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| type_species = {{extinct}}'''''Pterodactylus antiquus''''' |
| type_species = {{extinct}}'''''Pterodactylus antiquus''''' |
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| type_species_authority = [[Samuel Thomas von Sömmerring|Sömmerring]], 1812 |
| type_species_authority = [[Samuel Thomas von Sömmerring|Sömmerring]], 1812 |
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| synonyms = |
| synonyms = |
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|title=<small>Genus synonymy</small> |
|title=<small>Genus synonymy</small> |
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'''''Pterodactylus''''' (from [[Greek]] πτεροδάκτυλος, pterodaktulos, meaning "winged finger"<ref>{{Cite journal|last=Gudger|first=E.W.|date=Aug 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 [[euctenochasmatia]]n [[pterodactyloid]] [[pterosaur]], 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''''', the first pterosaur species to be named and identified as a flying [[reptile]]. |
'''''Pterodactylus''''' (from [[Greek]] πτεροδάκτυλος, pterodaktulos, meaning "winged finger"<ref>{{Cite journal|last=Gudger|first=E.W.|date=Aug 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 [[euctenochasmatia]]n [[pterodactyloid]] [[pterosaur]], 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''''', the first pterosaur species to be named and identified as a flying [[reptile]]. |
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[[Fossil]] remains of ''Pterodactylus'' have primarily been found in the [[Solnhofen limestone]] of [[Bavaria]], [[Germany]], which dates back to the [[Late Jurassic]] [[Period (geology)|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 ''Pterodactylus'' have primarily been found in the [[Solnhofen limestone]] of [[Bavaria]], [[Germany]], which dates back to the [[Late Jurassic]] [[Period (geology)|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> |
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Many studies conclude that ''Pterodactylus'' was a [[carnivore]] that probably preyed upon fish as well as other small animals. 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]]'', 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]]''. The classification of ''Pterodactylus'' has since been confusing to paleontologists. Some classify ''Pterodactylus'' within the clade [[Euctenochasmatia]], while others just consider it as a basal member of the suborder [[Pterodactyloidea]].<ref name="unwin2003"/><ref name=longrichetal2018/> |
Many studies conclude that ''Pterodactylus'' was a [[carnivore]] that probably preyed upon fish as well as other small animals. 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]]''. The classification of ''Pterodactylus'' has since been confusing to paleontologists. Some classify ''Pterodactylus'' within the clade [[Euctenochasmatia]], while others just consider it as a basal member of the suborder [[Pterodactyloidea]].<ref name="unwin2003"/><ref name=longrichetal2018/> |
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==History of discovery== |
==History of discovery== |
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[[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 [[Cosimo Alessandro Collini]], 1784]] |
[[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 [[Cosimo Alessandro Collini]], 1784]] |
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The [[type specimen]] of the animal now known as ''Pterodactylus antiquus'' was one of the first [[pterosaur]] fossils ever to be identified. The first ''Pterodactylus'' specimen was described by the Italian scientist [[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 "Naturalienkabinett", or nature [[Cabinet of curiosities|cabinet]] (a precursor to the modern concept of the [[natural history]] museum), in the palace of [[Charles Theodore, Elector of Bavaria]] at [[Mannheim]].<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 [[Pappenheim (state)|Friedrich Ferdinand zu Pappenheim]], probably around 1780, having been recovered from a [[lithographic limestone]] quarry in [[Eichstätt]].<ref name=brougham1844>Brougham, H.P. (1844). ''Dialogues on instinct; with analytical view of the researches on fossil osteology.'' Volume 19 of Knight's weekly vol.</ref> The actual date of the specimen's discovery and entry into the collection is unknown. It was not mentioned in a catalogue of the collection taken in 1767 and so 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 ''[[Pterodactylus micronyx]]'' was described in 1779 and possibly discovered earlier than the Mannheim specimen, but it was at first considered to be a fossil crustacean.<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_The_history_of_Late_Jurassic_pterosaurs_housed_in_Hungarian_collections_and_the_revision_of_the_holotype_of_Pterodactylus_micronyx_Meyer_1856_a_'Pester_Exemplar' }}</ref> |
The [[type specimen]] of the animal now known as ''Pterodactylus antiquus'' was one of the first [[pterosaur]] fossils ever to be identified. The first ''Pterodactylus'' specimen was described by the Italian scientist [[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 "Naturalienkabinett", or nature [[Cabinet of curiosities|cabinet]] (a precursor to the modern concept of the [[natural history]] museum), in the palace of [[Charles Theodore, Elector of Bavaria]] at [[Mannheim]].<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 [[Pappenheim (state)|Friedrich Ferdinand zu Pappenheim]], probably around 1780, having been recovered from a [[lithographic limestone]] quarry in [[Eichstätt]].<ref name=brougham1844>Brougham, H.P. (1844). ''Dialogues on instinct; with analytical view of the researches on fossil osteology.'' Volume 19 of Knight's weekly vol.</ref> The actual date of the specimen's discovery and entry into the collection is unknown. It was not mentioned in a catalogue of the collection taken in 1767 and so 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 ''[[Pterodactylus micronyx]]'' was described in 1779 and possibly discovered earlier than the Mannheim specimen, but it was at first considered to be a fossil crustacean.<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_The_history_of_Late_Jurassic_pterosaurs_housed_in_Hungarian_collections_and_the_revision_of_the_holotype_of_Pterodactylus_micronyx_Meyer_1856_a_'Pester_Exemplar' }}</ref> |
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[[File:Aquatic Pterodactylus.jpg|thumb|left|Wagler's 1830 restoration of an aquatic ''Pterodactylus'']] |
[[File:Aquatic Pterodactylus.jpg|thumb|left|Wagler's 1830 restoration of an aquatic ''Pterodactylus'']] |
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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">Collini, C A. (1784). "Sur quelques Zoolithes du Cabinet d’Histoire naturelle de S. A. S. E. Palatine & de Bavière, à Mannheim." ''Acta Theodoro-Palatinae Mannheim 5 Pars Physica'', pp. 58–103 (1 plate).</ref><ref name="taquet&padian2004">{{Cite journal | doi = 10.1016/j.crpv.2004.02.002 |author1=Taquet, P. |author2=Padian, K. |lastauthoramp=yes | 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 name="wagler1830">Wagler, J. (1830). ''Natürliches System der Amphibien'' Munich, 1830: 1–354.</ref> |
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">Collini, C A. (1784). "Sur quelques Zoolithes du Cabinet d’Histoire naturelle de S. A. S. E. Palatine & de Bavière, à Mannheim." ''Acta Theodoro-Palatinae Mannheim 5 Pars Physica'', pp. 58–103 (1 plate).</ref><ref name="taquet&padian2004">{{Cite journal | doi = 10.1016/j.crpv.2004.02.002 |author1=Taquet, P. |author2=Padian, K. |lastauthoramp=yes | 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 name="wagler1830">Wagler, J. (1830). ''Natürliches System der Amphibien'' Munich, 1830: 1–354.</ref> |
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[[File:Hermann pterodactylus restoration1.png|thumb|Hermann's original life restoration, the first of any pterosaur, 1800]] |
[[File:Hermann pterodactylus restoration1.png|thumb|Hermann's original life restoration, the first of any pterosaur, 1800]] |
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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 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"/> 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." However, contrary to Hermann, Cuvier was convinced the animal was a [[reptile]].<ref name="cuvier1801">{{Cite journal | author = Cuvier, G. | year = 1801 | title = [Reptile volant]. In: Extrait d'un ouvrage sur les espèces de quadrupèdes dont on a trouvé les ossemens dans l'intérieur de la terre | journal = Journal de Physique, de Chimie et d'Histoire Naturelle | volume = 52 | pages = 253–267 }}</ref> |
