Ornithocheiromorpha: Difference between revisions
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{{short description|Clade of pteranodontoid pterosaurs}} |
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{{Good article}} |
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{{Use mdy dates|date=March 2021}} |
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{{Automatic taxobox |
{{Automatic taxobox |
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| name = Ornithocheiromorphs |
| name = Ornithocheiromorphs |
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| fossil_range = [[Valanginian]]-[[Turonian]]<br>~{{fossil_range|140| |
| fossil_range = [[Valanginian]]-[[Turonian]]<br/>~{{fossil_range|140|92.5}} |
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| image = Tropeognathus mesembrinus MN 01.jpg |
| image = Tropeognathus mesembrinus MN 01.jpg |
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| image_caption = Reconstructed skeleton of ''[[Tropeognathus]]'' in the [[National Museum of Brazil]] |
| image_caption = Reconstructed skeleton of ''[[Tropeognathus]]'' in the [[National Museum of Brazil]] |
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| image2 = Coloborhynchus spielbergi2.jpg |
| image2 = Coloborhynchus spielbergi2.jpg |
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| image2_caption = Skeletal cast of ''[[ |
| image2_caption = Skeletal cast of ''[[Maaradactylus spielbergi]]'' in the [[Naturalis Biodiversity Center]] |
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| taxon = Ornithocheiromorpha |
| taxon = Ornithocheiromorpha |
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| authority = Andres ''et al.'', 2014 |
| authority = Andres ''et al.'', 2014 |
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| subdivision_ranks = Subgroups |
| subdivision_ranks = Subgroups |
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| subdivision = |
| subdivision = |
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* {{extinct}}''[[Draigwenia]]''? |
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* {{extinct}}''[[Prejanopterus]]''? |
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* {{extinct}}''[[Serradraco]]'' |
* {{extinct}}''[[Serradraco]]'' |
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* {{extinct}}''[[Unwindia]]'' |
* {{extinct}}''[[Unwindia]]'' |
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* {{extinct}}''[[Yixianopterus]]'' |
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* {{extinct}}[[Lonchodectidae]] |
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* {{extinct}}'''Lanceodontia''' |
* {{extinct}}'''Lanceodontia''' |
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** {{extinct}}[[Boreopteridae]] |
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** {{extinct}}[[Targaryendraconia]] |
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** {{extinct}}'''Istiodactyliformes''' |
** {{extinct}}'''Istiodactyliformes''' |
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*** {{extinct}}''[[Haopterus]]'' |
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*** {{extinct}}''[[Hongshanopterus]]'' |
*** {{extinct}}''[[Hongshanopterus]]'' |
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*** {{extinct}}''[[Linlongopterus]]'' |
*** {{extinct}}''[[Linlongopterus]]'' |
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*** {{extinct}}''[[Yixianopterus]]''<ref>{{Cite journal|last1=Jiang|first1=Shun-Xing|last2=Zhang|first2=Xin-Jun|last3=Cheng|first3=Xin|last4=Wang|first4=Xiao-Lin|year=2020|title=A new pteranodontoid pterosaur forelimb from the upper Yixian Formation, with a revision of ''Yixianopterus jingangshanensis''|journal=Vertebrata PalAsiatica|doi=10.19615/j.cnki.1000-3118.201124}}</ref> |
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*** {{extinct}}[[Lonchodectidae]] |
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*** {{extinct}}[[Mimodactylidae]] |
*** {{extinct}}[[Mimodactylidae]] |
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*** {{extinct}}[[Istiodactylidae]] |
*** {{extinct}}[[Istiodactylidae]] |
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** {{extinct}}''' |
** {{extinct}}'''Ornithocheiriformes''' |
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*** {{extinct}} |
*** {{extinct}}[[Boreopteridae]] |
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*** {{extinct}} |
*** {{extinct}}[[Ornithocheirae]] |
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*** {{extinct}}''[[Guidraco]]'' |
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*** {{extinct}}''[[Ludodactylus]]'' |
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*** {{extinct}}[[Hamipteridae]] |
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*** {{extinct}}'''[[Ornithocheirae]]''' |
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**** {{extinct}}[[Anhangueridae]] |
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**** {{extinct}}[[Ornithocheiridae]] |
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}} |
}} |
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'''Ornithocheiromorpha''' (from [[Ancient Greek]], meaning "bird hand form") is a group of [[pterosaurs]] within the [[suborder]] [[Pterodactyloidea]]. [[Fossil]] remains of this group date back from the [[Early Cretaceous|Early]] to [[Late Cretaceous]] [[Period (geology)|periods]] ([[Valanginian]] to [[Turonian]] stages), around 140 to |
'''Ornithocheiromorpha''' (from [[Ancient Greek]], meaning "bird hand form") is a group of [[pterosaurs]] within the [[suborder]] [[Pterodactyloidea]]. [[Fossil]] remains of this group date back from the [[Early Cretaceous|Early]] to [[Late Cretaceous]] [[Period (geology)|periods]] ([[Valanginian]] to [[Turonian]] stages), around 140 to 92.5 million years ago. Ornithocheiromorphs were discovered worldwide except [[Antarctica]], though most [[genera]] were recovered in [[Europe]], [[Asia]] and [[South America]].<ref>Barrett, P. M., Butler, R. J., Edwards, N. P., & Milner, A. R. (2008). Pterosaur distribution in time and space: an atlas. ''Zitteliana'': 61-107.[http://epub.ub.uni-muenchen.de/12007/1/zitteliana_2008_b28_05.pdf]</ref> They were the most diverse and successful pterosaurs during the Early Cretaceous, but throughout the Late Cretaceous they were replaced by [[pteranodontia]]ns and [[azhdarchoid]]s. The Ornithocheiromorpha was defined in 2014 by Andres and colleagues, and they made Ornithocheiromorpha the most inclusive [[clade]] containing ''Ornithocheirus'', but not ''[[Pteranodon]]''.<ref name=kryptodrakon>{{Cite journal | doi = 10.1016/j.cub.2014.03.030| title = The Earliest Pterodactyloid and the Origin of the Group| journal = Current Biology| year = 2014| last1 = Andres | first1 = B. | last2 = Clark | first2 = J. | last3 = Xu | first3 = X. | pmid=24768054 | volume=24 | issue = 9| pages=1011–6| doi-access = free }}</ref> |
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Ornithocheiromorphs are considered to be some of the largest animals to have ever flown. Members of this group are also regarded to have some of the largest pterosaur [[wingspan]]s, such as the one estimated for the huge ''[[Tropeognathus]]'', though still not as large as those estimated for the [[azhdarchid]]s, which may have reached up to {{convert|12|m|ft|sp=us}}.<ref name=witton2010>{{cite journal| |
Ornithocheiromorphs are considered to be some of the largest animals to have ever flown. Members of this group are also regarded to have some of the largest pterosaur [[wingspan]]s, such as the one estimated for the huge ''[[Tropeognathus]]'', though still not as large as those estimated for the [[azhdarchid]]s, which may have reached up to {{convert|12|m|ft|sp=us}}.<ref name=witton2010>{{cite journal|author=Witton, M.P. |author2=Martill, D.M. |author3=Loveridge, R.F. |year=2010|title=Clipping the Wings of Giant Pterosaurs: Comments on Wingspan Estimations and Diversity.|journal=Acta Geoscientica Sinica|volume=31|pages=79–81|url=https://www.researchgate.net/publication/287830761}}</ref> When ornithocheiromorphs first appeared, they were initially [[scavenger]]s, consisting in a more terrestrial setting, but their success had made them the top predators of the skies, as well as the most common type of [[Piscivore|fish-eating]] pterosaur throughout the early Late Cretaceous. Some [[paleontologists]] also consider ornithocheiromorphs an earlier step of evolution to the [[Pteranodontia|pteranodontians]], this is due to the similar flying techniques and flight locomotions, as well as their diet, which mainly consisted of fish, and therefore also hunted very similarly. Ornithocheiromorphs also flew like soaring birds, keeping their wings stretched and rarely flapping. |
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== History of research == |
== History of research == |
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=== Early discoveries === |
=== Early discoveries === |
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[[File:Pterodactylus giganteus.jpg|thumb|upright|left|Associated and referred bones of ''[[Pterodactylus]] giganteus'', now considered as ''[[Lonchodraco giganteus]]'']] |
[[File:Pterodactylus giganteus.jpg|thumb|upright|left|Associated and referred bones of ''[[Pterodactylus]] giganteus'', now considered as ''[[Lonchodraco giganteus]]'']] |
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The first specimens of ornithocheiromorphs were unearthed at a chalk pit near [[Burham]] in [[Kent]], [[England]]. In 1846, [[British people|British]] [[paleontologist]] [[James Scott Bowerbank]] named and described the remains found as ''[[Pterodactylus]] giganteus'', as it was common at that time to assign any new described pterosaur species to ''Pterodactylus''.<ref name=Bowerbank1846/> In the same chalk pit as ''P. giganteus'', two other pterosaur species were discovered. The first was named in 1851 by Bowerbank as ''Pterodactylus cuvieri'',<ref name="Bowerbank1851">{{cite journal | last1 = Bowerbank | first1 = J.S. | year = 1851 | title = On the pterodactyles of the Chalk Formation | url = https://zenodo.org/record/1447536| journal = Proceedings of the Zoological Society of London | volume = 19 |
The first specimens of ornithocheiromorphs were unearthed at a chalk pit near [[Burham]] in [[Kent]], [[England]]. In 1846, [[British people|British]] [[paleontologist]] [[James Scott Bowerbank]] named and described the remains found as ''[[Pterodactylus]] giganteus'', as it was common at that time to assign any new described pterosaur species to ''Pterodactylus''.<ref name=Bowerbank1846>{{cite journal | last1 = Bowerbank | first1 = J.S. | year = 1846 | title = On a new species of pterodactyl found in the Upper Chalk of Kent (''Pterodactylus giganteus'') | url =https://zenodo.org/record/1448505 | journal = Quarterly Journal of the Geological Society of London | volume = 2 | issue = 1–2| pages = 7–9 | doi=10.1144/gsl.jgs.1846.002.01-02.05| s2cid = 129389179 }}</ref> In the same chalk pit as ''P. giganteus'', two other pterosaur species were discovered. The first was named in 1851 by Bowerbank as ''Pterodactylus cuvieri'',<ref name="Bowerbank1851">{{cite journal | last1 = Bowerbank | first1 = J.S. | year = 1851 | title = On the pterodactyles of the Chalk Formation | url = https://zenodo.org/record/1447536| journal = Proceedings of the Zoological Society of London | volume = 19 | pages = 14–20 | doi=10.1111/j.1096-3642.1851.tb01125.x}}</ref> in honor of the prominent German [[naturalist]] and [[zoologist]] [[Georges Cuvier]], while the second was described in the same year by British paleontologist Sir [[Richard Owen]] as ''Pterodactylus compressirostris''.<ref name=Owen1851>Owen, R. (1851). Monograph on the fossil Reptilia of the Cretaceous Formations. ''The Palaeontographical Society'' '''5'''(11):1-118.</ref> ''P. compressirostris'' later became the type species of a newly created genus called ''[[Lonchodectes]]'' (meaning "[[lance]] biter") in a review by English paleontologist [[Reginald Walter Hooley]] in 1914.<ref name=HGS70/> Confusingly, this species was also long regarded, incorrectly, as the type species of ''Ornithocheirus''.<ref name=RH14>{{Cite journal|last=Hooley|first=Reginald Walter|date=1914|title=On the Ornithosaurian genus ''Ornithocheirus'', with a review of the specimens from the Cambridge Greensand in the Sedgwick Museum, Cambridge|url=https://zenodo.org/record/2207691|journal=Annals and Magazine of Natural History|language=en|volume=13|issue=78|pages=529–557|doi=10.1080/00222931408693521|issn=0374-5481}}</ref> |
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In 1861, further pterosaur specimens were found in the [[UK]], and were given the new species ''Pterodactylus simus'' by Owen.<ref name=History>{{cite journal|author=Martill, D.M.|year=2010|title=The early history of pterosaur discovery in Great Britain|journal=Geological Society of London, Special Publications|volume=343|issue=1|pages=287–311|doi=10.1144/SP343.18|s2cid=130116778|url=https://sp.lyellcollection.org/content/343/1/287}}</ref> British paleontologist [[Harry Govier Seeley]] then created the new genus ''Ornithocheirus'' for the new species in the same year, the generic name translating as "bird hand" is due to the notion of the time that pterosaurs were the ancestors of modern birds. |
In 1861, further pterosaur specimens were found in the [[UK]], and were given the new species ''Pterodactylus simus'' by Owen.<ref name=History>{{cite journal|author=Martill, D.M.|year=2010|title=The early history of pterosaur discovery in Great Britain|journal=Geological Society of London, Special Publications|volume=343|issue=1|pages=287–311|doi=10.1144/SP343.18|bibcode=2010GSLSP.343..287M |s2cid=130116778|url=https://sp.lyellcollection.org/content/343/1/287}}</ref> British paleontologist [[Harry Govier Seeley]] then created the new genus ''[[Ornithocheirus]]'' for the new species in the same year, the generic name translating as "bird hand" is due to the notion of the time that pterosaurs were the ancestors of modern birds. In 1870, Seeley reassigned the species ''Pterodactylus cuvieri'' as ''Ornithocheirus cuvieri''.<ref>Seeley, H.G., 1869, ''Index to the fossil remains of'' Aves, Ornithosauria, ''and'' Reptilia, ''from the Secondary System of Strata, arranged in the Woodwardian Museum of the University of Cambridge''. St. John's College, Cambridge '''8''': 143. {{doi|10.1080/00222937008696143}}</ref><ref name=HGS70>{{cite journal|author=Seeley, H.G.|year=1870|title=The Ornithosauria: an Elementary Study of the Bones of Pterodactyles|journal=Cambridge|page=113|url=https://books.google.com/books?id=0M8yAQAAMAAJ}}</ref> In 1874, Richard Owen proposed two new genera, ''[[Coloborhynchus]]'', meaning "maimed beak", and ''[[Criorhynchus]]'', meaning "ram beak". While ''Coloborhynchus'' consisted in a totally new type species, ''C. clavirostris'', as well as two other species reassigned from ''Ornithocheirus'', ''Criorhynchus'' consisted entirely of former ''Ornithocheirus'' species, including ''O. simus'', which was later reassigned by Owen as ''Criorhynchus simus''.<ref>Owen, R. 1874, ''Monograph on the fossil Reptilia of the Mesozoic Formations''. Palaeontographical Society, London, 14 pp</ref> |
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In 1869, Seeley reassigned ''Pterodactylus cuvieri'' into ''Ptenodactylus cuvieri'', but in 1870, Seeley had realized that the generic name ''Ptenodactylus'' had been considered a ''[[nomen nudum]]'', and therefore reassigned the species into ''Ornithocheirus cuvieri''. Between 1869 and 1870, Seeley reassigned various species of pterosaurs, including ''P. cuvieri'' into newly named species of ''[[Ornithocheirus]]'', therefore 27 in total.<ref>Seeley, H.G., 1869, ''Index to the fossil remains of'' Aves, Ornithosauria, ''and'' Reptilia, ''from the Secondary System of Strata, arranged in the Woodwardian Museum of the University of Cambridge''. St. John's College, Cambridge '''8''': 143. {{DOI|10.1080/00222937008696143}}</ref><ref name=HGS70>{{cite journal|author=Seeley, H.G.|year=1870|title=The Ornithosauria: an Elementary Study of the Bones of Pterodactyles|journal=Cambridge|page=113|url=https://books.google.com/books?id=0M8yAQAAMAAJ}}</ref> |
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Richard Owen wasn't pleased when Seeley published his conclusions in his 1870 book ''The Ornithosauria'', and he therefore considered the name ''Ornithocheirus'' to be inappropriate. In 1874, Owen created two new genera, ''[[Coloborhynchus]]'', meaning "maimed beak" and ''[[Criorhynchus]]'', meaning "ram beak". |
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While ''Coloborhynchus'' consisted in a totally new type species, ''C. clavirostris'', as well as two other species reassigned from ''Ornithocheirus'', ''Criorhynchus'' consisted entirely of former ''Ornithocheirus'' species, including the type species, ''O. simus'', which was later reassigned by Owen as ''Criorhynchus simus'', as well as 5 other species. In the same year, Owen also reassigned ''O. cuvieri'' into ''Coloborhynchus cuvieri''.<ref>Owen, R. 1874, ''Monograph on the fossil Reptilia of the Mesozoic Formations''. Palaeontographical Society, London, 14 pp</ref> Seeley later disagreed and didn't accept Owen's position, and he therefore assigned ''Ornithocheirus simus'' as the type species of ''Ornithocheirus'' in 1881. Seeley had also named a new separate species called ''[[Ornithocheirus buenzeli|O. bunzeli]]'' in the same year.<ref>{{cite journal|author=Seeley, H. G.|authorlink=Harry Seeley|year=1881|title=The Reptile Fauna of the Gosau Formation preserved in the Geological Museum of the University of Vienna|journal=Quarterly Journal of the Geological Society|volume=37|issue=1|pages=620–706|doi=10.1144/GSL.JGS.1881.037.01-04.49|s2cid=219235284|url=https://jgs.lyellcollection.org/content/37/1-4/620}}</ref> |
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In 1914, English paleontologist [[Reginald Walter Hooley]] kept the name ''Ornithocheirus'', but created two new genera: ''[[Lonchodectes]]'', meaning "[[lance]] biter", and ''[[Amblydectes]]'', meaning "blunt biter". This classification was rarely applied at the time, though.<ref name=RH14/> The species ''Pterodactylus compressirostris'' was formerly transferred to ''Ornithocheirus'' in 1870 by Seeley, but it became the type species of ''Lonchodectes'' in Hooley's 1914 review of ''Ornithocheirus''.<ref name=HGS70/> Confusingly, this species was also long regarded, incorrectly, as the type species of ''Ornithocheirus''.<ref name=RH14>{{Cite journal|last=Hooley|first=Reginald Walter|date=1914|title=On the Ornithosaurian genus ''Ornithocheirus'', with a review of the specimens from the Cambridge Greensand in the Sedgwick Museum, Cambridge|url=https://zenodo.org/record/2207691|journal=Annals and Magazine of Natural History|language=en|volume=13|issue=78|pages=529–557|doi=10.1080/00222931408693521|issn=0374-5481|via=}}</ref> |
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[[File:PZSL1851PlateReptilia04.png|thumb|left|[[Holotype]] of ''Pterodactylus cuvieri'', now known as ''[[Cimoliopterus]]'']] |
[[File:PZSL1851PlateReptilia04.png|thumb|left|[[Holotype]] of ''Pterodactylus cuvieri'', now known as ''[[Cimoliopterus]]'']] |
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In 2013, Brazilian paleontologists [[Taissa Rodrigues]] & [[Alexander Kellner]] made a deeper analysis on the species ''Pterodactylus cuvieri''. In the analysis, they stated that it needed a separate genus, and assigning it to ''Ornithocheirus'' was inappropriate, therefore, they created the new genus called ''[[Cimoliopterus]]'', with the new resulting combination ''Cimoliopterus cuvieri'' |
In 2013, Brazilian paleontologists [[Taissa Rodrigues]] & [[Alexander Kellner]] made a deeper analysis on the species ''Pterodactylus cuvieri''. In the analysis, they stated that it needed a separate genus, and assigning it to ''Ornithocheirus'' was inappropriate, therefore, they created the new genus called ''[[Cimoliopterus]]'', with the new resulting combination ''Cimoliopterus cuvieri''. In the same study, Rodrigues & Kellner also reviewed the species ''Pterodactylus giganteus'', and reassigned it to a newly created genus called ''[[Lonchodraco]]'', this resulted in a new combination called ''Lonchodraco giganteus''.<ref name="Rodrigues & Kellner 2013"/> |
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[[File:Istiodactylus.tif|thumb|Skull fragments of ''Ornithodesmus latidens'', which is currently known as ''[[Istiodactylus latidens]]'']] |
[[File:Istiodactylus.tif|thumb|Skull fragments of ''Ornithodesmus latidens'', which is currently known as ''[[Istiodactylus latidens]]'']] |
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In 1887, Seeley had described new fossil remains from the [[Isle of Wight]], an island off the coast of southern England. He thought it belonged to some kind of bird-like creature, which he named it ''[[Ornithodesmus cluniculus]]''.<ref name=seeley1887>{{cite journal|author=Seeley, H. G.| |
In 1887, Seeley had described new fossil remains from the [[Isle of Wight]], an island off the coast of southern England. He thought it belonged to some kind of bird-like creature, which he named it ''[[Ornithodesmus cluniculus]]''.<ref name=seeley1887>{{cite journal|author=Seeley, H. G.|author-link=Harry Seeley|year=1887|title=On a sacrum apparently indicating a new type of bird, ''Ornithodesmus cluniculus'' Seeley from the Wealden of Brook|journal=Quarterly Journal of the Geological Society of London|volume=43|issue=1–4|pages=206–211|doi=10.1144/GSL.JGS.1887.043.01-04.19|s2cid=129459937|url=https://zenodo.org/record/2017266}}</ref> Seeley also reported another specimen found on the same site. He then considered it another species of ''Ornithodesmus''. In 1901, Seeley named this new species as ''O. latidens'', meaning "wide tooth".<ref name="Dragons">{{cite book|author=Seeley, H. G.|title=Dragons of the Air: an Account of Extinct Flying Reptiles|orig-date=1901|year=2015|publisher=D. Appleton & Co.|location=New York|pages=173–175|isbn=978-1-4400-8494-2}}</ref> Later, Reginald Hooley discussed ''O. latidens'' in detail, based on specimens he had found, which led ''Ornithodesmus'' to be placed within a new family called Ornithodesmidae.<ref name=hooley1913>{{cite journal|author=Hooley, R. W.|author-link=Reginald Hooley|title=On the skeleton of ''Ornithodesmus latidens''; an ornithosaur from the Wealden Shales of Atherfield (Isle of Wight)|journal=Quarterly Journal of the Geological Society|year=1913|volume=69|issue=1–4|pages=372–422|doi=10.1144/GSL.JGS.1913.069.01-04.23|s2cid=128604856}}</ref> Paleontologist [[Charles William Andrews]] however, had expressed doubts as to whether ''O. latidens'' belonged in the genus ''Ornithodesmus'', as the vertebrae of the specimen of that genus was based on differed markedly from those of Hooley's specimen.{{Sfn|Witton|2013|pp=143–151}} |
