pg%3DPA2&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-51"><span class="mw-cite-backlink"><b><a href="#cite_ref-51">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFMünster1839" class="citation book cs1">Münster, Georg Graf zu (1839). <a rel="nofollow" class="external text" href="https://www.biodiversitylibrary.org/item/110811#page/696/mode/1up">"Über einige neue Versteinerungen in der lithographischen Schiefer von Baiern"</a>. <i>Neues Jahrbuch für Mineralogie, Geognosie, Geologie und Petrefaktenkunde</i>. Stuttgart: E. Schweizerbart's Verlagshandlung. pp. 676–682.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=%C3%9Cber+einige+neue+Versteinerungen+in+der+lithographischen+Schiefer+von+Baiern&rft.btitle=Neues+Jahrbuch+f%C3%BCr+Mineralogie%2C+Geognosie%2C+Geologie+und+Petrefaktenkunde&rft.place=Stuttgart&rft.pages=676-682&rft.pub=E.+Schweizerbart%27s+Verlagshandlung&rft.date=1839&rft.aulast=M%C3%BCnster&rft.aufirst=Georg+Graf+zu&rft_id=https%3A%2F%2Fwww.biodiversitylibrary.org%2Fitem%2F110811%23page%2F696%2Fmode%2F1up&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-vonmeyer1845-52"><span class="mw-cite-backlink"><b><a href="#cite_ref-vonmeyer1845_52-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFvon_Meyer1845" class="citation journal cs1">von Meyer, Hermann (1845). <a rel="nofollow" class="external text" href="https://www.biodiversitylibrary.org/item/110636#page/300/mode/1up">"System der fossilen Saurier"</a> [Taxonomy of fossil saurians]. <i>Neues Jahrbuch für Mineralogie, Geognosie, Geologie und Petrefaktenkunde</i> (in German). Stuttgart: E. Schweizerbart's Verlagshandlung: 278–285.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Neues+Jahrbuch+f%C3%BCr+Mineralogie%2C+Geognosie%2C+Geologie+und+Petrefaktenkunde&rft.atitle=System+der+fossilen+Saurier&rft.pages=278-285&rft.date=1845&rft.aulast=von+Meyer&rft.aufirst=Hermann&rft_id=https%3A%2F%2Fwww.biodiversitylibrary.org%2Fitem%2F110636%23page%2F300%2Fmode%2F1up&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-vonmeyer1846-53"><span class="mw-cite-backlink"><b><a href="#cite_ref-vonmeyer1846_53-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFvon_Meyer1846" class="citation journal cs1">von Meyer, Hermann (1846). <a rel="nofollow" class="external text" href="https://www.biodiversitylibrary.org/item/43708#page/17/mode/1up">"Pterodactylus (Rhamphorhynchus) gemmingi aus dem Kalkschiefer von Solenhofen"</a>. <i>Palaeontographica</i>. Cassel (published 1851). <b>1</b>: 1–20.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Palaeontographica&rft.atitle=Pterodactylus+%28Rhamphorhynchus%29+gemmingi+aus+dem+Kalkschiefer+von+Solenhofen&rft.volume=1&rft.pages=1-20&rft.date=1846&rft.aulast=von+Meyer&rft.aufirst=Hermann&rft_id=https%3A%2F%2Fwww.biodiversitylibrary.org%2Fitem%2F43708%23page%2F17%2Fmode%2F1up&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-vonmeyer1847-54"><span class="mw-cite-backlink"><b><a href="#cite_ref-vonmeyer1847_54-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFvon_Meyer1847" class="citation book cs1">von Meyer, Hermann (1847). <a rel="nofollow" class="external text" href="http://data.onb.ac.at/rep/10B61475"><i>Homoeosaurus maximiliani und Rhamphorhynchus (Pterodactylus) longicaudus: Zwei fossile Reptilien aus dem Kalkschiefer von Solenhofen</i></a> (in German). Frankfurt: S. Schmerber'schen buchhandlung.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Homoeosaurus+maximiliani+und+Rhamphorhynchus+%28Pterodactylus%29+longicaudus%3A+Zwei+fossile+Reptilien+aus+dem+Kalkschiefer+von+Solenhofen&rft.place=Frankfurt&rft.pub=S.+Schmerber%27schen+buchhandlung&rft.date=1847&rft.aulast=von+Meyer&rft.aufirst=Hermann&rft_id=http%3A%2F%2Fdata.onb.ac.at%2Frep%2F10B61475&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-Bowerbank1846-55"><span class="mw-cite-backlink"><b><a href="#cite_ref-Bowerbank1846_55-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFBowerbank1846" class="citation journal cs1">Bowerbank, J.S. (1846). <a rel="nofollow" class="external text" href="https://zenodo.org/record/1448505">"On a new species of pterodactyl found in the Upper Chalk of Kent (<i>Pterodactylus giganteus</i>)"</a>. <i>Quarterly Journal of the Geological Society of London</i>. <b>2</b> (1–2): 7–9. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1144%2Fgsl.jgs.1846.002.01-02.05">10.1144/gsl.jgs.1846.002.01-02.05</a>. <a href="/enwiki/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:129389179">129389179</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Quarterly+Journal+of+the+Geological+Society+of+London&rft.atitle=On+a+new+species+of+pterodactyl+found+in+the+Upper+Chalk+of+Kent+%28Pterodactylus+giganteus%29&rft.volume=2&rft.issue=1%E2%80%932&rft.pages=7-9&rft.date=1846&rft_id=info%3Adoi%2F10.1144%2Fgsl.jgs.1846.002.01-02.05&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A129389179%23id-name%3DS2CID&rft.aulast=Bowerbank&rft.aufirst=J.S.&rft_id=https%3A%2F%2Fzenodo.org%2Frecord%2F1448505&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-Bowerbank1851-56"><span class="mw-cite-backlink"><b><a href="#cite_ref-Bowerbank1851_56-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFBowerbank1851" class="citation journal cs1">Bowerbank, J.S. (1851). <a rel="nofollow" class="external text" href="https://zenodo.org/record/1447536">"On the pterodactyles of the Chalk Formation"</a>. <i>Proceedings of the Zoological Society of London</i>. <b>19</b>: 14–20. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Fj.1096-3642.1851.tb01125.x">10.1111/j.1096-3642.1851.tb01125.x</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Proceedings+of+the+Zoological+Society+of+London&rft.atitle=On+the+pterodactyles+of+the+Chalk+Formation&rft.volume=19&rft.pages=14-20&rft.date=1851&rft_id=info%3Adoi%2F10.1111%2Fj.1096-3642.1851.tb01125.x&rft.aulast=Bowerbank&rft.aufirst=J.S.&rft_id=https%3A%2F%2Fzenodo.org%2Frecord%2F1447536&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-Owen1851-57"><span class="mw-cite-backlink"><b><a href="#cite_ref-Owen1851_57-0">^</a></b></span> <span class="reference-text">Owen, R. (1851). Monograph on the fossil Reptilia of the Cretaceous Formations. <i>The Palaeontographical Society</i> <b>5</b>(11):1–118.</span>
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<li id="cite_note-:1-58"><span class="mw-cite-backlink">^ <a href="#cite_ref-:1_58-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:1_58-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFHooley1914" class="citation journal cs1">Hooley, Reginald Walter (1914). <a rel="nofollow" class="external text" href="https://zenodo.org/record/2207691">"On the Ornithosaurian genus <i>Ornithocheirus</i>, with a review of the specimens from the Cambridge Greensand in the Sedgwick Museum, Cambridge"</a>. <i>Annals and Magazine of Natural History</i>. <b>13</b> (78): 529–557. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1080%2F00222931408693521">10.1080/00222931408693521</a>. <a href="/enwiki/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="/enwiki//www.worldcat.org/issn/0374-5481">0374-5481</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Annals+and+Magazine+of+Natural+History&rft.atitle=On+the+Ornithosaurian+genus+Ornithocheirus%2C+with+a+review+of+the+specimens+from+the+Cambridge+Greensand+in+the+Sedgwick+Museum%2C+Cambridge&rft.volume=13&rft.issue=78&rft.pages=529-557&rft.date=1914&rft_id=info%3Adoi%2F10.1080%2F00222931408693521&rft.issn=0374-5481&rft.aulast=Hooley&rft.aufirst=Reginald+Walter&rft_id=https%3A%2F%2Fzenodo.org%2Frecord%2F2207691&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-S1869-62"><span class="mw-cite-backlink">^ <a href="#cite_ref-S1869_62-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-S1869_62-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFSeeley1869" class="citation journal cs1">Seeley, Harry Govier (1869). <a rel="nofollow" class="external text" href="https://www.biodiversitylibrary.org/part/59487">"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"</a>. <i>Annals and Magazine of Natural History</i>. <b>5</b> (27): 225–226. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1080%2F00222937008696143">10.1080/00222937008696143</a>. <a href="/enwiki/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="/enwiki//www.worldcat.org/issn/0374-5481">0374-5481</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Annals+and+Magazine+of+Natural+History&rft.atitle=Index+to+the+fossil+remains+of+Aves%2C+Ornithosauria%2C+and+Reptilia%2C+from+the+Secondary+System+of+Strata%2C+arranged+in+the+Woodwardian+Museum+of+the+University+of+Cambridge&rft.volume=5&rft.issue=27&rft.pages=225-226&rft.date=1869&rft_id=info%3Adoi%2F10.1080%2F00222937008696143&rft.issn=0374-5481&rft.aulast=Seeley&rft.aufirst=Harry+Govier&rft_id=https%3A%2F%2Fwww.biodiversitylibrary.org%2Fpart%2F59487&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-HGS70-63"><span class="mw-cite-backlink"><b><a href="#cite_ref-HGS70_63-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFSeeley,_H.G.1870" class="citation journal cs1">Seeley, H.G. (1870). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=0M8yAQAAMAAJ">"The Ornithosauria: an Elementary Study of the Bones of Pterodactyles"</a>. <i>Cambridge</i>: 112–128.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Cambridge&rft.atitle=The+Ornithosauria%3A+an+Elementary+Study+of+the+Bones+of+Pterodactyles&rft.pages=112-128&rft.date=1870&rft.au=Seeley%2C+H.G.&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3D0M8yAQAAMAAJ&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-66"><span class="mw-cite-backlink"><b><a href="#cite_ref-66">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFMarsh,_O.C.1871" class="citation journal cs1">Marsh, O.C. (1871). <a rel="nofollow" class="external text" href="http://oceansofkansas.com/Marsh71.htm">"Note on a new and gigantic species of Pterodactyle"</a>. <i>American Journal of Science</i>. 3. <b>1</b> (6): 472.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=American+Journal+of+Science&rft.atitle=Note+on+a+new+and+gigantic+species+of+Pterodactyle&rft.volume=1&rft.issue=6&rft.pages=472&rft.date=1871&rft.au=Marsh%2C+O.C.&rft_id=http%3A%2F%2Foceansofkansas.com%2FMarsh71.htm&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-History-67"><span class="mw-cite-backlink">^ <a href="#cite_ref-History_67-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-History_67-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-History_67-2"><sup><i><b>c</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFWitton,_M.P.2010" class="citation journal cs1">Witton, M.P. (2010). <a rel="nofollow" class="external text" href="https://www.researchgate.net/publication/258391482">"<i>Pteranodon</i> and beyond: The history of giant pterosaurs from 1870 onwards. Geological Society of London Special Publications"</a>. <b>343</b>: 313–323.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.atitle=Pteranodon+and+beyond%3A+The+history+of+giant+pterosaurs+from+1870+onwards.+Geological+Society+of+London+Special+Publications&rft.volume=343&rft.pages=313-323&rft.date=2010&rft.au=Witton%2C+M.P.&rft_id=https%3A%2F%2Fwww.researchgate.net%2Fpublication%2F258391482&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span> <span class="cs1-hidden-error error citation-comment">Cite journal requires <code class="cs1-code">|journal=</code> (<a href="/enwiki/wiki/Help:CS1_errors#missing_periodical" title="Help:CS1 errors">help</a>)</span></span>
