Ornithocheiromorpha

{{short description|Clade of pteranodontoid pterosaurs}} {{Good article}} {{Use mdy dates|date=March 2021}} {{Automatic taxobox | name = Ornithocheiromorphs | fossil_range = Valanginian-Turonian<br/>~{{fossil_range|140|92.5}} | image = Tropeognathus mesembrinus MN 01.jpg | image_caption = Reconstructed skeleton of ''Tropeognathus'' in the National Museum of Brazil | image_upright = 1.3 | image2 = Coloborhynchus spielbergi2.jpg | image2_caption = Skeletal cast of ''Anhanguera spielbergi'' in the Naturalis Biodiversity Center | image2_upright = 1.3 | taxon = Ornithocheiromorpha | authority = Andres et al., 2014 | subdivision_ranks = Subgroups | subdivision = * {{extinct}}''Serradraco'' * {{extinct}}''Unwindia'' * {{extinct}}'''Lonchodectidae''' <small>(relationships uncertain)</small> * {{extinct}}'''Lanceodontia''' <small>Andres et al., 2014</small> ** {{extinct}}''Draigwenia'' ** {{extinct}}''Linlongopterus'' ** {{extinct}}'''Istiodactyliformes''' <small>Kellner et al., 2019</small> *** {{extinct}}''Hongshanopterus'' *** {{extinct}}''Yixianopterus'' *** {{extinct}}'''Istiodactylidae''' *** {{extinct}}'''Mimodactylidae''' ** {{extinct}}'''Ornithocheiriformes''' <small>Kellner et al., 2019</small> *** {{extinct}}''Ornithocheirus'' *** {{extinct}}''Thapunngaka'' *** {{extinct}}'''Anhangueridae''' *** {{extinct}}'''Boreopteridae'''? *** {{extinct}}'''Hamipteridae''' *** {{extinct}}'''Targaryendraconia''' }} '''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 have been discovered worldwide except Antarctica, though most genera have been 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 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''.<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 | bibcode = 2014CBio...24.1011A}}</ref>

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 {{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 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 === [[File:Pterodactylus giganteus.jpg|thumb|upright|left|Associated and referred bones of ''Pterodactylus giganteus'', now considered as ''Lonchodraco giganteus'']] 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''.<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>

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|url-access=subscription}}</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> [[File:PZSL1851PlateReptilia04.png|thumb|Holotype of ''Pterodactylus cuvieri'', now known as ''Cimoliopterus'']] 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"/> [[File:Istiodactylus.tif|thumb|left|Skull fragments of ''Ornithodesmus latidens'', which is currently known as ''Istiodactylus latidens'']] 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|bibcode=1887QJGS...43..206S |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|bibcode=1913QJGS...69..372H |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}}

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''.<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>

=== Discoveries outside Europe === [[File:Ornithocheirus & Tropeognathus.jpg|thumb|left|Comparison between the holotypes of ''Tropeognathus mesembrinus'' and ''Ornithocheirus simus'' ]] Other important ornithocheiromorph discoveries include the anhanguerids ''Tropeognathus'' and ''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 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.<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, 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 | issue = 1 | pages = 23–36 | doi=10.1007/bf03022595| bibcode = 2001PalZ...75...23F | 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 | bibcode = 2013ZooK..308....1R }}</ref> [[File:Hamipterus-Paleozoological Museum of China.jpg|thumb|Skull of ''Hamipterus tianshanensis'']] 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 |bibcode=2014CBio...24.1323W }}</ref>

== Description == === Size === [[File:Istiodactylus size.jpg|thumb|Size of ''Istiodactylus latidens'' compared to a human]] 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|article-number=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"/>

=== Skull and crests === [[File:Anhangueria skulls.jpg|thumb|left|Skull comparison between different species of ornithocheiromorphs; notice their different structures]] 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>

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 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>

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|bibcode=2005CrRes..26..686S }}</ref>