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In 1993, the British paleontologists [[Stafford C. Howse]] and [[Andrew C. Milner]] concluded that the holotype sacrum and only specimen of ''O. cluniculus'' didn't belong to a pterosaur, but instead to a [[maniraptora]]n [[theropod]] dinosaur. They also pointed out that no detailed attempts had been made to compare the sacrum of ''O. cluniculus'' with those of pterosaurs, and that ''O. latidens'' had in effect been treated as the type species of the genus ''Ornithodesmus''.<ref name="Ornithodesmus'">{{cite journal|author=Howse, S. C. B.|author2=Milner, A. R.|title=''Ornithodesmus'' – a maniraptoran theropod dinosaur from the Lower Cretaceous of the Isle of Wight, England|journal=Palaeontology|year=1993|volume=36|pages=425–437|url=https://www.palass.org/publications/palaeontology-journal/archive/36/2/article_pp425-437}}</ref> Howse, Milner, and David Martill in 2001, moved "''O.''" ''latidens'' to a new genus called ''[[Istiodactylus]]''. They had also named a new family called [[Istiodactylidae]], with ''Istiodactylus'' as the only member.<ref name="PterosaursWight">{{cite book|author=Howse, S. C. B.|author2=Milner, A. R.|author3=Martill, D. M.|year=2001|pages=324–335|title=Dinosaurs of the Isle of Wight|place=London|publisher=The Palaeontological Association|series=Guide 10; Field Guides to Fossils|chapter=Pterosaurs|editor=Martill, D. M.|editor2=Naish, D.|editor-link2=Darren Naish|isbn=978-0-901702-72-2}}</ref> |
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Reginald Hooley later described two more specimens of ''O. latidens'' in 1913, which were collected in 1904 from the sea after a [[rockfall]] near [[Atherfield]] Point on the Isle of Wight. The first of the specimens was collected, and consisted of a skull, neck and trunk vertebrae, a shoulder blade, an [[ischium]], and parts of the [[forelimbs]]. The second specimen was collected in just one block, and it includes parts of the forelimbs and [[pectoral girdle]]. These two specimens represent the most complete remains of [[Cretaceous]] pterosaurs found in England. Hooley also discussed ''O. latidens'' in detail, which led ''Ornithodesmus'' to be placed within its own family, Ornithodesmidae.<ref name=hooley1913>{{cite journal|author=Hooley, R. W.|authorlink=Reginald Hooley|title=On the skeleton of ''Ornithodesmus latidens''; an ornithosaur from the Wealden Shales of Atherfield (Isle of Wight)|journal=Quarterly Journal of the Geological Society|year=1913|volume=69|issue=1–4|pages=372–422|doi=10.1144/GSL.JGS.1913.069.01-04.23|s2cid=128604856}}</ref> But then his article ended with a discussion wherein it was noted that the palaeontologist [[Charles William Andrews]] had expressed doubts as to whether ''O. latidens'' belonged in the genus ''Ornithodesmus'', as the vertebrae of the specimen that genus was based on differed markedly from those of Hooley's specimen. The American palaeontologist [[Samuel W. Williston]] subsequently reviewed Hooley's article, disagreeing with some of his conclusions about the anatomy and classification of the animal.<ref>{{cite journal|author=Wllliston, S. W.|authorlink=Samuel Wendell Williston|title=Reviews: the skeleton of ''Ornithodesmus latidens''|journal=The Journal of Geology|year=1913|volume=21|issue=8|pages=754–756|jstor=30058408|doi=10.1086/622124|bibcode=1913JG.....21..754W|doi-access=free}}</ref> After Hooley's [[monograph]], little was written about the animal for the rest of the 20th century, and no similar pterosaurs were found for decades.{{Sfn|Witton|2013|pp=143–151}} |
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In 1993 however, the British palaeontologists [[Stafford C. Howse]] and [[Andrew C. Milner]] concluded that the holotype sacrum and only specimen of ''O. cluniculus'' didn't belong to a pterosaur, but instead to a [[maniraptora]]n [[theropod]] dinosaur. They also pointed out that no detailed attempts had been made to compare the sacrum of ''O. cluniculus'' with those of pterosaurs, and that ''O. latidens'' had in effect been treated as the type species of the genus ''Ornithodesmus''. Now as a definite species of pterosaur, "''O.''" ''latidens'' thus required a new genus name.<ref name="Ornithodesmus'">{{cite journal|author=Howse, S. C. B.|author2=Milner, A. R.|title=''Ornithodesmus'' – a maniraptoran theropod dinosaur from the Lower Cretaceous of the Isle of Wight, England|journal=Palaeontology|year=1993|volume=36|pages=425–437|url=https://www.palass.org/publications/palaeontology-journal/archive/36/2/article_pp425-437}}</ref> Howse, Milner, and David Martill in 2001, moved "''O.''" ''latidens'' to a new genus called ''[[Istiodactylus]]''. The generic name means "sail finger", which refers to the wings of large pterosaurs. After moving "''O.''" ''latidens'' into ''Istiodactylus'', they had also named a new family called [[Istiodactylidae]], with ''Istiodactylus'' as the only member.<ref name="PterosaursWight">{{cite book|author=Howse, S. C. B.|author2=Milner, A. R.|author3=Martill, D. M.|year=2001|pages=324–335|title=Dinosaurs of the Isle of Wight|place=London|publisher=The Palaeontological Association|series=Guide 10; Field Guides to Fossils|chapter=Pterosaurs|editor=Martill, D. M.|editor2=Naish, D.|editorlink2=Darren Naish|isbn=978-0-901702-72-2}}</ref> |
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=== Discoveries outside Europe === |
=== Discoveries outside Europe === |
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[[File:Ornithocheirus & Tropeognathus.jpg|thumb|Comparison between the holotypes of ''[[Tropeognathus mesembrinus]]'' and ''[[Ornithocheirus simus]]'' ]] |
[[File:Ornithocheirus & Tropeognathus.jpg|thumb|Comparison between the holotypes of ''[[Tropeognathus mesembrinus]]'' and ''[[Ornithocheirus simus]]'' ]] |
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Other important ornithocheiromorph discoveries include the |
Other important ornithocheiromorph discoveries include the anhanguerids ''[[Tropeognathus]]'' and ''[[Anhanguera (pterosaur)|Anhanguera]]'' from the [[Romualdo Formation]] in [[Brazil]].<ref name="Wellnhofer1987">{{cite journal|author=Peter Wellnhofer|year=1987|title=New crested pterosaurs from the Lower Cretaceous of Brazil|journal=Mitteilungen der Bayerischen Staatssammlung für Paläontologie und historische Geologie|volume=27|pages=175–186|url=https://www.zobodat.at/pdf/Mitt-Bayer-Staatsslg-Pal-hist-Geol_27_0175-0186.pdf}}</ref><ref name=camposkellner1885b>Campos, D. de A., and Kellner, A. W. (1985). "Um novo exemplar de ''Anhanguera blittersdorffi'' (Reptilia, Pterosauria) da formação Santana, Cretaceo Inferior do Nordeste do Brasil." In Congresso Brasileiro de Paleontologia, Rio de Janeiro, Resumos, p. 13.</ref> ''Tropeognathus'' was described with its type species, ''T. mesembrinus'' in 1987 by [[Germans|German]] paleontologist [[Peter Wellnhofer]]. The generic name is derived from [[Greek language|Greek]] τρόπις, ''tropis'', meaning "keel", and γνάθος, ''gnathos'', meaning "jaw". The [[Specific name (zoology)|specific name]] is derived from [[Koine]] ''mesembrinos'', "of the noontide", simplified as "southern", in reference to the provenance from the [[Southern hemisphere]]. The description then led to an enormous taxonomic confusion.<ref name=wellnhofferencyclo>Wellnhofer, P. (1987). ''The Illustrated Encyclopedia of Pterosaurs''. New York: Barnes and Noble Books. pp. 124. {{ISBN|0-7607-0154-7}}.</ref> In 1989, [[Brazilians|Brazilian]] paleontologist Alexander Kellner considered it an ''Anhanguera mesembrinus'',<ref>{{cite journal|author=Kellner, A.W.A.|title=A new Edentate Pterosaur of the lower Cretaceous from the Araripe Basin, Northeast Brazil|year=1989|journal=Anais da Academia Brasileira de Ciências|pages=439–446|volume=61|s2cid=89420181}}</ref> then a ''Coloborhynchus mesembrinus'' by Veldmeijer in 1998,<ref>{{cite journal|author=Veldmeijer, A.J.|title=Pterosaurs from the Lower Cretaceous of Brazil in the Stuttgart Collection|journal=Geoscience and Engineering|year=1998|volume=327|pages=1–27|url=https://repository.tudelft.nl/islandora/object/uuid%3Aa77b585e-f57a-4a3f-a9ee-11534292c11b}}</ref> and then a ''Criorhynchus mesembrinus'' in 2001 by German paleontologist [[Michael Fastnacht]].<ref name=fastnacht2001>{{cite journal | last1 = Fastnacht | first1 = M | year = 2001 | title = First record of Coloborhynchus (Pterosauria) from the Santana Formation (Lower Cretaceous) of the Chapada do Araripe of Brazil | journal = Paläontologische Zeitschrift | volume = 75 | pages = 23–36 | doi=10.1007/bf03022595| s2cid = 128410270 }}</ref> ''T. mesembrinus'' was then considered a [[junior synonym]] of ''Ornithocheirus simus'' by British paleontologist [[David Unwin]] in 2001, but he then proposed an ''Ornithocheirus mesembrinus'' in 2003.<ref name=Unwin2001>Unwin, D.M., 2001, "An overview of the pterosaur assemblage from the Cambridge Greensand (Cretaceous) of Eastern England", ''Mitteilungen aus dem Museum für Naturkunde in Berlin, Geowissenschaftliche Reihe'' '''4''': 189–221</ref><ref name=Unwin2003/> In 2013 however, Taissa Rodrigues and Alexander Kellner concluded that ''Tropeognathus'' would be valid again, and containing only ''T. mesembrinus'', the type species.<ref name="Rodrigues & Kellner 2013">{{Cite journal | last1 = Rodrigues | first1 = T. | last2 = Kellner | first2 = A. | doi = 10.3897/zookeys.308.5559 | title = Taxonomic review of the ''Ornithocheirus'' complex (Pterosauria) from the Cretaceous of England | journal = ZooKeys | pages = 1–112 | year = 2013 | issue = 308 | pmid = 23794925| pmc = 3689139| doi-access = free }}</ref> |
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The discovery of ''Anhanguera'' in 1985 made some authors assign several species of ''Ornithocheirus'' to the new genus, this is due to the similarities of ''Anhanguera'' and ''Ornithocheirus''. One of which was ''Ornithocheirus cuvieri'', formerly known as ''Pterodactylus cuvieri'', was renamed by David Unwin in 2001 as ''Anhanguera cuvieri'', though in 2013 it has been considered a synonym of ''Cimoliopterus''.<ref name=Unwin2001/><ref name="Rodrigues & Kellner 2013"/> Other species of ''Anhanguera'' include ''A. piscator'', which is known from a nearly-complete skeleton, and was once proposed to belong to the genus ''Coloborhynchus'', but it has recently been placed back into ''Anhanguera'' by Andres and Myers (2013).<ref>Veldmeijer, A. J. (2003). Preliminary description of a skull and wing of a Brazilian Cretaceous (Santana Formation; Aptian-Albian) pterosaur (Pterodactyloidea) in the collection of the AMNH. ''PalArch's Journal of Vertebrate Palaeontology 0'', 1-14.</ref><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 | volume = 103 | issue = 3–4 | pages = 383–398 | year = 2013 | pmid = | pmc = }}</ref> Another species which is also known from a complete skeleton is ''A. spielbergi'', which was also originally considered as a species of ''Coloborhynchus'' in 2003, though in 2018, it has been reassigned to the genus ''[[Maaradactylus]]'', specifically to its own species, ''M. spielbergi''.<ref name="taxonomy">{{cite journal|last1=Pinheiro|first1=F.L.|last2=Rodrigues|first2=Taissa|date=2017|title=''Anhanguera'' taxonomy revisited: is our understanding of Santana Group pterosaur diversity biased by poor biological and stratigraphic control?|url=|journal=PeerJ|volume=5|page=e3285|doi=10.7717/peerj.3285|pmc=5420195|pmid=28484676}}</ref><ref>{{Cite journal|last1=Jacobs|first1=Megan L.|last2=Martill|first2=David M.|last3=Ibrahim|first3=Nizar|author-link3=Nizar Ibrahim|last4=Longrich|first4=Nick|year=2019|title=A new species of ''Coloborhynchus'' (Pterosauria, Ornithocheiridae) from the mid-Cretaceous of North Africa|url=https://sites.udmercy.edu/campusconnection/wp-content/uploads/sites/106/2018/10/New-species-of-Coloborhynchus-from-Africa.pdf|journal=Cretaceous Research|language=en|volume=95|pages=77–88|doi=10.1016/j.cretres.2018.10.018|issn=0195-6671|via=}}</ref> |
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[[File:Hamipterus-Paleozoological Museum of China.jpg|thumb |
[[File:Hamipterus-Paleozoological Museum of China.jpg|thumb|Skull of ''[[Hamipterus tianshanensis]]'']] |
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A discovery in Asia, specifically northwestern [[China]], was reported in 2006. The lake sediments allowed an exceptional preservation of fossils, and therefore paleontologists Qiu Zhanxiang and Wang Banyue started official excavations. Part of the findings consisted of dense concentrations of pterosaur bones, associated with soft tissues and eggs. |
A discovery in Asia, specifically northwestern [[China]], was reported in 2006. The lake sediments allowed an exceptional preservation of fossils, and therefore paleontologists Qiu Zhanxiang and Wang Banyue started official excavations. Part of the findings consisted of dense concentrations of pterosaur bones, associated with soft tissues and eggs. In 2014, a new species was named and described: ''[[Hamipterus tianshanensis]]''. It was named by Wang Xiaolin, Alexander Kellner, Jiang Shunxing, Wang Qiang, Ma Yingxia, Yahefujiang Paidoula, Cheng Xin, Taissa Rodrigues, Meng Xi, Zhang Jialiang, Li Ning, and Zhou Zhonghe. The generic name ''Hamipterus'' combines that of the [[Hami]] region, with the word ''pteron'', meaning "wing", and the specific name refers to the provenance from the [[Tian Shan]], a mountain range.<ref name="Andres2014">{{cite journal |author1=Xiaolin Wang |author2=Alexander W.A. Kellner |author3=Shunxing Jiang |author4=Qiang Wang |author5=Yingxia Ma |author6=Yahefujiang Paidoula |author7=Xin Cheng |author8=Taissa Rodrigues |author9=Xi Meng |author10=Jialiang Zhang |author11=Ning Li |author12=Zhonghe Zhou |year=2014 |title=Sexually dimorphic tridimensionally preserved pterosaurs and their eggs from China |journal=[[Current Biology]] |volume=24 |issue=12 |pages=1323–1330 |doi=10.1016/j.cub.2014.04.054 |pmid=24909325|doi-access=free }}</ref> |
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== Description == |
== Description == |
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=== Size === |
=== Size === |
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[[File:Istiodactylus size.jpg|thumb|Size of ''[[Istiodactylus latidens]]'' compared to a human]] |
[[File:Istiodactylus size.jpg|thumb|left|Size of ''[[Istiodactylus latidens]]'' compared to a human]] |
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Ornithocheiromorphs were large pterosaurs, with wingspans normally ranging between {{convert|3| |
Ornithocheiromorphs were large pterosaurs, with wingspans normally ranging between {{convert|3|and|6|m|ft|sp=us}}.<ref name=witton2010/> ''Istiodactylus'' for example, had a [[wingspan]] ranging from {{convert|4.3|to|5|m|ft|sp=us}}, with the most complete known skull estimated to have been about {{convert|45|cm|ft|sp=us}} in length, based on a long-lost fragment of its jaw reported in 2012.<ref name=witton2012>{{cite journal|author=Witton, M. P.|author-link=Mark P. Witton|title=New Insights into the Skull of ''Istiodactylus latidens'' (Ornithocheiroidea, Pterodactyloidea)|journal=PLOS ONE|year=2012|volume=7|issue=3|pages=e33170|doi=10.1371/journal.pone.0033170|pmid=22470442|pmc=3310040|bibcode=2012PLoSO...733170W|doi-access=free}}</ref> Though its jaws measured only {{convert|28.5|cm|in|sp=us}}, which was less than 80 percent of the skull's length.{{Sfn|Witton|2013|pp=143–151}} Anhanguerids and were typically larger than others of the group and were more successful within the [[food chain]] rather than other ornithocheiromorphs, one reason is because of their large size, for example, ''Tropeognathus mesembrinus'', had a normal wingspan of about {{convert|8.26|m|ft|sp=us}}, and {{convert|8.70|m|ft|sp=us}} as the maximum estimate.<ref name=kellneretal2013>{{Cite journal | last1 = Kellner | first1 = A. W. A. | last2 = Campos | first2 = D. A. | last3 = Sayão | first3 = J. M. | last4 = Saraiva | first4 = A. N. A. F. | last5 = Rodrigues | first5 = T. | last6 = Oliveira | first6 = G. | last7 = Cruz | first7 = L. A. | last8 = Costa | first8 = F. R. | last9 = Silva | first9 = H. P. | last10 = Ferreira | first10 = J. S. | title = The largest flying reptile from Gondwana: A new specimen of Tropeognathus cf. T. Mesembrinus Wellnhofer, 1987 (Pterodactyloidea, Anhangueridae) and other large pterosaurs from the Romualdo Formation, Lower Cretaceous, Brazil | doi = 10.1590/S0001-37652013000100009 | journal = Anais da Academia Brasileira de Ciências | volume = 85 | pages = 113–135 | year = 2013 | issue = 1 | pmid = 23538956| doi-access = free }}</ref> Another species which was impressively large is ''Coloborhynchus capito'', with a total skull length that could have been up to {{convert|75|cm|ft|sp=us}}, leading to an estimated wingspan of {{convert|7|m|ft|sp=us}}.<ref name=martill&unwin2011>Martill, D.M. and Unwin, D.M. (2011). "The world's largest toothed pterosaur, NHMUK R481, an incomplete rostrum of ''Coloborhynchus capito'' (Seeley 1870) from the Cambridge Greensand of England." ''Cretaceous Research'', (advance online publication). {{doi|10.1016/j.cretres.2011.09.003}}</ref> However, this species may belong to a different genus called ''[[Nicorhynchus]]''.<ref name="holgado2020"/> |
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=== Skull and crests === |
=== Skull and crests === |
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[[File:Anhangueria skulls.jpg|thumb |
[[File:Anhangueria skulls.jpg|thumb|Skull comparison between different [[species]] of ornithocheiromorphs; notice their different structures]] |
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Most |
Most anhanguerids bore distinctive convex "keeled" crests on their snout and underside of their [[mandible]], this was well developed in several genera such as ''Tropeognathus''.<ref name=Veldmeijer>Veldmeijer, A.J. (2006). "[http://igitur-archive.library.uu.nl/dissertations/2006-0201-200610/index.htm Toothed pterosaurs from the Santana Formation (Cretaceous; Aptian-Albian) of northeastern Brazil. A reappraisal on the basis of newly described material] {{Webarchive|url=https://web.archive.org/web/20120317052929/http://igitur-archive.library.uu.nl/dissertations/2006-0201-200610/index.htm |date=2012-03-17 }}." Tekst. - Proefschrift Universiteit Utrecht.</ref> The similar ''Anhanguera'' possessed jaws that were tapered in width, but expanded into a broad, spoon-shaped rosette at the tip. The jaws are distinguished from its relatives by several differences in the crest and teeth: unlike its close relatives ''Coloborhynchus'' and ''Ornithocheirus'', the crest on the upper jaw of ''Anhanguera'' didn't begin at the tip of the snout, therefore, it was set farther back on the skull.<ref name=kellner2000>Kellner, A.W.A. and Tomida, Y. (2000). "Description of a new species of ''Anhanguera'' (Pterodactyloidea) with comments on the pterosaur fauna from the Santana Formation (Aptian–Albian), northeastern Brazil." Tokyo, National Science Museum (''National Science Museum Monographs'', '''17''').</ref> |
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Other anhanguerids like ''[[Cearadactylus]]'' had its first preparations with many serious mistakes: the front of the snout and the lower jaws were confused leading to a reconstruction in which the anterior part of the head was upside down.<ref name="leonardi&borgomanero">Leonardi, G. & Borgomanero, G. (1985). "''Cearadactylus atrox'' nov. gen., nov. sp.: novo Pterosauria (Pterodactyloidea) da Chapada do Araripe, Ceara, Brasil." ''Resumos dos communicaçoes VIII Congresso bras. de Paleontologia e Stratigrafia'', '''27''': 75–80.</ref> Some of the teeth were extensively restored and enlarged until the wider front of the jaws showed very large and robust teeth projecting outward. With this arrangement, the maxilla was kinked, and its interlocking [[teeth]] suggested that ''Cearadactylus'' had a [[Piscivore|piscivourous]] diet, allowing the animal to keep hold of slippery fish.<ref name=ornithocheiroids/> Another smaller genus similar to ''Cearadactylus'' is ''[[Guidraco]]''. Its holotype [[skull]] has a length of {{convert|38|cm|in|sp=us}}, which makes it smaller than other genera. The skull is very elongated however, and a hollow profile is seen, but not very pointed, as the upper edge and the line of the jaw run nearly parallel over most of their length.<ref name=Guidraco>{{cite journal |author1=Xiaolin Wang |author2=Alexander W. A. Kellner |author3=Shunxing Jiang |author4=Xin Cheng |year=2012 |title=New toothed flying reptile from Asia: close similarities between early Cretaceous pterosaur faunas from China and Brazil |journal=Naturwissenschaften |volume=99 |issue= 4|pages= 249–57|doi=10.1007/s00114-012-0889-1 |pmid=22354475|bibcode=2012NW.....99..249W |s2cid=7323552 }}</ref> |
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Even though most ornithocheiromorphs didn't have a [[cranial crest]] like the closely related pteranodontids, there were some exceptions, this included ''[[Caulkicephalus]]'' and ''[[Ludodactylus]]''.<ref name=MFDB00>Frey, E., Martill, D., and Buchy, M. (2003). A new crested ornithocheirid from the Lower Cretaceous of northeastern Brazil and the unusual death of an unusual pterosaur. In: Buffetaut, E., and Mazin, J.-M. (eds.). ''Evolution and Palaeobiology of Pterosaurs''. ''Geological Society Special Publication'' '''217''':56-63. {{ISBN|1-86239-143-2}}.</ref> |