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<li id="cite_note-68"><span class="mw-cite-backlink"><b><a href="#cite_ref-68">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFCope1872" class="citation journal cs1">Cope, E. D. (1872). "On two new Ornithosaurians from Kansas". <i>Proceedings of the American Philosophical Society</i>. <b>12</b> (88): 420–422. <a href="/enwiki/wiki/JSTOR_(identifier)" class="mw-redirect" title="JSTOR (identifier)">JSTOR</a> <a rel="nofollow" class="external text" href="/enwiki//www.jstor.org/stable/981730">981730</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Proceedings+of+the+American+Philosophical+Society&rft.atitle=On+two+new+Ornithosaurians+from+Kansas&rft.volume=12&rft.issue=88&rft.pages=420-422&rft.date=1872&rft_id=%2F%2Fwww.jstor.org%2Fstable%2F981730%23id-name%3DJSTOR&rft.aulast=Cope&rft.aufirst=E.+D.&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-cope1875-69"><span class="mw-cite-backlink"><b><a href="#cite_ref-cope1875_69-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFCope1875" class="citation journal cs1">Cope, E. D. (1875). <a rel="nofollow" class="external text" href="https://www.biodiversitylibrary.org/item/125656#page/11/mode/1up">"The Vertebrata of the Cretaceous formations of the West"</a>. <i>Report, U. S. Geological Survey of the Territories (Hayden)</i>. <b>2</b>: 302 pp., 57 pls.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Report%2C+U.+S.+Geological+Survey+of+the+Territories+%28Hayden%29&rft.atitle=The+Vertebrata+of+the+Cretaceous+formations+of+the+West.&rft.volume=2&rft.pages=302+pp.%2C+57+pls&rft.date=1875&rft.aulast=Cope&rft.aufirst=E.+D.&rft_id=https%3A%2F%2Fwww.biodiversitylibrary.org%2Fitem%2F125656%23page%2F11%2Fmode%2F1up&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-marsh1876a-70"><span class="mw-cite-backlink"><b><a href="#cite_ref-marsh1876a_70-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFMarsh,_O.C.1876a" class="citation journal cs1">Marsh, O.C. (1876a). <a rel="nofollow" class="external text" href="https://zenodo.org/record/1450032">"Notice of a new sub-order of Pterosauria"</a>. <i>American Journal of Science</i>. Series 3. <b>11</b> (65): 507–509. <a href="/enwiki/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1876AmJS...11..507M">1876AmJS...11..507M</a>. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.2475%2Fajs.s3-11.66.507">10.2475/ajs.s3-11.66.507</a>. <a href="/enwiki/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:130203580">130203580</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=American+Journal+of+Science&rft.atitle=Notice+of+a+new+sub-order+of+Pterosauria&rft.volume=11&rft.issue=65&rft.pages=507-509&rft.date=1876&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A130203580%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.2475%2Fajs.s3-11.66.507&rft_id=info%3Abibcode%2F1876AmJS...11..507M&rft.au=Marsh%2C+O.C.&rft_id=https%3A%2F%2Fzenodo.org%2Frecord%2F1450032&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-SCB06-71"><span class="mw-cite-backlink">^ <a href="#cite_ref-SCB06_71-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-SCB06_71-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFBennett2006" class="citation journal cs1">Bennett, S.C. (2006). "Juvenile specimens of the pterosaur <i>Germanodactylus cristatus</i>, with a review of the genus". <i>Journal of Vertebrate Paleontology</i>. <b>26</b> (4): 872–878. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1671%2F0272-4634%282006%2926%5B872%3AJSOTPG%5D2.0.CO%3B2">10.1671/0272-4634(2006)26[872:JSOTPG]2.0.CO;2</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Vertebrate+Paleontology&rft.atitle=Juvenile+specimens+of+the+pterosaur+Germanodactylus+cristatus%2C+with+a+review+of+the+genus&rft.volume=26&rft.issue=4&rft.pages=872-878&rft.date=2006&rft_id=info%3Adoi%2F10.1671%2F0272-4634%282006%2926%5B872%3AJSOTPG%5D2.0.CO%3B2&rft.aulast=Bennett&rft.aufirst=S.C.&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-LoneStarPterosaurs-72"><span class="mw-cite-backlink"><b><a href="#cite_ref-LoneStarPterosaurs_72-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFAndresMyers2013" class="citation journal cs1">Andres, B.; Myers, T. S. (2013). "Lone Star Pterosaurs". <i>Earth and Environmental Science Transactions of the Royal Society of Edinburgh</i>. <b>103</b> (3–4): 383–398. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1017%2FS1755691013000303">10.1017/S1755691013000303</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Earth+and+Environmental+Science+Transactions+of+the+Royal+Society+of+Edinburgh&rft.atitle=Lone+Star+Pterosaurs&rft.volume=103&rft.issue=3%E2%80%934&rft.pages=383-398&rft.date=2013&rft_id=info%3Adoi%2F10.1017%2FS1755691013000303&rft.aulast=Andres&rft.aufirst=B.&rft.au=Myers%2C+T.+S.&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-bennett_2017_pteranodon_juv-73"><span class="mw-cite-backlink"><b><a href="#cite_ref-bennett_2017_pteranodon_juv_73-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFBennett2017" class="citation journal cs1">Bennett, S.C. (2017). "New smallest specimen of the pterosaur <i>Pteranodon</i> and ontogenetic niches in pterosaurs". <i>Journal of Paleontology</i>. <b>92</b> (2): 1–18. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1017%2Fjpa.2017.84">10.1017/jpa.2017.84</a>. <a href="/enwiki/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:90893067">90893067</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Paleontology&rft.atitle=New+smallest+specimen+of+the+pterosaur+Pteranodon+and+ontogenetic+niches+in+pterosaurs&rft.volume=92&rft.issue=2&rft.pages=1-18&rft.date=2017&rft_id=info%3Adoi%2F10.1017%2Fjpa.2017.84&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A90893067%23id-name%3DS2CID&rft.aulast=Bennett&rft.aufirst=S.C.&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-74"><span class="mw-cite-backlink"><b><a href="#cite_ref-74">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFArnold2014" class="citation book cs1">Arnold, Caroline (2014). <i>Pterosaurs: Rulers of the Skies in the Dinosaur Age</i>. <a href="/enwiki/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/enwiki/wiki/Special:BookSources/978-1-63083-412-8" title="Special:BookSources/978-1-63083-412-8"><bdi>978-1-63083-412-8</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Pterosaurs%3A+Rulers+of+the+Skies+in+the+Dinosaur+Age&rft.date=2014&rft.isbn=978-1-63083-412-8&rft.aulast=Arnold&rft.aufirst=Caroline&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-myths-75"><span class="mw-cite-backlink"><b><a href="#cite_ref-myths_75-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite id="CITEREFNaish,_Darren" class="citation web cs1">Naish, Darren. <a rel="nofollow" class="external text" href="http://www.pterosaur.net/myths.php">"Pterosaurs: Myths and Misconceptions"</a>. Pterosaur.net<span class="reference-accessdate">. Retrieved <span class="nowrap">June 18,</span> 2011</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Pterosaurs%3A+Myths+and+Misconceptions&rft.pub=Pterosaur.net&rft.au=Naish%2C+Darren&rft_id=http%3A%2F%2Fwww.pterosaur.net%2Fmyths.php&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-76"><span class="mw-cite-backlink"><b><a href="#cite_ref-76">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r999302996"/><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://www.imdb.com/title/tt0452668/">"Pterodactyl (2005)"</a>. <i>Sci Fi Channel</i>. IMDb<span class="reference-accessdate">. Retrieved <span class="nowrap">August 27,</span> 2005</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Sci+Fi+Channel&rft.atitle=Pterodactyl+%282005%29&rft_id=https%3A%2F%2Fwww.imdb.com%2Ftitle%2Ftt0452668%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3APterodactylus" class="Z3988"></span></span>
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<li id="cite_note-Letter_211-77"><span class="mw-cite-backlink"><b><a href="#cite_ref-Letter_211_77-0">^</a></b></span> <span class="reference-text"><i><a href="/enwiki/wiki/Letters_of_J._R._R._Tolkien" class="mw-redirect" title="Letters of J. R. R. Tolkien">Letters of J. R. R. Tolkien</a></i>, #211 to Rhona Beare, October 14, 1958</span>
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<li id="cite_note-T-78"><span class="mw-cite-backlink"><b><a href="#cite_ref-T_78-0">^</a></b></span> <span class="reference-text"><i><a href="/enwiki/wiki/Letters_of_J._R._R._Tolkien" class="mw-redirect" title="Letters of J. R. R. Tolkien">Letters of J. R. R. Tolkien</a></i>, #100 to <a href="/enwiki/wiki/Christopher_Tolkien" title="Christopher Tolkien">Christopher Tolkien</a>, May 29, 1945, expressing his "loathing" for the <a href="/enwiki/wiki/Royal_Air_Force" title="Royal Air Force">Royal Air Force</a>: "My sentiments are more or less those that <a href="/enwiki/wiki/Frodo" class="mw-redirect" title="Frodo">Frodo</a> would have had if he discovered some <a href="/enwiki/wiki/Hobbit" title="Hobbit">Hobbits</a> learning to ride Nazgûl-birds, 'for the liberation of the Shire'."</span>
</li>
</ol></div>
<h2><span class="mw-headline" id="External_links">External links</span><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Pterodactylus&action=edit&section=16" title="Edit section: External links">edit</a><span class="mw-editsection-bracket">]</span></span></h2>
<ul><li><a href="/enwiki/wiki/File:Commons-logo.svg" class="image"><img alt="" src="/upwiki/wikipedia/en/thumb/4/4a/Commons-logo.svg/12px-Commons-logo.svg.png" decoding="async" width="12" height="16" class="noviewer" srcset="/upwiki/wikipedia/en/thumb/4/4a/Commons-logo.svg/18px-Commons-logo.svg.png 1.5x, /upwiki/wikipedia/en/thumb/4/4a/Commons-logo.svg/24px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /></a> Media related to <a href="https://commons.wikimedia.org/wiki/Category:Pterodactylus" class="extiw" title="commons:Category:Pterodactylus"><span style=""><i>Pterodactylus</i></span></a> at Wikimedia Commons</li>
<li><a href="/enwiki/wiki/File:Wikispecies-logo.svg" class="image"><img alt="" src="/upwiki/wikipedia/commons/thumb/d/df/Wikispecies-logo.svg/14px-Wikispecies-logo.svg.png" decoding="async" width="14" height="16" class="noviewer" srcset="/upwiki/wikipedia/commons/thumb/d/df/Wikispecies-logo.svg/20px-Wikispecies-logo.svg.png 1.5x, /upwiki/wikipedia/commons/thumb/d/df/Wikispecies-logo.svg/27px-Wikispecies-logo.svg.png 2x" data-file-width="941" data-file-height="1103" /></a> Data related to <a href="https://species.wikimedia.org/wiki/Pterodactylus" class="extiw" title="wikispecies:Pterodactylus"><i>Pterodactylus</i></a> at Wikispecies</li></ul>
<div role="navigation" class="navbox" aria-labelledby="Pterosauria" style="padding:3px"><table class="nowraplinks hlist mw-collapsible expanded navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><style data-mw-deduplicate="TemplateStyles:r992953826">.mw-parser-output .navbar{display:inline;font-size:88%;font-weight:normal}.mw-parser-output .navbar-collapse{float:left;text-align:left}.mw-parser-output .navbar-boxtext{word-spacing:0}.mw-parser-output .navbar ul{display:inline-block;white-space:nowrap;line-height:inherit}.mw-parser-output .navbar-brackets::before{margin-right:-0.125em;content:"[ "}.mw-parser-output .navbar-brackets::after{margin-left:-0.125em;content:" ]"}.mw-parser-output .navbar li{word-spacing:-0.125em}.mw-parser-output .navbar-mini abbr{font-variant:small-caps;border-bottom:none;text-decoration:none;cursor:inherit}.mw-parser-output .navbar-ct-full{font-size:114%;margin:0 7em}.mw-parser-output .navbar-ct-mini{font-size:114%;margin:0 4em}.mw-parser-output .infobox .navbar{font-size:100%}.mw-parser-output .navbox .navbar{display:block;font-size:100%}.mw-parser-output .navbox-title .navbar{float:left;text-align:left;margin-right:0.5em}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/enwiki/wiki/Template:Pterosauria" title="Template:Pterosauria"><abbr title="View this template" style=";;background:none transparent;border:none;box-shadow:none;padding:0;">v</abbr></a></li><li class="nv-talk"><a href="/enwiki/wiki/Template_talk:Pterosauria" title="Template talk:Pterosauria"><abbr title="Discuss this template" style=";;background:none transparent;border:none;box-shadow:none;padding:0;">t</abbr></a></li><li class="nv-edit"><a class="external text" href="https://en.wikipedia.org/enwiki/w/index.php?title=Template:Pterosauria&action=edit"><abbr title="Edit this template" style=";;background:none transparent;border:none;box-shadow:none;padding:0;">e</abbr></a></li></ul></div><div id="Pterosauria" style="font-size:114%;margin:0 4em"><a href="/enwiki/wiki/Pterosaur" title="Pterosaur">Pterosauria</a></div></th></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks mw-collapsible mw-collapsed navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="*_Kingdom:_Animalia_*_Phylum:_Chordata_*_Class:_Sauropsida_*_Clade:_Avemetatarsalia" style="font-size:114%;margin:0 4em"><small><style data-mw-deduplicate="TemplateStyles:r886047488">.mw-parser-output .nobold{font-weight:normal}</style><span class="nobold">
