Pterodactyloidea

{{short description|Suborder of monofenestratan pterosaurs}} {{Automatic taxobox | name = Pterodactyloids | fossil_range = <br />Late Jurassic – Late Cretaceous, {{Fossil range|152|earliest=Bathonian|66}} <small>Possible Middle Jurassic - Bathonian record</small><ref name=martill2022>{{cite journal | url=https://www.sciencedirect.com/science/article/pii/S0016787822000402 | title=Did William Smith (1769–1839), the father of biostratigraphy, discover a Jurassic pterosaur tooth? | journal=Proceedings of the Geologists' Association | last1=Martill | first1=David M. | last2=Steel | first2=Lorna | last3=Smith | first3=Roy E. | year=2022 | volume=133 | issue=3 | pages=250–255 | doi=10.1016/j.pgeola.2022.05.001 | bibcode=2022PrGA..133..250M }}</ref><br /> | image = Pterodactylodiea Infobox Panoply.png | image_caption = Several members of Pterodactyloidea (top to bottom): ''Pteranodon'', ''Pterodaustro'', the skulls of several pterosaurs (''Guidraco'', ''Anhanguera'', ''Tupandactylus'', and an unnamed dsungaripterid), ''Quetzalcoatlus'', ''Aerodactylus'', and ''Maaradactylus'' | image_upright = 1.3 | taxon = Pterodactyloidea | authority = Plieninger, 1901 | subdivision_ranks = Subgroups | subdivision = * {{extinct}}''Diopecephalus'' * {{extinct}}''Kryptodrakon''? * {{extinct}}''Ningchengopterus'' <small>(''incertae sedis'')</small> * {{extinct}}''Pangupterus'' <small>(''incertae sedis'')</small> * {{extinct}}''Wenupteryx'' <small>(''incertae sedis'')</small> * {{extinct}}'''Archaeopterodactyloidea''' * {{extinct}}'''Eupterodactyloidea''' <small>Bennett, 1994</small> ** {{extinct}}''Herbstosaurus'' ** {{extinct}}'''Azhdarchoidea''' ** {{extinct}}'''Dsungaripteridae''' **{{extinct}}'''Germanodactylidae'''? ** {{extinct}}'''Pteranodontoidea''' | synonyms = *Dracochira <small>Haeckel, 1895</small> }}

'''Pterodactyloidea''' ({{IPA|/ˌtɛrəˈdæktl̩ɔɪdɪːə/|cat=no}}; derived from the Greek words ''πτερόν'' (''pterón'', for usual ''ptéryx'') "wing", and ''δάκτυλος'' (''dáktylos'') "finger")<ref>{{cite book |last1=Colbert |first1=Edwin H. (Edwin Harris) |last2=Knight |first2=Charles Robert |title=The dinosaur book: the ruling reptiles and their relatives |date=1951 |publisher=McGraw-Hill |location=New York |page=153 |url=https://archive.org/details/bookruli00colb/page/152/mode/2up}}</ref> is one of the two traditional suborders of pterosaurs ("wing lizards"), and contains the most derived members of this group of flying reptiles. They appeared during the middle Jurassic Period, and differ from the basal (though paraphyletic) rhamphorhynchoids by their short tails and long wing metacarpals (hand bones). The most advanced forms also lack teeth, and by the late Cretaceous, all known pterodactyloids were toothless.<ref>[https://zookeys.pensoft.net/article/3978/element/7/azhdarchidae/ Review of taxonomy, geographic distribution, and paleoenvironments of Azhdarchidae (Pterosauria) – ZooKeys]</ref> Many species had well-developed crests on the skull, a form of display taken to extremes in giant-crested forms like ''Nyctosaurus'' and ''Tupandactylus''. Pterodactyloids were the last surviving pterosaurs when the order became extinct at the end of the Cretaceous Period, together with the non-avian dinosaurs and most marine reptiles.

"Pterodactyl" is also a common term for pterodactyloid pterosaurs, though it can also be used to refer to ''Pterodactylus'' specifically. Well-known examples of pterodactyloids include ''Pterodactylus'', ''Pteranodon'', and ''Quetzalcoatlus''.