Even though most ornithocheiromorphs didn't have a [[cranial crest]] like the closely related pteranodontids, there were some exceptions, this included ''[[Caulkicephalus]]'' and ''[[Ludodactylus]]''.<ref name=MFDB00>Frey, E., Martill, D., and Buchy, M. (2003). A new crested ornithocheirid from the Lower Cretaceous of northeastern Brazil and the unusual death of an unusual pterosaur. In: Buffetaut, E., and Mazin, J.-M. (eds.). ''Evolution and Palaeobiology of Pterosaurs''. ''Geological Society Special Publication'' '''217''':56-63. {{ISBN|1-86239-143-2}}.</ref> ''Caulkicephalus'' had a rounded snout, very similar to that of ''Ornithocheirus'' and ''Anhanguera'', and therefore it is placed within either Anhangueridae or Ornithocheiridae, depending on the author.<ref name=Targaryendraco/><ref name=Caulkicephalus/> ''Caulkicephalus'' was also a large pterosaur, with wingspan estimates of around {{convert|5|m|ft|sp=us}}.<ref name=Caulkicephalus>{{cite journal|author=Steel, L., Martill, D.M., Unwin, D.M. and Winch, J. D.|year=2005|title=A new pterodactyloid pterosaur from the Wessex Formation (Lower Cretaceous) of the Isle of Wight, England.|journal=Cretaceous Research|volume=26|issue=4|pages=686–698|doi=10.1016/j.cretres.2005.03.005}}</ref> |
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=== Wings === |
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[[File:TropeognathusDB22.jpg|thumb|Life [[Paleoart|restorations]] of ''Tropeognathus'' in flight; notice their high [[Aspect ratio (aeronautics)|aspect ratio]]]] |
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Most pterosaurs of this group had extensive wing-membranes, which were distended by a long wing-finger and would have been covered in hair-like [[pycnofibres]].{{Sfn|Witton|2013|pp=51–52}} Like the modern-day [[albatross]], ornithocheiromorphs also had a high [[aspect ratio (aeronautics)|aspect ratio]] and low [[wing loading]]s, though this is typically in the larger and more advanced genera. Wing remains of the istiodactylid ''[[Nurhachius]]'' have been compared to those of modern [[soaring birds]], which fly with little flapping, and may have been ideal for low-energy soaring, which is necessary when searching for [[carrion]] or [[fish]].<ref name="Zhou2019"/> The wings of istiodactylids also seem to have been shorter compared to the fish-eating ornithocheirids, and unlike them, which were more suited for taking off and landing, istiodactylids may have preferred a more terrestrial setting. This is also seen in many inland soaring birds, which have a lower aspect ratio compared those that soar over the ocean. Huge genera such as ''Tropeognathus'' and ''Ornithocheirus'' are considered to have flown similar to modern-day albatrosses, which consisted on travelling very long distances and rarely flapping. This technique is mostly seen in later pterosaur locomotion, such as the related and mentioned pteranodontids, though not so common in the larger azhdarchids such as ''[[Hatzegopteryx]]'' and ''Quetzalcoatlus''.<ref name=martin2017/><ref name=witton2010/> |
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===Shoulder girdle=== |
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Ornithocheiromorphs had [[shoulder girdle]]s of strong structure, which transferred the forces of a rarely flapping flight to the [[thorax]]. |
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The shoulder girdle was probably covered by thick muscle layers, with the upper bone ([[shoulder blade]]) resulting in some kind of straight bar.{{sfn|Witton|2013|p=44}} The shoulder blade was connected to a lower bone called the [[coracoid]], which is relatively long in most ornithocheiromorphs. In the more advanced species, the shoulder is combined whole, with the scapulocoracoid, being almost vertically oriented, meaning that it fitted into a recess in the side of the notarium, while the coracoid likewise connected to the breastbone. Some species also had the shoulder joint moved from the shoulder blade to the coracoid. The breastbone is somewhat wide, and was formed by fused paired ''sterna''. At its rear, a row of belly ribs or [[gastralia]] was present, covering the entire belly.{{sfn|Wellnhofer|1991|p=52}} To the front of the breastbone, a long point called the ''cristospina'', jutted obliquely upwards, and the rear edge of the breastbone was the deepest point of the thorax. The absence of clavicles or interclavicles had made paleontologists wonder if pterosaurs evolved in different ways than other archosaurs.{{sfn|Witton|2013|p=32}}{{sfn|Wellnhofer|1991|p=52}} |
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=== Vertebrae === |
=== Vertebrae === |
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[[File:Istiodactylus bones.jpg|200px|thumb|upright|Illustrations of various ''Istiodactylus'' bones, including the [[neck vertebra]]e and [[notarium]]]] |
[[File:Istiodactylus bones.jpg|200px|thumb|upright|Illustrations of various ''Istiodactylus'' bones, including the [[neck vertebra]]e and [[notarium]]]] |
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The vertebral column of ornithocheiromorphs was heavily pneumatized by an extensive system of air sacs, leaving prominent pneumatic foraminae.<ref name="Buchmann2019">{{cite journal|last1=Buchmann|first1=R.|last2=dos Santos Avilla|first2=L.|last3=Rodrigues|first3=T.|date=October 25, 2019|title=Comparative analysis of the vertebral pneumatization in pterosaurs (Reptilia: Pterosauria) and extant birds (Avialae: Neornithes)|journal=[[PLoS One]] |volume=14|issue=10|pages=e0224165|doi=10.1371/journal.pone.0224165|pmid=31652295|s2cid=204909043}}</ref> The neck of ornithocheiromorphs was typically relatively long and robust, being longer than the torso in some derived clades.{{sfn|Wellnhofer|1991|p=54}} The neural spines of ornithocheiromorph cervical vertebrae were generally tall and spikelike.<ref name=Guidraco/><ref name="kellner2000"/> In some genera such as ''Tropeognathus'' and ''Istiodactylus'', up to six dorsal vertebrae are fused into a [[notarium]].{{Sfn|Witton|2013|pp=158}} In some genera such as ''Anhanguera'', four to seven sacral vertebrae are fused into a [[synsacrum]].<ref name="kellner2000"/> The tail is not well |
The vertebral column of ornithocheiromorphs was heavily pneumatized by an extensive system of air sacs, leaving prominent pneumatic foraminae.<ref name="Buchmann2019">{{cite journal|last1=Buchmann|first1=R.|last2=dos Santos Avilla|first2=L.|last3=Rodrigues|first3=T.|date=October 25, 2019|title=Comparative analysis of the vertebral pneumatization in pterosaurs (Reptilia: Pterosauria) and extant birds (Avialae: Neornithes)|journal=[[PLoS One]] |volume=14|issue=10|pages=e0224165|doi=10.1371/journal.pone.0224165|pmid=31652295|pmc=6814219|bibcode=2019PLoSO..1424165B |s2cid=204909043|doi-access=free}}</ref> The neck of ornithocheiromorphs was typically relatively long and robust, being longer than the torso in some derived clades.{{sfn|Wellnhofer|1991|p=54}} The neural spines of ornithocheiromorph cervical vertebrae were generally tall and spikelike.<ref name=Guidraco/><ref name="kellner2000"/> In some genera such as ''Tropeognathus'' and ''Istiodactylus'', up to six dorsal vertebrae are fused into a [[notarium]].{{Sfn|Witton|2013|pp=158}} In some genera such as ''Anhanguera'', four to seven sacral vertebrae are fused into a [[synsacrum]].<ref name="kellner2000"/> The tail is not well known in ornithocheiromorphs, however. ''Zhenyuanopterus'', which is known for having 13 caudal vertebrae, formed one of the longest tails of any pterodactyloid.<ref name=JL10>{{cite journal |last=Lü |first=J. |year=2010 |title=A new boreopterid pterodactyloid pterosaur from the Early Cretaceous Yixian Formation of Liaoning Province, northeastern China |journal=Acta Geologica Sinica |volume=24 |issue=2 |pages=241–246 |doi=10.1111/j.1755-6724.2010.00204.x|s2cid=140600398 }}</ref> ''Anhanguera'', another well-known genus, had a shorter tail, with broad caudal vertebrae that bore a "duplex" cross-section similar to ''Pteranodon''.<ref name="kellner2000"/> |
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=== Pelvic structure === |
=== Pelvic structure === |
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[[File:Anhanguera-santanae sacrum.jpg|thumb|left|A rotated [[pelvis]] of ''[[Anhanguera santanae]]'', showing its right side]] |
[[File:Anhanguera-santanae sacrum.jpg|thumb|left|A rotated [[pelvis]] of ''[[Anhanguera santanae]]'', showing its right side]] |
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The [[pelvis]] of ornithocheiromorphs was of moderate size compared to the body as a whole, similar to other [[ornithocheiroid]]s. The three [[pelvic bones]] were often fused, as seen in many species such as ''[[Anhanguera santanae]]'', the [[Ilium (bone)|ilium]] was long and low, and its front and rear blades projected horizontally beyond the edges of the lower pelvic bones.{{sfn|Wellnhofer|1991|p=55}} Despite the structure length, the processes of these rod-like forms indicate that the hindlimb muscles attached to them were limited in strength.{{sfn|Witton|2013|p=46}} The [[pubic bone]] was fused with the broad ischium into an ischiopubic blade, resulting in a narrow build. Sometimes, the blades of both sides were fused, closing the pelvis from below and forming the pelvic canal. The front of the pubic bones was also articulated with a unique structure, resulting in a pair of prepubic bones within. This formed a cusp covering the rear belly, and was located between the pelvis and the belly ribs. The [[hip joint]] of ornithocheiromorphs was not perforated and allowed considerable mobility to the leg, and suggests that it was vertical, as therefore had a function in breathing, compensating the relative rigidity of the chest cavity.{{sfn|Witton|2013|p=35}}{{sfn|Wellnhofer|1991|p=55}} |
The [[pelvis]] of ornithocheiromorphs was of moderate size compared to the body as a whole, similar to other [[ornithocheiroid]]s. The three [[pelvic bones]] were often fused, as seen in many species such as ''[[Anhanguera santanae]]'', the [[Ilium (bone)|ilium]] was long and low, and its front and rear blades projected horizontally beyond the edges of the lower pelvic bones.{{sfn|Wellnhofer|1991|p=55}} Despite the structure length, the processes of these rod-like forms indicate that the hindlimb muscles attached to them were limited in strength.{{sfn|Witton|2013|p=46}} The [[pubic bone]] was fused with the broad ischium into an ischiopubic blade, resulting in a narrow build. Sometimes, the blades of both sides were fused, closing the pelvis from below and forming the pelvic canal. The front of the pubic bones was also articulated with a unique structure, resulting in a pair of prepubic bones within. This formed a cusp covering the rear belly, and was located between the pelvis and the belly ribs. The [[hip joint]] of ornithocheiromorphs was not perforated and allowed considerable mobility to the leg, and suggests that it was vertical, as therefore had a function in breathing, compensating the relative rigidity of the chest cavity.{{sfn|Witton|2013|p=35}}{{sfn|Wellnhofer|1991|p=55}} |
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===Hindlimbs=== |
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The hindlimbs of ornithocheiromorphs were strongly built, yet relative to their wingspans, smaller than those of birds in terms of hindlimb-to-wingspan ratio. The hindlimbs were long in comparison to the torso length, and the thighbone was rather straight, with the head making only a small angle with the shaft.{{sfn|Wellnhofer|1991|p=56}}{{sfn|Witton|2013|p=35}} This implied the legs not being held vertically below the body, but were somewhat sprawling within.{{sfn|Wellnhofer|1991|p=56}} A tibiotarsus is mostly seen in many ornithocheiromorphs, this consisted on the shinbone ([[tibia]]) being fused with the upper ankle bones, resulting in a longer structure than the thighbone, which could attain a vertical position when walking.<ref name=padian1983/> Most of the toes of ornithocheiromorphs were long, though compared to earlier forms of pterosaurs, they were less robust, and were also connected to the ankle in a higher position than the other [[metatarsal]]s. Often curved, the mobile clawless fifth toe consisted of two phalanges, this is seen some specimens, which showed membranes between the toes, and was concluded that this allowed them to function as flight control surfaces.<ref name=wittonhabib2010/> |
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== Classification == |
== Classification == |
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[[File:Targaryendraco.jpg|thumb|left|upright|Holotype [[mandible]] of the |
[[File:Targaryendraco.jpg|thumb|left|upright|Holotype [[mandible]] of the recently named ''[[Targaryendraco]]'']] |
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Several studies show that ornithocheiromorphs were less derived than the toothless pteranodontids such as ''Pteranodon'', and based on the different evolutionary changes, they therefore need to be grouped in a different [[clade]] than ''Pteranodon'', though still within [[Pteranodontoidea]].<ref>Bennett, S. C. (1994). "Taxonomy and systematics of the Late Cretaceous pterosaur ''Pteranodon'' (Pterosauria, Pterodactyloidea)", ''Occasional Papers of the Museum of Natural History'', University of Kansas, Lawrence, '''169''': 1-70</ref |
Several studies show that ornithocheiromorphs were less derived than the toothless pteranodontids such as ''Pteranodon'', and based on the different evolutionary changes, they therefore need to be grouped in a different [[clade]] than ''Pteranodon'', though still within [[Pteranodontoidea]].<ref>Bennett, S. C. (1994). "Taxonomy and systematics of the Late Cretaceous pterosaur ''Pteranodon'' (Pterosauria, Pterodactyloidea)", ''Occasional Papers of the Museum of Natural History'', University of Kansas, Lawrence, '''169''': 1-70</ref> In 2003, David Unwin considered the family Istiodactylidae to group with the toothless Pteranodontidae, within the group [[Ornithocheiroidea]],<ref name=Unwin2003>{{cite journal|author=Unwin, D. M.|title=On the phylogeny and evolutionary history of pterosaurs|journal=Geological Society, London, Special Publications|date=2003|volume=217|issue=1|pages=139–190|doi=10.1144/GSL.SP.2003.217.01.11|bibcode=2003GSLSP.217..139U|citeseerx=10.1.1.924.5957|s2cid=86710955}}</ref> but Alexander Kellner however, grouped it with the toothed Anhangueridae instead, resulting in a more understandable change of evolution between the two toothed families.<ref name=Phylogeny>{{cite journal|author=Kellner, A. W. A.|author-link=Alexander Kellner|title=Pterosaur phylogeny and comments on the evolutionary history of the group|journal=Geological Society, London, Special Publications|year=2003|volume=217|issue=1|pages=105–137|doi=10.1144/GSL.SP.2003.217.01.10|bibcode=2003GSLSP.217..105K|s2cid=128892642}}</ref> |
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Brian Andres and colleagues found the families Istiodactylidae, Ornithocheiridae and Anhangueridae to form a group in 2014, which he called Lanceodontia, and consists of the more advanced ornithocheiromorphs. The clade however, excludes the more poorly known family Lonchodectidae, even though members of the family had previously been seen as some of the most derived forms of toothed pterosaurs. In their analysis, they also included the family Boreopteridae within the clade [[Anhangueria]], though placed in a more basal position, while also containing the genus ''Guidraco''.<ref name=kryptodrakon/> In 2018 however, Nicholas Longrich and colleagues found Boreopteridae outside Anhangueria as the sister taxon of the family Lonchodectidae, both groups placed as basal members of the Ornithocheiromorpha.<ref name=longrichetal2018/> |
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Studies on ''Lonchodectes'' and its family, [[Lonchodectidae]] (which only consisted of ''Lonchodectes''), has since been classified to many different clades, including [[Ctenochasmatoidea]] and [[Azhdarchoidea]]. More recent reclassifications show that Lonchodectidae was instead a closer relative of the ornithocheirids, but was given a basal placement because of uncertain fossil structures and other features seen uniquely in ''Lonchodectes''. The genus ''[[Prejanopterus]]'' was formerly considered a lonchodectid, but its placement is very uncertain, and some authors just consider it a basal [[archaeopterodactyloid]].<ref name=Averianov2020>{{cite journal|first=A.O. |last=Averianov |year=2020 |title=Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea) |journal=Proceedings of the Zoological Institute RAS |volume=324 |issue=1 |pages=41–55 |doi=10.31610/trudyzin/2020.324.1.41|doi-access=free }}</ref> Another genus, ''Lonchodraco'', is also in a dubious placement, Rodrigues & Kellner made a family called [[Lonchodraconidae]] containing only ''Lonchodraco'', but it was then reclassified to Lonchodectidae along with ''Lonchodectes'', and therefore synonymizing the two families.<ref name="Rodrigues & Kellner 2013"/> |
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=== Relationships === |
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Brian Andres and colleagues found the families Istiodactylidae, Ornithocheiridae and Anhangueridae to form a group in 2014, which he called Lanceodontia, and consists of the more advanced ornithocheiromorphs. The clade however, excludes the more poorly known family Lonchodectidae, even though members of the family had previously been seen as some of the most derived forms of toothed pterosaurs. In their analysis, they also included the family Boreopteridae within the clade Anhangueria, though placed in a more basal position, while also containing the genus ''Guidraco''.<ref name=kryptodrakon/> In 2018 however, Nicholas Longrich and colleagues found Boreopteridae outside Anhangueria as the sister taxon of the family Lonchodectidae, both groups placed as basal members of the Ornithocheiromorpha.<ref name=longrichetal2018/> |
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There are competing theories of ornithocheiromorph [[phylogeny]] (evolutionary relationships). Below is [[cladogram]] following a topology recovered by Brian Andres, using the most recent iteration of his data set (Andres, 2021).<ref name="andres2021">Andres, B. (2021) Phylogenetic systematics of ''Quetzalcoatlus'' Lawson 1975 (Pterodactyloidea: Azhdarchoidea). ''Journal of Vertebrate Paleontology'', '''41''':sup1, 203-217. DOI: 10.1080/02724634.2020.1801703 https://www.tandfonline.com/doi/full/10.1080/02724634.2020.1801703</ref> |
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{{clade| style=font-size:90%;line-height:85% |
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|label1= [[Pteranodontoidea]] |
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|1={{clade |
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|1=[[Pteranodontia]][[File:Pteranodon longiceps mmartyniuk wiki.png|50 px]] |
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|label2='''Ornithocheiromorpha''' |
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|2={{clade |
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|1={{clade |
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|1=''[[Serradraco]]'' |
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|2=''[[Aussiedraco]]'' }} |
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|2={{clade |
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|label1= |
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|1={{clade |
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|1=''[[Hongshanopterus]]'' |
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|2={{clade |
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|1=''[[Targaryendraco]]'' |
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|2=''[[Lonchodectes]]''[[File:Lonchodectes-concepts.png|50 px]] }} }} |
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|2={{clade |
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|1={{clade |
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|1=''[[Ikrandraco]]'' |
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|2=''[[Lonchodraco]]'' }} |
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|label2='''Lanceodontia''' |
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|2={{clade |
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|1=[[Istiodactylidae]] |
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|label2='''Ornithocheiriformes''' |
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|2={{clade |
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|1=[[Boreopteridae]] |
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|2={{clade |
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|1=''[[Hamipterus]]'' |
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|2={{clade |
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|1={{clade |
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|1=''[[Brasileodactylus]]'' |
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|2=''[[Barbosania]]'' |
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|3=''[[Cearadactylus]]'' }} |
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|2={{clade |
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|label1=[[Ornithocheirae]] |
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|1={{clade |
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|1=[[Ornithocheiridae]] |
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|2=[[Anhangueridae]][[File:LiaoningopterusDB flipped.jpg|30 px]] }} |
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|2={{clade |
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|1=''[[Ludodactylus]]''[[File:Ludodactylus.jpg|40 px]] |
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|2={{clade |
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|1=''[[Guidraco]]'' |
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|2={{clade |