<ul><li>Kingdom: <a href="/enwiki/wiki/Animal" title="Animal">Animalia</a></li>
<li>Phylum: <a href="/enwiki/wiki/Chordate" title="Chordate">Chordata</a></li>
<li>Class: <a href="/enwiki/wiki/Sauropsida" title="Sauropsida">Sauropsida</a></li>
<li><i>Clade</i>: <a href="/enwiki/wiki/Avemetatarsalia" title="Avemetatarsalia">Avemetatarsalia</a></li></ul>
</span></small></div></th></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="Avemetatarsalia" scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Avemetatarsalia" title="Avemetatarsalia">Avemetatarsalia</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><ul><li>see <a href="/enwiki/wiki/Template:Avemetatarsalia" title="Template:Avemetatarsalia">Avemetatarsalia</a></li></ul></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Pterosaur" title="Pterosaur">Pterosauria</a></th><td class="navbox-list navbox-even" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><ul><li>see below↓</li></ul></li></ul>
</div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks mw-collapsible mw-collapsed navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="col" class="navbox-title" colspan="3"><div id="Pterosauria" style="font-size:114%;margin:0 4em"><a href="/enwiki/wiki/Pterosaur" title="Pterosaur">Pterosauria</a></div></th></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Preondactylia" class="mw-redirect" title="Preondactylia">Preondactylia</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Austriadactylus" title="Austriadactylus">Austriadactylus</a></i></li>
<li><i><a href="/enwiki/wiki/Preondactylus" title="Preondactylus">Preondactylus</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Caviramidae" title="Caviramidae">Caviramidae</a>?</th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Carniadactylus" title="Carniadactylus">Carniadactylus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Caviramus" title="Caviramus">Caviramus</a></i></li>
<li><i><a href="/enwiki/wiki/Raeticodactylus" title="Raeticodactylus">Raeticodactylus</a></i>?</li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Austriadraconidae" class="mw-redirect" title="Austriadraconidae">Austriadraconidae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Arcticodactylus" title="Arcticodactylus">Arcticodactylus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Austriadraco" title="Austriadraco">Austriadraco</a></i></li>
<li><i><a href="/enwiki/wiki/Seazzadactylus" title="Seazzadactylus">Seazzadactylus</a></i></li></ul>
</div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><div style="position:absolute;-webkit-transform:translate(-50%,-50%) rotate(270deg);-moz-transform:translate(-50%,-50%) rotate(270deg);-ms-transform:translate(-50%,-50%) rotate(270deg);-o-transform:translate(-50%,-50%) rotate(270deg);transform:translate(-50%,-50%) rotate(270deg);"><a href="/enwiki/wiki/Eopterosauria" title="Eopterosauria">Eopterosauria</a>?</div></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Austriadraco" title="Austriadraco">Austriadraco</a></i>?</li>
<li><i><a href="/enwiki/wiki/Peteinosaurus" title="Peteinosaurus">Peteinosaurus</a></i>?</li>
<li><b><a href="/enwiki/wiki/Preondactylia" class="mw-redirect" title="Preondactylia">Preondactylia</a></b>?</li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Eudimorphodontoidea" class="mw-redirect" title="Eudimorphodontoidea">Eudimorphodontoidea</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Raeticodactylidae" title="Raeticodactylidae">Raeticodactylidae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Caviramus" title="Caviramus">Caviramus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Raeticodactylus" title="Raeticodactylus">Raeticodactylus</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Eudimorphodontidae" title="Eudimorphodontidae">Eudimorphodontidae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Arcticodactylus" title="Arcticodactylus">Arcticodactylus</a></i>?</li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Eudimorphodontinae" class="mw-redirect" title="Eudimorphodontinae">Eudimorphodontinae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Carniadactylus" title="Carniadactylus">Carniadactylus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Eudimorphodon" title="Eudimorphodon">Eudimorphodon</a></i></li></ul>
</div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><div style="position:absolute;-webkit-transform:translate(-50%,-50%) rotate(270deg);-moz-transform:translate(-50%,-50%) rotate(270deg);-ms-transform:translate(-50%,-50%) rotate(270deg);-o-transform:translate(-50%,-50%) rotate(270deg);transform:translate(-50%,-50%) rotate(270deg);"><a href="/enwiki/wiki/Zambellisauria" class="mw-redirect" title="Zambellisauria">Zambellisauria</a>?</div></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Peteinosaurus" title="Peteinosaurus">Peteinosaurus</a></i>?</li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><div style="position:absolute;-webkit-transform:translate(-50%,-50%) rotate(270deg);-moz-transform:translate(-50%,-50%) rotate(270deg);-ms-transform:translate(-50%,-50%) rotate(270deg);-o-transform:translate(-50%,-50%) rotate(270deg);transform:translate(-50%,-50%) rotate(270deg);"><a href="/enwiki/wiki/Macronychoptera" class="mw-redirect" title="Macronychoptera">Macronychoptera</a></div></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="Dimorphodontidae" scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Dimorphodontidae" title="Dimorphodontidae">Dimorphodontidae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Caelestiventus" title="Caelestiventus">Caelestiventus</a></i></li>
<li><i><a href="/enwiki/wiki/Dimorphodon" title="Dimorphodon">Dimorphodon</a></i></li>
<li><i><a href="/enwiki/wiki/Parapsicephalus" title="Parapsicephalus">Parapsicephalus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Peteinosaurus" title="Peteinosaurus">Peteinosaurus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Rhamphinion" title="Rhamphinion">Rhamphinion</a></i>?</li></ul>
</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><div style="position:absolute;-webkit-transform:translate(-50%,-50%) rotate(270deg);-moz-transform:translate(-50%,-50%) rotate(270deg);-ms-transform:translate(-50%,-50%) rotate(270deg);-o-transform:translate(-50%,-50%) rotate(270deg);transform:translate(-50%,-50%) rotate(270deg);"><a href="/enwiki/wiki/Lonchognatha" class="mw-redirect" title="Lonchognatha">Lonchognatha</a>?</div></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Eudimorphodon" title="Eudimorphodon">Eudimorphodon</a></i>?</li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><div style="position:absolute;-webkit-transform:translate(-50%,-50%) rotate(270deg);-moz-transform:translate(-50%,-50%) rotate(270deg);-ms-transform:translate(-50%,-50%) rotate(270deg);-o-transform:translate(-50%,-50%) rotate(270deg);transform:translate(-50%,-50%) rotate(270deg);"><a href="/enwiki/wiki/Novialoidea" title="Novialoidea">Novialoidea</a></div></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="Campylognathoididae" scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Campylognathoididae" class="mw-redirect" title="Campylognathoididae">Campylognathoididae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Bergamodactylus" title="Bergamodactylus">Bergamodactylus</a></i></li>
<li><i><a href="/enwiki/wiki/Campylognathoides" title="Campylognathoides">Campylognathoides</a></i></li></ul>
</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><div style="position:absolute;-webkit-transform:translate(-50%,-50%) rotate(270deg);-moz-transform:translate(-50%,-50%) rotate(270deg);-ms-transform:translate(-50%,-50%) rotate(270deg);-o-transform:translate(-50%,-50%) rotate(270deg);transform:translate(-50%,-50%) rotate(270deg);"><a href="/enwiki/wiki/Breviquartossa" class="mw-redirect" title="Breviquartossa">Breviquartossa</a></div></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Rhamphorhynchidae" title="Rhamphorhynchidae">Rhamphorhynchidae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Klobiodon" title="Klobiodon">Klobiodon</a></i></li>
<li><i><a href="/enwiki/wiki/Parapsicephalus" title="Parapsicephalus">Parapsicephalus</a></i>?</li>
<li><b><a href="/enwiki/wiki/Scaphognathidae" class="mw-redirect" title="Scaphognathidae">Scaphognathinae</a></b>?</li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Rhamphorhynchinae" class="mw-redirect" title="Rhamphorhynchinae">Rhamphorhynchinae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Angustinaripterus" title="Angustinaripterus">Angustinaripterus</a></i></li>
<li><i><a href="/enwiki/wiki/Bellubrunnus" title="Bellubrunnus">Bellubrunnus</a></i></li>
<li><i><a href="/enwiki/wiki/Cacibupteryx" title="Cacibupteryx">Cacibupteryx</a></i></li>
<li><i><a href="/enwiki/wiki/Dorygnathus" title="Dorygnathus">Dorygnathus</a></i></li>
<li><i><a href="/enwiki/wiki/Harpactognathus" title="Harpactognathus">Harpactognathus</a></i></li>
<li><i><a href="/enwiki/wiki/Nesodactylus" title="Nesodactylus">Nesodactylus</a></i></li>
<li><i><a href="/enwiki/wiki/Orientognathus" title="Orientognathus">Orientognathus</a></i></li>
<li><i><a href="/enwiki/wiki/Qinglongopterus" title="Qinglongopterus">Qinglongopterus</a></i></li>
<li><i><a href="/enwiki/wiki/Rhamphorhynchus" title="Rhamphorhynchus">Rhamphorhynchus</a></i></li>
<li><i><a href="/enwiki/wiki/Sericipterus" title="Sericipterus">Sericipterus</a></i></li></ul>
</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Digibrevisauria" class="mw-redirect" title="Digibrevisauria">Digibrevisauria</a>?</th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Scaphognathidae" class="mw-redirect" title="Scaphognathidae">Scaphognathidae</a>?</th><td class="navbox-list navbox-even" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Fenghuangopterus" title="Fenghuangopterus">Fenghuangopterus</a></i></li>
<li><i><a href="/enwiki/wiki/Jianchangnathus" title="Jianchangnathus">Jianchangnathus</a></i></li>
<li><i><a href="/enwiki/wiki/Jianchangopterus" title="Jianchangopterus">Jianchangopterus</a></i></li>
<li><i><a href="/enwiki/wiki/Scaphognathus" title="Scaphognathus">Scaphognathus</a></i></li>
<li><i><a href="/enwiki/wiki/Sordes" title="Sordes">Sordes</a></i>?</li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Pterodactylomorpha" class="mw-redirect" title="Pterodactylomorpha">Pterodactylomorpha</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><ul><li>see below↓</li></ul></li></ul>
</div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td><td class="noviewer navbox-image" rowspan="1" style="width:1px;padding:0px 0px 0px 2px"><div><span style="display: inline-block; -moz-transform: scaleX(-1); -ms-transform: scaleX(-1); -o-transform: scaleX(-1); -webkit-transform: scaleX(-1); transform: scaleX(-1);"><a href="/enwiki/wiki/File:Campylogn_DB.jpg" class="image" title="Campylognathoides liasicus"><img alt="Campylognathoides liasicus" src="/upwiki/wikipedia/commons/thumb/1/1a/Campylogn_DB.jpg/140px-Campylogn_DB.jpg" decoding="async" width="140" height="81" srcset="/upwiki/wikipedia/commons/thumb/1/1a/Campylogn_DB.jpg/210px-Campylogn_DB.jpg 1.5x, /upwiki/wikipedia/commons/thumb/1/1a/Campylogn_DB.jpg/280px-Campylogn_DB.jpg 2x" data-file-width="1000" data-file-height="580" /></a></span>