In 2014, fossils from the Shishugou Formation of China were classified as the most basal pterodactyloid yet found, ''Kryptodrakon''. At a minimum age of about 161 my, it is about 5 million years older than the oldest previously known confirmed specimens.<ref name=kryptodrakon/> Some later studies have found ''Kryptodrakon'' to be a non-pterodactyloid.<ref name=martinsilverstone2024>{{Cite journal |last1=Martin-Silverstone |first1=Elizabeth |last2=Unwin |first2=David M. |last3=Cuff |first3=Andrew R. |last4=Brown |first4=Emily E. |last5=Allington-Jones |first5=Lu |last6=Barrett |first6=Paul M. |date=2024-02-05 |title=A new pterosaur from the Middle Jurassic of Skye, Scotland and the early diversification of flying reptiles |journal=Journal of Vertebrate Paleontology |volume=43 |issue=4 |language=en |doi=10.1080/02724634.2023.2298741 |issn=0272-4634|doi-access=free |article-number=e2298741 }}</ref> Previously, a fossil jaw recovered from the Middle Jurassic (Bathonian) Stonesfield Slate formation was considered the oldest known, but further examination suggested it belonged to a teleosaurid instead of a pterosaur.<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| bibcode = 2014CBio...24.1011A| doi-access = free }}</ref><ref>{{cite journal | last1 = Buffetaut | first1 = E. | last2 = Jeffrey | first2 = P. | year = 2012 | title = A ctenochasmatid pterosaur from the Stonesfield Slate (Bathonian, Middle Jurassic) of Oxfordshire, England | url = | journal = Geological Magazine | volume = 149| issue = 3| pages = 552–556| doi = 10.1017/S0016756811001154 | bibcode = 2012GeoM..149..552B }}</ref> In 2018 Michael O'Sullivan and David Martill described a partial synsacrum from the Stonesfield Slate identified as possibly pterodactyloid, though they noted it could also be a wukongopterid. If correctly identified, it would be the oldest pterodactyloid fossil known.<ref name=Martill18>{{cite journal | last1 = O'Sullivan | first1 = Michael | last2 = Martill | first2 = David M. | year = 2018 | title = Pterosauria of the Great Oolite Group (Bathonian, Middle Jurassic) of Oxfordshire and Gloucestershire, England | url = | journal = Acta Palaeontologica Polonica | volume = 63| issue = | page = | doi = 10.4202/app.00490.2018 }}</ref> In 2022 Martill and colleagues described a likely ctenochasmatid tooth, also from Stonesfield.<ref name=martill2022/>

==Classification== Pterodactyloidea is traditionally considered to be the group of short-tailed pterosaurs with long wrists (metacarpus), compared with the relatively long tails and short wrist bones of basal pterosaurs ("rhamphorhynchoids"). In 2004, Kevin Padian formally defined Pterodactyloidea as an apomorphy-based clade containing those species possessing a metacarpal at least 80% of the length of the humerus, homologous with that of ''Pterodactylus''. This definition was adopted by the PhyloCode in 2020.<ref>de Queiroz. K., Cantino. P. D., Gauthier. J. A. eds. (2020). ''Phylonyms: A Companion to the PhyloCode''. CRC Press Boca Raton, FL</ref>

===Subgroups=== [[File:Lusognathus.png|thumb|left|Reconstruction of ''Lusognathus'', a ctenochasmatoid]] A subgroup of pterodactyloids, called the '''Lophocratia''', was named by David Unwin in 2003. Unwin defined the group as the most recent common ancestor of ''Pterodaustro guinazui'' and ''Quetzalcoatlus northropi'', and all its descendants.<ref name="unwin2003">Unwin, D. M., (2003). "On the phylogeny and evolutionary history of pterosaurs." Pp. 139–190. in Buffetaut, E. & Mazin, J.-M., (eds.) (2003). ''Evolution and Palaeobiology of Pterosaurs''. Geological Society of London, Special Publications 217, London, 1–347.</ref> This group was named for the presence of a head crest in most known species, though this feature has since been found in more primitive pterosaurs and was probably an ancestral feature for all pterodactyloids.<ref name=witton>{{cite book|first=Mark|last=Witton|title=Pterosaurs: Natural History, Evolution, Anatomy|publisher=Princeton University Press|year=2013|isbn=978-0-691-15061-1}}</ref>