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 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"/> 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." However, contrary to Hermann, Cuvier was convinced the animal was a [[reptile]].<ref name="cuvier1801">{{Cite journal | author = Cuvier, G. | year = 1801 | title = [Reptile volant]. In: Extrait d'un ouvrage sur les espèces de quadrupèdes dont on a trouvé les ossemens dans l'intérieur de la terre | journal = Journal de Physique, de Chimie et d'Histoire Naturelle | volume = 52 | pages = 253–267 }}</ref> |
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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. 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 a "ptero-dactyle" and refuted a hypothesis by [[Johann Friedrich Blumenbach]] that it would have been a shore bird.<ref name="cuvier1809"/> |
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. 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 a "ptero-dactyle" and refuted a hypothesis by [[Johann Friedrich Blumenbach]] that it would have been a shore bird.<ref name="cuvier1809"/> |
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[[File:Pterodactylus antiquus soemmerring.png|thumb|Von Soemmerring's incorrect 1817 restoration of ''[[Pterodactylus brevirostris|P. brevirostris]]'']] |
[[File:Pterodactylus antiquus soemmerring.png|thumb|Von Soemmerring's incorrect 1817 restoration of ''[[Pterodactylus brevirostris|P. brevirostris]]'']] |
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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 27 December 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", ''Denkschriften der königlichen bayerischen Akademie der Wissenschaften | url = | journal = , München: mathematisch-physikalische Classe | volume = 3 | issue = | pages = 89–158 }}</ref> The animal was described as being both a mammal, 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 name=cuvier1812>Cuvier, G. (1812). ''Recherches sur les ossemens fossiles''. I ed. p. 24, tab. 31</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 Soemmerring 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 Söemmerring 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 | author = Sömmering, T. v. | year = 1817 | title = Über einen Ornithocephalus brevirostris der Vorwelt | journal = Denkschr. KGL. Bayer Akad. Wiss., Math.phys. Cl. | volume = 6 |pages = 89–104 }}</ref> Söemmerring 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>Padian, K. (1987). "The case of the bat-winged pterosaur. Typological taxonomy and the influence of pictorial representation on scientific perception", pp. 65–81 in: Czerkas, S. J. and Olson, E. C., eds. ''Dinosaurs past and present. An exhibition and symposium organized by the Natural History Museum of Los Angeles County. Volume 2''. Natural History Museum of Los Angeles County and University of Washington Press, Seattle and London</ref> |
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 27 December 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", ''Denkschriften der königlichen bayerischen Akademie der Wissenschaften | url = | journal = , München: mathematisch-physikalische Classe | volume = 3 | issue = | pages = 89–158 }}</ref> The animal was described as being both a mammal, 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 name=cuvier1812>Cuvier, G. (1812). ''Recherches sur les ossemens fossiles''. I ed. p. 24, tab. 31</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 Soemmerring 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 Söemmerring 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 | author = Sömmering, T. v. | year = 1817 | title = Über einen Ornithocephalus brevirostris der Vorwelt | journal = Denkschr. KGL. Bayer Akad. Wiss., Math.phys. Cl. | volume = 6 |pages = 89–104 }}</ref> Söemmerring 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>Padian, K. (1987). "The case of the bat-winged pterosaur. Typological taxonomy and the influence of pictorial representation on scientific perception", pp. 65–81 in: Czerkas, S. J. and Olson, E. C., eds. ''Dinosaurs past and present. An exhibition and symposium organized by the Natural History Museum of Los Angeles County. Volume 2''. Natural History Museum of Los Angeles County and University of Washington Press, Seattle and London</ref> |
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==Description== |
==Description== |
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[[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]] |
[[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]] |
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''Pterodactylus'' is known from over 30 [[fossil]] specimens, and though most of those are 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|abbr=off|sp=us}} (the only known adult specimen is represented by an isolated skull).<ref name=BennettPZ/> Other "species" were once thought to be smaller. 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> |
''Pterodactylus'' is known from over 30 [[fossil]] specimens, and though most of those are 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|abbr=off|sp=us}} (the only known adult specimen is represented by an isolated skull).<ref name=BennettPZ/> Other "species" were once thought to be smaller. 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> |
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The skulls of adult ''Pterodactylus'' were long and thin with about 90 narrow, conical teeth. The teeth extended back from the tips of both jaws, and became smaller farther away from the jaw tips (unlike some relatives, where teeth were absent in the upper jaw tip and were relatively uniform in size). The teeth extended farther back into the jaw than in close relatives, as some were present below the front of the ''nasoantorbital fenestra'', the largest opening in the skull.<ref name=BennettPZ/> Unlike related species, the skull and jaws were straight, not curved upwards.<ref name="jouve2004"/> |
The skulls of adult ''Pterodactylus'' were long and thin with about 90 narrow, conical teeth. The teeth extended back from the tips of both jaws, and became smaller farther away from the jaw tips (unlike some relatives, where teeth were absent in the upper jaw tip and were relatively uniform in size). The teeth extended farther back into the jaw than in close relatives, as some were present below the front of the ''nasoantorbital fenestra'', the largest opening in the skull.<ref name=BennettPZ/> Unlike related species, the skull and jaws were straight, not curved upwards.<ref name="jouve2004"/> |
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[[File:Pterodactylus BMMS7 life.png|thumb|Life restoration of BMMS 7, the largest known ''Pterodactylus'' specimen. The shape of the soft tissue crest is based on BSP 1929.]] |
[[File:Pterodactylus BMMS7 life.png|thumb|Life restoration of BMMS 7, the largest known ''Pterodactylus'' specimen. The shape of the soft tissue crest is based on BSP 1929.]] |
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''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]] (the largest opening in the skull) 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}}</ref><ref name="frey&martill1998">{{Cite journal |author1=Frey, E. |author2=Martill, D.M. |
''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]] (the largest opening in the skull) 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}}</ref><ref name="frey&martill1998">{{Cite journal |author1=Frey, E. |author2=Martill, D.M. |lastauthoramp=yes | 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> Bennett (2013) 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''. This is BSP 1929 I 18, which has an occipital lappet similar to the proposed ''Aerodactylus'' definition. This specimen also has a small triangular soft tissue crest with the peak of the crest positioned above the eyes.<ref name=BennettPZ /> |
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==Paleobiology== |
==Paleobiology== |
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===Life history=== |
===Life history=== |
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[[File:Pterodactylus spectabilis 4.JPG|thumb|left|Juvenile specimen of ''P. antiquus'']] |
[[File:Pterodactylus spectabilis 4.JPG|thumb|left|Juvenile specimen of ''P. antiquus'']] |