<p><span style="display: inline-block; -moz-transform: scaleX(-1); -ms-transform: scaleX(-1); -o-transform: scaleX(-1); -webkit-transform: scaleX(-1); transform: scaleX(-1);"><a href="/enwiki/wiki/File:ScaphognDB.jpg" class="image" title="Scaphognathus crassirostris"><img alt="Scaphognathus crassirostris" src="/upwiki/wikipedia/commons/thumb/0/06/ScaphognDB.jpg/140px-ScaphognDB.jpg" decoding="async" width="140" height="97" srcset="/upwiki/wikipedia/commons/thumb/0/06/ScaphognDB.jpg/210px-ScaphognDB.jpg 1.5x, /upwiki/wikipedia/commons/thumb/0/06/ScaphognDB.jpg/280px-ScaphognDB.jpg 2x" data-file-width="2284" data-file-height="1587" /></a></span>
</p>
<a href="/enwiki/wiki/File:Dorygn_DB.jpg" class="image" title="Dorygnathus banthensis"><img alt="Dorygnathus banthensis" src="/upwiki/wikipedia/commons/thumb/2/20/Dorygn_DB.jpg/140px-Dorygn_DB.jpg" decoding="async" width="140" height="106" srcset="/upwiki/wikipedia/commons/thumb/2/20/Dorygn_DB.jpg/210px-Dorygn_DB.jpg 1.5x, /upwiki/wikipedia/commons/thumb/2/20/Dorygn_DB.jpg/280px-Dorygn_DB.jpg 2x" data-file-width="1545" data-file-height="1173" /></a></div></td></tr></tbody></table><div></div></td></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks mw-collapsible mw-collapsed navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="col" class="navbox-title" colspan="3"><div id="Pterodactylomorpha" style="font-size:114%;margin:0 4em"><a href="/enwiki/wiki/Pterodactylomorpha" class="mw-redirect" title="Pterodactylomorpha">Pterodactylomorpha</a></div></th></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="Pterodactylomorpha" scope="row" class="navbox-group" style="width:1%"><div style="position:absolute;-webkit-transform:translate(-50%,-50%) rotate(270deg);-moz-transform:translate(-50%,-50%) rotate(270deg);-ms-transform:translate(-50%,-50%) rotate(270deg);-o-transform:translate(-50%,-50%) rotate(270deg);transform:translate(-50%,-50%) rotate(270deg);"><a href="/enwiki/wiki/Pterodactylomorpha" class="mw-redirect" title="Pterodactylomorpha">Pterodactylomorpha</a></div></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Allkaruen" title="Allkaruen">Allkaruen</a></i></li>
<li><i><a href="/enwiki/wiki/Sordes" title="Sordes">Sordes</a></i>?</li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><div style="position:absolute;-webkit-transform:translate(-50%,-50%) rotate(270deg);-moz-transform:translate(-50%,-50%) rotate(270deg);-ms-transform:translate(-50%,-50%) rotate(270deg);-o-transform:translate(-50%,-50%) rotate(270deg);transform:translate(-50%,-50%) rotate(270deg);"><a href="/enwiki/wiki/Monofenestrata" title="Monofenestrata">Monofenestrata</a></div></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Archaeoistiodactylus" title="Archaeoistiodactylus">Archaeoistiodactylus</a></i></li>
<li><i><a href="/enwiki/wiki/Normannognathus" title="Normannognathus">Normannognathus</a></i></li>
<li><b><a href="/enwiki/wiki/Anurognathidae" title="Anurognathidae">Anurognathidae</a></b>?</li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Darwinoptera" class="mw-redirect" title="Darwinoptera">Darwinoptera</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Pterorhynchus" title="Pterorhynchus">Pterorhynchus</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Wukongopteridae" title="Wukongopteridae">Wukongopteridae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Kunpengopterus" title="Kunpengopterus">Kunpengopterus</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Wukongopterinae" class="mw-redirect" title="Wukongopterinae">Wukongopterinae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Cuspicephalus" title="Cuspicephalus">Cuspicephalus</a></i></li>
<li><i><a href="/enwiki/wiki/Darwinopterus" title="Darwinopterus">Darwinopterus</a></i></li>
<li><i><a href="/enwiki/wiki/Wukongopterus" title="Wukongopterus">Wukongopterus</a></i></li></ul>
</div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><div style="position:absolute;-webkit-transform:translate(-50%,-50%) rotate(270deg);-moz-transform:translate(-50%,-50%) rotate(270deg);-ms-transform:translate(-50%,-50%) rotate(270deg);-o-transform:translate(-50%,-50%) rotate(270deg);transform:translate(-50%,-50%) rotate(270deg);"><a href="/enwiki/wiki/Pterodactyliformes" class="mw-redirect" title="Pterodactyliformes">Pterodactyliformes</a></div></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Changchengopterus" title="Changchengopterus">Changchengopterus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Douzhanopterus" title="Douzhanopterus">Douzhanopterus</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><div style="position:absolute;-webkit-transform:translate(-50%,-50%) rotate(270deg);-moz-transform:translate(-50%,-50%) rotate(270deg);-ms-transform:translate(-50%,-50%) rotate(270deg);-o-transform:translate(-50%,-50%) rotate(270deg);transform:translate(-50%,-50%) rotate(270deg);"><a href="/enwiki/wiki/Caelidracones" title="Caelidracones">Caelidracones</a></div></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Anurognathidae" title="Anurognathidae">Anurognathidae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Mesadactylus" title="Mesadactylus">Mesadactylus</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Anurognathinae" class="mw-redirect" title="Anurognathinae">Anurognathinae</a></th><td class="navbox-list navbox-even" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Anurognathus" title="Anurognathus">Anurognathus</a></i></li>
<li><i><a href="/enwiki/wiki/Dendrorhynchoides" title="Dendrorhynchoides">Dendrorhynchoides</a></i>?</li>
<li><i><a href="/enwiki/wiki/Jeholopterus" title="Jeholopterus">Jeholopterus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Luopterus" title="Luopterus">Luopterus</a></i></li>
<li><i><a href="/enwiki/wiki/Vesperopterylus" title="Vesperopterylus">Vesperopterylus</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Batrachognathinae" class="mw-redirect" title="Batrachognathinae">Batrachognathinae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Batrachognathus" title="Batrachognathus">Batrachognathus</a></i></li>
<li><i><a href="/enwiki/wiki/Dendrorhynchoides" title="Dendrorhynchoides">Dendrorhynchoides</a></i>?</li>
<li><i><a href="/enwiki/wiki/Jeholopterus" title="Jeholopterus">Jeholopterus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Sinomacrops" title="Sinomacrops">Sinomacrops</a></i></li></ul>
</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Pterodactyloidea" title="Pterodactyloidea">Pterodactyloidea</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Dermodactylus" title="Dermodactylus">Dermodactylus</a></i></li>
<li><i><a href="/enwiki/wiki/Eurolimnornis" title="Eurolimnornis">Eurolimnornis</a></i></li>
<li><i><a href="/enwiki/wiki/Herbstosaurus" title="Herbstosaurus">Herbstosaurus</a></i></li>
<li><i><a href="/enwiki/wiki/Kryptodrakon" title="Kryptodrakon">Kryptodrakon</a></i></li>
<li><i><a href="/enwiki/wiki/Ningchengopterus" title="Ningchengopterus">Ningchengopterus</a></i></li>
<li><i><a href="/enwiki/wiki/Pangupterus" title="Pangupterus">Pangupterus</a></i></li>
<li><i><a href="/enwiki/wiki/Samrukia" title="Samrukia">Samrukia</a></i>?</li>
<li><i><a href="/enwiki/wiki/Wenupteryx" title="Wenupteryx">Wenupteryx</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Lophocratia" class="mw-redirect" title="Lophocratia">Lophocratia</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><ul><li>see below↓</li></ul></li></ul>
</div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td><td class="noviewer navbox-image" rowspan="1" style="width:1px;padding:0px 0px 0px 2px"><div><a href="/enwiki/wiki/File:Jeholopterus_BW.jpg" class="image" title="Jeholopterus ninchengensis"><img alt="Jeholopterus ninchengensis" src="/upwiki/wikipedia/commons/thumb/b/b2/Jeholopterus_BW.jpg/140px-Jeholopterus_BW.jpg" decoding="async" width="140" height="88" srcset="/upwiki/wikipedia/commons/thumb/b/b2/Jeholopterus_BW.jpg/210px-Jeholopterus_BW.jpg 1.5x, /upwiki/wikipedia/commons/thumb/b/b2/Jeholopterus_BW.jpg/280px-Jeholopterus_BW.jpg 2x" data-file-width="800" data-file-height="502" /></a>
<a href="/enwiki/wiki/File:Kryptodrakon.jpeg" class="image" title="Kryptodrakon progenitor"><img alt="Kryptodrakon progenitor" src="/upwiki/wikipedia/commons/thumb/3/37/Kryptodrakon.jpeg/140px-Kryptodrakon.jpeg" decoding="async" width="140" height="118" srcset="/upwiki/wikipedia/commons/thumb/3/37/Kryptodrakon.jpeg/210px-Kryptodrakon.jpeg 1.5x, /upwiki/wikipedia/commons/thumb/3/37/Kryptodrakon.jpeg/280px-Kryptodrakon.jpeg 2x" data-file-width="1500" data-file-height="1265" /></a></div></td></tr></tbody></table><div></div></td></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks mw-collapsible uncollapsed navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="col" class="navbox-title" colspan="3"><div id="Lophocratia" style="font-size:114%;margin:0 4em"><a href="/enwiki/wiki/Lophocratia" class="mw-redirect" title="Lophocratia">Lophocratia</a></div></th></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="Archaeopterodactyloidea" scope="row" class="navbox-group" style="width:1%"><div style="position:absolute;-webkit-transform:translate(-50%,-50%) rotate(270deg);-moz-transform:translate(-50%,-50%) rotate(270deg);-ms-transform:translate(-50%,-50%) rotate(270deg);-o-transform:translate(-50%,-50%) rotate(270deg);transform:translate(-50%,-50%) rotate(270deg);"><a href="/enwiki/wiki/Archaeopterodactyloidea" title="Archaeopterodactyloidea">Archaeopterodactyloidea</a></div></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Prejanopterus" title="Prejanopterus">Prejanopterus</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Germanodactylidae" title="Germanodactylidae">Germanodactylidae</a></th><td class="navbox-list navbox-even" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Altmuehlopterus" title="Altmuehlopterus">Altmuehlopterus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Germanodactylus" title="Germanodactylus">Germanodactylus</a></i></li>
<li><i><a href="/enwiki/wiki/Normannognathus" title="Normannognathus">Normannognathus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Tendaguripterus" title="Tendaguripterus">Tendaguripterus</a></i>?</li></ul>
</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><div style="position:absolute;-webkit-transform:translate(-50%,-50%) rotate(270deg);-moz-transform:translate(-50%,-50%) rotate(270deg);-ms-transform:translate(-50%,-50%) rotate(270deg);-o-transform:translate(-50%,-50%) rotate(270deg);transform:translate(-50%,-50%) rotate(270deg);"><a href="/enwiki/wiki/Euctenochasmatia" title="Euctenochasmatia">Euctenochasmatia</a></div></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Diopecephalus" title="Diopecephalus">Diopecephalus</a></i></li>
<li><i><a class="mw-selflink selflink">Pterodactylus</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><div style="position:absolute;-webkit-transform:translate(-50%,-50%) rotate(270deg);-moz-transform:translate(-50%,-50%) rotate(270deg);-ms-transform:translate(-50%,-50%) rotate(270deg);-o-transform:translate(-50%,-50%) rotate(270deg);transform:translate(-50%,-50%) rotate(270deg);"><a href="/enwiki/wiki/Ctenochasmatoidea" title="Ctenochasmatoidea">Ctenochasmatoidea</a></div></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Gallodactylidae" title="Gallodactylidae">Gallodactylidae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Cycnorhamphus" title="Cycnorhamphus">Cycnorhamphus</a></i></li>
<li><i><a href="/enwiki/wiki/Normannognathus" title="Normannognathus">Normannognathus</a></i>?</li>
<li><b><a href="/enwiki/wiki/Moganopterinae" class="mw-redirect" title="Moganopterinae">Moganopterinae</a></b>?</li></ul>