Another subgroup within Lophocratia is '''Eupterodactyloidea''' (meaning "true Pterodactyloidea"). Eupterodactyloidea was named by S. Christopher Bennett in 1994 as an infraorder of the suborder Pterodactyloidea. Bennett defined it as an apomorphy-based clade.<ref name="Bennett94">{{cite journal|author=S. Christopher Bennett|year=1994|title=Taxonomy and systematics of the Late Cretaceous pterosaur ''Pteranodon'' (Pterosauria, Pterodactyloidea)|url=https://www.biodiversitylibrary.org/page/4467188#page/5/mode/1up|journal=Occasional Papers of the Natural History Museum of the University of Kansas|volume=169|pages=1–70}}</ref> However, in 2010, Brian Andres re-defined the group as a stem-based taxon in his dissertation,<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 |access-date=2012-06-05 |archive-date=2013-10-02 |archive-url=https://web.archive.org/web/20131002080824/http://gradworks.umi.com/34/40/3440534.html }} [https://www.proquest.com/docview/846734901 A preview that shows the cladogram without clade names]</ref> and then formalized the definition in 2014 as all pterosaurs more closely related to ''Pteranodon longiceps'' than to ''Pterodactylus antiquus''.<ref name=kryptodrakon/> The slightly more exclusive group Ornithocheiroidea was re-defined in 2003 by Alexander Kellner. He defined it as the least inclusive clade containing ''Anhanguera blittersdorffi'', ''Pteranodon longiceps'', ''Dsungaripterus weii'', and ''Quetzalcoatlus northropi''. Ornithocheiroidea has often been used for a much more exclusive group including only the branch of traditional ornithocheirid pterosaurs, though this use has since fallen out of favor by many researchers after years of competing definitions for the various pterodactyloid clades. The compromise definitions by Andres and others have since become more widely adopted.