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|1=''[[Cimoliopterus]]''[[File:Cimoliopterus dunni.jpg|40 px]] |
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|2={{clade |
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|1=''[[Camposipterus]]'' |
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|2=''[[Aetodactylus]]'' |
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}} }} }} }} }} }} }} }} }} }} }} }} }} }} |
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Below is a [[cladogram]] showing the results of a [[phylogenetic]] analysis presented by Longrich ''et al.'' (2018). In the analysis, they placed the genus ''[[Hongshanopterus]]'' as a basal member.<ref name=longrichetal2018>Longrich, N.R., Martill, D.M., and Andres, B. (2018). [http://journals.plos.org/plosbiology/article?id=10.1371/journal.pbio.2001663 "Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary."] ''PLoS Biology'', '''16'''(3): e2001663. {{doi|10.1371/journal.pbio.2001663}}</ref> |
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=== Phylogeny === |
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Below is a [[cladogram]] showing the results of a [[phylogenetic]] analysis presented by Longrich ''et al.'' (2018). In the analysis, they placed the genus ''[[Hongshanopterus]]'' as a basal member, and also found ''Guidraco'' as the sister taxon of the more derived ''Ludodactylus'', meaning that ''Guidraco'' is placed in a more derived position, which contradicts the earlier analysis by Andres ''et al.'' in 2014. Longrich ''et al.'' also included the ornithocheirid ''[[Siroccopteryx]]'' in their analysis, more precisely as the sister taxon of ''Coloborhynchus''.<ref name=longrichetal2018>Longrich, N.R., Martill, D.M., and Andres, B. (2018). [http://journals.plos.org/plosbiology/article?id=10.1371/journal.pbio.2001663 "Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary."] ''PLoS Biology'', '''16'''(3): e2001663. {{doi|10.1371/journal.pbio.2001663}}</ref> |
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[[File:Ludodactylus.jpg|thumb|Life restoration of a ''[[Ludodactylus]]'' in flight]] |
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{{clade| style=font-size:90%;line-height:85% |
{{clade| style=font-size:90%;line-height:85% |
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|label1='''Ornithocheiromorpha''' |
|label1='''Ornithocheiromorpha''' |
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Line 126: | Line 149: | ||
|2=''[[Lonchodectes]]'' |
|2=''[[Lonchodectes]]'' |
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}} |
}} |
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|2=[[Boreopteridae]] |
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|2={{clade |
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|1=''[[Boreopterus]]'' |
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|2=''[[Zhenyuanopterus]]'' |
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}} |
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}} |
}} |
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|label2= '''Lanceodontia''' |
|label2= '''Lanceodontia''' |
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|2={{clade |
|2={{clade |
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| |
|1=[[Istiodactylidae]] |
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|label2='''Ornithocheiriformes''' |
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|1={{clade |
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|1=''[[Nurhachius]]'' |
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|2={{clade |
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|1=''[[Liaoxipterus]]'' |
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|2=''[[Istiodactylus]]'' |
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}} }} |
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|2={{clade |
|2={{clade |
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|1={{clade |
|1={{clade |
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|1=''[[Aetodactylus]]'' |
|1=''[[Aetodactylus]]'' |
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|2=''[[Cimoliopterus]]'' |
|2=''[[Cimoliopterus]]''[[File:Cimoliopterus dunni.jpg|40 px]] |
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}} |
}} |
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|label2= |
|label2= [[Anhangueria]] |
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|2={{clade |
|2={{clade |
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|1={{clade |
|1={{clade |
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|1=''[[Guidraco]]'' |
|1=''[[Guidraco]]'' |
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|2=''[[Ludodactylus]]'' |
|2=''[[Ludodactylus]]''[[File:Ludodactylus.jpg|40 px]] |
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}} |
}} |
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|2={{clade |
|2={{clade |
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Line 159: | Line 173: | ||
|label2=[[Ornithocheirae]] |
|label2=[[Ornithocheirae]] |
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|2={{clade |
|2={{clade |
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| |
|2=[[Anhangueridae]][[File:LiaoningopterusDB flipped.jpg|30 px]] |
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|1=[[Ornithocheiridae]] |
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|1={{clade |
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}} }} }} }} }} }} }} }} }} |
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|1=''[[Anhanguera (pterosaur)|Anhanguera]]'' |
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|2=''[[Liaoningopterus]]'' |
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}} |
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|label2=[[Ornithocheiridae]] |
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|2={{clade |
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|1=''[[Tropeognathus]]'' |
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|2={{clade |
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|1=''[[Ornithocheirus]]'' |
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|2={{clade |
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|1=''[[Siroccopteryx]]'' |
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|2=''[[Coloborhynchus]]'' |
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}} }} }} }} }} }} }} }} }} }} }} }} |
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In 2019, several new species of ornithocheiromorphs were found, and the former species ''Ornithocheirus wiedenrothi'' was renamed as ''[[Targaryendraco]] wiedenrothi''.<ref name=Targaryendraco>Rodrigo V. Pêgas, Borja Holgado & Maria Eduarda C. Leal (2019) On ''Targaryendraco wiedenrothi'' gen. nov. (Pterodactyloidea, Pteranodontoidea, Lanceodontia) and recognition of a new cosmopolitan lineage of Cretaceous toothed pterodactyloids, Historical Biology, {{doi|10.1080/08912963.2019.1690482}} |
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[[File:Holotypes of Ferrodraco, Mythunga and Aussiedraco.jpg|thumb|Holotype skull and mandible of the ornithocheirids ''[[Ferrodraco]]'' (A) and ''[[Mythunga]]'' (B), and holotype mandible of the targaryendraconian ''[[Aussiedraco]]'' (C)]] |
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</ref> The description of ''[[Iberodactylus]]'' in [[Spain]] also made some paleontologists reclassify the genus ''Hamipterus'' in a newly named family called [[Hamipteridae]].<ref name="Holgado2019"/> The ornithocheirid ''Cimoliopterus'' was also reclassified as well, and it is currently grouped with ''[[Aetodactylus]]'' and ''[[Camposipterus]]'' in the clade [[Targaryendraconia]], specifically to its own family, the [[Cimoliopteridae]].<ref name=Targaryendraco/> However, an analysis by Jacobs ''et al.'' (2019) recovers both ''Camposipterus'' and ''Cimoliopterus'' within the Ornithocheiridae again, using a new data matrix not including ''Targaryendraco.''<ref name=J2019>{{cite journal|author=Jacobs, M.L. |author2=Martill, D.M. |author3=Ibrahim, N. |author4=Longrich, N. |year=2019|title=A new species of ''Coloborhynchus'' (Pterosauria, Ornithocheiridae) from the mid-Cretaceous of North Africa|journal=Cretaceous Research|volume=95|pages=77–88|doi=10.1016/j.cretres.2018.10.018|s2cid=134439172 |url=https://sites.udmercy.edu/campusconnection/wp-content/uploads/sites/106/2018/10/New-species-of-Coloborhynchus-from-Africa.pdf}}</ref> The previously recovered basal [[eupterodactyloid]] ''[[Haopterus]]'' was reclassified due to the description of ''[[Mimodactylus]]'', and is placed in a new family called [[Mimodactylidae]].<ref name="Kellner2019">Kellner, Alexander W. A.; Caldwell, Michael W.; Holgado, Borja; Vecchia, Fabio M. Dalla; Nohra, Roy; Sayão, Juliana M.; Currie, Philip J. (2019). "First complete pterosaur from the Afro-Arabian continent: insight into pterodactyloid diversity". ''Scientific Reports''. '''9'''(1). {{doi|10.1038/s41598-019-54042-z}}.</ref> However, a more recent analysis using the data in the description of ''Mimodactylus'' has found ''Haopterus'' as a basal member of the more inclusive group Istiodactyliformes.<ref name=Luchibang>{{cite journal |author1=David W. E. Hone |author2=Adam J. Fitch |author3=Feimin Ma |author4=Xing Xu |year=2020 |title=An unusual new genus of istiodactylid pterosaur from China based on a near complete specimen |journal=Palaeontologia Electronica |volume=23 |issue=1 |pages=Article number 23(1):a09 |doi=10.26879/1015 |doi-access=free }}</ref> Many recent analyses have also recovered several ornithocheirids, including ''Tropeognathus'', ''Coloborhynchus'', and ''Caulkicephalus'' within the family Anhangueridae, meaning that they were more closely related to ''Anhanguera'' than to ''Ornithocheirus''.<ref name="Holgado2019">Borja Holgado, Rodrigo V. Pêgas, José Ignacio Canudo, Josep Fortuny, Taissa Rodrigues, Julio Company & Alexander W.A. Kellner, 2019, "On a new crested pterodactyloid from the Early Cretaceous of the Iberian Peninsula and the radiation of the clade Anhangueria", ''Scientific Reports'' '''9''': 4940. {{doi|10.1038/s41598-019-41280-4}}</ref><ref name=Targaryendraco/><ref name=Luchibang/><ref name="Kellner2019"/><ref name="holgado2020">{{cite journal |last1=Holgado |first1=B. |last2=Pêgas |first2=R.V. |year=2020 |title=A taxonomic and phylogenetic review of the anhanguerid pterosaur group Coloborhynchinae and the new clade Tropeognathinae |journal=Acta Palaeontologica Polonica |volume=65 |doi=10.4202/app.00751.2020|doi-access=free }}</ref> |
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In 2019, new species of ornithocheiromorphs were found, and the former species ''Ornithocheirus wiedenrothi'' was renamed as ''[[Targaryendraco]] wiedenrothi''.<ref name=Targaryendraco>Rodrigo V. Pêgas, Borja Holgado & Maria Eduarda C. Leal (2019) On ''Targaryendraco wiedenrothi'' gen. nov. (Pterodactyloidea, Pteranodontoidea, Lanceodontia) and recognition of a new cosmopolitan lineage of Cretaceous toothed pterodactyloids, Historical Biology, {{DOI|10.1080/08912963.2019.1690482}} |
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</ref> The discovery of ''[[Iberodactylus]]'' in [[Spain]] also made paleontologists reclassify the genus ''Hamipterus'' in a newly named family called [[Hamipteridae]].<ref name="Holgado2019"/> The ornithocheirid ''Cimoliopterus'' was also reclassified as well, and it is currently grouped with ''[[Aetodactylus]]''<ref name=TSM10>{{cite journal |last=Myers |first=Timothy S. |year=2010 |title=A new ornithocheirid pterosaur from the Upper Cretaceous (Cenomanian–Turonian) Eagle Ford Group of Texas |journal=Journal of Vertebrate Paleontology |volume=30 |issue=1 |pages=280–287 |doi=10.1080/02724630903413099 |s2cid=130367018 |url=http://pdfserve.informaworld.com/929241__918962831.pdf }}</ref> and ''[[Camposipterus]]'' in the clade [[Targaryendraconia]], specifically to its own family, the [[Cimoliopteridae]].<ref name=Targaryendraco/> However, an analysis by Jacobs ''et al.'' (2019) recovers both ''Camposipterus'' and ''Cimoliopterus'' within the Ornithocheiridae again.<ref name=J2019>{{cite journal|authors=Jacobs, M.L., Martill, D.M., Ibrahim, N., Longrich, N.|year=2019|title=A new species of ''Coloborhynchus'' (Pterosauria, Ornithocheiridae) from the mid-Cretaceous of North Africa|journal=Cretaceous Research|volume=95|pages=77–88|doi=10.1016/j.cretres.2018.10.018|url=https://sites.udmercy.edu/campusconnection/wp-content/uploads/sites/106/2018/10/New-species-of-Coloborhynchus-from-Africa.pdf}}</ref> The basal [[eupterodactyloid]] ''[[Haopterus]]'' was reclassified due to the discovery of ''[[Mimodactylus]]'', and it's placed as the sister taxon of it in a new family called [[Mimodactylidae]].<ref name="Kellner2019">Kellner, Alexander W. A.; Caldwell, Michael W.; Holgado, Borja; Vecchia, Fabio M. Dalla; Nohra, Roy; Sayão, Juliana M.; Currie, Philip J. (2019). "First complete pterosaur from the Afro-Arabian continent: insight into pterodactyloid diversity". ''Scientific Reports''. '''9'''(1). {{doi|10.1038/s41598-019-54042-z}}.</ref> However, a more recent analysis had found ''Haopterus'' as a basal member of the more inclusive group Istiodactyliformes.<ref>{{cite journal |author1=David W. E. Hone |author2=Adam J. Fitch |author3=Feimin Ma |author4=Xing Xu |year=2020 |title=An unusual new genus of istiodactylid pterosaur from China based on a near complete specimen |journal=Palaeontologia Electronica |volume=23 |issue=1 |pages=Article number 23(1):a09 |doi=10.26879/1015 |doi-access=free }}</ref> |
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The cladogram below is a topology recovered by Pentland ''et al.'' in 2019. In the analysis, they found a slightly different set than the previous analysis by Longrich ''et al.'' (2018); they recovered Boreopteridae (identified as Boreopterinae in the analysis) within the Lanceodontia as sister taxon to the Anhangueria. Pentland ''et al.'' also included the genera ''[[Ferrodraco]]'' and ''[[Mythunga]]'' in their analysis, where they placed them within the subfamily [[Ornithocheirinae]] as sister taxa.<ref name="Pentland2019"/> |
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{{clade| style=font-size:90%;line-height:85% |
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|label1='''Ornithocheiromorpha''' |
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|1={{clade |
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|1=''[[Hongshanopterus]]'' |
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|2={{clade |
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|1=''[[Lonchodectes]]'' |
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|label2= '''Lanceodontia''' |
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|2={{clade |
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|label1=[[Istiodactylidae]] |
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|1={{clade |
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|1=''[[Nurhachius]]'' |
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|2={{clade |
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|1=''[[Longchengpterus]]'' |
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|2={{clade |
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|1=''[[Liaoxipterus]]'' |
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|2=''[[Istiodactylus]]'' |
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}} }} }} |
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|2={{clade |
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|label1=[[Boreopterinae]] |
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|1={{clade |
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|1=''[[Boreopterus]]'' |
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|2=''[[Zhenyuanopterus]]'' |
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}} |
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|label2= '''Anhangueria''' |
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|2={{clade |
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|1=''[[Guidraco]]'' |
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|2={{clade |
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|1={{clade |
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|1={{clade |
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|1=''[[Brasileodactylus]]'' |
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|2=''[[Ludodactylus]]'' |
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}} |
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|2={{clade |
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|1=''[[Cearadactylus]]'' |
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|label2=[[Ornithocheirae]] |
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|2={{clade |
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|label1=[[Anhangueridae]] |
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|1={{clade |
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|1=''[[Liaoningopterus]]'' |
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|2=''[[Anhanguera (pterosaur)|Anhanguera]]'' |
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}} |
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|label2=[[Ornithocheiridae]] |
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|2={{clade |
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|1=''[[Tropeognathus]]'' |
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|label2=[[Ornithocheirinae]] |
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|2={{clade |
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|1=''[[Coloborhynchus]]'' |
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|2={{clade |
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|1=''[[Ornithocheirus]]'' |
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|2={{clade |
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|1=''[[Ferrodraco]]'' |
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|2=''[[Mythunga]]'' |
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}} }} }} }} }} }} }} }} }} }} }} }} }} }} |
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== Paleobiology == |
== Paleobiology == |
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=== Diet and feeding === |
=== Diet and feeding === |
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[[File:Istiodactylus scavenging.tif|thumb|upright|left|A hypothetical recreation of a group of ''Istiodactylus'' feeding on a carcass of a [[stegosaur]]]] |
[[File:Istiodactylus scavenging.tif|thumb|upright|left|A hypothetical recreation of a group of ''Istiodactylus'' feeding on a carcass of a [[stegosaur]]]] |
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Ornithocheiromorphs were originally regarded as [[piscivorous]] creatures, feeding mainly on small and mid-sized fish.<ref name=ornithocheiroids/> Some paleontologists even suggested details on how these pterosaurs caught fish, some of which included dipping their [[beak]]s close to the water for prey.<ref name=ornithocheiroids/> Hooley for example, found that the beak of the well known ''Istiodactylus'' was similar to those of birds such as [[heron]]s, [[stork]]s, and [[Skimmer (bird)|skimmer]]s, and suggested that ''Istiodactylus'' probably fed on fish, this was mainly based on his 1913 jaw reconstruction of the animal.<ref name=hooley1913/> In 1991, Peter Wellnhofer compared the jaw endings of ''Istiodactylus'' with those of a [[duck]], but he then noticed that it wasn't a "duck-billed pterosaur" or anything similar, even though it was popularly called that way.<ref name="Wellnhofer">{{cite book|author=Wellnhofer, P.|authorlink=Peter Wellnhofer|year=1991 |pages=114–116|title=The Illustrated Encyclopedia of Pterosaurs|publisher=Crescent Books|isbn=978-0-517-03701-0|location=New York}}</ref> Other diet suggestions for ''Istiodactylus'' were mostly based on the broad snout remains, as well as the very distinctive teeth; leading to the thought of a more terrestrial life and hunting methods, which is far different from the lifestyle of later genera such as the huge ''Tropeognathus''. This conclusion was later followed by Howse and colleagues suggesting that the strong, yet bizarre teeth of ''Istiodactylus'' were more suited for scavenging carcasses rather than catching fish, this led the idea of a different feeding technique, which consisted in bitting and twisting the skull to remove large chunks out of the carcass, very different from the initial suggestions of how early ornithocheiromorphs fed. Witton in 2012 found that the teeth of ''Istiodactylus'' were unlike the recurved and enlarged teeth seen in the more derived ornithocheirids such as ''Ornithocheirus'', he instead pointed out that it was more "razor-edged" and better suited for carrion rather than fish.<ref name=witton2012/> |
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Another ornithocheiromorph that possessed similar features to ''Istiodactylus'' is ''[[Liaoxipterus]]'', which is known from a skull with several unique traits, including numerous peg-like teeth. The shape of its teeth indicated that ''Liaoxipterus'' was a possible [[insectivore]],<ref name=liaoxipterus_hyoid>{{cite journal|doi=10.3975/cagsb.2015.03.13|url=http://www.cagsbulletin.com/dqxben/ch/reader/view_abstract.aspx?flag=1&file_no=20150313&journal_id=dqxbcn|title=The Hyoid Apparatus of ''Liaoxipterus brachycephalus'' (Pterosauria) and Its Implications for Food-catching Behavior}}</ref> though this conclusion isn't considered entirely accurate.<ref>{{cite journal|author=Jiang S, Li Z, Cheng X, Wang X.|year=2020|title=The first pterosaur basihyal, shedding light on the evolution and function of pterosaur hyoid apparatuses.|journal=PeerJ|volume=8|pages=292|doi=10.7717/peerj.8292|pmid=31934505|pmc=6951291}}</ref> ''Liaoxipterus'' was also smaller in size compared to ''Istiodactylus'' and other ornithocheiromorphs, this is perhaps an indication that the more primitive members of this group (istiodactylids in this case) were smaller compared to more derived and advanced families such as the lonchodectids, ornithocheirids and targaryendraconians. This can also be seen in the size comparison between ornithocheirids and istiodactylids, with the wingspan of ''Tropeognathus'' for example, measuring almost double that of ''Istiodactylus''.<ref name="Wellnhofer1987"/><ref name=witton2012/> |