</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Aurorazhdarchia" class="mw-redirect" title="Aurorazhdarchia">Aurorazhdarchia</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Aerodactylus" title="Aerodactylus">Aerodactylus</a></i></li>
<li><b><a href="/enwiki/wiki/Gallodactylidae" title="Gallodactylidae">Gallodactylidae</a></b>?</li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Aurorazhdarchidae" class="mw-redirect" title="Aurorazhdarchidae">Aurorazhdarchidae</a></th><td class="navbox-list navbox-even" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Ardeadactylus" title="Ardeadactylus">Ardeadactylus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Aurorazhdarcho" title="Aurorazhdarcho">Aurorazhdarcho</a></i></li>
<li><i><a href="/enwiki/wiki/Huanhepterus" title="Huanhepterus">Huanhepterus</a></i>?</li></ul>
</div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><div style="position:absolute;-webkit-transform:translate(-50%,-50%) rotate(270deg);-moz-transform:translate(-50%,-50%) rotate(270deg);-ms-transform:translate(-50%,-50%) rotate(270deg);-o-transform:translate(-50%,-50%) rotate(270deg);transform:translate(-50%,-50%) rotate(270deg);"><a href="/enwiki/wiki/Ctenochasmatidae" title="Ctenochasmatidae">Ctenochasmatidae</a></div></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Ardeadactylus" title="Ardeadactylus">Ardeadactylus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Cathayopterus" title="Cathayopterus">Cathayopterus</a></i></li>
<li><i><a href="/enwiki/wiki/Elanodactylus" title="Elanodactylus">Elanodactylus</a></i></li>
<li><i><a href="/enwiki/wiki/Forfexopterus" title="Forfexopterus">Forfexopterus</a></i></li>
<li><i><a href="/enwiki/wiki/Gladocephaloideus" title="Gladocephaloideus">Gladocephaloideus</a></i></li>
<li><i><a href="/enwiki/wiki/Kepodactylus" title="Kepodactylus">Kepodactylus</a></i></li>
<li><i><a href="/enwiki/wiki/Liaodactylus" title="Liaodactylus">Liaodactylus</a></i></li>
<li><i><a href="/enwiki/wiki/Otogopterus" title="Otogopterus">Otogopterus</a></i></li>
<li><i><a href="/enwiki/wiki/Pterofiltrus" title="Pterofiltrus">Pterofiltrus</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Moganopterinae" class="mw-redirect" title="Moganopterinae">Moganopterinae</a>?</th><td class="navbox-list navbox-even" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Feilongus" title="Feilongus">Feilongus</a></i></li>
<li><i><a href="/enwiki/wiki/Moganopterus" title="Moganopterus">Moganopterus</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Gnathosaurinae" class="mw-redirect" title="Gnathosaurinae">Gnathosaurinae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Gnathosaurus" title="Gnathosaurus">Gnathosaurus</a></i></li>
<li><i><a href="/enwiki/wiki/Huanhepterus" title="Huanhepterus">Huanhepterus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Plataleorhynchus" title="Plataleorhynchus">Plataleorhynchus</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Ctenochasmatinae" class="mw-redirect" title="Ctenochasmatinae">Ctenochasmatinae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Ctenochasma" title="Ctenochasma">Ctenochasma</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Pterodaustrini" class="mw-redirect" title="Pterodaustrini">Pterodaustrini</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Beipiaopterus" title="Beipiaopterus">Beipiaopterus</a></i></li>
<li><i><a href="/enwiki/wiki/Eosipterus" title="Eosipterus">Eosipterus</a></i></li>
<li><i><a href="/enwiki/wiki/Gegepterus" title="Gegepterus">Gegepterus</a></i></li>
<li><i><a href="/enwiki/wiki/Pterodaustro" title="Pterodaustro">Pterodaustro</a></i></li></ul>
</div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="Eupterodactyloidea" scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Eupterodactyloidea" title="Eupterodactyloidea">Eupterodactyloidea</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Altmuehlopterus" title="Altmuehlopterus">Altmuehlopterus</a></i>?</li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Ornithocheiroidea" title="Ornithocheiroidea">Ornithocheiroidea</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><ul><li>see below↓</li></ul></li></ul>
</div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td><td class="noviewer navbox-image" rowspan="1" style="width:1px;padding:0px 0px 0px 2px"><div><span style="display: inline-block; -moz-transform: scaleX(-1); -ms-transform: scaleX(-1); -o-transform: scaleX(-1); -webkit-transform: scaleX(-1); transform: scaleX(-1);"><a href="/enwiki/wiki/File:Pterodactylus_BMMS7_life.png" class="image" title="Pterodactylus antiquus"><img alt="Pterodactylus antiquus" src="/upwiki/wikipedia/commons/thumb/a/a0/Pterodactylus_BMMS7_life.png/140px-Pterodactylus_BMMS7_life.png" decoding="async" width="140" height="66" srcset="/upwiki/wikipedia/commons/thumb/a/a0/Pterodactylus_BMMS7_life.png/210px-Pterodactylus_BMMS7_life.png 1.5x, /upwiki/wikipedia/commons/thumb/a/a0/Pterodactylus_BMMS7_life.png/280px-Pterodactylus_BMMS7_life.png 2x" data-file-width="931" data-file-height="438" /></a></span>
<a href="/enwiki/wiki/File:Plataleorhynchus.jpg" class="image" title="Plataleorhynchus streptorophorodon"><img alt="Plataleorhynchus streptorophorodon" src="/upwiki/wikipedia/commons/thumb/2/24/Plataleorhynchus.jpg/140px-Plataleorhynchus.jpg" decoding="async" width="140" height="78" srcset="/upwiki/wikipedia/commons/thumb/2/24/Plataleorhynchus.jpg/210px-Plataleorhynchus.jpg 1.5x, /upwiki/wikipedia/commons/thumb/2/24/Plataleorhynchus.jpg/280px-Plataleorhynchus.jpg 2x" data-file-width="1500" data-file-height="831" /></a></div></td></tr></tbody></table><div></div></td></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks mw-collapsible mw-collapsed navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="col" class="navbox-title" colspan="3"><div id="Ornithocheiroidea" style="font-size:114%;margin:0 4em"><a href="/enwiki/wiki/Ornithocheiroidea" title="Ornithocheiroidea">Ornithocheiroidea</a></div></th></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="Ornithocheiroidea" scope="row" class="navbox-group" style="width:1%"><div style="position:absolute;-webkit-transform:translate(-50%,-50%) rotate(270deg);-moz-transform:translate(-50%,-50%) rotate(270deg);-ms-transform:translate(-50%,-50%) rotate(270deg);-o-transform:translate(-50%,-50%) rotate(270deg);transform:translate(-50%,-50%) rotate(270deg);"><a href="/enwiki/wiki/Ornithocheiroidea" title="Ornithocheiroidea">Ornithocheiroidea</a></div></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Piksi" title="Piksi">Piksi</a></i>?</li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><div style="position:absolute;-webkit-transform:translate(-50%,-50%) rotate(270deg);-moz-transform:translate(-50%,-50%) rotate(270deg);-ms-transform:translate(-50%,-50%) rotate(270deg);-o-transform:translate(-50%,-50%) rotate(270deg);transform:translate(-50%,-50%) rotate(270deg);"><a href="/enwiki/wiki/Tapejaroidea" title="Tapejaroidea">Tapejaroidea</a>?</div></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><b><a href="/enwiki/wiki/Dsungaripteridae" title="Dsungaripteridae">Dsungaripteridae</a></b>?</li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><div style="position:absolute;-webkit-transform:translate(-50%,-50%) rotate(270deg);-moz-transform:translate(-50%,-50%) rotate(270deg);-ms-transform:translate(-50%,-50%) rotate(270deg);-o-transform:translate(-50%,-50%) rotate(270deg);transform:translate(-50%,-50%) rotate(270deg);"><a href="/enwiki/wiki/Azhdarchoidea" title="Azhdarchoidea">Azhdarchoidea</a></div></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Argentinadraco" title="Argentinadraco">Argentinadraco</a></i></li>
<li><i><a href="/enwiki/wiki/Leptostomia" title="Leptostomia">Leptostomia</a></i>?</li>
<li><i><a href="/enwiki/wiki/Montanazhdarcho" title="Montanazhdarcho">Montanazhdarcho</a></i></li>
<li><i><a href="/enwiki/wiki/Ornithostoma" title="Ornithostoma">Ornithostoma</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><div style="position:absolute;-webkit-transform:translate(-50%,-50%) rotate(270deg);-moz-transform:translate(-50%,-50%) rotate(270deg);-ms-transform:translate(-50%,-50%) rotate(270deg);-o-transform:translate(-50%,-50%) rotate(270deg);transform:translate(-50%,-50%) rotate(270deg);"><a href="/enwiki/wiki/Tapejaromorpha" title="Tapejaromorpha">Tapejaromorpha</a></div></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Bennettazhia" title="Bennettazhia">Bennettazhia</a></i></li>
<li><i><a href="/enwiki/wiki/Keresdrakon" title="Keresdrakon">Keresdrakon</a></i></li>
<li><i><a href="/enwiki/wiki/Nemicolopterus" title="Nemicolopterus">Nemicolopterus</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Tapejaridae" title="Tapejaridae">Tapejaridae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Afrotapejara" title="Afrotapejara">Afrotapejara</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Thalassodrominae" title="Thalassodrominae">Thalassodrominae</a>?</th><td class="navbox-list navbox-even" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Aerotitan" title="Aerotitan">Aerotitan</a></i>?</li>
<li><i><a href="/enwiki/wiki/Alanqa" title="Alanqa">Alanqa</a></i>?</li>
<li><i><a href="/enwiki/wiki/Banguela" title="Banguela">Banguela</a></i>?</li>
<li><i><a href="/enwiki/wiki/Lacusovagus" title="Lacusovagus">Lacusovagus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Thalassodromeus" title="Thalassodromeus">Thalassodromeus</a></i></li>
<li><i><a href="/enwiki/wiki/Tupuxuara" title="Tupuxuara">Tupuxuara</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Sinopterinae" class="mw-redirect" title="Sinopterinae">Sinopterinae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Bakonydraco" title="Bakonydraco">Bakonydraco</a></i>?</li>
<li><i><a href="/enwiki/wiki/Eopteranodon" title="Eopteranodon">Eopteranodon</a></i>?</li>
<li><i><a href="/enwiki/wiki/Huaxiapterus" title="Huaxiapterus">Huaxiapterus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Sinopterus" title="Sinopterus">Sinopterus</a></i></li>
<li><i><a href="/enwiki/wiki/Wightia_declivirostris" title="Wightia declivirostris">Wightia</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Tapejarinae" class="mw-redirect" title="Tapejarinae">Tapejarinae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Aymberedactylus" title="Aymberedactylus">Aymberedactylus</a></i></li>
<li><i><a href="/enwiki/wiki/Caupedactylus" title="Caupedactylus">Caupedactylus</a></i></li>
<li><i><a href="/enwiki/wiki/Eopteranodon" title="Eopteranodon">Eopteranodon</a></i>?</li>
<li><i><a href="/enwiki/wiki/Huaxiapterus" title="Huaxiapterus">Huaxiapterus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Sinopterus" title="Sinopterus">Sinopterus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Vectidraco" title="Vectidraco">Vectidraco</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Tapejarini" class="mw-redirect" title="Tapejarini">Tapejarini</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Bakonydraco" title="Bakonydraco">Bakonydraco</a></i>?</li>
<li><i><a href="/enwiki/wiki/Caiuajara" title="Caiuajara">Caiuajara</a></i></li>
<li><i><a href="/enwiki/wiki/Europejara" title="Europejara">Europejara</a></i></li>
<li><i><a href="/enwiki/wiki/Tapejara_wellnhoferi" title="Tapejara wellnhoferi">Tapejara</a></i></li>
<li><i><a href="/enwiki/wiki/Tupandactylus" title="Tupandactylus">Tupandactylus</a></i></li></ul>
</div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><div style="position:absolute;-webkit-transform:translate(-50%,-50%) rotate(270deg);-moz-transform:translate(-50%,-50%) rotate(270deg);-ms-transform:translate(-50%,-50%) rotate(270deg);-o-transform:translate(-50%,-50%) rotate(270deg);transform:translate(-50%,-50%) rotate(270deg);"><a href="/enwiki/wiki/Neoazhdarchia" class="mw-redirect" title="Neoazhdarchia">Neoazhdarchia</a></div></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Cretornis" title="Cretornis">Cretornis</a></i>?</li>