Ornithocheiriform teeth were heterodont, and they have sufficient differences that individual teeth can be distinguished between whether they belonged to the rostral, medial, or the posterior portion of the tooth row. The rostral teeth had a subcircular cross-section and their crowns curved more lingually than distally. The medial teeth shared the curvature of the anterior teeth but were more elliptical in cross-section. The posterior teeth were oval in cross-section and curved more distally than lingually.<ref>{{Cite journal |last1=Ciaffi |first1=Alessio |last2=Bellardini |first2=Flavio |date=2024 |title=Pterosaur teeth from the Southern Neuquén Basin (Patagonia, Argentina): new insights on the reconstruction of ornithocheiriform dental anatomy |url=https://www.app.pan.pl/article/item/app011222023.html |journal=Acta Palaeontologica Polonica |volume=69 |doi=10.4202/app.01122.2023 |access-date=23 September 2025 |via=Acta Palaeontologica Polonica}}</ref>

=== Postcrania === [[File:Anhanguerid_AMNH.jpg|thumb|left|Skeleton of ''Anhanguera'' showing neck and torso vertebrae]] 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|article-number=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|bibcode=2010AcGlS..84..241L |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"/>

[[File:Anhanguera-santanae sacrum.jpg|thumb|A rotated pelvis of ''Anhanguera santanae'', showing its right side]] 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.{{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}}

== 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.<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>

===Subgroups=== [[File:Mimodactylus_skeletal.png|thumb|Skeletal reconstruction of ''Mimodactylus'', a mimodactylid istiodactyliform]] Brian Andres and colleagues found the families Istiodactylidae, Ornithocheiridae, and Anhangueridae to form a group in 2014, which he called '''Lanceodontia'''. The clade includes most families of ornithocheiromorphs, however they excluded the poorly-known family Lonchodectidae from membership in this clade. 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/>

Within Lanceodontia, there is a more restrictive clade called '''Ornithocheirae''', which has a more convoluted taxonomic history. It was named by Harry Seeley in 1870 as a family that contains ''Ornithocheirus'' and its relatives. The name was emended to Ornithocheiridae, to match the requirements of the ICZN Code that a family-ranked clade should end with an "-idae" suffix. Brian Andres (2010) in his review of pterosaur phylogeny, defined the name Ornithocheirae phylogenetically, as a node-based taxon consisting of the last common ancestor of ''Anhanguera'' and ''Ornithocheirus'' and all its descendants.<ref name=BBA10/> Thus Ornithocheirae is defined to include two families, the Anhangueridae and the Ornithocheiridae, following the opinion of Alexander Kellner<ref name=Kellner03>Kellner, A. W. A., (2003): Pterosaur phylogeny and comments on the evolutionary history of the group. pp. 105-137. &mdash; ''in'' Buffetaut, E. & Mazin, J.-M., (eds.): ''Evolution and Palaeobiology of Pterosaurs''. Geological Society of London, Special Publications 217, London, 1-347</ref> and Andres that these families should not be synonymized based on their original phylogenetical definitions.<ref name=BBA10>{{cite book |last=Andres |first=Brian Blake |year=2010 |title=Systematics of the Pterosauria |url=http://gradworks.umi.com/34/40/3440534.html |publisher=Yale University |page=366}} [https://www.proquest.com/docview/846734901 A preview that shows the cladogram without clade names]</ref> Subsequent studies in 2019 have found Ornithocheirae to be a more inclusive group containing both Anhangueria (see below) and Targaryendraconia. [[File:Targaryendraco.jpg|thumb|left|Mandible of ''Targaryendraco'', a targaryendraconian anhanguerian]] Another constituent clade of Ornithocheiromorpha is '''Anhangueria'''. Anhangueria was named by paleontologists Taissa Rodrigues and Alexander Kellner in a review of ''Ornithocheirus'' species in 2013, they defined the clade as a branch-based taxon consisting of all pteranodontoids more closely related to ''Anhanguera blittersdorffi'' than to ''Istiodactylus latidens'' and ''Cimoliopterus cuvieri''.<ref name="Rodrigues & Kellner 2013"/> Anhangueria originally only contained the genera ''Brasileodactylus'', ''Camposipterus'', ''Cearadactylus'', ''Ludodactylus'' as well as the family Anhangueridae,<ref name="Rodrigues & Kellner 2013"/> however, recent analyses had recovered the family Hamipteridae within this clade as well.<ref name="Holgado2019"/><ref name="Mimodactylus"/><ref name="Targaryendraco"/><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> In 2014, paleontologist Brian Andres and colleagues assigned more groups and genera within this clade, this included ''Guidraco'', the subfamily Boreopterinae, and the clade Ornithocheirae, which was further divided into the families Anhangueridae and Ornithocheiridae.<ref name=kryptodrakon/> This topology was later followed by a few other studies.<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|doi-access=free}}</ref><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>