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[[File:OrnitocheiridsDB.jpg|thumb|''[[Cimoliopterus]]'' (right) stealing prey from a ''[[Lonchodectes]]'' (left), both were [[Derived (phylogenetics)|derived]] members of this group and possible [[Piscivore|fish hunters]]]] |
[[File:OrnitocheiridsDB.jpg|thumb|''[[Cimoliopterus]]'' (right) stealing prey from a ''[[Lonchodectes]]'' (left), both were [[Derived (phylogenetics)|derived]] members of this group and possible [[Piscivore|fish hunters]]]] |
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Ornithocheiromorphs were originally regarded as [[piscivorous]] creatures, feeding mainly on small and mid-sized fish.<ref name=ornithocheiroids/> Some paleontologists even suggested details on how these pterosaurs caught fish, some of which included dipping their [[beak]]s close to the water for prey.<ref name=ornithocheiroids/> Hooley for example, found that the beak of the well known ''Istiodactylus'' was similar to those of birds such as [[heron]]s, [[stork]]s, and [[Skimmer (bird)|skimmer]]s, and suggested that ''Istiodactylus'' probably fed on fish, this was mainly based on his 1913 jaw reconstruction of the animal.<ref name=hooley1913/> In 1991, Peter Wellnhofer compared the jaw endings of ''Istiodactylus'' with those of a [[duck]], but he then noticed that it wasn't a "duck-billed pterosaur" or anything similar, even though it was popularly called that way.<ref name="Wellnhofer">{{cite book|author=Wellnhofer, P.|author-link=Peter Wellnhofer|year=1991 |pages=114–116|title=The Illustrated Encyclopedia of Pterosaurs|publisher=Crescent Books|isbn=978-0-517-03701-0|location=New York}}</ref> An analysis by Witton in 2012 found that the teeth of ''Istiodactylus'' were unlike the recurved and enlarged teeth seen in the more derived ornithocheirids such as ''Ornithocheirus'', he instead pointed out that it was more "razor-edged" and better suited for carrion rather than fish.<ref name=witton2012/> |
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Ornithocheirids like ''Tropeognathus'' and ''Coloborhynchus'' are considered to be fish-eaters, and had longer and sharper teeth compared to the more rounded teeth of ''Istiodactylus'', though this is still sometimes disputed.<ref name=ornithocheiroids>{{Cite web|url=https://gsa.confex.com/gsa/2017AM/webprogram/Paper305496.html|title=Abstract: DIET OF ORNITHOCHEIROID PTEROSAURS INFERRED FROM STABLE CARBON ISOTOPE ANALYSIS OF TOOTH ENAMEL (GSA Annual Meeting in Seattle, Washington, USA - 2017)|website=gsa.confex.com}}</ref> Another difference that can be seen in more primitive ornithocheiromorphs their eyes being proportionally smaller compared to the assumed predatory and more advanced ornithocheirids, this again adds to the fact that the more [[primitive (phylogenetics)|primitive]] groups (istiodactylids) were most likely scavengers, and later got more successful within the food chain, leading the later, more advanced groups (ornithocheirids and lonchodectids) to be the [[dominance (ecology)|dominant]] fish hunters during the early Late Cretaceous.<ref name=ornithocheiroids/><ref>{{Cite journal|last1=Jacobs|first1=Megan L.|last2=Martill|first2=David M.|last3=Unwin|first3=David M.|last4=Ibrahim|first4=Nizar|author-link4=Nizar Ibrahim|last5=Zouhri|first5=Samir|last6=Longrich|first6=Nicholas R.|year=2020|title=New toothed pterosaurs (Pterosauria: Ornithocheiridae) from the middle Cretaceous Kem Kem beds of Morocco and implications for pterosaur palaeobiogeography and diversity|url=http://www.sciencedirect.com/science/article/pii/S0195667119303258|journal=Cretaceous Research|language=en|volume=110|pages=104413|doi=10.1016/j.cretres.2020.104413|issn=0195-6671|via=}}</ref> |
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Another ornithocheiromorph that possessed similar features to ''Istiodactylus'' is ''[[Liaoxipterus]]'', which is known from a skull with several unique traits, including numerous peg-like teeth. The shape of its teeth indicated that ''Liaoxipterus'' was a possible [[insectivore]],<ref name=liaoxipterus_hyoid>{{cite journal|doi=10.3975/cagsb.2015.03.13|url=http://www.cagsbulletin.com/dqxben/ch/reader/view_abstract.aspx?flag=1&file_no=20150313&journal_id=dqxbcn|title=The Hyoid Apparatus of ''Liaoxipterus brachycephalus'' (Pterosauria) and Its Implications for Food-catching Behavior}}</ref> though this conclusion isn't considered entirely accurate.<ref>{{cite journal|author=Jiang S, Li Z, Cheng X, Wang X.|year=2020|title=The first pterosaur basihyal, shedding light on the evolution and function of pterosaur hyoid apparatuses.|journal=PeerJ|volume=8|page=292|doi=10.7717/peerj.8292|pmid=31934505|pmc=6951291 |doi-access=free }}</ref> |
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==== Teeth enamels ==== |
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A 2017 study confirmed that most ornithocheirids were fish-eaters, though in a deeper analysis, paleontologists found carbon isotopes within several teeth, and later compared the [[tooth enamel|enamel]] values of the ornithocheirid teeth between the aquatic and terrestrial consumers near the fossil site.<ref name=ornithocheiroids/> Pterosaur enamels from the [[Twin Mountains Formation]] were shown with lower enamel values in comparison to the aquatic consumers that lived near, suggesting a more terrestrial diet, this is mostly due to the similarly low enamel values found in carnivorous theropods such as ''[[Acrocanthosaurus]]'', which mainly had a terrestrial diet. The enamels found in the [[Paw Paw Formation]] however, were described with higher values, in comparison to the nearby terrestrial consumers. Analyzing these enamel remains, paleontologists found out that the values seen were more comparable to the aquatic consumer values rather the terrestrial ones. These analyses are considered a very useful reference to the evolutionary changes between the earlier and later species of ornithocheirids, as well as other ornithocheiromorphs. The diet changes were also taken into account, with earlier genera such as ''Coloborhynchus'' having a varied diet, preying partially on fish, while also hunting on land. The later genera such as ''[[Uktenadactylus]]'' however, fed almost entirely on fish, and rarely hunted on land.<ref name=ornithocheiroids/> This can be demonstrated with the age difference between the Twin Mountains Formation and the Paw Paw Formation. The enamel remains found in the Twin Mountains Formation dated back to the earlier Aptian stage, while the remains found in the Paw Paw Formation dated back to the later Albian stage, resulting in a slight diet transition between more primitive and more derived genera.<ref name=ornithocheiroids/> |
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Anhanguerids like ''Tropeognathus'' and ''Coloborhynchus'' are considered to be fish-eaters, and had longer and sharper teeth compared to the more rounded teeth of ''Istiodactylus'', though this is still sometimes disputed.<ref name=ornithocheiroids>{{Cite web|url=https://gsa.confex.com/gsa/2017AM/webprogram/Paper305496.html|title=Abstract: DIET OF ORNITHOCHEIROID PTEROSAURS INFERRED FROM STABLE CARBON ISOTOPE ANALYSIS OF TOOTH ENAMEL (GSA Annual Meeting in Seattle, Washington, USA - 2017)|website=gsa.confex.com}}</ref> Another difference that can be seen in more primitive ornithocheiromorphs their eyes being proportionally smaller compared to the assumed predatory and more advanced anhanguerids, this again adds to the fact that the more [[primitive (phylogenetics)|primitive]] groups were most likely scavengers, and later got more successful within the food chain, leading the later, more advanced groups to be the [[dominance (ecology)|dominant]] fish hunters during the early Late Cretaceous.<ref>{{Cite journal|last1=Jacobs|first1=Megan L.|last2=Martill|first2=David M.|last3=Unwin|first3=David M.|last4=Ibrahim|first4=Nizar|author-link4=Nizar Ibrahim|last5=Zouhri|first5=Samir|last6=Longrich|first6=Nicholas R.|year=2020|title=New toothed pterosaurs (Pterosauria: Ornithocheiridae) from the middle Cretaceous Kem Kem beds of Morocco and implications for pterosaur palaeobiogeography and diversity|url=http://www.sciencedirect.com/science/article/pii/S0195667119303258|journal=Cretaceous Research|language=en|volume=110|page=104413|doi=10.1016/j.cretres.2020.104413|s2cid=214542129 |issn=0195-6671}}</ref> |
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=== Respiration === |
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[[File:Pterosaurs.jpg|thumb|upright|Diagrams of three different [[pterosaur]] [[genera]], including an [[Anhangueridae|anhanguerid]] (bottom); notice the breathing motion (top two) and internal [[air sac]] system (bottom two)]] |
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Pterosaur studies in 2009 showed that several pterosaurs, including ornithocheiromorphs, had a lung-and-air-sac system and a precisely controlled skeletal breathing pump, which supports a flow-through pulmonary ventilation model in pterosaurs, analogous to that of birds. This breathing method is also seen is carnivorous dinosaurs such as ''[[Aerosteon]]'', but there are currently no studies showing its relationships with pterosaurs.<ref name=Sereno>{{cite journal |authors=Sereno, P.C., Martinez,R.N., Wilson, J.A., Varricchio, D.J., Alcober, O.A., and Larsson, H.C.E. |year=2008 |title=Evidence for Avian Intrathoracic Air Sacs in a New Predatory Dinosaur from Argentina |journal=PLOS ONE |volume=3 |issue=9 |pages=e3303 |doi=10.1371/journal.pone.0003303 |pmid=18825273 |pmc=2553519 |editor1-last=Kemp |editor1-first=Tom |bibcode=2008PLoSO...3.3303S }}</ref> A [[wikt:subcutaneous|subcutaneous]] air sac system is present in at least some species of pterodactyloids, and this technique would have given a further reduced density to the living animal when breathing.<ref name=claessensetal2009>{{cite journal |vauthors=Claessens LP, O'Connor PM, Unwin DM |title=Respiratory evolution facilitated the origin of pterosaur flight and aerial gigantism |journal=PLOS ONE |volume=4 |issue=2 |pages=e4497 |year=2009 |pmid=19223979 |pmc=2637988 |doi=10.1371/journal.pone.0004497 |editor1-last=Sereno |editor1-first=Paul |bibcode=2009PLoSO...4.4497C}}</ref> Similar to modern crocodilians, many ornithocheiromorph genera appeared to have had some kind of [[Reptile#Respiratory system|hepatic piston]], regarding their shoulder-pectoral girdles, which were suggested to be too inflexible to move the sternum. This can also be seen in modern-day birds, which similarly possess strong gastralia, and therefore paleontologists erect the idea that their [[respiratory system]] had characteristics comparable to primitive archosaurs.<ref>{{cite journal | last1 = Geist | first1 = N. | last2 = Hillenius | first2 = W. | last3 = Frey | first3 = E. | last4 = Jones | first4 = T. | last5 = Elgin | first5 = R. | year = 2014 | title = Breathing in a box: Constraints on lung ventilation in giant pterosaurs | url = | journal = The Anatomical Record | volume = 297 | issue = 12| pages = 2233–2253 | doi = 10.1002/ar.22839 | pmid = 24357452| s2cid = 27659270 }}</ref> |
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=== Locomotion and flight === |
=== Locomotion and flight === |
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[[File:IstiodactylusDB3.jpg|thumb|left|Restoration of a flying ''Istiodactylus''; notice its high aspect ratio]] |
[[File:IstiodactylusDB3.jpg|thumb|left|Restoration of a flying ''Istiodactylus''; notice its high aspect ratio]] |
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Ornithocheiromorphs, like other pterosaurs, are considered to have been skilled fliers as well as swift at moving on the ground. Footprints from several species show that most pterosaurs did not sprawl their limbs to a large degree, as in modern reptiles, but rather held the limbs relatively erect when walking, like [[dinosaur]]s. While footprints are yet to be known, it is likely that ornithocheiromorphs also walked erect.<ref name=wittonhabib2010>{{cite journal | last1 = Witton | first1 = M.P. | last2 = Habib | first2 = M.B. | year = 2010 | title = On the Size and Flight Diversity of Giant Pterosaurs, the Use of Birds as Pterosaur Analogues and Comments on Pterosaur Flightlessness | journal = PLOS ONE | volume = 5 | issue = 11| page = e13982 | doi = 10.1371/journal.pone.0013982 | pmid=21085624 | pmc=2981443| bibcode = 2010PLoSO...513982W }}</ref> Compared to other earlier pterosaurs such as rhamphorhynchids, ornithocheiromorphs had unusually uneven [[limb (anatomy)|limb]] proportions, with the forelimbs resulting in a much longer scale compared to the hind limbs. Their close relatives, the pteranodontids, were also found with similar features, though they more likely flew like modern-day |
Ornithocheiromorphs, like other pterosaurs, are considered to have been skilled fliers as well as swift at moving on the ground. Footprints from several species show that most pterosaurs did not sprawl their limbs to a large degree, as in modern reptiles, but rather held the limbs relatively erect when walking, like [[dinosaur]]s. While footprints are yet to be known, it is likely that ornithocheiromorphs also walked erect.<ref name=wittonhabib2010>{{cite journal | last1 = Witton | first1 = M.P. | last2 = Habib | first2 = M.B. | year = 2010 | title = On the Size and Flight Diversity of Giant Pterosaurs, the Use of Birds as Pterosaur Analogues and Comments on Pterosaur Flightlessness | journal = PLOS ONE | volume = 5 | issue = 11| page = e13982 | doi = 10.1371/journal.pone.0013982 | pmid=21085624 | pmc=2981443| bibcode = 2010PLoSO...513982W | doi-access = free }}</ref> Compared to other earlier pterosaurs such as rhamphorhynchids, ornithocheiromorphs had unusually uneven [[limb (anatomy)|limb]] proportions, with the forelimbs resulting in a much longer scale compared to the hind limbs. Their close relatives, the pteranodontids, were also found with similar features, though they more likely flew like modern-day [[albatross]]es rather than anything else. Paleontologists also suggest that they most likely spend long stretches of time sea fishing, traveling very long distances without flapping while at the same time flying close the surface of the water with exploited wind speed, and without the necessity of [[thermal]]s.<ref name=padian1983>{{Cite journal | author = Padian, K. | year = 1983 | title = A functional analysis of flying and walking in pterosaurs | journal = Paleobiology | volume = 9 | issue = 3| pages = 218–239| doi = 10.1017/S009483730000765X| s2cid = 88434056 }}</ref> This would likely have required them to use unique modes of locomotion compared to other pterosaurs, this can already be seen in earlier evolutions such as ''Istiodactylus'' and ''[[Nurhachius]]'', with powerful musculature attachments and well-developed pectoral and upper arm bones.<ref name=witton2012/><ref name="Zhou2019">Zhou X., Pêgas R.V., Leal M.E.C. & Bonde N. 2019. ''Nurhachius luei'', a new istiodactylid pterosaur (Pterosauria, Pterodactyloidea) from the Early Cretaceous Jiufotang Formation of Chaoyang City, Liaoning Province (China) and comments on the Istiodactylidae". ''PeerJ'' '''7''':e7688. {{DOI: 10.7717/peerj.7688}}</ref> It is also possible that ornithocheiromorphs ran (but not walked) bipedally, or that they used a hopping gait.<ref name=wittonhabib2010/> Many pterosaur researchers like Mike Habib have noted that the limb proportions of some ornithocheiromorphs such as ''Anhanguera'' are consistent with hopping, though the scavenging istiodactylids are probably still the best examples of pterosaurs with a more terrestrial setting.<ref name=habibblog>Habib, M. (2011). "[http://h2vp.blogspot.com/2011/09/dinosaur-revolution-anhanguera.html Dinosaur Revolution: ''Anhanguera''.]" ''H2VP: Paleobiomechanics''. Weblog entry, September 20, 2011. Accessed September 28, 2011: http://h2vp.blogspot.com/2011/09/dinosaur-revolution-anhanguera.html</ref><ref name=hooley1913/> |
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== Paleoecology == |
== Paleoecology == |
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[[File:Araripe Basin map - formations and resources.jpg|thumb|250px|[[Geologic map|Geological map]] of the [[Araripe Basin]], with the extent of the [[Santana Group]] shown in dark blue]] |
[[File:Araripe Basin map - formations and resources.jpg|thumb|250px|[[Geologic map|Geological map]] of the [[Araripe Basin]], with the extent of the [[Santana Group]] shown in dark blue]] |
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[[File:20130825 Brazil Rio de Janeiro 0144.jpg|thumb|Mounted skeleton of ''[[Irritator]]'' with a possible anhanguerid in its jaws]] |
[[File:20130825 Brazil Rio de Janeiro 0144.jpg|thumb|Mounted skeleton of ''[[Irritator]]'' with a possible anhanguerid in its jaws]] |
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Even though the ornithocheiromorphs were discovered worldwide, most of them were concentrated in specific places. One of which is the fossil site called Romualdo Formation, which contains an impressive amount of pterosaur fossils. It is a diverse [[Lagerstätte]] in the larger geologic group called the [[Santana Group]] (sometimes called the Santana Formation), which is located in the [[Araripe Basin]] of northeastern Brazil, and dates back around 111 and 108 million years ago, during the [[Albian]] stage of the Early Cretaceous. The formation includes many species of ''Anhanguera'',<ref name=kellner2000/> several fossil remains of basal ornithocheiromorphs, including ''[[Brasileodactylus]]'', ''Cearadactylus'' and ''[[Unwindia]]'', as well as the anhanguerids ''Tropeognathus'',<ref name="Wellnhofer1987"/> ''Coloborhynchus'', ''[[Maaradactylus]]'', and ''[[Araripesaurus]]''.<ref name=fastnacht2001/> These pterosaur genera were just some of the many recovered from the site, which also include the [[thalassodromine]]s ''[[Tupuxuara]]'' and ''[[Thalassodromeus]]'',<ref name=":0">{{cite journal |last1=Pêgas |first1=R. V. |last2=Costa |first2=F. R. |last3=Kellner |first3=A. W. A. |title=New Information on the osteology and a taxonomic revision of The genus ''Thalassodromeus'' (Pterodactyloidea, Tapejaridae, Thalassodrominae) |journal=Journal of Vertebrate Paleontology |date=2018 |volume=38 |issue=2 |pages=e1443273 |doi=10.1080/02724634.2018.1443273|s2cid=90477315 }}</ref> as well as the [[tapejarid]] ''[[Tapejara (pterosaur)|Tapejara]]''.<ref name=KC07>{{cite journal |last=Kellner |first=A.W.A. |author2=Campos, D.A. |year=2007 |title=Short note on the ingroup relationships of the Tapejaridae (Pterosauria, Pterodactyloidea |journal=Boletim do Museu Nacional |volume=75 |pages=1–14 }}</ref> Some other creatures from the formation include the theropods ''[[Irritator]]'', ''[[Mirischia]]'' and ''[[Santanaraptor]]'', and the [[crocodylomorph]] ''[[Araripesuchus]]''. The formation also includes several turtle remains, with some specimens referring to ''[[Santanachelys]]'', ''[[Cearachelys]]'' and ''[[Araripemys]]''.<ref>{{cite web|last1=Zug|first1=George R.|title=Turtle: Origin and evolution|url=http://www.britannica.com/animal/turtle-reptile/Origin-and-evolution#ref917622|website=Encyclopædia Britannica|access-date=September 18, 2015}}</ref> A few fish remains were also found within the Romualdo Formation, some of which were referred to ''[[Brannerion]]'', ''[[Rhinobatos]]'', ''[[Rhacolepis]]'', ''[[Tharrhias]]'' and ''[[Tribodus]]''. The Santana Group also consists of another Lagerstätte called the [[Crato Formation]], which is not as diverse as the Romualdo Formation, but its fossil remains are still considered important.<ref name=martilletal2007>Martill, D.M., Bechly, G. and Loveridge, R.F. (2007). ''The Crato fossil beds of Brazil: window into an ancient world.'' Cambridge University Press. {{ISBN|0-521-85867-4}}, {{ISBN|978-0-521-85867-0}}</ref> This fossil site underlies the Romualdo Formation, and dates back around 115 and 113 million years ago during the [[Aptian]] stage, meaning that its fossil content is of older age. Similarly, the Crato Formation also contained several species of pterosaurs, including the basal lanceodontians ''Ludodactylus''<ref name=MFDB00/> and ''Brasileodactylus'', as well as the ornithocheirid ''[[Arthurdactylus]]''. Other pterosaur genera include the tapejarids ''[[Tupandactylus]]'' and ''[[Aymberedactylus]]'',<ref name=KC07/> as well as the [[chaoyangopterid]] ''[[Lacusovagus]]''. The formation also contains other creatures such as the [[enantiornithine]] ''[[Cratoavis]]'', the [[neosuchia]]n ''[[Susisuchus]]'', and several species of fish, including ''[[Belonostomus]]'', ''[[Calamopleurus]]'',<ref>{{cite journal |author=Peter L. Forey & Lance Grande |year=1998 |title=An African twin to the Brazilian ''Calamopleurus'' (Actinopterygii: Amiidae) |journal=[[Zoological Journal of the Linnean Society]] |volume=123 |issue=2 |pages=179–195 |doi=10.1111/j.1096-3642.1998.tb01299.x|doi-access=free }}</ref> ''[[Cladocyclus]]'',<ref name=FWCladocyclus>[https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=35248 ''Cladocyclus''] at [[Fossilworks]].org</ref> ''[[Dastilbe]]''<ref name="sepkoskidb">{{cite journal|last=Sepkoski |first=Jack |title=A compendium of fossil marine animal genera |journal=Bulletins of American Paleontology |volume=364 |page=560 |date=2002 |url=http://strata.ummp.lsa.umich.edu/jack/showgenera.php?taxon=611&rank=class |access-date=February 27, 2009 |archive-url=https://web.archive.org/web/20110723131237/http://strata.ummp.lsa.umich.edu/jack/showgenera.php?taxon=611&rank=class |archive-date=July 23, 2011 }}</ref> and ''[[Lepidotes]]''.<ref name=Lepidotes>{{cite journal|last=Woodward|first=A.S.|year=1895|title=Catalogue of the Fossil Fishes in the British Museum (Natural History)|location=British Museum of Natural History Department of Geology|volume=2|pages=77–119|doi=10.5962/bhl.title.61854|url=https://www.biodiversitylibrary.org/item/125733|doi-access=free|hdl=2027/coo1.ark:/13960/t3jx39x2t|hdl-access=free}}</ref> These fish genera were suggested to be prey for the pterosaurs that lived in the formation, but fossil remains are limited, so the subject is still controversial.<ref name=ornithocheiroids/> |