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Like other pterosaurs (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 ''Pterodactylus'' species, ''P. antiquus''.<ref name="jouve2004"/> |
Like other pterosaurs (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 ''Pterodactylus'' species, ''P. antiquus''.<ref name="jouve2004"/> |
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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), 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 some specimens).<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), 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 some specimens).<ref name="jouve2004"/> |
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''Pterodactylus'' specimens can be divided into two distinct year classes. In the first year class, the skulls are only 15-45 mm (0.59-1.77 inches) in length. The second year class is characterized by skulls 55-95 mm (2.16-3.74 inches) long, but still immature. 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/> |
''Pterodactylus'' specimens can be divided into two distinct year classes. In the first year class, the skulls are only 15-45 mm (0.59-1.77 inches) in length. The second year class is characterized by skulls 55-95 mm (2.16-3.74 inches) long, but still immature. 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/> |
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===Growth and breeding seasons=== |
===Growth and breeding seasons=== |
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[[File:Pterodactylus grandipelvis Teylers 2.JPG|thumb|Fossil pelvis of a large specimen, now referred to the dubious species ''P. grandipelvis'']] |
[[File:Pterodactylus grandipelvis Teylers 2.JPG|thumb|Fossil pelvis of a large specimen, now referred to the dubious species ''P. grandipelvis'']] |
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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>Wellnhofer, P. (1970). ''Die Pterodactyloidea (Pterosauria) der Oberjura-Plattenkalke Siiddeutschlands.'' Bayerische Akademie der Wissenschaften, Mathematisch-Wissenschaftlichen Klasse, Abhandlungen, '''141''': 133 pp.</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/> |
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>Wellnhofer, P. (1970). ''Die Pterodactyloidea (Pterosauria) der Oberjura-Plattenkalke Siiddeutschlands.'' Bayerische Akademie der Wissenschaften, Mathematisch-Wissenschaftlichen Klasse, Abhandlungen, '''141''': 133 pp.</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/> |
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===Daily activity patterns=== |
===Daily activity patterns=== |
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Comparisons between the [[sclerotic ring|scleral ring]]s 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 '' |
Comparisons between the [[sclerotic ring|scleral ring]]s 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}}</ref> |
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==Paleoecology== |
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[[File:Pterodactylus holotype fly mmartyniuk.png|thumb|left|Hypothetical restoration of ''Pterodactylus'' in its environment]] |
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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 diverse [[Lagerstätte]] that contains a wide range of different creatures, including highly detailed [[Fossil|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>Bennett, S. C. (2004). "New information on the pterosaur ''Scaphognathus crassirostris'' and the pterosaurian cervical series", ''Journal of Vertebrate Paleontology'', 24(Supplement to #3):38A</ref> several [[gallodactylid]]s such as ''Aerodactylus'',<ref name=bennett1996a>{{cite journal | last1 = Bennett | first1 = S.C. | year = 1996 | title = Year-classes of pterosaurs from the Solnhofen Limestone of Germany: Taxonomic and Systematic Implications | url = | journal = Journal of Vertebrate Paleontology | volume = 16 | issue = 3| pages = 432–444 | doi=10.1080/02724634.1996.10011332}}</ref> ''[[Ardeadactylus]]'', ''Aurorazhdarcho'' and ''[[Cycnorhamphus]]'',<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 |page=246}}</ref> 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 | url = | 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 euctenochasmatian ''[[Diopecephalus]]''.<ref>{{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|pages=125–147|doi=10.1144/SP455.12|url=https://eprints.soton.ac.uk/423063/1/Vidovic_Martill_2017_Taxonomy_of_Diopecephalus_and_Germanodactylus_AM_with_Figures.pdf|bibcode=2018GSLSP.455..125V}}</ref> 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|veditors=Fastovsky DE, Weishampel DB |vauthors=Fastovsky DE, Weishampel DB |pages=265–299|publisher=Cambridge University Press|isbn=978-0-521-81172-9|edition=2nd}}</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 Societé Géologique de France |volume=8 |issue=4 |pages=592–595 |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 |issue=|pages=37–52|doi=|url=}}</ref> [[Crocodylomorph]] specimens were widely distributed within the fossil site, most were assigned to the [[metriorhynchid]] genera ''[[Cricosaurus]]'', ''[[Dakosaurus]]'', ''[[Geosaurus]]'' and ''[[Rhacheosaurus]]''. These genera are colloquially called as marine or sea crocodiles due to their similar built.<ref name=hypercarnivory>De Andrade, M. B., Young, M. T., Desojo, J. B., & Brusatte, S. L. (2010). The evolution of extreme hypercarnivory in Metriorhynchidae (Mesoeucrocodylia: Thalattosuchia) based on evidence from microscopic denticle morphology. Journal of Vertebrate Paleontology, 30(5), 1451-1465.</ref><ref>Young MT, Brusatte SL, de Andrade MB, Desojo JB, Beatty BL, et al. (2012) The Cranial Osteology and Feeding Ecology of the Metriorhynchid Crocodylomorph Genera ''Dakosaurus'' and ''Plesiosuchus'' from the Late Jurassic of Europe. PLoS ONE 7(9): e44985. doi:10.1371/journal.pone.0044985</ref> The [[rhynchocephalia]]n genera ''[[Homoeosaurus]]'' and ''[[Pleurosaurus]]'' were also found in the formation alongside several turtles such as ''[[Eurysternum]]'' and ''[[Paleomedusa]]''.<ref name=anatomy>Dupret, V. (2004). The pleurosaurs: anatomy and phylogeny. ''Revue de Paléobiologie'', '''9''': 61-80.[https://scholar.google.com/scholar?hl=en&q=The+pleurosaurs+%3A+anatomy+and+phylogeny&btnG=&as_sdt=1%2C31&as_sdtp=]</ref><ref name="WGJ">{{cite journal|author=Walter G. 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=2015-10-01}}</ref> Fossils of the [[ichthyosaur]] ''[[Aegirosaurus]]'' were also present in the site,<ref name=aegirosaurus>Bardet, N., & Fernández, M. (2000). A new ichthyosaur from the Upper Jurassic lithographic limestones of Bavaria. Journal Information, 74(3).</ref> as well as fish remains, with many specimens assigned to [[ray-finned fish]]es such as the [[halecomorph]]s ''[[Lepidotes]]'',<ref name="lamberstable-semionotidae">"Table 1; Halecostomi; Semionotidae" in Lambers (1999) pg. 216.</ref> ''[[Propterus]]'',<ref name=Marine/> ''[[Gyrodus]]'', ''[[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 |accessdate=2009-02-27 |url-status=dead |archiveurl=https://web.archive.org/web/20110723131237/http://strata.ummp.lsa.umich.edu/jack/showgenera.php?taxon=611&rank=class |archivedate=July 23, 2011 }}</ref> and ''[[Ophiopsis]]'',<ref>Lane, J. A. & Ebert, M., 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. Journal of Vertebrate Paleontology, 35 (1): e883238. doi: 10.1080/02724634.2014.883238</ref> the [[Pachycormidae|pachycormids]] ''[[Asthenocormus]]'' and ''[[Hypsocormus]]'',<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 [[teleosts]] ''[[Aspidorhynchus]]'' and ''[[Thrissops]]''.<ref name=FT12>{{cite journal |last=Frey |first=E. |author2=and Tischlinger, 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 |url=http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0031945|bibcode = 2012PLoSO...7E1945F |pmid=22412850 |pmc=3296705}}</ref><ref>Orvar Nybelin, "Versuch einer taxonomischen revision der jurassischen Fischgattung Thrissops Agassiz", ''Nature'' (1964)</ref> |
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==Classification== |
==Classification== |
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[[File:Pterodactylus AMNH.jpg|thumb|A ''P. antiquus'' specimen (AMNH 1942) showing muscle impressions]] |
[[File:Pterodactylus AMNH.jpg|thumb|A ''P. antiquus'' specimen (AMNH 1942) showing muscle impressions]] |