In 2019, Alexander Kellner and colleagues described the new ornithocheiromorph ''Mimodactylus'', and used their analysis to name a new clade, '''Istiodactyliformes'''. They defined this clade as the most inclusive clade containing ''Istiodactylus'' but not ''Anhanguera''. Under their analysis, this new clade included the family Istiodactylidae, Mimodactylidae, and the genus ''Hongshanopterus''.<ref name=kellner2019 /> Andres and colleagues later named a clade '''Ornithocheiriformes''' and defined it in opposition to Istiodactyliformes. Its given definition was the most inclusive clade containing ''Anhanguera'', but not ''Istiodactylus''.<ref name="andres2021" />

Some of these clades have been recovered by various authors since their definition. A table summarizing all of these clades is shown below. {| class="wikitable sortable" align="center" width="100%" ! Name ! Named by ! Definition ! Notes |- | '''Anhangueria''' | Rodrigues & Kellner, 2013<ref name="Rodrigues & Kellner 2013" /> | Most-inclusive clade containing ''Anhanguera'', but not ''Istiodactylus'' or ''Cimoliopterus'' | May be a parent taxon of,<ref name=longrichetal2018 /> belong inside of,<ref name="holgado2020" /> or be synonymous with Ornithocheirae<ref name="andres2021" /> depending on the analysis |- | '''Anhangueroidea''' | Pêgas, 2025<ref name="Pegas2025">{{cite journal |doi=10.26879/20 |title=On the systematics and phylogenetic nomenclature of the Ornithocheiriformes (Pterosauria, Pteranodontoidea) |date=2025 |last1=Pêgas |first1=R.V. |journal=Palaeontologia Electronica }}</ref> | Apomorphy-based clade containing all pterosaurs with the reflected palatal tip synapomorphic with ''Anhanguera'' provided that this clade does not contain ''Ornithocheirus''<ref name="Pegas2025" /> | |- | '''Istiodactyliformes''' | Kellner et al., 2019<ref name=kellner2019>{{cite journal |doi=10.1038/s41598-019-54042-z |title=First complete pterosaur from the Afro-Arabian continent: Insight into pterodactyloid diversity |date=2019 |last1=Kellner |first1=Alexander W. A. |last2=Caldwell |first2=Michael W. |last3=Holgado |first3=Borja |last4=Vecchia |first4=Fabio M. Dalla |last5=Nohra |first5=Roy |last6=Sayão |first6=Juliana M. |last7=Currie |first7=Philip J. |journal=Scientific Reports |volume=9 |issue=1 |page=17875 |pmid=31784545 |pmc=6884559 |bibcode=2019NatSR...917875K }}</ref> | Most-inclusive clade containing ''Istiodactylus'', but not ''Anhanguera'' | May include Boreopteridae and/or Lonchodectidae; may be synonymous with Istiodactylidae, if all non-istiodactylid lanceodontians are ornithocheiriformes<ref>{{cite journal |doi=10.1080/02724634.2021.1946068 |title=A new species of crested pterosaur (Pterodactyloidea, Anhangueridae) from the Lower Cretaceous (Upper Albian) of Richmond, North West Queensland, Australia |date=2021 |last1=Richards |first1=Timothy M. |last2=Stumkat |first2=Paul E. |last3=Salisbury |first3=Steven W. |journal=Journal of Vertebrate Paleontology |volume=41 |issue=3 |bibcode=2021JVPal..41E6068R }}</ref> |- | '''Lanceodontia''' | Andres et al., 2014<ref name=kryptodrakon/> | Least-inclusive clade containing both ''Anhanguera'' and ''Istiodactylus'' | May be a synonym of Ornithocheiromorpha if it includes Boreopteridae and Lonchodectidae; may include both, neither, or one of the two families |- | '''Ornithocheirae''' | Seeley, 1870<ref name=HGS70 /> | Least-inclusive clade including both ''Anhanguera'' and ''Ornithocheirus'' | May be more or less exclusive than Anhangueria, depending on the position of ''Ornithocheirus'' |- | '''Ornithocheiriformes''' | Andres et al., 2021<ref name="andres2021" /> | Most-inclusive clade containing both ''Ornithocheirus'', but not ''Istiodactylus'' | May or may not include Boreopteridae |- |}