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Ornithocheiromorphs were present since the Early Cretaceous ([[Valanginian]] stage), and evolved throughout the rest of the period. Studies on the ornithocheirids ''Tropeognathus''<ref name="Wellnhofer1987"/> and ''Anhanguera'',<ref name=kellner2000/> suggested that ornithocheiromorphs also expanded their distribution, while also evolving specialized features. The significant adaptions of the ornithocheiromorphs had also made them the top pterosaurs in almost every formation by the early Late Cretaceous, including the ones they were rare in, such as the [[Blesa Formation]] in Spain, where only ''Iberodactylus'' has been found.<ref name="Holgado2019">Borja Holgado, Rodrigo V. Pêgas, José Ignacio Canudo, Josep Fortuny, Taissa Rodrigues, Julio Company & Alexander W.A. Kellner, 2019, "On a new crested pterodactyloid from the Early Cretaceous of the Iberian Peninsula and the radiation of the clade Anhangueria", ''Scientific Reports'' '''9''': 4940. {{DOI|10.1038/s41598-019-41280-4}}</ref> |
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Even though the ornithocheiromorphs were discovered worldwide, most of them were concentrated in specific places. One specific fossil site containing an impressive number of pterosaurs is the Romualdo Formation, a diverse [[Lagerstätte]] in the larger geologic group called the [[Santana Group]] (sometimes called the Santana Formation), which is located in the [[Araripe Basin]] of northeastern Brazil, and dates back 111-108 million years ago during the [[Albian]] stage of the Early Cretaceous. The formation includes many species of ''Anhanguera'',<ref name=kellner2000/> several fossil remains of basal ornithocheiromorphs, including ''[[Brasileodactylus]]'', ''Cearadactylus'' and ''[[Unwindia]]'', as well as the ornithocheirids ''Tropeognathus'',<ref name="Wellnhofer1987"/> ''Coloborhynchus'', ''Maaradactylus'', ''[[Araripesaurus]]'' and ''[[Barbosania]]''.<ref name=fastnacht2001/> These pterosaur genera were just some of the many recovered from the site, which also include the [[thalassodromid]]s ''[[Tupuxuara]]'' and ''[[Thalassodromeus]]'',<ref name=":0">{{cite journal |last1=Pêgas |first1=R. V. |last2=Costa |first2=F. R. |last3=Kellner |first3=A. W. A. |title=New Information on the osteology and a taxonomic revision of The genus ''Thalassodromeus'' (Pterodactyloidea, Tapejaridae, Thalassodrominae) |journal=Journal of Vertebrate Paleontology |date=2018 |volume=38 |issue=2 |pages=e1443273 |doi=10.1080/02724634.2018.1443273|s2cid=90477315 }}</ref> as well as the [[tapejarid]] ''[[Tapejara (pterosaur)|Tapejara]]'',<ref name=KC07>{{cite journal |last=Kellner |first=A.W.A. |author2=Campos, D.A. |year=2007 |title=Short note on the ingroup relationships of the Tapejaridae (Pterosauria, Pterodactyloidea |journal=Boletim do Museu Nacional |volume=75 |pages=1–14 }}</ref> the basal pteranodontoid ''[[Santanadactylus]]'' and the dubious ''[[Araripedactylus]]''. Some other creatures from the formation include the theropods ''[[Irritator]]'', ''[[Mirischia]]'' and ''[[Santanaraptor]]'', and the [[crocodylomorph]] ''[[Araripesuchus]]''. The formation also includes several turtle remains, with some specimens referring to ''[[Santanachelys]]'', ''[[Cearachelys]]'' and ''[[Araripemys]]''.<ref>{{cite web|last1=Zug|first1=George R.|title=Turtle: Origin and evolution|url=http://www.britannica.com/animal/turtle-reptile/Origin-and-evolution#ref917622|website=Encyclopædia Britannica|accessdate=18 September 2015}}</ref> A few fish remains were also found within the Romualdo Formation, some of which were referred to ''[[Brannerion]]'', ''[[Rhinobatos]]'', ''[[Rhacolepis]]'', ''[[Tharrhias]]'' and ''[[Tribodus]]''. The Santana Group also consists of another Lagerstätte called the [[Crato Formation]], which is not as diverse as the Romualdo Formation, but its fossil remains are still considered important.<ref name=martilletal2007>Martill, D.M., Bechly, G. and Loveridge, R.F. (2007). ''The Crato fossil beds of Brazil: window into an ancient world.'' Cambridge University Press. {{ISBN|0-521-85867-4}}, {{ISBN|978-0-521-85867-0}}</ref> This fossil site underlies the Romualdo Formation, and dates back around 115-113 million years ago during the [[Aptian]] stage, meaning that its fossil content is of older age. Similarly, the Crato Formation also contained several species of pterosaurs, including the basal lanceodontians ''Ludodactylus''<ref name=MFDB00/> and ''Brasileodactylus'', as well as the ornithocheirid ''[[Arthurdactylus]]''. Other pterosaur genera include the tapejarids ''[[Tupandactylus]]'' and ''[[Aymberedactylus]]'',<ref name=KC07/> as well as the [[chaoyangopterid]] ''[[Lacusovagus]]''. The formation also contains other creatures such as the [[enantiornithine]] ''[[Cratoavis]]'', the [[neosuchia]]n ''[[Susisuchus]]'', and several species of fish, including ''[[Belonostomus]]'', ''[[Calamopleurus]]'',<ref>{{cite journal |author=Peter L. Forey & Lance Grande |year=1998 |title=An African twin to the Brazilian ''Calamopleurus'' (Actinopterygii: Amiidae) |journal=[[Zoological Journal of the Linnean Society]] |volume=123 |issue=2 |pages=179–195 |doi=10.1111/j.1096-3642.1998.tb01299.x|doi-access=free }}</ref> ''[[Cladocyclus]]'',<ref name=FWCladocyclus>[http://fossilworks.org/bridge.pl?a=taxonInfo&taxon_no=35248 ''Cladocyclus''] at [[Fossilworks]].org</ref> ''[[Dastilbe]]''<ref name="sepkoskidb">{{cite journal|last=Sepkoski |first=Jack |title=A compendium of fossil marine animal genera |journal=Bulletins of American Paleontology |volume=364 |pages=560 |date=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=2011-07-23 }}</ref> and ''[[Lepidotes]]''.<ref name=Lepidotes>{{cite journal|last=Woodward|first=A.S.|year=1895|title=Catalogue of the Fossil Fishes in the British Museum (Natural History)|location=British Museum of Natural History Department of Geology|volume=2|pages=77–119|doi=10.5962/bhl.title.61854}}</ref> These fish genera were suggested to be prey for the pterosaurs that lived in the formation, but fossil remains are limited, so the subject is still controversial.<ref name=ornithocheiroids/> |
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Another important fossil site is the [[Wessex Formation]] in the Isle of Wight, near the coast of England, which dates back 140 |
Another important fossil site is the [[Wessex Formation]] in the Isle of Wight, near the coast of England, which dates back around 140 and 125 million years ago ([[Berriasian]] to [[Barremian]] stages). The formation doesn't contain many fossil remains of pterosaurs compared to the Romualdo Formation, but it is still a very important site. Fossil remains of the istiodactylid ''Istiodactylus'',<ref name=witton2012/> the anhanguerid ''Caulkicephalus'',<ref name=Caulkicephalus/> and the tapejarid ''[[Wightia (pterosaur)|Wightia]]'' were found.<ref>{{Cite journal|last1=Martill|first1=David M.|last2=Green|first2=Mick|last3=Smith|first3=Roy|last4=Jacobs|first4=Megan|last5=Winch|first5=John|date=April 2020|title=First tapejarid pterosaur from the Wessex Formation (Wealden Group: Lower Cretaceous, Barremian) of the United Kingdom|journal=Cretaceous Research|volume=113|language=en|page=104487|doi=10.1016/j.cretres.2020.104487|s2cid=219099220|url=https://researchportal.port.ac.uk/portal/en/publications/first-tapejarid-pterosaur-from-the-wessex-formation-wealden-group-lower-cretaceous-barremian-of-the-united-kingdom(0c4f61db-81ab-468b-b075-905c358dac8c).html}}</ref> The formation is also known for several theropods, including the spinosaurid ''[[Baryonyx]]'', the [[tyrannosauroid]] ''[[Eotyrannus]]'', the [[dromaeosaurid]] ''Ornithodesmus'', the [[compsognathid]] ''[[Aristosuchus]]'' as well as the [[allosauroid]] ''[[Neovenator]]''.<ref name="Charig Milner 1986">{{cite journal |doi=10.1038/324359a0 |pmid=3785404 |title=''Baryonyx'', a remarkable new theropod dinosaur |journal=Nature |volume=324 |issue=6095 |pages=359–361 |year=1986 |last1=Charig |first1=A. J. |last2=Milner |first2=A. C. |bibcode=1986Natur.324..359C|s2cid=4343514 }}</ref><ref>{{cite journal | last1 = Hutt | first1 = S. | last2 = Simmonds | first2 = K. | last3 = Hullman | first3 = G. | year = 1990 | title = Predatory dinosaurs from the Isle of Wight | journal = Proceedings of the Isle of Wight Natural History and Archaeological Society | volume = 9 | pages = 137–146 }}</ref> Different types of herbivorous dinosaurs like ''[[Iguanodon]]'', ''[[Polacanthus]]'', ''[[Ornithopsis]]'', ''[[Mantellisaurus]]'' and ''[[Hypsilophodon]]'' were also found within the fossil site.<ref name="iguanodonts2010carpenter">{{cite journal | last1 = Carpenter | first1 = K. | last2 = Ishida | first2 = Y. | year = 2010 | title = Early and "Middle" Cretaceous Iguanodonts in Time and Space | journal = Journal of Iberian Geology | volume = 36 | issue = 2| pages = 145–164 | doi = 10.5209/rev_JIGE.2010.v36.n2.3| doi-access = free }}</ref><ref name=hulke1873>{{cite wikisource | last1=Hulke | first1=J. W. | title=Contribution to the Anatomy of Hypsilophodon Foxii | wslink=Quarterly Journal of the Geological Society of London/Volume 29/Contribution to the Anatomy of Hypsilophodon Foxii | work=Quarterly Journal of the Geological Society of London | volume=29 | year=1873 | doi=10.1144/GSL.JGS.1873.029.01-02.46 | pages=522–532 | firsticon=yes | noicon=yes }}</ref> Some other animals from the formation include the neosuchian ''[[Bernissartia]]'', sea turtles such as ''[[Helochelydra]]'' and ''[[Brodiechelys]]'', the cartilaginous fish ''[[Hybodus]]'', [[ray-finned fish]]es such as ''Belonostomus'', ''[[Caturus]]'', ''Lepidotes'' and ''[[Scheenstia]]'', as well as the [[mammal]]s ''[[Eobaatar]]'', ''[[Loxaulax]]'' and ''[[Yaverlestes]]''.<ref name=Lepidotes/><ref>{{Cite journal|last=Sweetman|first=Steven C.|date=September 2009|title=A New Species of the Plagiaulacoid Multituberculate Mammal Eobaatarfrom the Early Cretaceous of Southern Britain|journal=Acta Palaeontologica Polonica|volume=54|issue=3|pages=373–384|doi=10.4202/app.2008.0003|s2cid=53975359|issn=0567-7920|doi-access=free}}</ref><ref name="BGS lexicon">{{Cite web |url=http://www.bgs.ac.uk/lexicon/lexicon.cfm?pub=WSEX |title=Wessex Formation |series=The BGS Lexicon of Named Rock Units |publisher=[[British Geological Survey]]}}</ref> |
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[[File:Ferrodraco.png|thumb|left|Reconstruction of the skeleton of ''[[Ferrodraco]]'' on a diagram showing the known material, and based on the related ''Tropeognathus mesembrinus'']] |
[[File:Ferrodraco.png|thumb|left|Reconstruction of the skeleton of ''[[Ferrodraco]]'' on a diagram showing the known material, and based on the related ''Tropeognathus mesembrinus'']] |
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A few fossils reported from the [[Toolebuc Formation]] and [[Winton Formation]] are believed to be from some of the most derived ornithocheiromorphs, due to the age of the fossil remains, which dated back to the Albian and [[Cenomanian]] stages of the Cretaceous, and some are even believed to belong to the Turonian stage. The Toolebuc Formation includes several remains of ornithocheiromorphs which are now referred to the genera ''[[Aussiedraco]]'' and ''Mythunga''.<ref name=Aussiedraco>{{cite journal |last=Kellner |first=Alexander W.A. |author2=Taissa Rodrigues |author3=Fabiana R. Costa |year=2011 |title=Short note on a pteranodontoid pterosaur (Pterodactyloidea) from western Queensland, Australia |url=http://www.scielo.br/pdf/aabc/v83n1/v83n1a18.pdf |journal=Anais da Academia Brasileira de Ciências |volume=83 |issue=1 |pages=301–308 |doi=10.1590/S0001-37652011000100018 |pmid=21437387}}</ref><ref name=MT08>{{cite journal |last=Molnar |first=Ralph E. |author2=Thulborn, R.A. |year=2008 |title=An incomplete pterosaur skull from the Cretaceous of north-central Queensland, Australia |journal=Arquivos do Museu Nacional, Rio de Janeiro |volume=65 |issue=4 |pages=461–470 }}</ref> The formation also includes several herbivorous dinosaurs such as the [[ornithopod]] ''[[Muttaburrasaurus]]'' and the [[ankylosaur]] ''[[Kunbarrasaurus]]''.<ref name=iguanocolossus>{{cite journal | last1 = McDonald | first1 = A.T. | last2 = Kirkland | first2 = J.I. | last3 = DeBlieux | first3 = D.D. | last4 = Madsen | first4 = S.K. | last5 = Cavin | first5 = J. | last6 = Milner | first6 = A.R.C. | last7 = Panzarin | first7 = L. | year = 2010 | title = New Basal Iguanodonts from the Cedar Mountain Formation of Utah and the Evolution of Thumb-Spiked Dinosaurs |
A few fossils reported from the [[Toolebuc Formation]] and [[Winton Formation]] are believed to be from some of the most derived ornithocheiromorphs, due to the age of the fossil remains, which dated back to the Albian and [[Cenomanian]] stages of the Cretaceous, and some are even believed to belong to the Turonian stage. The Toolebuc Formation includes several remains of ornithocheiromorphs which are now referred to the genera ''[[Aussiedraco]]'' and ''Mythunga''.<ref name=Aussiedraco>{{cite journal |last=Kellner |first=Alexander W.A. |author2=Taissa Rodrigues |author3=Fabiana R. Costa |year=2011 |title=Short note on a pteranodontoid pterosaur (Pterodactyloidea) from western Queensland, Australia |url=http://www.scielo.br/pdf/aabc/v83n1/v83n1a18.pdf |journal=Anais da Academia Brasileira de Ciências |volume=83 |issue=1 |pages=301–308 |doi=10.1590/S0001-37652011000100018 |pmid=21437387}}</ref><ref name=MT08>{{cite journal |last=Molnar |first=Ralph E. |author2=Thulborn, R.A. |year=2008 |title=An incomplete pterosaur skull from the Cretaceous of north-central Queensland, Australia |journal=Arquivos do Museu Nacional, Rio de Janeiro |volume=65 |issue=4 |pages=461–470 }}</ref> The formation also includes several herbivorous dinosaurs such as the [[ornithopod]] ''[[Muttaburrasaurus]]'' and the [[ankylosaur]] ''[[Kunbarrasaurus]]''.<ref name=iguanocolossus>{{cite journal | last1 = McDonald | first1 = A.T. | last2 = Kirkland | first2 = J.I. | last3 = DeBlieux | first3 = D.D. | last4 = Madsen | first4 = S.K. | last5 = Cavin | first5 = J. | last6 = Milner | first6 = A.R.C. | last7 = Panzarin | first7 = L. | year = 2010 | title = New Basal Iguanodonts from the Cedar Mountain Formation of Utah and the Evolution of Thumb-Spiked Dinosaurs | journal = PLOS ONE | volume = 5| issue = 11| page = e14075 | doi = 10.1371/journal.pone.0014075 | pmid = 21124919 | pmc=2989904| bibcode = 2010PLoSO...514075M | doi-access = free }}</ref><ref name="Leahy2015">{{Cite journal|author1=Lucy G. Leahey |author2=Ralph E. Molnar |author3=Kenneth Carpenter |author4=Lawrence M. Witmer |author5=Steven W. Salisbury |year=2015 |title=Cranial osteology of the ankylosaurian dinosaur formerly known as ''Minmi'' sp. (Ornithischia: Thyreophora) from the Lower Cretaceous Allaru Mudstone of Richmond, Queensland, Australia |journal=PeerJ |volume=3 |pages=e1475 |doi=10.7717/peerj.1475 |pmid=26664806 |pmc=4675105 |doi-access=free }}</ref> Fossil remains of marine animals were also uncovered within the fossil site, and some specimens of which belong to the [[ichthyosaur]] ''[[Platypterygius]]'', the [[pliosaurid]] ''[[Kronosaurus]]'' and the [[elasmosaurid]] ''[[Eromangasaurus]]''.<ref name=Gardiner1931>{{cite journal |last1=Gardiner |first1=J. Stanley |title=The Harvard Museum Expedition to Australia |journal=Nature |date=September 1931 |volume=128 |issue=3228 |pages=457–458 |doi=10.1038/128457c0 |bibcode=1931Natur.128..457G |s2cid=29715877 }}</ref> Turtle remains from turtles that were proposed to be prey for pterosaurs were also found within the Toolebuc Formation, this included the genera ''[[Bouliachelys]]'', ''[[Cratochelone]]'' and ''[[Notochelone]]''.<ref name=ornithocheiroids/> The Winton Formation consisted on a more terrestrial environment, containing several [[sauropod]] dinosaurs like ''[[Austrosaurus]]'', ''[[Diamantinasaurus]]'', ''[[Savannasaurus]]'' and ''[[Wintonotitan]]'' as well as large carnivorous dinosaurs such as ''[[Australovenator]]''<ref name= Whiteetal2013>{{Cite journal | last1 = White | first1 = M. A. | last2 = Falkingham | first2 = P. L. | last3 = Cook | first3 = A. G. | last4 = Hocknull | first4 = S. A. | last5 = Elliott | first5 = D. A. | title = Morphological comparisons of metacarpal I for ''Australovenator wintonensis'' and ''Rapator ornitholestoides'': Implications for their taxonomic relationships | doi = 10.1080/03115518.2013.770221 | journal = Alcheringa: An Australasian Journal of Palaeontology | pages = 435–441 | year = 2013 | volume=37| issue = 4 | s2cid = 82672110 }}</ref> and crocodylomorphs like ''[[Isisfordia]]''. The formation's only pterosaur is the derived genus ''Ferrodraco'',<ref name="Pentland2019">{{Cite journal|last1=Pentland|first1=Adele H.|last2=Poropat|first2=Stephen F.|last3=Tischler|first3=Travis R.|last4=Sloan|first4=Trish|last5=Elliott|first5=Robert A.|last6=Elliott|first6=Harry A.|last7=Elliott|first7=Judy A.|last8=Elliott|first8=David A.|date=December 2019|title=''Ferrodraco lentoni'' gen. et sp. nov., a new ornithocheirid pterosaur from the Winton Formation (Cenomanian–lower Turonian) of Queensland, Australia|journal=Scientific Reports|volume=9|issue=1|page=13454|doi=10.1038/s41598-019-49789-4|pmid=31582757|pmc=6776501|bibcode=2019NatSR...913454P|issn=2045-2322}}</ref> which is also considered as one of the last ornithocheiromorphs, and a close relative of ''Mythunga''.<ref name=MT08/> |
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''Lonchodraco'', formerly known as ''Pterodactylus giganteus'',<ref name=Bowerbank1846>{{cite journal | last1 = Bowerbank | first1 = J.S. | year = 1846 | title = On a new species of pterodactyl found in the Upper Chalk of Kent (''Pterodactylus giganteus'') | url =https://zenodo.org/record/1448505 | journal = Quarterly Journal of the Geological Society of London | volume = 2 | issue = 1–2| pages = 7–9 | doi=10.1144/gsl.jgs.1846.002.01-02.05| s2cid = 129389179 }}</ref> lived throughout the late Cenomanian and early Turonian stages, and ''Cimoliopterus'' is also considered to be from the same time. ''Lonchodraco'', along with ''Lonchodectes'', are both classified as basal members of Ornithocheiromorpha, due to complex [[taxonomy (biology)|taxonomy]] issues, despite their young ages.<ref name="Martill2017">{{cite journal|last1=Rigal|first1=S.|last2=Martill|first2=D. M.|last3=Sweetman|first3=S. C.|title=A new pterosaur specimen from the Upper Tunbridge Wells Sand Formation (Cretaceous, Valanginian) of southern England and a review of ''Lonchodectes sagittirostris'' (Owen 1874)|journal=Geological Society, London, Special Publications|date=2017|volume=455|pages=221–232|doi=10.1144/SP455.5 |s2cid=133080548|url=https://www.researchgate.net/publication/313966182}}</ref> |
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== See also == |
== See also == |
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== Further reading == |
== Further reading == |
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*{{cite book|last=Wellnhofer|first=Peter|title=The Illustrated Encyclopedia of Pterosaurs: An illustrated natural history of the flying reptiles of the Mesozoic Era|publisher=Crescent Books|year=1991|isbn= |
*{{cite book|last=Wellnhofer|first=Peter|title=The Illustrated Encyclopedia of Pterosaurs: An illustrated natural history of the flying reptiles of the Mesozoic Era|publisher=Crescent Books|year=1991|isbn=0-517-03701-7}} |
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*{{cite book|last=Witton|first=Mark|title=Pterosaurs: Natural History, Evolution, Anatomy|publisher=Princeton University Press|year=2013|isbn=978- |
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Revision as of 08:21, 18 November 2024
Ornithocheiromorphs | |
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Reconstructed skeleton of Tropeognathus in the National Museum of Brazil | |
Skeletal cast of Maaradactylus spielbergi in the Naturalis Biodiversity Center | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Order: | †Pterosauria |
Suborder: | †Pterodactyloidea |
Clade: | †Pteranodontoidea |
Clade: | †Ornithocheiromorpha Andres et al., 2014 |
Subgroups | |
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Ornithocheiromorpha (from Ancient Greek, meaning "bird hand form") is a group of pterosaurs within the suborder Pterodactyloidea. Fossil remains of this group date back from the Early to Late Cretaceous periods (Valanginian to Turonian stages), around 140 to 92.5 million years ago. Ornithocheiromorphs were discovered worldwide except Antarctica, though most genera were recovered in Europe, Asia and South America.[2] They were the most diverse and successful pterosaurs during the Early Cretaceous, but throughout the Late Cretaceous they were replaced by pteranodontians and azhdarchoids. The Ornithocheiromorpha was defined in 2014 by Andres and colleagues, and they made Ornithocheiromorpha the most inclusive clade containing Ornithocheirus, but not Pteranodon.[3]
Ornithocheiromorphs are considered to be some of the largest animals to have ever flown. Members of this group are also regarded to have some of the largest pterosaur wingspans, such as the one estimated for the huge Tropeognathus, though still not as large as those estimated for the azhdarchids, which may have reached up to 12 meters (39 ft).[4] When ornithocheiromorphs first appeared, they were initially scavengers, consisting in a more terrestrial setting, but their success had made them the top predators of the skies, as well as the most common type of fish-eating pterosaur throughout the early Late Cretaceous. Some paleontologists also consider ornithocheiromorphs an earlier step of evolution to the pteranodontians, this is due to the similar flying techniques and flight locomotions, as well as their diet, which mainly consisted of fish, and therefore also hunted very similarly. Ornithocheiromorphs also flew like soaring birds, keeping their wings stretched and rarely flapping.