Within Eupterodactyloidea, there is a large clade - '''Ornithocheiroidea'''. The name Ornithocheiroidea was originally defined as an apomorphy-based taxon by Christopher Bennett in 1994. It was given a relationship-based definition in 2003 by Alexander Kellner, who defined it as the least inclusive clade containing ''Anhanguera blittersdorffi'', ''Pteranodon longiceps'', ''Dsungaripterus weii'', and ''Quetzalcoatlus northropi''.<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> Later that year, David Unwin suggested a more restrictive definition, in which the clade only contains ''Pteranodon longiceps'', ''Istiodactylus latidens'', and their descentants.<ref name=Unwin03>Unwin, D. M., (2003): On the phylogeny and evolutionary history of pterosaurs. pp. 139-190. &mdash; ''in'' Buffetaut, E. & Mazin, J.-M., (eds.): ''Evolution and Palaeobiology of Pterosaurs''. Geological Society of London, Special Publications 217, London, 1-347 {{doi|10.1144/GSL.SP.2003.217.01.11}}</ref> Brian Andres (2008, 2010, 2014) in his analyses, defined Ornithocheiroidea using the definition of Kellner (2003) to avoid confusion with similarly defined groups, like Pteranodontoidea.<ref name=BBA10 /><ref name="Andres2014">{{cite journal |author1=Andres, B. |author2=Clark, J. |author3=Xu, X. |year=2014 |title=The Earliest Pterodactyloid and the Origin of the Group |journal=Current Biology |volume= 24|issue=9 |pages= 1011–6|url=https://www.cell.com/current-biology/pdfExtended/S0960-9822(14)00322-4|doi=10.1016/j.cub.2014.03.030 |pmid=24768054|bibcode=2014CBio...24.1011A |doi-access=free }}</ref> In 2019, a phylogenetic analysis conducted by Kellner and colleagues had recovered Ornithocheiroidea as the sister taxon of the Archaeopterodactyloidea, and consisting of the clades Tapejaroidea and Pteranodontoidea.<ref name="Kellner2019">{{Cite journal|last1=Kellner|first1=Alexander W. A.|last2=Weinschütz|first2=Luiz C.|last3=Holgado|first3=Borja|last4=Bantim|first4=Renan A. M.|last5=Sayão|first5=Juliana M.|date=August 19, 2019|title=A new toothless pterosaur (Pterodactyloidea) from Southern Brazil with insights into the paleoecology of a Cretaceous desert|journal=Anais da Academia Brasileira de Ciências|language=en|volume=91|issue=suppl 2|article-number=e20190768|doi=10.1590/0001-3765201920190768|pmid=31432888|issn=0001-3765|doi-access=free}}</ref> Several recent studies have followed this or a similar concept.<ref name="Mimodactylus">{{cite journal |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. |title=First complete pterosaur from the Afro-Arabian continent: insight into pterodactyloid diversity |journal=Scientific Reports |date=2019 |volume=9 |issue=1 |page=17875 |doi=10.1038/s41598-019-54042-z|pmid=31784545 |pmc=6884559 |bibcode=2019NatSR...917875K |doi-access=free }}</ref><ref name="Holgado2019">{{cite journal | last1 = Holgado | first1 = Borja | last2 = Pêgas | first2 = Rodrigo V. | last3 = Ignacio Canudo | first3 = José | last4 = Fortuny | first4 = Josep | last5 = Rodrigues | first5 = Taissa | last6 = Company | first6 = Julio | last7 = Kellner | first7 = Alexander W.A. | year = 2019 | title = On a new crested pterodactyloid from the Early Cretaceous of the Iberian Peninsula and the radiation of the clade Anhangueria | url = | journal = Scientific Reports | volume = 9 | issue = 1| article-number = 4940 | doi = 10.1038/s41598-019-41280-4 | pmid = 30894614 | pmc = 6426928 | bibcode = 2019NatSR...9.4940H }}</ref><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> [[File:Kariridraco_by_Júlia_d’Oliveira.jpg|thumb|left|Reconstruction of ''Kariridraco'', a terrestrial azhdarchoid]] However, not all of the subgroups of pterodactyloids are universally accepted. One controversial taxon is '''Tapejaroidea'''. Tapejaroidea was named by paleontologist Alexander Kellner from Brazil in 1996,<ref name="kellner96"/><ref>{{cite book|last1=Buffetaut|first1=Eric|last2=Mazin|first2=Jean-Michel|title=Evolution and Palaeobiology of pterosaurs|publisher=Geological Society of London|year=2003|isbn=978-1-86239-143-7}}</ref> and in 2003 it was given a phylogenetic definition by Kellner himself as the most recent common ancestor of ''Dsungaripterus'', ''Tapejara'' and ''Quetzalcoatlus'', and all their descendants. Tapejaroidea, in Kellner's 2003 study, was recovered as the sister taxon of the Pteranodontoidea, both within the group Ornithocheiroidea, and consisting of the groups Dsungaripteridae and Azhdarchoidea.<ref name="Kellner2003">{{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> However, in a phylogenetic analysis made by Jaime Headden and Hebert Bruno Nascimento Campos in 2014, Tapejaroidea was recovered within the Azhdarchoidea, as a clade comprising the families Tapejaridae and Thalassodromidae.<ref name="Banguela">{{Cite journal|author=Jaime A. Headden and Hebert B.N. Campos |year=2014 |title=An unusual edentulous pterosaur from the Early Cretaceous Romualdo Formation of Brazil |journal=Historical Biology: An International Journal of Paleobiology |volume=27 |issue=7 |pages=815–826 |doi=10.1080/08912963.2014.904302 |s2cid=129306469 }}</ref> More recently, the original definition of Tapejaroidea has been used in a number of phylogenetic analyses conducted in 2019 and 2020, meaning that Tapejaroidea and Pteranodontoidea were once again recovered as the sister taxa and within the larger Ornithocheiroidea.<ref name="Holgado2019"/><ref name="Mimodactylus"/><ref name="Kellner2019" /><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> The cladogram below represents the phylogenetic analysis conducted by Kellner and colleagues in 2019, where they recovered Tapejaroidea as the more inclusive group containing both the Dsungaripteridae and the Azhdarchoidea.<ref name="Kellner2019"/>