<li><i><a href="/enwiki/wiki/Microtuban" title="Microtuban">Microtuban</a></i></li>
<li><i><a href="/enwiki/wiki/Xericeps" title="Xericeps">Xericeps</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Dsungaripteromorpha" class="mw-redirect" title="Dsungaripteromorpha">Dsungaripteromorpha</a>?</th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><b><a href="/enwiki/wiki/Thalassodrominae" title="Thalassodrominae">Thalassodromidae</a></b>/<b><a href="/enwiki/wiki/Thalassodrominae" title="Thalassodrominae">Thalassodrominae</a></b>?</li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Dsungaripteridae" title="Dsungaripteridae">Dsungaripteridae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Banguela" title="Banguela">Banguela</a></i>?</li>
<li><i><a href="/enwiki/wiki/Lonchognathosaurus" title="Lonchognathosaurus">Lonchognathosaurus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Noripterus" title="Noripterus">Noripterus</a></i></li>
<li><i><a href="/enwiki/wiki/Ordosipterus" title="Ordosipterus">Ordosipterus</a></i></li>
<li><i><a href="/enwiki/wiki/Puntanipterus" title="Puntanipterus">Puntanipterus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Tendaguripterus" title="Tendaguripterus">Tendaguripterus</a></i>?</li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Dsungaripterinae" class="mw-redirect" title="Dsungaripterinae">Dsungaripterinae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Domeykodactylus" title="Domeykodactylus">Domeykodactylus</a></i></li>
<li><i><a href="/enwiki/wiki/Dsungaripterus" title="Dsungaripterus">Dsungaripterus</a></i></li></ul>
</div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Neopterodactyloidea" class="mw-redirect" title="Neopterodactyloidea">Neopterodactyloidea</a></th><td class="navbox-list navbox-even" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Eoazhdarcho" title="Eoazhdarcho">Eoazhdarcho</a></i></li>
<li><i><a href="/enwiki/wiki/Radiodactylus" title="Radiodactylus">Radiodactylus</a></i></li></ul>
</div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Chaoyangopteridae" title="Chaoyangopteridae">Chaoyangopteridae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Apatorhamphus" title="Apatorhamphus">Apatorhamphus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Eoazhdarcho" title="Eoazhdarcho">Eoazhdarcho</a></i>?</li>
<li><i><a href="/enwiki/wiki/Lacusovagus" title="Lacusovagus">Lacusovagus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Microtuban" title="Microtuban">Microtuban</a></i>?</li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Chaoyangopterinae" class="mw-redirect" title="Chaoyangopterinae">Chaoyangopterinae</a></th><td class="navbox-list navbox-even" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Chaoyangopterus" title="Chaoyangopterus">Chaoyangopterus</a></i></li>
<li><i><a href="/enwiki/wiki/Jidapterus" title="Jidapterus">Jidapterus</a></i></li>
<li><i><a href="/enwiki/wiki/Shenzhoupterus" title="Shenzhoupterus">Shenzhoupterus</a></i></li></ul>
</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Azhdarchidae" title="Azhdarchidae">Azhdarchidae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Aerotitan" title="Aerotitan">Aerotitan</a></i>?</li>
<li><i><a href="/enwiki/wiki/Alanqa" title="Alanqa">Alanqa</a></i>?</li>
<li><i><a href="/enwiki/wiki/Albadraco" title="Albadraco">Albadraco</a></i></li>
<li><i><a href="/enwiki/wiki/Aralazhdarcho" title="Aralazhdarcho">Aralazhdarcho</a></i></li>
<li><i><a href="/enwiki/wiki/Azhdarcho" title="Azhdarcho">Azhdarcho</a></i></li>
<li><i><a href="/enwiki/wiki/Bogolubovia" title="Bogolubovia">Bogolubovia</a></i></li>
<li><i><a href="/enwiki/wiki/Cryodrakon" title="Cryodrakon">Cryodrakon</a></i></li>
<li><i><a href="/enwiki/wiki/Eurazhdarcho" title="Eurazhdarcho">Eurazhdarcho</a></i></li>
<li><i><a href="/enwiki/wiki/Mistralazhdarcho" title="Mistralazhdarcho">Mistralazhdarcho</a></i></li>
<li><i><a href="/enwiki/wiki/Montanazhdarcho" title="Montanazhdarcho">Montanazhdarcho</a></i>?</li>
<li><i><a href="/enwiki/wiki/Navajodactylus" title="Navajodactylus">Navajodactylus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Palaeocursornis" title="Palaeocursornis">Palaeocursornis</a></i></li>
<li><i><a href="/enwiki/wiki/Phosphatodraco" title="Phosphatodraco">Phosphatodraco</a></i></li>
<li><i><a href="/enwiki/wiki/Tethydraco" title="Tethydraco">Tethydraco</a></i>?</li>
<li><i><a href="/enwiki/wiki/Volgadraco" title="Volgadraco">Volgadraco</a></i>?</li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Quetzalcoatlinae" class="mw-redirect" title="Quetzalcoatlinae">Quetzalcoatlinae</a></th><td class="navbox-list navbox-even" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Arambourgiania" title="Arambourgiania">Arambourgiania</a></i></li>
<li><i><a href="/enwiki/wiki/Hatzegopteryx" title="Hatzegopteryx">Hatzegopteryx</a></i></li>
<li><i><a href="/enwiki/wiki/Quetzalcoatlus" title="Quetzalcoatlus">Quetzalcoatlus</a></i></li>
<li><i><a href="/enwiki/wiki/Zhejiangopterus" title="Zhejiangopterus">Zhejiangopterus</a></i></li></ul>
</div></td></tr></tbody></table><div></div></td></tr></tbody></table><div>
</div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="Pteranodontoidea" scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Pteranodontoidea" title="Pteranodontoidea">Pteranodontoidea</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><ul><li>see below↓</li></ul></li></ul>
</div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td><td class="noviewer navbox-image" rowspan="1" style="width:1px;padding:0px 0px 0px 2px"><div><span style="display: inline-block; -moz-transform: scaleX(-1); -ms-transform: scaleX(-1); -o-transform: scaleX(-1); -webkit-transform: scaleX(-1); transform: scaleX(-1);"><a href="/enwiki/wiki/File:Bakonydraco_as_tapejarid_DB.jpg" class="image" title="Bakonydraco galaczi"><img alt="Bakonydraco galaczi" src="/upwiki/wikipedia/commons/thumb/0/0c/Bakonydraco_as_tapejarid_DB.jpg/140px-Bakonydraco_as_tapejarid_DB.jpg" decoding="async" width="140" height="101" srcset="/upwiki/wikipedia/commons/thumb/0/0c/Bakonydraco_as_tapejarid_DB.jpg/210px-Bakonydraco_as_tapejarid_DB.jpg 1.5x, /upwiki/wikipedia/commons/thumb/0/0c/Bakonydraco_as_tapejarid_DB.jpg/280px-Bakonydraco_as_tapejarid_DB.jpg 2x" data-file-width="1684" data-file-height="1214" /></a></span>
<p><span style="display: inline-block; -moz-transform: scaleX(-1); -ms-transform: scaleX(-1); -o-transform: scaleX(-1); -webkit-transform: scaleX(-1); transform: scaleX(-1);"><a href="/enwiki/wiki/File:TapimpDB.jpg" class="image" title="Tupandactylus imperator"><img alt="Tupandactylus imperator" src="/upwiki/wikipedia/commons/thumb/c/c3/TapimpDB.jpg/140px-TapimpDB.jpg" decoding="async" width="140" height="65" srcset="/upwiki/wikipedia/commons/thumb/c/c3/TapimpDB.jpg/210px-TapimpDB.jpg 1.5x, /upwiki/wikipedia/commons/thumb/c/c3/TapimpDB.jpg/280px-TapimpDB.jpg 2x" data-file-width="2262" data-file-height="1058" /></a></span>
</p>
<span style="display: inline-block; -moz-transform: scaleX(-1); -ms-transform: scaleX(-1); -o-transform: scaleX(-1); -webkit-transform: scaleX(-1); transform: scaleX(-1);"><a href="/enwiki/wiki/File:Quetzalcoatlus07.jpg" class="image" title="Quetzalcoatlus"><img alt="Quetzalcoatlus" src="/upwiki/wikipedia/commons/thumb/4/42/Quetzalcoatlus07.jpg/140px-Quetzalcoatlus07.jpg" decoding="async" width="140" height="85" srcset="/upwiki/wikipedia/commons/thumb/4/42/Quetzalcoatlus07.jpg/210px-Quetzalcoatlus07.jpg 1.5x, /upwiki/wikipedia/commons/thumb/4/42/Quetzalcoatlus07.jpg/280px-Quetzalcoatlus07.jpg 2x" data-file-width="900" data-file-height="545" /></a></span></div></td></tr></tbody></table><div></div></td></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks mw-collapsible mw-collapsed navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="col" class="navbox-title" colspan="3"><div id="Pteranodontoidea" style="font-size:114%;margin:0 4em"><a href="/enwiki/wiki/Pteranodontoidea" title="Pteranodontoidea">Pteranodontoidea</a></div></th></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="Pteranodontoidea" scope="row" class="navbox-group" style="width:1%"><div style="position:absolute;-webkit-transform:translate(-50%,-50%) rotate(270deg);-moz-transform:translate(-50%,-50%) rotate(270deg);-ms-transform:translate(-50%,-50%) rotate(270deg);-o-transform:translate(-50%,-50%) rotate(270deg);transform:translate(-50%,-50%) rotate(270deg);"><a href="/enwiki/wiki/Pteranodontoidea" title="Pteranodontoidea">Pteranodontoidea</a></div></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Santanadactylus" title="Santanadactylus">Santanadactylus</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Pteranodontia" title="Pteranodontia">Pteranodontia</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Pteranodontidae" title="Pteranodontidae">Pteranodontidae</a></th><td class="navbox-list navbox-even" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Dawndraco" title="Dawndraco">Dawndraco</a></i></li>
<li><i><a href="/enwiki/wiki/Geosternbergia" title="Geosternbergia">Geosternbergia</a></i></li>
<li><i><a href="/enwiki/wiki/Pteranodon" title="Pteranodon">Pteranodon</a></i></li>
<li><i><a href="/enwiki/wiki/Tethydraco" title="Tethydraco">Tethydraco</a></i>?</li>
<li><i><a href="/enwiki/wiki/Volgadraco" title="Volgadraco">Volgadraco</a></i>?</li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Nyctosauridae" title="Nyctosauridae">Nyctosauridae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Alamodactylus" title="Alamodactylus">Alamodactylus</a></i></li>
<li><i><a href="/enwiki/wiki/Alcione_elainus" title="Alcione elainus">Alcione</a></i></li>
<li><i><a href="/enwiki/wiki/Barbaridactylus" title="Barbaridactylus">Barbaridactylus</a></i></li>
<li><i><a href="/enwiki/wiki/Cretornis" title="Cretornis">Cretornis</a></i>?</li>
<li><i><a href="/enwiki/wiki/Muzquizopteryx" title="Muzquizopteryx">Muzquizopteryx</a></i></li>
<li><i><a href="/enwiki/wiki/Nyctosaurus" title="Nyctosaurus">Nyctosaurus</a></i></li>
<li><i><a href="/enwiki/wiki/Simurghia" title="Simurghia">Simurghia</a></i></li>
<li><i><a href="/enwiki/wiki/Volgadraco" title="Volgadraco">Volgadraco</a></i>?</li></ul>
</div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><div style="position:absolute;-webkit-transform:translate(-50%,-50%) rotate(270deg);-moz-transform:translate(-50%,-50%) rotate(270deg);-ms-transform:translate(-50%,-50%) rotate(270deg);-o-transform:translate(-50%,-50%) rotate(270deg);transform:translate(-50%,-50%) rotate(270deg);"><a href="/enwiki/wiki/Ornithocheiromorpha" title="Ornithocheiromorpha">Ornithocheiromorpha</a></div></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Serradraco" title="Serradraco">Serradraco</a></i></li>
<li><i><a href="/enwiki/wiki/Unwindia" title="Unwindia">Unwindia</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Lonchodectidae" title="Lonchodectidae">Lonchodectidae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Ikrandraco" title="Ikrandraco">Ikrandraco</a></i>?</li>
<li><i><a href="/enwiki/wiki/Lonchodectes" title="Lonchodectes">Lonchodectes</a></i></li>
<li><i><a href="/enwiki/wiki/Lonchodraco" title="Lonchodraco">Lonchodraco</a></i>?</li></ul>
</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><div style="position:absolute;-webkit-transform:translate(-50%,-50%) rotate(270deg);-moz-transform:translate(-50%,-50%) rotate(270deg);-ms-transform:translate(-50%,-50%) rotate(270deg);-o-transform:translate(-50%,-50%) rotate(270deg);transform:translate(-50%,-50%) rotate(270deg);"><a href="/enwiki/wiki/Lanceodontia" class="mw-redirect" title="Lanceodontia">Lanceodontia</a></div></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Lonchodectidae" title="Lonchodectidae">Lonchodraconidae</a></th><td class="navbox-list navbox-even" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Ikrandraco" title="Ikrandraco">Ikrandraco</a></i></li>