History of research
Early discoveries
The first specimens of ornithocheiromorphs were unearthed at a chalk pit near Burham in Kent, England. In 1846, British paleontologist James Scott Bowerbank named and described the remains found as Pterodactylus giganteus, as it was common at that time to assign any new described pterosaur species to Pterodactylus.[5] In the same chalk pit as P. giganteus, two other pterosaur species were discovered. The first was named in 1851 by Bowerbank as Pterodactylus cuvieri,[6] in honor of the prominent German naturalist and zoologist Georges Cuvier, while the second was described in the same year by British paleontologist Sir Richard Owen as Pterodactylus compressirostris.[7] P. compressirostris later became the type species of a newly created genus called Lonchodectes (meaning "lance biter") in a review by English paleontologist Reginald Walter Hooley in 1914.[8] Confusingly, this species was also long regarded, incorrectly, as the type species of Ornithocheirus.[9]
In 1861, further pterosaur specimens were found in the UK, and were given the new species Pterodactylus simus by Owen.[10] British paleontologist Harry Govier Seeley then created the new genus Ornithocheirus for the new species in the same year, the generic name translating as "bird hand" is due to the notion of the time that pterosaurs were the ancestors of modern birds. In 1870, Seeley reassigned the species Pterodactylus cuvieri as Ornithocheirus cuvieri.[11][8] In 1874, Richard Owen proposed two new genera, Coloborhynchus, meaning "maimed beak", and Criorhynchus, meaning "ram beak". While Coloborhynchus consisted in a totally new type species, C. clavirostris, as well as two other species reassigned from Ornithocheirus, Criorhynchus consisted entirely of former Ornithocheirus species, including O. simus, which was later reassigned by Owen as Criorhynchus simus.[12]
In 2013, Brazilian paleontologists Taissa Rodrigues & Alexander Kellner made a deeper analysis on the species Pterodactylus cuvieri. In the analysis, they stated that it needed a separate genus, and assigning it to Ornithocheirus was inappropriate, therefore, they created the new genus called Cimoliopterus, with the new resulting combination Cimoliopterus cuvieri. In the same study, Rodrigues & Kellner also reviewed the species Pterodactylus giganteus, and reassigned it to a newly created genus called Lonchodraco, this resulted in a new combination called Lonchodraco giganteus.[13]
In 1887, Seeley had described new fossil remains from the Isle of Wight, an island off the coast of southern England. He thought it belonged to some kind of bird-like creature, which he named it Ornithodesmus cluniculus.[14] Seeley also reported another specimen found on the same site. He then considered it another species of Ornithodesmus. In 1901, Seeley named this new species as O. latidens, meaning "wide tooth".[15] Later, Reginald Hooley discussed O. latidens in detail, based on specimens he had found, which led Ornithodesmus to be placed within a new family called Ornithodesmidae.[16] Paleontologist Charles William Andrews however, had expressed doubts as to whether O. latidens belonged in the genus Ornithodesmus, as the vertebrae of the specimen of that genus was based on differed markedly from those of Hooley's specimen.[17]
In 1993, the British paleontologists Stafford C. Howse and Andrew C. Milner concluded that the holotype sacrum and only specimen of O. cluniculus didn't belong to a pterosaur, but instead to a maniraptoran theropod dinosaur. They also pointed out that no detailed attempts had been made to compare the sacrum of O. cluniculus with those of pterosaurs, and that O. latidens had in effect been treated as the type species of the genus Ornithodesmus.[18] Howse, Milner, and David Martill in 2001, moved "O." latidens to a new genus called Istiodactylus. They had also named a new family called Istiodactylidae, with Istiodactylus as the only member.[19]
Discoveries outside Europe
Other important ornithocheiromorph discoveries include the anhanguerids Tropeognathus and Anhanguera from the Romualdo Formation in Brazil.[20][21] Tropeognathus was described with its type species, T. mesembrinus in 1987 by German paleontologist Peter Wellnhofer. The generic name is derived from Greek τρόπις, tropis, meaning "keel", and γνάθος, gnathos, meaning "jaw". The specific name is derived from Koine mesembrinos, "of the noontide", simplified as "southern", in reference to the provenance from the Southern hemisphere. The description then led to an enormous taxonomic confusion.[22] In 1989, Brazilian paleontologist Alexander Kellner considered it an Anhanguera mesembrinus,[23] then a Coloborhynchus mesembrinus by Veldmeijer in 1998,[24] and then a Criorhynchus mesembrinus in 2001 by German paleontologist Michael Fastnacht.[25] T. mesembrinus was then considered a junior synonym of Ornithocheirus simus by British paleontologist David Unwin in 2001, but he then proposed an Ornithocheirus mesembrinus in 2003.[26][27] In 2013 however, Taissa Rodrigues and Alexander Kellner concluded that Tropeognathus would be valid again, and containing only T. mesembrinus, the type species.[13]
A discovery in Asia, specifically northwestern China, was reported in 2006. The lake sediments allowed an exceptional preservation of fossils, and therefore paleontologists Qiu Zhanxiang and Wang Banyue started official excavations. Part of the findings consisted of dense concentrations of pterosaur bones, associated with soft tissues and eggs. In 2014, a new species was named and described: Hamipterus tianshanensis. It was named by Wang Xiaolin, Alexander Kellner, Jiang Shunxing, Wang Qiang, Ma Yingxia, Yahefujiang Paidoula, Cheng Xin, Taissa Rodrigues, Meng Xi, Zhang Jialiang, Li Ning, and Zhou Zhonghe. The generic name Hamipterus combines that of the Hami region, with the word pteron, meaning "wing", and the specific name refers to the provenance from the Tian Shan, a mountain range.[28]
Description
Size
Ornithocheiromorphs were large pterosaurs, with wingspans normally ranging between 3 and 6 meters (9.8 and 19.7 ft).[4] Istiodactylus for example, had a wingspan ranging from 4.3 to 5 meters (14 to 16 ft), with the most complete known skull estimated to have been about 45 centimeters (1.48 ft) in length, based on a long-lost fragment of its jaw reported in 2012.[29] Though its jaws measured only 28.5 centimeters (11.2 in), which was less than 80 percent of the skull's length.[17] Anhanguerids and were typically larger than others of the group and were more successful within the food chain rather than other ornithocheiromorphs, one reason is because of their large size, for example, Tropeognathus mesembrinus, had a normal wingspan of about 8.26 meters (27.1 ft), and 8.70 meters (28.5 ft) as the maximum estimate.[30] Another species which was impressively large is Coloborhynchus capito, with a total skull length that could have been up to 75 centimeters (2.46 ft), leading to an estimated wingspan of 7 meters (23 ft).[31] However, this species may belong to a different genus called Nicorhynchus.[32]
Skull and crests
Most anhanguerids bore distinctive convex "keeled" crests on their snout and underside of their mandible, this was well developed in several genera such as Tropeognathus.[33] The similar Anhanguera possessed jaws that were tapered in width, but expanded into a broad, spoon-shaped rosette at the tip. The jaws are distinguished from its relatives by several differences in the crest and teeth: unlike its close relatives Coloborhynchus and Ornithocheirus, the crest on the upper jaw of Anhanguera didn't begin at the tip of the snout, therefore, it was set farther back on the skull.[34]
Other anhanguerids like Cearadactylus had its first preparations with many serious mistakes: the front of the snout and the lower jaws were confused leading to a reconstruction in which the anterior part of the head was upside down.[35] Some of the teeth were extensively restored and enlarged until the wider front of the jaws showed very large and robust teeth projecting outward. With this arrangement, the maxilla was kinked, and its interlocking teeth suggested that Cearadactylus had a piscivourous diet, allowing the animal to keep hold of slippery fish.[36] Another smaller genus similar to Cearadactylus is Guidraco. Its holotype skull has a length of 38 centimeters (15 in), which makes it smaller than other genera. The skull is very elongated however, and a hollow profile is seen, but not very pointed, as the upper edge and the line of the jaw run nearly parallel over most of their length.[37]
Even though most ornithocheiromorphs didn't have a cranial crest like the closely related pteranodontids, there were some exceptions, this included Caulkicephalus and Ludodactylus.[38] Caulkicephalus had a rounded snout, very similar to that of Ornithocheirus and Anhanguera, and therefore it is placed within either Anhangueridae or Ornithocheiridae, depending on the author.[39][40] Caulkicephalus was also a large pterosaur, with wingspan estimates of around 5 meters (16 ft).[40]
Vertebrae
The vertebral column of ornithocheiromorphs was heavily pneumatized by an extensive system of air sacs, leaving prominent pneumatic foraminae.[41] The neck of ornithocheiromorphs was typically relatively long and robust, being longer than the torso in some derived clades.[42] The neural spines of ornithocheiromorph cervical vertebrae were generally tall and spikelike.[37][34] In some genera such as Tropeognathus and Istiodactylus, up to six dorsal vertebrae are fused into a notarium.[43] In some genera such as Anhanguera, four to seven sacral vertebrae are fused into a synsacrum.[34] The tail is not well known in ornithocheiromorphs, however. Zhenyuanopterus, which is known for having 13 caudal vertebrae, formed one of the longest tails of any pterodactyloid.[44] Anhanguera, another well-known genus, had a shorter tail, with broad caudal vertebrae that bore a "duplex" cross-section similar to Pteranodon.[34]
Pelvic structure
The pelvis of ornithocheiromorphs was of moderate size compared to the body as a whole, similar to other ornithocheiroids. The three pelvic bones were often fused, as seen in many species such as Anhanguera santanae, the ilium was long and low, and its front and rear blades projected horizontally beyond the edges of the lower pelvic bones.[45] Despite the structure length, the processes of these rod-like forms indicate that the hindlimb muscles attached to them were limited in strength.[46] The pubic bone was fused with the broad ischium into an ischiopubic blade, resulting in a narrow build. Sometimes, the blades of both sides were fused, closing the pelvis from below and forming the pelvic canal. The front of the pubic bones was also articulated with a unique structure, resulting in a pair of prepubic bones within. This formed a cusp covering the rear belly, and was located between the pelvis and the belly ribs. The hip joint of ornithocheiromorphs was not perforated and allowed considerable mobility to the leg, and suggests that it was vertical, as therefore had a function in breathing, compensating the relative rigidity of the chest cavity.[47][45]
Classification
Several studies show that ornithocheiromorphs were less derived than the toothless pteranodontids such as Pteranodon, and based on the different evolutionary changes, they therefore need to be grouped in a different clade than Pteranodon, though still within Pteranodontoidea.[48] In 2003, David Unwin considered the family Istiodactylidae to group with the toothless Pteranodontidae, within the group Ornithocheiroidea,[27] but Alexander Kellner however, grouped it with the toothed Anhangueridae instead, resulting in a more understandable change of evolution between the two toothed families.[49]
Brian Andres and colleagues found the families Istiodactylidae, Ornithocheiridae and Anhangueridae to form a group in 2014, which he called Lanceodontia, and consists of the more advanced ornithocheiromorphs. The clade however, excludes the more poorly known family Lonchodectidae, even though members of the family had previously been seen as some of the most derived forms of toothed pterosaurs. In their analysis, they also included the family Boreopteridae within the clade Anhangueria, though placed in a more basal position, while also containing the genus Guidraco.[3] In 2018 however, Nicholas Longrich and colleagues found Boreopteridae outside Anhangueria as the sister taxon of the family Lonchodectidae, both groups placed as basal members of the Ornithocheiromorpha.[50]
Relationships
There are competing theories of ornithocheiromorph phylogeny (evolutionary relationships). Below is cladogram following a topology recovered by Brian Andres, using the most recent iteration of his data set (Andres, 2021).[51]
Pteranodontoidea |
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Below is a cladogram showing the results of a phylogenetic analysis presented by Longrich et al. (2018). In the analysis, they placed the genus Hongshanopterus as a basal member.[50]
Ornithocheiromorpha | |
In 2019, several new species of ornithocheiromorphs were found, and the former species Ornithocheirus wiedenrothi was renamed as Targaryendraco wiedenrothi.[39] The description of Iberodactylus in Spain also made some paleontologists reclassify the genus Hamipterus in a newly named family called Hamipteridae.[52] The ornithocheirid Cimoliopterus was also reclassified as well, and it is currently grouped with Aetodactylus and Camposipterus in the clade Targaryendraconia, specifically to its own family, the Cimoliopteridae.[39] However, an analysis by Jacobs et al. (2019) recovers both Camposipterus and Cimoliopterus within the Ornithocheiridae again, using a new data matrix not including Targaryendraco.[53] The previously recovered basal eupterodactyloid Haopterus was reclassified due to the description of Mimodactylus, and is placed in a new family called Mimodactylidae.[54] However, a more recent analysis using the data in the description of Mimodactylus has found Haopterus as a basal member of the more inclusive group Istiodactyliformes.[55] Many recent analyses have also recovered several ornithocheirids, including Tropeognathus, Coloborhynchus, and Caulkicephalus within the family Anhangueridae, meaning that they were more closely related to Anhanguera than to Ornithocheirus.[52][39][55][54][32]
Paleobiology
Diet and feeding
Ornithocheiromorphs were originally regarded as piscivorous creatures, feeding mainly on small and mid-sized fish.[36] Some paleontologists even suggested details on how these pterosaurs caught fish, some of which included dipping their beaks close to the water for prey.[36] Hooley for example, found that the beak of the well known Istiodactylus was similar to those of birds such as herons, storks, and skimmers, and suggested that Istiodactylus probably fed on fish, this was mainly based on his 1913 jaw reconstruction of the animal.[16] In 1991, Peter Wellnhofer compared the jaw endings of Istiodactylus with those of a duck, but he then noticed that it wasn't a "duck-billed pterosaur" or anything similar, even though it was popularly called that way.[56] An analysis by Witton in 2012 found that the teeth of Istiodactylus were unlike the recurved and enlarged teeth seen in the more derived ornithocheirids such as Ornithocheirus, he instead pointed out that it was more "razor-edged" and better suited for carrion rather than fish.[29]
Another ornithocheiromorph that possessed similar features to Istiodactylus is Liaoxipterus, which is known from a skull with several unique traits, including numerous peg-like teeth. The shape of its teeth indicated that Liaoxipterus was a possible insectivore,[57] though this conclusion isn't considered entirely accurate.[58]
Anhanguerids like Tropeognathus and Coloborhynchus are considered to be fish-eaters, and had longer and sharper teeth compared to the more rounded teeth of Istiodactylus, though this is still sometimes disputed.[36] Another difference that can be seen in more primitive ornithocheiromorphs their eyes being proportionally smaller compared to the assumed predatory and more advanced anhanguerids, this again adds to the fact that the more primitive groups were most likely scavengers, and later got more successful within the food chain, leading the later, more advanced groups to be the dominant fish hunters during the early Late Cretaceous.[59]
Locomotion and flight
Ornithocheiromorphs, like other pterosaurs, are considered to have been skilled fliers as well as swift at moving on the ground. Footprints from several species show that most pterosaurs did not sprawl their limbs to a large degree, as in modern reptiles, but rather held the limbs relatively erect when walking, like dinosaurs. While footprints are yet to be known, it is likely that ornithocheiromorphs also walked erect.[60] Compared to other earlier pterosaurs such as rhamphorhynchids, ornithocheiromorphs had unusually uneven limb proportions, with the forelimbs resulting in a much longer scale compared to the hind limbs. Their close relatives, the pteranodontids, were also found with similar features, though they more likely flew like modern-day albatrosses rather than anything else. Paleontologists also suggest that they most likely spend long stretches of time sea fishing, traveling very long distances without flapping while at the same time flying close the surface of the water with exploited wind speed, and without the necessity of thermals.[61] This would likely have required them to use unique modes of locomotion compared to other pterosaurs, this can already be seen in earlier evolutions such as Istiodactylus and Nurhachius, with powerful musculature attachments and well-developed pectoral and upper arm bones.[29][62] It is also possible that ornithocheiromorphs ran (but not walked) bipedally, or that they used a hopping gait.[60] Many pterosaur researchers like Mike Habib have noted that the limb proportions of some ornithocheiromorphs such as Anhanguera are consistent with hopping, though the scavenging istiodactylids are probably still the best examples of pterosaurs with a more terrestrial setting.[63][16]
Paleoecology
Even though the ornithocheiromorphs were discovered worldwide, most of them were concentrated in specific places. One of which is the fossil site called Romualdo Formation, which contains an impressive amount of pterosaur fossils. It is a diverse Lagerstätte in the larger geologic group called the Santana Group (sometimes called the Santana Formation), which is located in the Araripe Basin of northeastern Brazil, and dates back around 111 and 108 million years ago, during the Albian stage of the Early Cretaceous. The formation includes many species of Anhanguera,[34] several fossil remains of basal ornithocheiromorphs, including Brasileodactylus, Cearadactylus and Unwindia, as well as the anhanguerids Tropeognathus,[20] Coloborhynchus, Maaradactylus, and Araripesaurus.[25] These pterosaur genera were just some of the many recovered from the site, which also include the thalassodromines Tupuxuara and Thalassodromeus,[64] as well as the tapejarid Tapejara.[65] Some other creatures from the formation include the theropods Irritator, Mirischia and Santanaraptor, and the crocodylomorph Araripesuchus. The formation also includes several turtle remains, with some specimens referring to Santanachelys, Cearachelys and Araripemys.[66] A few fish remains were also found within the Romualdo Formation, some of which were referred to Brannerion, Rhinobatos, Rhacolepis, Tharrhias and Tribodus. The Santana Group also consists of another Lagerstätte called the Crato Formation, which is not as diverse as the Romualdo Formation, but its fossil remains are still considered important.[67] This fossil site underlies the Romualdo Formation, and dates back around 115 and 113 million years ago during the Aptian stage, meaning that its fossil content is of older age. Similarly, the Crato Formation also contained several species of pterosaurs, including the basal lanceodontians Ludodactylus[38] and Brasileodactylus, as well as the ornithocheirid Arthurdactylus. Other pterosaur genera include the tapejarids Tupandactylus and Aymberedactylus,[65] as well as the chaoyangopterid Lacusovagus. The formation also contains other creatures such as the enantiornithine Cratoavis, the neosuchian Susisuchus, and several species of fish, including Belonostomus, Calamopleurus,[68] Cladocyclus,[69] Dastilbe[70] and Lepidotes.[71] These fish genera were suggested to be prey for the pterosaurs that lived in the formation, but fossil remains are limited, so the subject is still controversial.[36]
Another important fossil site is the Wessex Formation in the Isle of Wight, near the coast of England, which dates back around 140 and 125 million years ago (Berriasian to Barremian stages). The formation doesn't contain many fossil remains of pterosaurs compared to the Romualdo Formation, but it is still a very important site. Fossil remains of the istiodactylid Istiodactylus,[29] the anhanguerid Caulkicephalus,[40] and the tapejarid Wightia were found.[72] The formation is also known for several theropods, including the spinosaurid Baryonyx, the tyrannosauroid Eotyrannus, the dromaeosaurid Ornithodesmus, the compsognathid Aristosuchus as well as the allosauroid Neovenator.[73][74] Different types of herbivorous dinosaurs like Iguanodon, Polacanthus, Ornithopsis, Mantellisaurus and Hypsilophodon were also found within the fossil site.[75][76] Some other animals from the formation include the neosuchian Bernissartia, sea turtles such as Helochelydra and Brodiechelys, the cartilaginous fish Hybodus, ray-finned fishes such as Belonostomus, Caturus, Lepidotes and Scheenstia, as well as the mammals Eobaatar, Loxaulax and Yaverlestes.[71][77][78]
A few fossils reported from the Toolebuc Formation and Winton Formation are believed to be from some of the most derived ornithocheiromorphs, due to the age of the fossil remains, which dated back to the Albian and Cenomanian stages of the Cretaceous, and some are even believed to belong to the Turonian stage. The Toolebuc Formation includes several remains of ornithocheiromorphs which are now referred to the genera Aussiedraco and Mythunga.[79][80] The formation also includes several herbivorous dinosaurs such as the ornithopod Muttaburrasaurus and the ankylosaur Kunbarrasaurus.[81][82] Fossil remains of marine animals were also uncovered within the fossil site, and some specimens of which belong to the ichthyosaur Platypterygius, the pliosaurid Kronosaurus and the elasmosaurid Eromangasaurus.[83] Turtle remains from turtles that were proposed to be prey for pterosaurs were also found within the Toolebuc Formation, this included the genera Bouliachelys, Cratochelone and Notochelone.[36] The Winton Formation consisted on a more terrestrial environment, containing several sauropod dinosaurs like Austrosaurus, Diamantinasaurus, Savannasaurus and Wintonotitan as well as large carnivorous dinosaurs such as Australovenator[84] and crocodylomorphs like Isisfordia. The formation's only pterosaur is the derived genus Ferrodraco,[85] which is also considered as one of the last ornithocheiromorphs, and a close relative of Mythunga.[80]
See also
References
- ^ Jiang, Shun-Xing; Zhang, Xin-Jun; Cheng, Xin; Wang, Xiao-Lin (2020). "A new pteranodontoid pterosaur forelimb from the upper Yixian Formation, with a revision of Yixianopterus jingangshanensis". Vertebrata PalAsiatica. doi:10.19615/j.cnki.1000-3118.201124.
- ^ Barrett, P. M., Butler, R. J., Edwards, N. P., & Milner, A. R. (2008). Pterosaur distribution in time and space: an atlas. Zitteliana: 61-107.[1]
- ^ a b Andres, B.; Clark, J.; Xu, X. (2014). "The Earliest Pterodactyloid and the Origin of the Group". Current Biology. 24 (9): 1011–6. doi:10.1016/j.cub.2014.03.030. PMID 24768054.
- ^ a b Witton, M.P.; Martill, D.M.; Loveridge, R.F. (2010). "Clipping the Wings of Giant Pterosaurs: Comments on Wingspan Estimations and Diversity". Acta Geoscientica Sinica. 31: 79–81.
- ^ Bowerbank, J.S. (1846). "On a new species of pterodactyl found in the Upper Chalk of Kent (Pterodactylus giganteus)". Quarterly Journal of the Geological Society of London. 2 (1–2): 7–9. doi:10.1144/gsl.jgs.1846.002.01-02.05. S2CID 129389179.
- ^ Bowerbank, J.S. (1851). "On the pterodactyles of the Chalk Formation". Proceedings of the Zoological Society of London. 19: 14–20. doi:10.1111/j.1096-3642.1851.tb01125.x.
- ^ Owen, R. (1851). Monograph on the fossil Reptilia of the Cretaceous Formations. The Palaeontographical Society 5(11):1-118.
- ^ a b Seeley, H.G. (1870). "The Ornithosauria: an Elementary Study of the Bones of Pterodactyles". Cambridge: 113.
- ^ Hooley, Reginald Walter (1914). "On the Ornithosaurian genus Ornithocheirus, with a review of the specimens from the Cambridge Greensand in the Sedgwick Museum, Cambridge". Annals and Magazine of Natural History. 13 (78): 529–557. doi:10.1080/00222931408693521. ISSN 0374-5481.
- ^ Martill, D.M. (2010). "The early history of pterosaur discovery in Great Britain". Geological Society of London, Special Publications. 343 (1): 287–311. Bibcode:2010GSLSP.343..287M. doi:10.1144/SP343.18. S2CID 130116778.