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The [[genus]] now known as ''Pterodactylus'' was originally named ''Petro-Dactyle'' by Cuvier in 1809,<ref name="cuvier1809">{{Cite journal | author = Cuvier, 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 }}</ref> though this was a typographical error, later corrected by him to ''Ptéro-Dactyle''.<ref name="taquet&padian2004"/> In 1812, Samuel Thomas von Sömmerring named the same specimen ''Ornithocephalus antiquus''. The genus name was emended to the current ''Pterodactylus'' by [[Constantine Samuel Rafinesque]] in 1815. Unaware of Rafinesque's publication, Cuvier himself in 1819 again emended the genus name,<ref>Cuvier, G., 1819, (''Pterodactylus longirostris'') in ''Isis von Oken'', 1126 und 1788, Jena</ref> but the specific name he then gave, ''longirostris'', has to give precedence to von Soemmerring's ''antiquus''. In 1888 [[Richard Lydekker]] designated ''Pterodactylus antiquus'' the [[type species]]. The original specimen is the [[holotype]] of the genus, BSP No. AS.I.739. |
The [[genus]] now known as ''Pterodactylus'' was originally named ''Petro-Dactyle'' by Cuvier in 1809,<ref name="cuvier1809">{{Cite journal | author = Cuvier, 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 }}</ref> though this was a typographical error, later corrected by him to ''Ptéro-Dactyle''.<ref name="taquet&padian2004"/> In 1812, Samuel Thomas von Sömmerring named the same specimen ''Ornithocephalus antiquus''. The genus name was emended to the current ''Pterodactylus'' by [[Constantine Samuel Rafinesque]] in 1815. Unaware of Rafinesque's publication, Cuvier himself in 1819 again emended the genus name,<ref>Cuvier, G., 1819, (''Pterodactylus longirostris'') in ''Isis von Oken'', 1126 und 1788, Jena</ref> but the specific name he then gave, ''longirostris'', has to give precedence to von Soemmerring's ''antiquus''. In 1888 [[Richard Lydekker]] designated ''Pterodactylus antiquus'' the [[type species]]. The original specimen is the [[holotype]] of the genus, BSP No. AS.I.739. |
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[[Christian Erich Hermann von Meyer|Hermann von Meyer]], in 1830, 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. This group has more recently been given several competing definitions.<ref name="unwin2003"/><ref name="kellner2003">Kellner, A.W.A. (2003). "Pterosaur phylogeny and comments on the evolutionary history of the group", pp. 105–137 ''in'' Buffetaut, E. and Mazin, J.-M., (eds.) (2003), ''Evolution and Palaeobiology of Pterosaurs''. Geological Society of London, Special Publications 217, London: 1–347.</ref> |
[[Christian Erich Hermann von Meyer|Hermann von Meyer]], in 1830, 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. This group has more recently been given several competing definitions.<ref name="unwin2003"/><ref name="kellner2003">Kellner, A.W.A. (2003). "Pterosaur phylogeny and comments on the evolutionary history of the group", pp. 105–137 ''in'' Buffetaut, E. and Mazin, J.-M., (eds.) (2003), ''Evolution and Palaeobiology of Pterosaurs''. Geological Society of London, Special Publications 217, London: 1–347.</ref> |
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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 ctenochasmatoids may have been more closely related to the more advanced [[Ornithocheiroidea|ornithocheiroids]], 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=Altmuehlopterus>Vidovic, S.U. and Martill, D.M. (2017). The taxonomy and phylogeny of ''Diopecephalus kochi'' (Wagner, 1837) and "''Germanodactylus rhamphastinus''" (Wagner, 1851). In Hone, D. W. E., Witton, M. P. &Martill, D. M. (eds) ''New Perspectives on Pterosaur Palaeobiology''. Geological Society, London, Special Publications, 455 {{doi|10.1144/SP455.12}}</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 ctenochasmatoids may have been more closely related to the more advanced [[Ornithocheiroidea|ornithocheiroids]], 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=Altmuehlopterus>Vidovic, S.U. and Martill, D.M. (2017). The taxonomy and phylogeny of ''Diopecephalus kochi'' (Wagner, 1837) and "''Germanodactylus rhamphastinus''" (Wagner, 1851). In Hone, D. W. E., Witton, M. P. &Martill, D. M. (eds) ''New Perspectives on Pterosaur Palaeobiology''. Geological Society, London, Special Publications, 455 {{doi|10.1144/SP455.12}}</ref> |
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{{clade| style=font-size:80%;line-height:80% |
{{clade| style=font-size:80%;line-height:80% |
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|label1=[[Pterodactyloidea]] |
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|1={{clade |
|1={{clade |
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|1=''[[Eosipterus yangi]]'' |
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|2={{clade |
|2={{clade |
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|1='''''Pterodactylus antiquus'''''[[File:Pterodactylus BMMS7 life.png|50 px]] |
|1='''''Pterodactylus antiquus'''''[[File:Pterodactylus BMMS7 life.png|50 px]] |
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|label2=[[Lophocratia]] |
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|2={{clade |
|2={{clade |
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|1={{clade |
|1={{clade |
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|1=''[[Diopecephalus kochi]]'' |
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|label2=[[Ctenochasmatoidea]] |
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|2={{clade |
|2={{clade |
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|label1=[[Gallodactylidae]] |
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|1={{clade |
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|1=''[[Aerodactylus scolopaciceps]]''[[File:Aerodactylus MCZ 1505.png|50 px]] |
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|2={{clade |
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|1=[[Aurorazhdarchidae]] |
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|2=[[Gallodactylidae]] }} }} |
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|2=[[Ctenochasmatidae]][[File:Pterodaustro BW.jpg|50px]] }} }} |
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|2={{clade |
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|1=''[[Altmuehlopterus ramphastinus]]''[[File:Altmuehlopterus DB.jpg|50 px]] |
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|label2=[[Dsungaripteroidea]] |
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|2={{clade |
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|1=''[[Germanodactylus cristatus]]'' |
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|2={{clade |
|2={{clade |
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|1=''[[Elanodactylus prolatus]]'' |
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|2=[[Ornithocheiroidea]][[File:Pteranodon longiceps mmartyniuk wiki.png|50 px]][[File:Quetzalcoatlus07.jpg|50px]]}} }} }} }} }} }} }} |
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|2={{clade |
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|1=''[[Elanodactylus prolatus]]'' |
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|2=[[Ornithocheiroidea]][[File:Pteranodon longiceps mmartyniuk wiki.png|50 px]][[File:Quetzalcoatlus07.jpg|50px]]}} }} }} }} }} }} }} |
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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, 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 | url = http://journals.plos.org/plosbiology/article?id=10.1371/journal.pbio.2001663 | journal = PLoS Biology | volume = 16 | issue = 3| page = e2001663 | doi = 10.1371/journal.pbio.2001663 | pmid = 29534059 | pmc = 5849296 }}</ref> |
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, 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 | url = http://journals.plos.org/plosbiology/article?id=10.1371/journal.pbio.2001663 | journal = PLoS Biology | volume = 16 | issue = 3| page = e2001663 | doi = 10.1371/journal.pbio.2001663 | pmid = 29534059 | pmc = 5849296 }}</ref> |
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{{clade| style=font-size:90%;line-height:90% |
{{clade| style=font-size:90%;line-height:90% |
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|label1=[[Archaeopterodactyloidea]] |
|label1=[[Archaeopterodactyloidea]] |
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|1={{clade |
|1={{clade |
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|label1=[[Germanodactylidae]] |
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|1={{clade |
|1={{clade |
