Editing Pterodactylus (section)

==Discovery and history==
[[File:Pterodactylus holotype Collini 1784.jpg|thumb|left|Original copper engraving of the ''P. antiquus'' [[holotype]] by [[Egid Verhelst the Younger|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 genus ''[[Aurorazhdarcho]]'' 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 |bibcode=2004CRPal...3..157T}}</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 invading forces of the [[French Consulate]] and sent to 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 |volume=1815 |edition=L'Imprimerie de Jean Barravecchia |page=75 |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="Cuvier1819">{{Cite book |last=Cuvier |first=G. |url=https://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="Cuvier1819" /> 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 />