- ^ Seeley, H.G., 1869, Index to the fossil remains of Aves, Ornithosauria, and Reptilia, from the Secondary System of Strata, arranged in the Woodwardian Museum of the University of Cambridge. St. John's College, Cambridge 8: 143. doi:10.1080/00222937008696143
- ^ Owen, R. 1874, Monograph on the fossil Reptilia of the Mesozoic Formations. Palaeontographical Society, London, 14 pp
- ^ a b Rodrigues, T.; Kellner, A. (2013). "Taxonomic review of the Ornithocheirus complex (Pterosauria) from the Cretaceous of England". ZooKeys (308): 1–112. doi:10.3897/zookeys.308.5559. PMC 3689139. PMID 23794925.
- ^ Seeley, H. G. (1887). "On a sacrum apparently indicating a new type of bird, Ornithodesmus cluniculus Seeley from the Wealden of Brook". Quarterly Journal of the Geological Society of London. 43 (1–4): 206–211. doi:10.1144/GSL.JGS.1887.043.01-04.19. S2CID 129459937.
- ^ Seeley, H. G. (2015) [1901]. Dragons of the Air: an Account of Extinct Flying Reptiles. New York: D. Appleton & Co. pp. 173–175. ISBN 978-1-4400-8494-2.
- ^ a b c Hooley, R. W. (1913). "On the skeleton of Ornithodesmus latidens; an ornithosaur from the Wealden Shales of Atherfield (Isle of Wight)". Quarterly Journal of the Geological Society. 69 (1–4): 372–422. doi:10.1144/GSL.JGS.1913.069.01-04.23. S2CID 128604856.
- ^ a b Witton 2013, pp. 143–151.
- ^ Howse, S. C. B.; Milner, A. R. (1993). "Ornithodesmus – a maniraptoran theropod dinosaur from the Lower Cretaceous of the Isle of Wight, England". Palaeontology. 36: 425–437.
- ^ Howse, S. C. B.; Milner, A. R.; Martill, D. M. (2001). "Pterosaurs". In Martill, D. M.; Naish, D. (eds.). Dinosaurs of the Isle of Wight. Guide 10; Field Guides to Fossils. London: The Palaeontological Association. pp. 324–335. ISBN 978-0-901702-72-2.
- ^ a b Peter Wellnhofer (1987). "New crested pterosaurs from the Lower Cretaceous of Brazil" (PDF). Mitteilungen der Bayerischen Staatssammlung für Paläontologie und historische Geologie. 27: 175–186.
- ^ Campos, D. de A., and Kellner, A. W. (1985). "Um novo exemplar de Anhanguera blittersdorffi (Reptilia, Pterosauria) da formação Santana, Cretaceo Inferior do Nordeste do Brasil." In Congresso Brasileiro de Paleontologia, Rio de Janeiro, Resumos, p. 13.
- ^ Wellnhofer, P. (1987). The Illustrated Encyclopedia of Pterosaurs. New York: Barnes and Noble Books. pp. 124. ISBN 0-7607-0154-7.
- ^ Kellner, A.W.A. (1989). "A new Edentate Pterosaur of the lower Cretaceous from the Araripe Basin, Northeast Brazil". Anais da Academia Brasileira de Ciências. 61: 439–446. S2CID 89420181.
- ^ Veldmeijer, A.J. (1998). "Pterosaurs from the Lower Cretaceous of Brazil in the Stuttgart Collection". Geoscience and Engineering. 327: 1–27.
- ^ a b Fastnacht, M (2001). "First record of Coloborhynchus (Pterosauria) from the Santana Formation (Lower Cretaceous) of the Chapada do Araripe of Brazil". Paläontologische Zeitschrift. 75: 23–36. doi:10.1007/bf03022595. S2CID 128410270.
- ^ Unwin, D.M., 2001, "An overview of the pterosaur assemblage from the Cambridge Greensand (Cretaceous) of Eastern England", Mitteilungen aus dem Museum für Naturkunde in Berlin, Geowissenschaftliche Reihe 4: 189–221
- ^ a b Unwin, D. M. (2003). "On the phylogeny and evolutionary history of pterosaurs". Geological Society, London, Special Publications. 217 (1): 139–190. Bibcode:2003GSLSP.217..139U. CiteSeerX 10.1.1.924.5957. doi:10.1144/GSL.SP.2003.217.01.11. S2CID 86710955.
- ^ Xiaolin Wang; Alexander W.A. Kellner; Shunxing Jiang; Qiang Wang; Yingxia Ma; Yahefujiang Paidoula; Xin Cheng; Taissa Rodrigues; Xi Meng; Jialiang Zhang; Ning Li; Zhonghe Zhou (2014). "Sexually dimorphic tridimensionally preserved pterosaurs and their eggs from China". Current Biology. 24 (12): 1323–1330. doi:10.1016/j.cub.2014.04.054. PMID 24909325.
- ^ a b c d Witton, M. P. (2012). "New Insights into the Skull of Istiodactylus latidens (Ornithocheiroidea, Pterodactyloidea)". PLOS ONE. 7 (3): e33170. Bibcode:2012PLoSO...733170W. doi:10.1371/journal.pone.0033170. PMC 3310040. PMID 22470442.
- ^ Kellner, A. W. A.; Campos, D. A.; Sayão, J. M.; Saraiva, A. N. A. F.; Rodrigues, T.; Oliveira, G.; Cruz, L. A.; Costa, F. R.; Silva, H. P.; Ferreira, J. S. (2013). "The largest flying reptile from Gondwana: A new specimen of Tropeognathus cf. T. Mesembrinus Wellnhofer, 1987 (Pterodactyloidea, Anhangueridae) and other large pterosaurs from the Romualdo Formation, Lower Cretaceous, Brazil". Anais da Academia Brasileira de Ciências. 85 (1): 113–135. doi:10.1590/S0001-37652013000100009. PMID 23538956.
- ^ Martill, D.M. and Unwin, D.M. (2011). "The world's largest toothed pterosaur, NHMUK R481, an incomplete rostrum of Coloborhynchus capito (Seeley 1870) from the Cambridge Greensand of England." Cretaceous Research, (advance online publication). doi:10.1016/j.cretres.2011.09.003
- ^ a b Holgado, B.; Pêgas, R.V. (2020). "A taxonomic and phylogenetic review of the anhanguerid pterosaur group Coloborhynchinae and the new clade Tropeognathinae". Acta Palaeontologica Polonica. 65. doi:10.4202/app.00751.2020.
- ^ Veldmeijer, A.J. (2006). "Toothed pterosaurs from the Santana Formation (Cretaceous; Aptian-Albian) of northeastern Brazil. A reappraisal on the basis of newly described material Archived 2012-03-17 at the Wayback Machine." Tekst. - Proefschrift Universiteit Utrecht.
- ^ a b c d e Kellner, A.W.A. and Tomida, Y. (2000). "Description of a new species of Anhanguera (Pterodactyloidea) with comments on the pterosaur fauna from the Santana Formation (Aptian–Albian), northeastern Brazil." Tokyo, National Science Museum (National Science Museum Monographs, 17).
- ^ Leonardi, G. & Borgomanero, G. (1985). "Cearadactylus atrox nov. gen., nov. sp.: novo Pterosauria (Pterodactyloidea) da Chapada do Araripe, Ceara, Brasil." Resumos dos communicaçoes VIII Congresso bras. de Paleontologia e Stratigrafia, 27: 75–80.
- ^ a b c d e f "Abstract: DIET OF ORNITHOCHEIROID PTEROSAURS INFERRED FROM STABLE CARBON ISOTOPE ANALYSIS OF TOOTH ENAMEL (GSA Annual Meeting in Seattle, Washington, USA - 2017)". gsa.confex.com.
- ^ a b Xiaolin Wang; Alexander W. A. Kellner; Shunxing Jiang; Xin Cheng (2012). "New toothed flying reptile from Asia: close similarities between early Cretaceous pterosaur faunas from China and Brazil". Naturwissenschaften. 99 (4): 249–57. Bibcode:2012NW.....99..249W. doi:10.1007/s00114-012-0889-1. PMID 22354475. S2CID 7323552.
- ^ a b Frey, E., Martill, D., and Buchy, M. (2003). A new crested ornithocheirid from the Lower Cretaceous of northeastern Brazil and the unusual death of an unusual pterosaur. In: Buffetaut, E., and Mazin, J.-M. (eds.). Evolution and Palaeobiology of Pterosaurs. Geological Society Special Publication 217:56-63. ISBN 1-86239-143-2.
- ^ a b c d Rodrigo V. Pêgas, Borja Holgado & Maria Eduarda C. Leal (2019) On Targaryendraco wiedenrothi gen. nov. (Pterodactyloidea, Pteranodontoidea, Lanceodontia) and recognition of a new cosmopolitan lineage of Cretaceous toothed pterodactyloids, Historical Biology, doi:10.1080/08912963.2019.1690482
- ^ a b c Steel, L., Martill, D.M., Unwin, D.M. and Winch, J. D. (2005). "A new pterodactyloid pterosaur from the Wessex Formation (Lower Cretaceous) of the Isle of Wight, England". Cretaceous Research. 26 (4): 686–698. doi:10.1016/j.cretres.2005.03.005.
{{cite journal}}
: CS1 maint: multiple names: authors list (link) - ^ Buchmann, R.; dos Santos Avilla, L.; Rodrigues, T. (October 25, 2019). "Comparative analysis of the vertebral pneumatization in pterosaurs (Reptilia: Pterosauria) and extant birds (Avialae: Neornithes)". PLoS One. 14 (10): e0224165. Bibcode:2019PLoSO..1424165B. doi:10.1371/journal.pone.0224165. PMC 6814219. PMID 31652295. S2CID 204909043.
- ^ Wellnhofer 1991, p. 54.
- ^ Witton 2013, pp. 158.
- ^ Lü, J. (2010). "A new boreopterid pterodactyloid pterosaur from the Early Cretaceous Yixian Formation of Liaoning Province, northeastern China". Acta Geologica Sinica. 24 (2): 241–246. doi:10.1111/j.1755-6724.2010.00204.x. S2CID 140600398.
- ^ a b Wellnhofer 1991, p. 55.
- ^ Witton 2013, p. 46.
- ^ Witton 2013, p. 35.
- ^ Bennett, S. C. (1994). "Taxonomy and systematics of the Late Cretaceous pterosaur Pteranodon (Pterosauria, Pterodactyloidea)", Occasional Papers of the Museum of Natural History, University of Kansas, Lawrence, 169: 1-70
- ^ Kellner, A. W. A. (2003). "Pterosaur phylogeny and comments on the evolutionary history of the group". Geological Society, London, Special Publications. 217 (1): 105–137. Bibcode:2003GSLSP.217..105K. doi:10.1144/GSL.SP.2003.217.01.10. S2CID 128892642.
- ^ a b Longrich, N.R., Martill, D.M., and Andres, B. (2018). "Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary." PLoS Biology, 16(3): e2001663. doi:10.1371/journal.pbio.2001663
- ^ Andres, B. (2021) Phylogenetic systematics of Quetzalcoatlus Lawson 1975 (Pterodactyloidea: Azhdarchoidea). Journal of Vertebrate Paleontology, 41:sup1, 203-217. DOI: 10.1080/02724634.2020.1801703 https://www.tandfonline.com/doi/full/10.1080/02724634.2020.1801703
- ^ a b Borja Holgado, Rodrigo V. Pêgas, José Ignacio Canudo, Josep Fortuny, Taissa Rodrigues, Julio Company & Alexander W.A. Kellner, 2019, "On a new crested pterodactyloid from the Early Cretaceous of the Iberian Peninsula and the radiation of the clade Anhangueria", Scientific Reports 9: 4940. doi:10.1038/s41598-019-41280-4
- ^ Jacobs, M.L.; Martill, D.M.; Ibrahim, N.; Longrich, N. (2019). "A new species of Coloborhynchus (Pterosauria, Ornithocheiridae) from the mid-Cretaceous of North Africa" (PDF). Cretaceous Research. 95: 77–88. doi:10.1016/j.cretres.2018.10.018. S2CID 134439172.
- ^ a b Kellner, Alexander W. A.; Caldwell, Michael W.; Holgado, Borja; Vecchia, Fabio M. Dalla; Nohra, Roy; Sayão, Juliana M.; Currie, Philip J. (2019). "First complete pterosaur from the Afro-Arabian continent: insight into pterodactyloid diversity". Scientific Reports. 9(1). doi:10.1038/s41598-019-54042-z.
- ^ a b David W. E. Hone; Adam J. Fitch; Feimin Ma; Xing Xu (2020). "An unusual new genus of istiodactylid pterosaur from China based on a near complete specimen". Palaeontologia Electronica. 23 (1): Article number 23(1):a09. doi:10.26879/1015.
- ^ Wellnhofer, P. (1991). The Illustrated Encyclopedia of Pterosaurs. New York: Crescent Books. pp. 114–116. ISBN 978-0-517-03701-0.
- ^ "The Hyoid Apparatus of Liaoxipterus brachycephalus (Pterosauria) and Its Implications for Food-catching Behavior". doi:10.3975/cagsb.2015.03.13.
{{cite journal}}
: Cite journal requires|journal=
(help) - ^ Jiang S, Li Z, Cheng X, Wang X. (2020). "The first pterosaur basihyal, shedding light on the evolution and function of pterosaur hyoid apparatuses". PeerJ. 8: 292. doi:10.7717/peerj.8292. PMC 6951291. PMID 31934505.
{{cite journal}}
: CS1 maint: multiple names: authors list (link) - ^ Jacobs, Megan L.; Martill, David M.; Unwin, David M.; Ibrahim, Nizar; Zouhri, Samir; Longrich, Nicholas R. (2020). "New toothed pterosaurs (Pterosauria: Ornithocheiridae) from the middle Cretaceous Kem Kem beds of Morocco and implications for pterosaur palaeobiogeography and diversity". Cretaceous Research. 110: 104413. doi:10.1016/j.cretres.2020.104413. ISSN 0195-6671. S2CID 214542129.
- ^ a b Witton, M.P.; Habib, M.B. (2010). "On the Size and Flight Diversity of Giant Pterosaurs, the Use of Birds as Pterosaur Analogues and Comments on Pterosaur Flightlessness". PLOS ONE. 5 (11): e13982. Bibcode:2010PLoSO...513982W. doi:10.1371/journal.pone.0013982. PMC 2981443. PMID 21085624.
- ^ Padian, K. (1983). "A functional analysis of flying and walking in pterosaurs". Paleobiology. 9 (3): 218–239. doi:10.1017/S009483730000765X. S2CID 88434056.
- ^ Zhou X., Pêgas R.V., Leal M.E.C. & Bonde N. 2019. Nurhachius luei, a new istiodactylid pterosaur (Pterosauria, Pterodactyloidea) from the Early Cretaceous Jiufotang Formation of Chaoyang City, Liaoning Province (China) and comments on the Istiodactylidae". PeerJ 7:e7688. {{DOI: 10.7717/peerj.7688}}
- ^ Habib, M. (2011). "Dinosaur Revolution: Anhanguera." H2VP: Paleobiomechanics. Weblog entry, September 20, 2011. Accessed September 28, 2011: http://h2vp.blogspot.com/2011/09/dinosaur-revolution-anhanguera.html
- ^ Pêgas, R. V.; Costa, F. R.; Kellner, A. W. A. (2018). "New Information on the osteology and a taxonomic revision of The genus Thalassodromeus (Pterodactyloidea, Tapejaridae, Thalassodrominae)". Journal of Vertebrate Paleontology. 38 (2): e1443273. doi:10.1080/02724634.2018.1443273. S2CID 90477315.
- ^ a b Kellner, A.W.A.; Campos, D.A. (2007). "Short note on the ingroup relationships of the Tapejaridae (Pterosauria, Pterodactyloidea". Boletim do Museu Nacional. 75: 1–14.
- ^ Zug, George R. "Turtle: Origin and evolution". Encyclopædia Britannica. Retrieved September 18, 2015.
- ^ Martill, D.M., Bechly, G. and Loveridge, R.F. (2007). The Crato fossil beds of Brazil: window into an ancient world. Cambridge University Press. ISBN 0-521-85867-4, ISBN 978-0-521-85867-0
- ^ Peter L. Forey & Lance Grande (1998). "An African twin to the Brazilian Calamopleurus (Actinopterygii: Amiidae)". Zoological Journal of the Linnean Society. 123 (2): 179–195. doi:10.1111/j.1096-3642.1998.tb01299.x.
- ^ Cladocyclus at Fossilworks.org
- ^ Sepkoski, Jack (2002). "A compendium of fossil marine animal genera". Bulletins of American Paleontology. 364: 560. Archived from the original on July 23, 2011. Retrieved February 27, 2009.
- ^ a b Woodward, A.S. (1895). "Catalogue of the Fossil Fishes in the British Museum (Natural History)". 2. British Museum of Natural History Department of Geology: 77–119. doi:10.5962/bhl.title.61854. hdl:2027/coo1.ark:/13960/t3jx39x2t.
{{cite journal}}
: Cite journal requires|journal=
(help) - ^ Martill, David M.; Green, Mick; Smith, Roy; Jacobs, Megan; Winch, John (April 2020). "First tapejarid pterosaur from the Wessex Formation (Wealden Group: Lower Cretaceous, Barremian) of the United Kingdom". Cretaceous Research. 113: 104487. doi:10.1016/j.cretres.2020.104487. S2CID 219099220.
- ^ Charig, A. J.; Milner, A. C. (1986). "Baryonyx, a remarkable new theropod dinosaur". Nature. 324 (6095): 359–361. Bibcode:1986Natur.324..359C. doi:10.1038/324359a0. PMID 3785404. S2CID 4343514.
- ^ Hutt, S.; Simmonds, K.; Hullman, G. (1990). "Predatory dinosaurs from the Isle of Wight". Proceedings of the Isle of Wight Natural History and Archaeological Society. 9: 137–146.
- ^ Carpenter, K.; Ishida, Y. (2010). "Early and "Middle" Cretaceous Iguanodonts in Time and Space". Journal of Iberian Geology. 36 (2): 145–164. doi:10.5209/rev_JIGE.2010.v36.n2.3.
- ^ Hulke, J. W. (1873). "Contribution to the Anatomy of Hypsilophodon Foxii". Quarterly Journal of the Geological Society of London. Vol. 29. pp. 522–532. doi:10.1144/GSL.JGS.1873.029.01-02.46 – via Wikisource.
- ^ Sweetman, Steven C. (September 2009). "A New Species of the Plagiaulacoid Multituberculate Mammal Eobaatarfrom the Early Cretaceous of Southern Britain". Acta Palaeontologica Polonica. 54 (3): 373–384. doi:10.4202/app.2008.0003. ISSN 0567-7920. S2CID 53975359.
- ^ "Wessex Formation". The BGS Lexicon of Named Rock Units. British Geological Survey.
- ^ Kellner, Alexander W.A.; Taissa Rodrigues; Fabiana R. Costa (2011). "Short note on a pteranodontoid pterosaur (Pterodactyloidea) from western Queensland, Australia" (PDF). Anais da Academia Brasileira de Ciências. 83 (1): 301–308. doi:10.1590/S0001-37652011000100018. PMID 21437387.
- ^ a b Molnar, Ralph E.; Thulborn, R.A. (2008). "An incomplete pterosaur skull from the Cretaceous of north-central Queensland, Australia". Arquivos do Museu Nacional, Rio de Janeiro. 65 (4): 461–470.
- ^ McDonald, A.T.; Kirkland, J.I.; DeBlieux, D.D.; Madsen, S.K.; Cavin, J.; Milner, A.R.C.; Panzarin, L. (2010). "New Basal Iguanodonts from the Cedar Mountain Formation of Utah and the Evolution of Thumb-Spiked Dinosaurs". PLOS ONE. 5 (11): e14075. Bibcode:2010PLoSO...514075M. doi:10.1371/journal.pone.0014075. PMC 2989904. PMID 21124919.
- ^ Lucy G. Leahey; Ralph E. Molnar; Kenneth Carpenter; Lawrence M. Witmer; Steven W. Salisbury (2015). "Cranial osteology of the ankylosaurian dinosaur formerly known as Minmi sp. (Ornithischia: Thyreophora) from the Lower Cretaceous Allaru Mudstone of Richmond, Queensland, Australia". PeerJ. 3: e1475. doi:10.7717/peerj.1475. PMC 4675105. PMID 26664806.
- ^ Gardiner, J. Stanley (September 1931). "The Harvard Museum Expedition to Australia". Nature. 128 (3228): 457–458. Bibcode:1931Natur.128..457G. doi:10.1038/128457c0. S2CID 29715877.
- ^ White, M. A.; Falkingham, P. L.; Cook, A. G.; Hocknull, S. A.; Elliott, D. A. (2013). "Morphological comparisons of metacarpal I for Australovenator wintonensis and Rapator ornitholestoides: Implications for their taxonomic relationships". Alcheringa: An Australasian Journal of Palaeontology. 37 (4): 435–441. doi:10.1080/03115518.2013.770221. S2CID 82672110.
- ^ Pentland, Adele H.; Poropat, Stephen F.; Tischler, Travis R.; Sloan, Trish; Elliott, Robert A.; Elliott, Harry A.; Elliott, Judy A.; Elliott, David A. (December 2019). "Ferrodraco lentoni gen. et sp. nov., a new ornithocheirid pterosaur from the Winton Formation (Cenomanian–lower Turonian) of Queensland, Australia". Scientific Reports. 9 (1): 13454. Bibcode:2019NatSR...913454P. doi:10.1038/s41598-019-49789-4. ISSN 2045-2322. PMC 6776501. PMID 31582757.
Further reading
- Wellnhofer, Peter (1991). The Illustrated Encyclopedia of Pterosaurs: An illustrated natural history of the flying reptiles of the Mesozoic Era. Crescent Books. ISBN 0-517-03701-7.
- Witton, Mark (2013). Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press. ISBN 978-0-691-15061-1.
External links
- Data related to Ornithocheiromorpha at Wikispecies