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|1=''[[Germanodactylus cristatus]]'' |
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|2=''[[Germanodactylus rhamphastinus]]''[[File:Altmuehlopterus DB.jpg|50px]] }} |
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|label2=[[Euctenochasmatia]] |
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|2={{clade |
|2={{clade |
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|1='''''Pterodactylus antiquus'''''[[File:Pterodactylus BMMS7 life.png|50px]] |
|1='''''Pterodactylus antiquus'''''[[File:Pterodactylus BMMS7 life.png|50px]] |
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|label2=[[Ctenochasmatoidea]] |
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|2={{clade |
|2={{clade |
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|label1=[[Gallodactylidae]] |
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|1={{clade |
|1={{clade |
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|1=''[[Cycnorhamphus suevicus]]'' |
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|2=''[[Normannognathus wellnhoferi]]'' }} |
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|2=[[Ctenochasmatidae]][[File:Pterodaustro BW.jpg|50px]] |
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}} }} }} }} |
}} }} }} }} |
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===Formerly assigned species=== |
===Formerly assigned species=== |
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[[File:Pterodactylus kochi 1.JPG|thumb|upright|Specimen of the species ''[[Pterodactylus kochi]]'', [[Bürgermeister Müller Museum]]]] |
[[File:Pterodactylus kochi 1.JPG|thumb|upright|Specimen of the species ''[[Pterodactylus kochi]]'', [[Bürgermeister Müller Museum]]]] |
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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 typical "[[wastebasket taxon]]". Even after clearly different forms had later been given their own generic name, new species would be created from the very productive late Jurassic German sites, often based on only slightly different material. |
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 typical "[[wastebasket taxon]]". Even after clearly different forms had later been given their own generic name, new species would be created from the very productive late Jurassic German sites, often based on only slightly different material. |
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Around 1980, 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 '' |
Around 1980, 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 ''Ctenochasma'' species,<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. Christopher |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 /> |
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===Controversial species=== |
===Controversial species=== |
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[[File:Bsp as xix.png|thumb|Juvenile type specimen of ''Pterodactylus kochi'', now referred to ''[[Diopecephalus kochi]]'']] |
[[File:Bsp as xix.png|thumb|Juvenile type specimen of ''Pterodactylus kochi'', now referred to ''[[Diopecephalus kochi]]'']] |
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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}}</ref> More recent studies of pterosaur relationships have found |
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}}</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 = 1 | year = 2013 | pmid = | pmc = }}</ref> |
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<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 /> |
<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 /> |
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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}}</ref><ref>{{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|pages=125–147|doi=10.1144/SP455.12|url=https://eprints.soton.ac.uk/423063/1/Vidovic_Martill_2017_Taxonomy_of_Diopecephalus_and_Germanodactylus_AM_with_Figures.pdf|bibcode=2018GSLSP.455..125V}}</ref> |
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}}</ref><ref>{{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|pages=125–147|doi=10.1144/SP455.12|url=https://eprints.soton.ac.uk/423063/1/Vidovic_Martill_2017_Taxonomy_of_Diopecephalus_and_Germanodactylus_AM_with_Figures.pdf|bibcode=2018GSLSP.455..125V}}</ref> |
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In 2017, Bennett challenged this hypothesis. He claimed that while Vidovic and Martill had identified real differences between the 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 | url = | journal = Journal of Paleontology | volume = | issue = | pages = 1–18| doi = 10.1017/jpa.2017.84 }}</ref> |
In 2017, Bennett challenged this hypothesis. He claimed that while Vidovic and Martill had identified real differences between the 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 | url = | journal = Journal of Paleontology | volume = | issue = | pages = 1–18| doi = 10.1017/jpa.2017.84 }}</ref> |
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===List of former species and ''nomina dubia''=== |
===List of former species and ''nomina dubia''=== |
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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 ''[[Nomen dubium|nomina dubia]]'' (meaning "doubtful names"). The following list includes names that |
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 ''[[Nomen dubium|nomina dubia]]'' (meaning "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.<ref>{{cite book|last=Caroline|first=Arnold|title=Pterosaurs: Rulers of the Skies in the Dinosaur Age|year=2014|isbn=978-1-63083-412-8|ref=harv}}</ref> |
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|[[1825 in paleontology|1825]] |
|[[1825 in paleontology|1825]] |
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|''[[Nomen dubium]]'', possible synonym of ''[[Rhamphorhynchus muensteri]]'' |
|''[[Nomen dubium]]'', possible synonym of ''[[Rhamphorhynchus muensteri]]'' |
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|''Pterodactylus macronyx'' |
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|[[William Buckland|Buckland]] |
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|[[1829 in paleontology|1829]] |
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|Reclassified as ''[[Dimorphodon macronyx]]'' |
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|''Pterodactylus banthensis'' |
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|Theodori |
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|[[1830 in paleontology|1830]] |
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|Reclassified as ''[[Dorygnathus banthensis]]'' |
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|''Pterodactylus crassirostris'' |
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|Goldfuss |
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|[[1831 in paleontology|1831]] |
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|Reclassified as ''[[Scaphognathus crassirostris]]'' |
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|Reclassified as ''[[Diopecephalus kochi]]'' |
|Reclassified as ''[[Diopecephalus kochi]]'' |
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|Name correction from ''Ornithocephalus kochi''; former synonym of ''Pterodactylus antiquus'' |
|Name correction from ''Ornithocephalus kochi''; former synonym of ''Pterodactylus antiquus'' |
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|''Pterodactylus lavateri'' |
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|von Meyer |
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|[[1839 in paleontology|1839]] |
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|Synonym of ''Rhamphorhynchus muensteri'' |
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|''Pterodactylus longicaudus'' |
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|Münster |
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|[[1838 in paleontology|1838]] |
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|Reclassified as ''[[Rhamphorhynchus longicaudus]]'' |
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|''Pterodactylus meyeri'' |
|''Pterodactylus meyeri'' |
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|''Pterodactylus gemmingi'' |
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|''Ornithocephalus meyeri'' |
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|Münster |
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|(Muenster) Wagner |
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|[[1846 in paleontology|1846]] |