=== Relationships === [[File:Haliskia_Life_Restoration.png|thumb|Life reconstruction of ''Haliskia'', an anhanguerid]] There are competing theories of ornithocheiromorph phylogeny (evolutionary relationships). 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/>

{{clade| style=font-size:80%;line-height:80% |label1='''Ornithocheiromorpha''' |1={{clade |1=''Hongshanopterus'' |2={{clade |1={{clade |1={{clade |1='''Lonchodectidae''' |2='''Boreopteridae''' }} |label2='''Lanceodontia''' |2={{clade |1='''Istiodactylidae''' |label2='''Ornithocheiriformes''' |2={{clade |1='''Cimoliopteridae''' |label2='''Anhangueria''' |2={{clade |1={{clade |1=''Guidraco'' |2=''Ludodactylus'' }} |2={{clade |1={{clade |1=''Cearadactylus'' |2=''Brasileodactylus'' }} |label2='''Ornithocheirae''' |2={{clade |2='''Anhangueridae''' |1='''Ornithocheiridae''' }} }} }} }} }} }} }} }} }} thumb|left|Life reconstruction of ''Mimodactylus'', one of many new ornithocheiromorphs recognized in 2019 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}} </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|bibcode=2019CrRes..95...77J |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="Mimodactylus">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="Mimodactylus"/><ref name="holgado2020"/>

Later, in 2019, a different phylogenetic analysis, this time conducted by Borja Holgado and colleagues, focused on derived pterodactyloids with particular emphasis on anhanguerians. In this analysis, Anhangueria consisted of ''Camposipterus'' and the families Anhangueridae and Hamipteridae, the clade Ornithocheirae was recovered as a more inclusive group consisting of ''Ornithocheirus'' and ''Cimoliopterus'' as basal members, as well as the Anhangueria in the most derived position. This classification is supported by the expansion of the premaxillary tip (a spoon-like expansion at the end of the snout) with a high jaw end.<ref name="Holgado2019"/> Many analyses afterwards have followed this concept,<ref name="Mimodactylus"/><ref name="Jiang2020">{{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|url=http://www.ivpp.cas.cn/cbw/gjzdwxb/pressonline/202011/P020210203354269995268.pdf|doi=10.19615/j.cnki.1000-3118.201124}}</ref> although some had recovered ''Camposipterus'' outside the Anhangueria (within the clade Targaryendraconia), but still within the Ornithocheirae.<ref name="Targaryendraco"/><ref name="holgado2020"/>

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>

{{clade| style=font-size:80%;line-height:80% |label1='''Ornithocheiromorpha''' |1={{clade |1={{clade |1=''Serradraco'' |2=''Aussiedraco'' }} |2={{clade |label1= |1={{clade |1=''Hongshanopterus'' |2={{clade |1=''Targaryendraco'' |2=''Lonchodectes''}} }} |2={{clade |1='''Lonchodraconidae''' |label2='''Lanceodontia''' |2={{clade |label1='''Istiodactyliformes''' |1='''Istiodactylidae''' |label2='''Ornithocheiriformes''' |2={{clade |1='''Boreopteridae''' |2={{clade |1=''Hamipterus'' |2={{clade |1={{clade |1=''Brasileodactylus'' |2=''Barbosania'' |3=''Cearadactylus'' }} |2={{clade |label2='''Ornithocheirae''' / '''Anhangueria''' |2={{clade |1='''Ornithocheiridae''' |2='''Anhangueridae'''}} |1='''Cimoliopteridae'''}} }} }} }} }} }} }} }} }} [[File:Lingyuanopterus_Head_Recon.png|thumb|Life reconstruction of ''Lingyuanopterus'', an istiodactylid]] In their description of ''Lingyuanopterus'' in 2022, Xu Yizhi and colleagues recovered the following phylogeny based on the earlier work of Borja Holgado and Rodrigo Pêgas.