<li><i><a href="/enwiki/wiki/Lonchodraco" title="Lonchodraco">Lonchodraco</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Istiodactyliformes" class="mw-redirect" title="Istiodactyliformes">Istiodactyliformes</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Hongshanopterus" title="Hongshanopterus">Hongshanopterus</a></i></li>
<li><i><a href="/enwiki/wiki/Linlongopterus" title="Linlongopterus">Linlongopterus</a></i></li>
<li><i><a href="/enwiki/wiki/Yixianopterus" title="Yixianopterus">Yixianopterus</a></i></li>
<li><b><a href="/enwiki/wiki/Lonchodectidae" title="Lonchodectidae">Lonchodraconidae</a></b>?</li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Mimodactylidae" title="Mimodactylidae">Mimodactylidae</a></th><td class="navbox-list navbox-even" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Haopterus" title="Haopterus">Haopterus</a></i></li>
<li><i><a href="/enwiki/wiki/Mimodactylus" title="Mimodactylus">Mimodactylus</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Istiodactylidae" title="Istiodactylidae">Istiodactylidae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Longchengpterus" title="Longchengpterus">Longchengpterus</a></i></li>
<li><i><a href="/enwiki/wiki/Luchibang" title="Luchibang">Luchibang</a></i></li>
<li><i><a href="/enwiki/wiki/Nurhachius" title="Nurhachius">Nurhachius</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Istiodactylinae" class="mw-redirect" title="Istiodactylinae">Istiodactylinae</a></th><td class="navbox-list navbox-even" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Istiodactylus" title="Istiodactylus">Istiodactylus</a></i></li>
<li><i><a href="/enwiki/wiki/Liaoxipterus" title="Liaoxipterus">Liaoxipterus</a></i></li></ul>
</div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Boreopteridae" title="Boreopteridae">Boreopteridae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Boreopterus" title="Boreopterus">Boreopterus</a></i></li>
<li><i><a href="/enwiki/wiki/Zhenyuanopterus" title="Zhenyuanopterus">Zhenyuanopterus</a></i></li></ul>
</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><div style="position:absolute;-webkit-transform:translate(-50%,-50%) rotate(270deg);-moz-transform:translate(-50%,-50%) rotate(270deg);-ms-transform:translate(-50%,-50%) rotate(270deg);-o-transform:translate(-50%,-50%) rotate(270deg);transform:translate(-50%,-50%) rotate(270deg);"><a href="/enwiki/wiki/Ornithocheirae" title="Ornithocheirae">Ornithocheirae</a></div></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Ornithocheiridae" title="Ornithocheiridae">Ornithocheiridae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Araripesaurus" title="Araripesaurus">Araripesaurus</a></i></li>
<li><i><a href="/enwiki/wiki/Arthurdactylus" title="Arthurdactylus">Arthurdactylus</a></i></li>
<li><i><a href="/enwiki/wiki/Camposipterus" title="Camposipterus">Camposipterus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Caulkicephalus" title="Caulkicephalus">Caulkicephalus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Cimoliopterus" title="Cimoliopterus">Cimoliopterus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Tropeognathus" title="Tropeognathus">Tropeognathus</a></i>?</li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Ornithocheirinae" class="mw-redirect" title="Ornithocheirinae">Ornithocheirinae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Coloborhynchus" title="Coloborhynchus">Coloborhynchus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Ferrodraco" title="Ferrodraco">Ferrodraco</a></i>?</li>
<li><i><a href="/enwiki/wiki/Mythunga" title="Mythunga">Mythunga</a></i>?</li>
<li><i><a href="/enwiki/wiki/Ornithocheirus" title="Ornithocheirus">Ornithocheirus</a></i></li>
<li><i><a href="/enwiki/wiki/Siroccopteryx" title="Siroccopteryx">Siroccopteryx</a></i>?</li>
<li><i><a href="/enwiki/wiki/Uktenadactylus" title="Uktenadactylus">Uktenadactylus</a></i>?</li></ul>
</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Targaryendraconia" title="Targaryendraconia">Targaryendraconia</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Cimoliopteridae" class="mw-redirect" title="Cimoliopteridae">Cimoliopteridae</a></th><td class="navbox-list navbox-even" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Aetodactylus" title="Aetodactylus">Aetodactylus</a></i></li>
<li><i><a href="/enwiki/wiki/Camposipterus" title="Camposipterus">Camposipterus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Cimoliopterus" title="Cimoliopterus">Cimoliopterus</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Targaryendraconidae" class="mw-redirect" title="Targaryendraconidae">Targaryendraconidae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Aussiedraco" title="Aussiedraco">Aussiedraco</a></i></li>
<li><i><a href="/enwiki/wiki/Barbosania" title="Barbosania">Barbosania</a></i></li>
<li><i><a href="/enwiki/wiki/Targaryendraco" title="Targaryendraco">Targaryendraco</a></i></li></ul>
</div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><div style="position:absolute;-webkit-transform:translate(-50%,-50%) rotate(270deg);-moz-transform:translate(-50%,-50%) rotate(270deg);-ms-transform:translate(-50%,-50%) rotate(270deg);-o-transform:translate(-50%,-50%) rotate(270deg);transform:translate(-50%,-50%) rotate(270deg);"><a href="/enwiki/wiki/Anhangueria" title="Anhangueria">Anhangueria</a></div></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Brasileodactylus" title="Brasileodactylus">Brasileodactylus</a></i></li>
<li><b><a href="/enwiki/wiki/Ornithocheiridae" title="Ornithocheiridae">Ornithocheiridae</a></b>?</li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Hamipteridae" title="Hamipteridae">Hamipteridae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Hamipterus" title="Hamipterus">Hamipterus</a></i></li>
<li><i><a href="/enwiki/wiki/Iberodactylus" title="Iberodactylus">Iberodactylus</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Anhangueridae" title="Anhangueridae">Anhangueridae</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Tropeognathinae" class="mw-redirect" title="Tropeognathinae">Tropeognathinae</a>?</th><td class="navbox-list navbox-even" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Ferrodraco" title="Ferrodraco">Ferrodraco</a></i>?</li>
<li><i><a href="/enwiki/wiki/Mythunga" title="Mythunga">Mythunga</a></i>?</li>
<li><i><a href="/enwiki/wiki/Siroccopteryx" title="Siroccopteryx">Siroccopteryx</a></i>?</li>
<li><i><a href="/enwiki/wiki/Tropeognathus" title="Tropeognathus">Tropeognathus</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Coloborhynchinae" class="mw-redirect" title="Coloborhynchinae">Coloborhynchinae</a>?</th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Aerodraco" title="Aerodraco">Aerodraco</a></i></li>
<li><i><a href="/enwiki/wiki/Coloborhynchus" title="Coloborhynchus">Coloborhynchus</a></i></li>
<li><i><a href="/enwiki/wiki/Nicorhynchus" title="Nicorhynchus">Nicorhynchus</a></i></li>
<li><i><a href="/enwiki/wiki/Siroccopteryx" title="Siroccopteryx">Siroccopteryx</a></i>?</li>
<li><i><a href="/enwiki/wiki/Uktenadactylus" title="Uktenadactylus">Uktenadactylus</a></i></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Anhanguerinae" class="mw-redirect" title="Anhanguerinae">Anhanguerinae</a></th><td class="navbox-list navbox-even" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><i><a href="/enwiki/wiki/Anhanguera_(pterosaur)" title="Anhanguera (pterosaur)">Anhanguera</a></i></li>
<li><i><a href="/enwiki/wiki/Caulkicephalus" title="Caulkicephalus">Caulkicephalus</a></i>?</li>
<li><i><a href="/enwiki/wiki/Cearadactylus" title="Cearadactylus">Cearadactylus</a></i></li>
<li><i><a href="/enwiki/wiki/Guidraco" title="Guidraco">Guidraco</a></i></li>
<li><i><a href="/enwiki/wiki/Liaoningopterus" title="Liaoningopterus">Liaoningopterus</a></i></li>
<li><i><a href="/enwiki/wiki/Ludodactylus" title="Ludodactylus">Ludodactylus</a></i></li>
<li><i><a href="/enwiki/wiki/Maaradactylus" title="Maaradactylus">Maaradactylus</a></i></li></ul>
</div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td><td class="noviewer navbox-image" rowspan="1" style="width:1px;padding:0px 0px 0px 2px"><div><span style="display: inline-block; -moz-transform: scaleX(-1); -ms-transform: scaleX(-1); -o-transform: scaleX(-1); -webkit-transform: scaleX(-1); transform: scaleX(-1);"><a href="/enwiki/wiki/File:Pteranodon_longiceps_mmartyniuk_wiki.png" class="image" title="Pteranodon longiceps"><img alt="Pteranodon longiceps" src="/upwiki/wikipedia/commons/thumb/1/1f/Pteranodon_longiceps_mmartyniuk_wiki.png/140px-Pteranodon_longiceps_mmartyniuk_wiki.png" decoding="async" width="140" height="136" srcset="/upwiki/wikipedia/commons/thumb/1/1f/Pteranodon_longiceps_mmartyniuk_wiki.png/210px-Pteranodon_longiceps_mmartyniuk_wiki.png 1.5x, /upwiki/wikipedia/commons/thumb/1/1f/Pteranodon_longiceps_mmartyniuk_wiki.png/280px-Pteranodon_longiceps_mmartyniuk_wiki.png 2x" data-file-width="500" data-file-height="484" /></a></span>
<p><a href="/enwiki/wiki/File:Nyctosaurus_DB.jpg" class="image" title="Nyctosaurus gracilis"><img alt="Nyctosaurus gracilis" src="/upwiki/wikipedia/commons/thumb/8/84/Nyctosaurus_DB.jpg/120px-Nyctosaurus_DB.jpg" decoding="async" width="120" height="137" srcset="/upwiki/wikipedia/commons/thumb/8/84/Nyctosaurus_DB.jpg/180px-Nyctosaurus_DB.jpg 1.5x, /upwiki/wikipedia/commons/thumb/8/84/Nyctosaurus_DB.jpg/240px-Nyctosaurus_DB.jpg 2x" data-file-width="1500" data-file-height="1713" /></a>
</p>
<a href="/enwiki/wiki/File:Ludodactylus.jpg" class="image" title="Ludodactylus sibbicki"><img alt="Ludodactylus sibbicki" src="/upwiki/wikipedia/commons/thumb/b/b6/Ludodactylus.jpg/150px-Ludodactylus.jpg" decoding="async" width="150" height="115" srcset="/upwiki/wikipedia/commons/thumb/b/b6/Ludodactylus.jpg/225px-Ludodactylus.jpg 1.5x, /upwiki/wikipedia/commons/thumb/b/b6/Ludodactylus.jpg/300px-Ludodactylus.jpg 2x" data-file-width="2166" data-file-height="1666" /></a></div></td></tr></tbody></table><div></div></td></tr></tbody></table></div>
<div role="navigation" class="navbox" aria-labelledby="Fins,_limbs_and_wings" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="3"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r992953826"/><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/enwiki/wiki/Template:Fins,_limbs_and_wings" title="Template:Fins, limbs and wings"><abbr title="View this template" style=";;background:none transparent;border:none;box-shadow:none;padding:0;">v</abbr></a></li><li class="nv-talk"><a href="/enwiki/wiki/Template_talk:Fins,_limbs_and_wings" title="Template talk:Fins, limbs and wings"><abbr title="Discuss this template" style=";;background:none transparent;border:none;box-shadow:none;padding:0;">t</abbr></a></li><li class="nv-edit"><a class="external text" href="https://en.wikipedia.org/enwiki/w/index.php?title=Template:Fins,_limbs_and_wings&action=edit"><abbr title="Edit this template" style=";;background:none transparent;border:none;box-shadow:none;padding:0;">e</abbr></a></li></ul></div><div id="Fins,_limbs_and_wings" style="font-size:114%;margin:0 4em"><a href="/enwiki/wiki/Fin" title="Fin">Fins</a>, <a href="/enwiki/wiki/Limb_(anatomy)" title="Limb (anatomy)">limbs</a> and <a href="/enwiki/wiki/Wing" title="Wing">wings</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Fin" title="Fin">Fins</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><a href="/enwiki/wiki/Aquatic_locomotion" title="Aquatic locomotion">Aquatic locomotion</a></li>
<li><a href="/enwiki/wiki/Cephalopod_fin" title="Cephalopod fin">Cephalopod fin</a></li>
<li><a href="/enwiki/wiki/Fish_locomotion" title="Fish locomotion">Fish locomotion</a></li>
<li><a href="/enwiki/wiki/Fin_and_flipper_locomotion" title="Fin and flipper locomotion">Fin and flipper locomotion</a></li>
<li><a href="/enwiki/wiki/Caudal_fin" class="mw-redirect" title="Caudal fin">Caudal fin</a></li>