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|Synonym of '' |
|Synonym of ''Rhamphorhynchus muensteri'' |
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|Incorrect name correction from ''Pterodactylus meyeri'' |
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|- |
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|''Pterodactylus giganteus'' |
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|[[James Scott Bowerbank|Bowerbank]] |
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| Quenstedt |
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|[[1846 in paleontology|1846]] |
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|Reclassified as ''[[Lonchodraco giganteus]]'' |
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|''Pterodactylus cuvieri'' |
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|Bowerbank |
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|[[1851 in paleontology|1851]] |
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|Reclassified as ''[[Cimoliopterus cuvieri]]'' |
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|''Pterodactylus compressirostris'' |
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|[[Richard Owen|Owen]] |
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|[[1851 in paleontology|1851]] |
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|Reclassified as ''[[Lonchodectes compressirostris]]'' |
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|- |
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|''Pterodactylus suevicus'' |
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|Quenstedt |
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|[[1855 in paleontology|1855]] |
|[[1855 in paleontology|1855]] |
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|Reclassified as ''[[Cycnorhamphus suevicus]]'' |
|Reclassified as ''[[Cycnorhamphus suevicus]]'' |
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|- |
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|''Pterodactylus hirundinaceus'' |
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|Wagner |
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|[[1857 in paleontology|1857]] |
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|Synonym ''Rhamphorhynchus muensteri'' |
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|Reclassified as ''[[Ctenochasma elegans]]'' |
|Reclassified as ''[[Ctenochasma elegans]]'' |
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|''Pterodactylus simus'' |
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|[[Richard Owen|Owen]] |
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|[[1861 in paleontology|1861]] |
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|Reclassified as ''[[Ornithocheirus simus]]'' |
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|''Pterodactylus oweni'' |
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|[[Othniel Marsh|Marsh]] |
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|[[1871 in paleontology|1871]] |
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|''Nomen dubium'' |
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|Reassigned as ''Pterodactylus occidentalis'' prior to being preoccupied |
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|- |
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|''Pterodactylus ingens'' |
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|Marsh |
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|[[1872 in paleontology|1872]] |
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|''Nomen dubium'' |
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|Reclassified as ''[[Pteranodon]] ingens'' |
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|- |
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|''Pterodactylus occidentalis'' |
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|Marsh |
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|[[1872 in paleontology|1872]] |
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|''Nomen dubium'' |
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|Reclassified from ''Pterodactylus oweni''; Reclassified as ''Pteranodon occidentalis'' |
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|- |
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|''Pterodactylus velox'' |
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|Marsh |
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|[[1872 in paleontology|1872]] |
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|''Nomen dubium'' |
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|Reclassified as ''Pteranodon velox'' |
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|- |
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|''Pterodactylus suprajurensis'' |
|''Pterodactylus suprajurensis'' |
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|- |
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|''Pterodactylus manseli'' |
|''Pterodactylus manseli'' |
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|Owen |
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|[[1874 in paleontology|1874]] |
|[[1874 in paleontology|1874]] |
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|''Nomen dubium'' |
|''Nomen dubium'' |
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|[[1874 in paleontology|1874]] |
|[[1874 in paleontology|1874]] |
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|''Nomen dubium'' |
|''Nomen dubium'' |
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|- |
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|''Pterodactylus umbrosus'' |
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|[[Edward Cope|Cope]] |
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|[[1874 in paleontology|1874]] |
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|''Nomen dubium'' |
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|Reclassified from ''[[Ornithochirus umbrosus]]'' |
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|- |
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|''Pterodactylus cristatus'' |
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|[[Carl Wiman|Wiman]] |
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|[[1925 in paleontology|1925]] |
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|Reclassified as ''[[Germanodactylus cristatus]]'' |
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==See also== |
==See also== |
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{{Portal|Paleontology}} |
{{Portal|Paleontology}} |
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* [[List of pterosaur genera]] |
* [[List of pterosaur genera]] |
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* [[Timeline of pterosaur research]] |
* [[Timeline of pterosaur research]] |
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==References== |
==References== |
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{{Reflist|35em}} |
{{Reflist|35em}} |
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{{Commons category|Pterodactylus}} |
{{Commons category|Pterodactylus}} |
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{{Pterosauria|Ar.}} |
{{Pterosauria|Ar.}} |
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{{fins, limbs and wings}} |
{{fins, limbs and wings}} |
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{{Portal bar|Dinosaurs|Germany|Biology}} |
{{Portal bar|Dinosaurs|Germany|Biology}} |
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{{Taxonbar|from=Q309490}} |
{{Taxonbar|from=Q309490}} |
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{{Authority control}} |
{{Authority control}} |
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[[Category:Late Jurassic pterosaurs of Europe]] |
[[Category:Late Jurassic pterosaurs of Europe]] |
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[[Category:Ctenochasmatoids]] |
[[Category:Ctenochasmatoids]] |
Revision as of 13:48, 7 June 2020
Pterodactylus Temporal range: Early Tithonian,
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Sub-adult type specimen of P. antiquus, Bavarian State Collection for Palaeontology and Geology | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Order: | †Pterosauria |
Suborder: | †Pterodactyloidea |
Infraorder: | †Archaeopterodactyloidea |
Clade: | †Euctenochasmatia Bonaparte, 1838 |
Genus: | †Pterodactylus Cuvier, 1809 |
Type species | |
†Pterodactylus antiquus Sömmerring, 1812
| |
Synonyms | |
Genus synonymy
Species synonymy
|