{{clade| style=font-size:80%;line-height:80% |label1='''Lanceodontia''' |1={{clade |label1='''Istiodactyliformes''' |1={{clade sequential |1='''Lonchodectidae''' |2='''Mimodactylidae''' |3=''Hongshanopterus'' |4='''Istiodactylidae''' }} |label2='''Ornithocheiriformes''' |2={{clade |1='''Boreopteridae''' |label2='''Ornithocheirae''' |2={{clade |1=''Ornithocheirus'' |2={{clade |1='''Targaryendraconia''' |label2='''Anhangueria''' |2={{clade |1='''Anhangueridae''' |2='''Hamipteridae''' }} }} }} }} }} }}

== Paleobiology == === Diet and feeding === [[File:Istiodactylus scavenging.tif|thumb|upright|left|A hypothetical recreation of a group of ''Istiodactylus'' feeding on a carcass of a stegosaur]] 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 beaks 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 herons, storks, and skimmers, 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/> [[File:OrnitocheiridsDB.jpg|thumb|''Cimoliopterus'' (right) stealing prey from a ''Lonchodectes'' (left), both were derived members of this group and possible fish hunters]] 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|article-number=e8292 |doi=10.7717/peerj.8292|pmid=31934505|pmc=6951291 |doi-access=free }}</ref>

Anhanguerids like ''Tropeognathus'' 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 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.<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|article-number=104413|doi=10.1016/j.cretres.2020.104413|bibcode=2020CrRes.11004413J |s2cid=214542129 |issn=0195-6671|url-access=subscription}}</ref>

In teeth microwear analyses, ''Coloborhynchus'' appears to have fed on terrestrial vertebrates and invertebrates.<ref>Bestwick, Jordan; Unwin, David M.; Butler, Richard J.; Purnell, Mark A. (28 October 2020). "Dietary diversity and evolution of the earliest flying vertebrates revealed by dental microwear texture analysis". Nature Communications. 11 (1): 5293. Bibcode:2020NatCo..11.5293B. doi:10.1038/s41467-020-19022-2. PMC 7595196. PMID 33116130.</ref>

=== Locomotion and flight === thumb|left|Restoration of a flying ''Istiodactylus''; notice its high aspect ratio 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.<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| article-number = 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 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.<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| bibcode = 1983Pbio....9..218P | 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/>

== Paleoecology == [[File:Araripe Basin map - formations and resources.jpg|thumb|400px|Geological map of the Araripe Basin, with the extent of the Santana Group shown in dark blue]] 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 thalassodromines ''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 |article-number=e1443273 |doi=10.1080/02724634.2018.1443273|bibcode=2018JVPal..38E3273P |s2cid=90477315 }}</ref> as well as the tapejarid ''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> [[File:20130825 Brazil Rio de Janeiro 0144.jpg|thumb|left|Mounted skeleton of ''Irritator'' with a possible anhanguerid in its jaws]] 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 neosuchian ''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/>

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'' 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|article-number=104487|doi=10.1016/j.cretres.2020.104487|bibcode=2020CrRes.11304487M |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 fishes such as ''Belonostomus'', ''Caturus'', ''Lepidotes'' and ''Scheenstia'', as well as the mammals ''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> [[File:Ferrodraco.png|thumb|Reconstruction of the skeleton of ''Ferrodraco'' on a diagram showing the known material, and based on the related ''Tropeognathus mesembrinus'']] A few fossils reported from the Toolebuc Formation and Winton Formation are believed to be from some of the last 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| article-number = 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 |article-number=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 | bibcode = 2013Alch...37..435W | s2cid = 82672110 | url = http://researchonline.rvc.ac.uk/id/eprint/8122/ }}</ref> and crocodylomorphs like ''Isisfordia''. The formation's only pterosaur is the derived genus ''Ferrodraco'',<ref name="Pentland2019" /> which is also considered as one of the last ornithocheiromorphs, and a close relative of ''Mythunga''.<ref name=MT08/>

== See also == * Pterosaur size

== References == {{Reflist}}

== Further reading == *{{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}} *{{cite book|last=Witton|first=Mark|title=Pterosaurs: Natural History, Evolution, Anatomy|publisher=Princeton University Press|year=2013|isbn=978-0-691-15061-1}}

== External links == * {{Wikispecies-inline}}

{{Pterosauria|P.}} {{Taxonbar|from=Q21008934}}

Category:Pteranodontoidea Category:Reptile clades Category:Cretaceous reptiles