<li><a href="/enwiki/wiki/Dorsal_fin" title="Dorsal fin">Dorsal fin</a></li>
<li><a href="/enwiki/wiki/Fish_fin" title="Fish fin">Fish fin</a></li>
<li><a href="/enwiki/wiki/Flipper_(anatomy)" title="Flipper (anatomy)">Flipper</a></li>
<li><a href="/enwiki/wiki/Sarcopterygii" title="Sarcopterygii">Lobe-finned fish</a></li>
<li><a href="/enwiki/wiki/Actinopterygii" title="Actinopterygii">Ray-finned fish</a></li>
<li><a href="/enwiki/wiki/Pectoral_fins" class="mw-redirect" title="Pectoral fins">Pectoral fins</a></li>
<li><a href="/enwiki/wiki/Pelvic_fin" title="Pelvic fin">Pelvic fin</a></li></ul>
</div></td><td class="noviewer navbox-image" rowspan="5" style="width:1px;padding:0px 0px 0px 2px"><div><a href="/enwiki/wiki/File:Crossopterygii_fins_tetrapod_legs.svg" class="image" title="Fin and limb"><img alt="Fin and limb" src="/upwiki/wikipedia/commons/thumb/5/55/Crossopterygii_fins_tetrapod_legs.svg/140px-Crossopterygii_fins_tetrapod_legs.svg.png" decoding="async" width="140" height="123" srcset="/upwiki/wikipedia/commons/thumb/5/55/Crossopterygii_fins_tetrapod_legs.svg/210px-Crossopterygii_fins_tetrapod_legs.svg.png 1.5x, /upwiki/wikipedia/commons/thumb/5/55/Crossopterygii_fins_tetrapod_legs.svg/280px-Crossopterygii_fins_tetrapod_legs.svg.png 2x" data-file-width="1093" data-file-height="957" /></a></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Limb_(anatomy)" title="Limb (anatomy)">Limbs</a></th><td class="navbox-list navbox-even" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><a href="/enwiki/wiki/Limb_development" title="Limb development">Limb development</a></li>
<li><a href="/enwiki/wiki/Comparative_foot_morphology" title="Comparative foot morphology">Limb morphology</a>
<ul><li><a href="/enwiki/wiki/Digitigrade" title="Digitigrade">digitigrade</a></li>
<li><a href="/enwiki/wiki/Plantigrade" title="Plantigrade">plantigrade</a></li>
<li><a href="/enwiki/wiki/Unguligrade" class="mw-redirect" title="Unguligrade">unguligrade</a></li>
<li><a href="/enwiki/wiki/Uniped" class="mw-redirect" title="Uniped">uniped</a></li>
<li><a href="/enwiki/wiki/Bipedalism" title="Bipedalism">biped</a></li>
<li><a href="/enwiki/wiki/Facultative_biped" class="mw-redirect" title="Facultative biped">facultative biped</a></li>
<li><a href="/enwiki/wiki/Tripedal" class="mw-redirect" title="Tripedal">triped</a></li>
<li><a href="/enwiki/wiki/Quadrupedalism" title="Quadrupedalism">quadruped</a></li></ul></li>
<li><a href="/enwiki/wiki/Arthropod_leg" title="Arthropod leg">Arthropod</a></li>
<li><a href="/enwiki/wiki/Cephalopod_limb" title="Cephalopod limb">Cephalopod</a></li>
<li><a href="/enwiki/wiki/Tetrapod#Limbs" title="Tetrapod">Tetrapod</a>
<ul><li><a href="/enwiki/wiki/Dactyly" title="Dactyly">dactyly</a></li></ul></li>
<li><a href="/enwiki/wiki/Digit_(anatomy)" title="Digit (anatomy)">Digit</a></li>
<li><a href="/enwiki/wiki/Webbed_foot" title="Webbed foot">Webbed foot</a></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Wing" title="Wing">Wings</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><a href="/enwiki/wiki/Flying_and_gliding_animals" title="Flying and gliding animals">Flying and gliding animals</a></li>
<li><a href="/enwiki/wiki/Bat_wing_development" title="Bat wing development">Bat wing</a></li>
<li><a href="/enwiki/wiki/Bird_wing" class="mw-redirect" title="Bird wing">Bird wing</a>
<ul><li><a href="/enwiki/wiki/Keel_(bird_anatomy)" title="Keel (bird anatomy)">keel</a></li>
<li><a href="/enwiki/wiki/Bird_anatomy#Skeletal_system" title="Bird anatomy">skeleton</a></li>
<li><a href="/enwiki/wiki/Flight_feather" title="Flight feather">feathers</a></li></ul></li>
<li><a href="/enwiki/wiki/Insect_wing" title="Insect wing">Insect wing</a></li>
<li><a href="/enwiki/wiki/Pterosaur#Wings" title="Pterosaur">Pterosaur wing</a></li>
<li><a href="/enwiki/wiki/Wingspan" title="Wingspan">Wingspan</a></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Evolution</th><td class="navbox-list navbox-even" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><a href="/enwiki/wiki/Evolution_of_fish" title="Evolution of fish">Evolution of fish</a></li>
<li><a href="/enwiki/wiki/Evolution_of_tetrapods" title="Evolution of tetrapods">Evolution of tetrapods</a></li>
<li><a href="/enwiki/wiki/Evolution_of_birds" title="Evolution of birds">Evolution of birds</a></li>
<li><a href="/enwiki/wiki/Origin_of_birds" title="Origin of birds">Origin of birds</a></li>
<li><a href="/enwiki/wiki/Origin_of_avian_flight" title="Origin of avian flight">Origin of avian flight</a></li>
<li><a href="/enwiki/wiki/Evolution_of_cetaceans" title="Evolution of cetaceans">Evolution of cetaceans</a></li>
<li><a href="/enwiki/wiki/Comparative_anatomy" title="Comparative anatomy">Comparative anatomy</a></li>
<li><a href="/enwiki/wiki/Convergent_evolution" title="Convergent evolution">Convergent evolution</a></li>
<li><a href="/enwiki/wiki/Analogy_(biology)" class="mw-redirect" title="Analogy (biology)">Analogous structures</a></li>
<li><a href="/enwiki/wiki/Homology_(biology)" title="Homology (biology)">Homologous structures</a></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related</th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><a href="/enwiki/wiki/Animal_locomotion" title="Animal locomotion">Animal locomotion</a></li>
<li><a href="/enwiki/wiki/Gait" title="Gait">Gait</a></li>
<li><a href="/enwiki/wiki/Robot_locomotion" title="Robot locomotion">Robot locomotion</a></li>
<li><a href="/enwiki/wiki/Samara_(fruit)" title="Samara (fruit)">Samara</a></li>
<li><a href="/enwiki/wiki/Terrestrial_locomotion" title="Terrestrial locomotion">Terrestrial locomotion</a></li>
<li><a href="/enwiki/wiki/Tradeoffs_for_locomotion_in_air_and_water" title="Tradeoffs for locomotion in air and water">Tradeoffs for locomotion in air and water</a></li>
<li><a href="/enwiki/wiki/Rotating_locomotion_in_living_systems" title="Rotating locomotion in living systems">Rotating locomotion</a></li>
<li><a href="/enwiki/wiki/Undulatory_locomotion" title="Undulatory locomotion">Undulatory locomotion</a></li></ul>
</div></td></tr></tbody></table></div>
<div class="noprint metadata navbox" role="navigation" aria-label="Portals" style="font-weight:bold;padding:0.4em 2em"><ul style="margin:0.1em 0 0"><li style="display:inline"><span style="display:inline-block;white-space:nowrap"><span style="margin:0 0.5em"><a href="/enwiki/wiki/File:Allosaurus_Jardin_des_Plantes.png" class="image"><img alt="Allosaurus Jardin des Plantes.png" src="/upwiki/wikipedia/commons/thumb/1/1b/Allosaurus_Jardin_des_Plantes.png/21px-Allosaurus_Jardin_des_Plantes.png" decoding="async" width="21" height="21" srcset="/upwiki/wikipedia/commons/thumb/1/1b/Allosaurus_Jardin_des_Plantes.png/32px-Allosaurus_Jardin_des_Plantes.png 1.5x, /upwiki/wikipedia/commons/thumb/1/1b/Allosaurus_Jardin_des_Plantes.png/42px-Allosaurus_Jardin_des_Plantes.png 2x" data-file-width="1000" data-file-height="1000" /></a></span><a href="/enwiki/wiki/Portal:Paleontology" title="Portal:Paleontology">Paleontology portal</a></span></li><li style="display:inline"><span style="display:inline-block;white-space:nowrap"><span style="margin:0 0.5em"><a href="/enwiki/wiki/File:Flag_of_Germany.svg" class="image"><img alt="Flag of Germany.svg" src="/upwiki/wikipedia/en/thumb/b/ba/Flag_of_Germany.svg/24px-Flag_of_Germany.svg.png" decoding="async" width="24" height="14" srcset="/upwiki/wikipedia/en/thumb/b/ba/Flag_of_Germany.svg/36px-Flag_of_Germany.svg.png 1.5x, /upwiki/wikipedia/en/thumb/b/ba/Flag_of_Germany.svg/48px-Flag_of_Germany.svg.png 2x" data-file-width="1000" data-file-height="600" /></a></span><a href="/enwiki/wiki/Portal:Germany" title="Portal:Germany">Germany portal</a></span></li><li style="display:inline"><span style="display:inline-block;white-space:nowrap"><span style="margin:0 0.5em"><a href="/enwiki/wiki/File:Issoria_lathonia.jpg" class="image"><img alt="Issoria lathonia.jpg" src="/upwiki/wikipedia/commons/thumb/2/2d/Issoria_lathonia.jpg/24px-Issoria_lathonia.jpg" decoding="async" width="24" height="17" srcset="/upwiki/wikipedia/commons/thumb/2/2d/Issoria_lathonia.jpg/36px-Issoria_lathonia.jpg 1.5x, /upwiki/wikipedia/commons/thumb/2/2d/Issoria_lathonia.jpg/48px-Issoria_lathonia.jpg 2x" data-file-width="629" data-file-height="445" /></a></span><a href="/enwiki/wiki/Portal:Biology" title="Portal:Biology">Biology portal</a></span></li></ul></div>
<div role="navigation" class="navbox" aria-labelledby="Taxon_identifiers" style="padding:3px"><table class="nowraplinks hlist navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th id="Taxon_identifiers" scope="row" class="navbox-group" style="width:1%;text-align: left;"><a href="/enwiki/wiki/Help:Taxon_identifiers" title="Help:Taxon identifiers">Taxon identifiers</a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><span style="white-space:nowrap;"><a href="/enwiki/wiki/Wikidata" title="Wikidata">Wikidata</a>: <span class="uid"><span class="external"><a href="https://www.wikidata.org/wiki/Q309490" class="extiw" title="wikidata:Q309490">Q309490</a></span></span></span></li>
<li><span style="white-space:nowrap;"><a href="/enwiki/wiki/Wikispecies" title="Wikispecies">Wikispecies</a>: <span class="uid"><span class="external"><a href="https://species.wikimedia.org/wiki/Pterodactylus" class="extiw" title="wikispecies:Pterodactylus">Pterodactylus</a></span></span></span></li>
<li><span style="white-space:nowrap;"><a href="/enwiki/wiki/Fossilworks" title="Fossilworks">Fossilworks</a>: <span class="uid"><a rel="nofollow" class="external text" href="http://fossilworks.org/bridge.pl?a=taxonInfo&taxon_no=38494">38494</a></span></span></li></ul>
</div></td></tr></tbody></table></div>
<div role="navigation" class="navbox authority-control" aria-labelledby="Authority_control_frameless_&#124;text-top_&#124;10px_&#124;alt=Edit_this_at_Wikidata_&#124;link=https&#58;//www.wikidata.org/wiki/Q309490#identifiers&#124;Edit_this_at_Wikidata" style="padding:3px"><table class="nowraplinks hlist navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th id="Authority_control_frameless_&#124;text-top_&#124;10px_&#124;alt=Edit_this_at_Wikidata_&#124;link=https&#58;//www.wikidata.org/wiki/Q309490#identifiers&#124;Edit_this_at_Wikidata" scope="row" class="navbox-group" style="width:1%"><a href="/enwiki/wiki/Help:Authority_control" title="Help:Authority control">Authority control</a> <a href="https://www.wikidata.org/wiki/Q309490#identifiers" title="Edit this at Wikidata"><img alt="Edit this at Wikidata" src="/upwiki/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/10px-OOjs_UI_icon_edit-ltr-progressive.svg.png" decoding="async" width="10" height="10" style="vertical-align: text-top" srcset="/upwiki/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/15px-OOjs_UI_icon_edit-ltr-progressive.svg.png 1.5x, /upwiki/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/20px-OOjs_UI_icon_edit-ltr-progressive.svg.png 2x" data-file-width="20" data-file-height="20" /></a></th><td class="navbox-list navbox-odd" style="text-align:left;border-left-width:2px;border-left-style:solid;width:100%;padding:0px"><div style="padding:0em 0.25em">
<ul><li><span class="nowrap"><a href="/enwiki/wiki/LCCN_(identifier)" class="mw-redirect" title="LCCN (identifier)">LCCN</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://id.loc.gov/authorities/subjects/sh94002837">sh94002837</a></span></span></li>
<li><span class="nowrap"><a href="/enwiki/wiki/MA_(identifier)" class="mw-redirect" title="MA (identifier)">MA</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://academic.microsoft.com/v2/detail/2776325289">2776325289</a></span></span></li></ul>
</div></td></tr></tbody></table></div>
<p class="mw-empty-elt">
</p>
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Whether or not the change was made through a Tor exit node (tor_exit_node) | false |
Unix timestamp of change (timestamp) | 1620311147 |