Pterodactylus (from Greek πτεροδάκτυλος, pterodaktulos, meaning "winged finger"[2]) is an extinct genus of euctenochasmatian pterodactyloid pterosaur, whose members are commonly known as pterodactyls ( /ˌtɛrəˈdæktɪlz/). It is thought to contain only a single species, Pterodactylus antiquus, the first pterosaur species 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.[3]
Many studies conclude that Pterodactylus was a carnivore that probably preyed upon fish as well as other small animals. 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 keratinous 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. The classification of Pterodactylus has since been confusing to paleontologists. Some classify Pterodactylus within the clade Euctenochasmatia, while others just consider it as a basal member of the suborder Pterodactyloidea.[4][5]
History of discovery
The type specimen of the animal now known as Pterodactylus antiquus was one of the first pterosaur fossils ever to be identified. The first Pterodactylus specimen was described by the Italian scientist 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 "Naturalienkabinett", or nature cabinet (a precursor to the modern concept of the natural history museum), in the palace of Charles Theodore, Elector of Bavaria at Mannheim.[6] The specimen had been given to the collection by Count Friedrich Ferdinand zu Pappenheim, probably around 1780, having been recovered from a lithographic limestone quarry in Eichstätt.[7] The actual date of the specimen's discovery and entry into the collection is unknown. It was not mentioned in a catalogue of the collection taken in 1767 and so 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 Pterodactylus micronyx was described in 1779 and possibly discovered earlier than the Mannheim specimen, but it was at first considered to be a fossil crustacean.[8]
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.[9][10] 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 plesiosaurs, ichthyosaurs, and monotremes), in the class Gryphi, between birds and mammals.[11]
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 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 commissars 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 bats. 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.[10] 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." However, contrary to Hermann, Cuvier was convinced the animal was a reptile.[12]
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. 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 a "ptero-dactyle" and refuted a hypothesis by Johann Friedrich Blumenbach that it would have been a shore bird.[13]
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 27 December 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.[14] The animal was described as being both a mammal, 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.[15] 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 Soemmerring 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[16]). He provided a restoration of the skeleton, the first one published for any pterosaur.[10] This restoration was very inaccurate, von Söemmerring mistaking the long metacarpals 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 blades.[17] Söemmerring 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.[18]
Description
Pterodactylus is known from over 30 fossil specimens, and though most of those are juveniles, many preserve complete skeletons.[16][19] Pterodactylus antiquus was a relatively small pterosaur, with an estimated adult wingspan of about 1.04 meters (3 feet 5 inches) (the only known adult specimen is represented by an isolated skull).[16] Other "species" were once thought to be smaller. 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).[20]
The skulls of adult Pterodactylus were long and thin with about 90 narrow, conical teeth. The teeth extended back from the tips of both jaws, and became smaller farther away from the jaw tips (unlike some relatives, where teeth were absent in the upper jaw tip and were relatively uniform in size). The teeth extended farther back into the jaw than in close relatives, as some were present below the front of the nasoantorbital fenestra, the largest opening in the skull.[16] Unlike related species, the skull and jaws were straight, not curved upwards.[21]
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 (the largest opening in the skull) 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.[16][22] Bennett (2013) 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.[16] 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.[16] 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. This is BSP 1929 I 18, which has an occipital lappet similar to the proposed Aerodactylus definition. This specimen also has a small triangular soft tissue crest with the peak of the crest positioned above the eyes.[16]
Paleobiology
Life history
Like other pterosaurs (notably 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 Pterodactylus species, P. antiquus.[21]
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), and the teeth have a relatively broad base.[19] The teeth of other P. antiquus specimens are both narrower and more numerous (up to 90 teeth are present in some specimens).[21]
Pterodactylus specimens can be divided into two distinct year classes. In the first year class, the skulls are only 15-45 mm (0.59-1.77 inches) in length. The second year class is characterized by skulls 55-95 mm (2.16-3.74 inches) long, but still immature. 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.[19]
Growth and breeding seasons
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.[19][23] 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 crocodilians, rather than the rapid growth of modern birds.[19]
Daily activity patterns
Comparisons between the scleral rings of Pterodactylus antiquus and modern birds and reptiles suggest that it may have been diurnal. This may also indicate niche partitioning with contemporary pterosaurs inferred to be nocturnal, such as Ctenochasma and Rhamphorhynchus.[24]
Paleoecology
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.[3] The Solnhofen Limestone is diverse Lagerstätte that contains a wide range of different creatures, including highly detailed fossilized imprints of soft bodied organisms such as jellyfishes. Abundant specimens of pterosaurs similar to Pterodactylus were also found within the formation, these include the rhamphorhynchids Rhamphorhynchus and Scaphognathus,[25] several gallodactylids such as Aerodactylus,[19] Ardeadactylus, Aurorazhdarcho and Cycnorhamphus,[6] the ctenochasmatids Ctenochasma[26] and Gnathosaurus, the anurognathid Anurognathus, the germanodactylid Germanodactylus, as well as the basal euctenochasmatian Diopecephalus.[27] Fossil remains of the dinosaurs 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.[28] Various lizard remains were also found alongside those of Pterodactylus, with several specimens assigned to Ardeosaurus, Bavarisaurus and Eichstaettisaurus.[29][30] Crocodylomorph specimens were widely distributed within the fossil site, most were assigned to the metriorhynchid genera Cricosaurus, Dakosaurus, Geosaurus and Rhacheosaurus. These genera are colloquially called as marine or sea crocodiles due to their similar built.[31][32] The rhynchocephalian genera Homoeosaurus and Pleurosaurus were also found in the formation alongside several turtles such as Eurysternum and Paleomedusa.[33][34] Fossils of the ichthyosaur Aegirosaurus were also present in the site,[35] as well as fish remains, with many specimens assigned to ray-finned fishes such as the halecomorphs Lepidotes,[36] Propterus,[37] Gyrodus, Caturus[37] and Ophiopsis,[38] the pachycormids Asthenocormus and Hypsocormus,[39] as well as the teleosts Aspidorhynchus and Thrissops.[40][41]
Classification
The genus now known as Pterodactylus was originally named Petro-Dactyle by Cuvier in 1809,[13] though this was a typographical error, later corrected by him to Ptéro-Dactyle.[10] In 1812, Samuel Thomas von Sömmerring named the same specimen Ornithocephalus antiquus. The genus name was emended to the current Pterodactylus by Constantine Samuel Rafinesque in 1815. Unaware of Rafinesque's publication, Cuvier himself in 1819 again emended the genus name,[42] but the specific name he then gave, longirostris, has to give precedence to von Soemmerring's antiquus. In 1888 Richard Lydekker designated Pterodactylus antiquus the type species. The original specimen is the holotype of the genus, BSP No. AS.I.739.
Hermann von Meyer, in 1830, used the name Pterodactyli to contain Pterodactylus and other pterosaurs known at the time. This was emended to the family Pterodactylidae by Prince Charles Lucien Bonaparte in 1838. This group has more recently been given several competing definitions.[4][43]
Beginning in 2014, researchers Steven Vidovic and David Martill constructed an analysis in which several pterosaurs traditionally thought of as archaeopterodactyloids closely related to the ctenochasmatoids may have been more closely related to the more advanced ornithocheiroids, 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.[44]
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, they placed Pterodactylus within the clade Euctenochasmatia, resulting in a more derived position.[5]
Formerly assigned species
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 typical "wastebasket taxon". Even after clearly different forms had later been given their own generic name, new species would be created from the very productive late Jurassic German sites, often based on only slightly different material.
Around 1980, 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.[21] 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 subulatus or one of the Ctenochasma species,[20][21] though after additional research Bennett assigned it to the genus Aurorazhdarcho.[16] 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.[4] 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.[45] "P." longicollum was eventually made the type species of a separate genus Ardeadactylus.[16]
Controversial species
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.[4] 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.[46] [45] 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.[21] 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.[16]
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.[47][48]
In 2017, Bennett challenged this hypothesis. He claimed that while Vidovic and Martill had identified real differences between the 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.[49]
List of former species and nomina dubia
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 nomina dubia (meaning "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.[50]
Name | Author | Year | Status | Notes |
---|---|---|---|---|
Pterodactylus grandis | Cuvier | 1825 | Nomen dubium, possible synonym of Rhamphorhynchus muensteri | |
Pterodactylus macronyx | Buckland | 1829 | Reclassified as Dimorphodon macronyx | |
Pterodactylus banthensis | Theodori | 1830 | Reclassified as Dorygnathus banthensis | |
Pterodactylus crassirostris | Goldfuss | 1831 | Reclassified as Scaphognathus crassirostris | |
Pterodactylus kochi | (Wagner) | (1837) | Reclassified as Diopecephalus kochi | Name correction from Ornithocephalus kochi; former synonym of Pterodactylus antiquus |
Pterodactylus lavateri | von Meyer | 1839 | Synonym of Rhamphorhynchus muensteri | |
Pterodactylus longicaudus | Münster | 1838 | Reclassified as Rhamphorhynchus longicaudus | |
Pterodactylus meyeri | Muenster | 1842 | Synonym of Diopecephalus kochi | |
Pterodactylus gemmingi | Münster | 1846 | Synonym of Rhamphorhynchus muensteri | |
Pterodactylus giganteus | Bowerbank | 1846 | Reclassified as Lonchodraco giganteus | |
Pterodactylus cuvieri | Bowerbank | 1851 | Reclassified as Cimoliopterus cuvieri | |
Pterodactylus compressirostris | Owen | 1851 | Reclassified as Lonchodectes compressirostris | |
Pterodactylus suevicus | Quenstedt | 1855 | Reclassified as Cycnorhamphus suevicus | |
Pterodactylus hirundinaceus | Wagner | 1857 | Synonym Rhamphorhynchus muensteri | |
Pterodactylus grandipelvis | von Meyer | 1860 | Nomen dubium | |
Pterodactylus cerinensis | von Meyer | 1860 | Nomen dubium | |
Pterodactylus elegans | Wagner | 1861 | Reclassified as Ctenochasma elegans | |
Pterodactylus simus | Owen | 1861 | Reclassified as Ornithocheirus simus | |
Pterodactylus oweni | Marsh | 1871 | Nomen dubium | Reassigned as Pterodactylus occidentalis prior to being preoccupied |
Pterodactylus ingens | Marsh | 1872 | Nomen dubium | Reclassified as Pteranodon ingens |
Pterodactylus occidentalis | Marsh | 1872 | Nomen dubium | Reclassified from Pterodactylus oweni; Reclassified as Pteranodon occidentalis |
Pterodactylus velox | Marsh | 1872 | Nomen dubium | Reclassified as Pteranodon velox |
Pterodactylus suprajurensis | Sauvage | 1873 | Nomen dubium | |
Pterodactylus manseli | Owen | 1874 | Nomen dubium | |
Pterodactylus pleydelli | Owen | 1874 | Nomen dubium | |
Pterodactylus umbrosus | Cope | 1874 | Nomen dubium | Reclassified from Ornithochirus umbrosus |
Pterodactylus cristatus | Wiman | 1925 | Reclassified as Germanodactylus cristatus | |
Pterodactylus arningi | Reck | 1931 | Nomen dubium | |
Pterodactylus maximus | Reck | 1931 | Nomen dubium |
See also
References
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{{cite book}}
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