Another controversial clade is '''Dsungaripteroidea'''. The Dsungaripteroidea was defined in 2003 by David Unwin. Unwin made Dsungaripteroidea the most inclusive clade containing both ''Dsungaripterus weii'' and ''Germanodactylus cristatus''.<ref name="unwin2003"/> Unwin at that time considered those two species to be close relatives. However, more recent studies have shown ''Germanodactylus'' to be much more primitive, either an archaeopterodactyloid or a primitive member of the Eupterodactyloidea. This makes Dsungaripteroidea a much larger group. Alexander Kellner in 2003 defined Dsungaripteroidea very differently as the group containing the last common ancestor of ''Nyctosaurus'' and ''Quetzalcoatlus'', and all its descendants. However, subsequent recent analysis use the name Ornithocheiroidea instead of Dsungaripteroidea for this definition.<ref name="Andres2014"/><ref name="upchurch15">{{cite journal|last1 = Upchurch | first1 = P.|last2=Andres|first2=B.B.|last3=Butler|first3=R.J.|last4=Barrett|first4=P.M.|year=2015|title=An analysis of pterosaurian biogeography: implications for the evolutionary history and fossil record quality of the first flying vertebrates|journal=Historical Biology|volume=27|issue=6|doi=10.1080/08912963.2014.939077|pages=697–717|pmc=4536946|pmid=26339122| bibcode = 2015HBio...27..697U}}</ref><ref name="Longrich2018">{{cite journal | last1 = Longrich | first1 = Nicholas R. | last2 = Martill | first2 = David M. | last3 = Andres | first3 = Brian | year = 2018 | title = Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary | journal = PLOS Biology | volume = 16 | issue = 3| article-number = e2001663 | doi = 10.1371/journal.pbio.2001663 | doi-access = free | pmid = 29534059 | pmc = 5849296 }}</ref> {| class="wikitable sortable" align="center" width="100%" ! Name ! Named by ! Definition ! Notes |- | '''Archaeopterodactyloidea''' | Kellner, 1996<ref name=kellner96 /> | Least-inclusive clade containing ''Pterodactylus'', ''Ctenochasma'', and ''Gallodactylus'' | May include all other subclades of pterodactyloids if ctenochasmatids are more closely related to Eupterodactyloidea (see below) than either are to ''Pterodactylus'' |- | '''Azhdarchoidea''' | Unwin, 1995<ref>Unwin, D. M. 1995. Preliminary results of a phylogenetic analysis of the Pterosauria (Diapsida: Archosauria); pp. 69–72 in A. Sun and Y. Wang (eds.), Sixth Symposium on Mesozoic Terrestrial Ecosystems and Biota, Short Papers. China Ocean Press, Beijing.</ref> | Least-inclusive clade containing both ''Quetzalcoatlus'' and ''Tapejara'' | |- | '''Dsungaripteroidea''' | Unwin, 2003<ref name=unwin2003 /> | Least-inclusive clade containing both ''Dsungaripterus'' and ''Germanodactylus'' | May be synonymous with Lophocratia if ''Germanodactylus'' is a member of Archaeopterodactyloidea; alternatively defined as the least-inclusive clade containing both ''Quetzalcoatlus'' and ''Nyctosaurus'' (possibly synonymous with Ornithocheiroidea) |- | '''Eupterodactyloidea''' | Bennett, 1994<ref name="Bennett94"/> | Most-inclusive clade containing ''Pteranodon'' but not ''Pterodactylus'' | Possibly synonymous with Ornithocheiroidea |- | '''Lophocratia''' | Unwin, 2003<ref name="unwin2003"/> | Least-inclusive clade containing both ''Pterodaustro'' and ''Quetzalcoatlus'' | |- | '''Ornithocheiroidea''' | Seeley, 1870<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> | Least-inclusive clade containing ''Anhanguera'', ''Pteranodon'', ''Dsungaripterus'', and ''Quetzalcoatlus'' | Previously defined as the least-inclusive clade containing both ''Pteranodon'' and ''Istiodactylus'' (this clade is now called Pteranodontoidea - see below) |- | '''Ornithocheiromorpha''' | Andres ''et al.'', 2014<ref name=kryptodrakon/> | Most-inclusive clade containing ''Ornithocheirus'' but not ''Pteranodon'' | |- | '''Pteranodontia''' | Marsh, 1887<ref>{{cite journal | last1 = Marsh | first1 = O. C. | year = 1876 | title = Notice of a new sub-order of Pterosauria | url = | journal = American Journal of Science | volume = 11 | issue = | pages = 507–509 }}</ref><ref name=unwin2003 /> | Least-inclusive clade containing both ''Pteranodon'' and ''Nyctosaurus'' | May be synonymous with Pteranodontoidea if ''Pteranodon'' is more closely related to Ornithocheiromorpha than either are to ''Nyctosaurus'' |- | '''Pteranodontoidea''' | Kellner, 1996<ref name=kellner96 /> | Least-inclusive clade containing both ''Pteranodon'' and ''Istiodactylus'' | |- | '''Tapejaroidea''' | Kellner, 1996<ref name=kellner96>{{cite journal|last=Kellner|first=A.W.A.|year=1996|title=Description of new material of Tapejaridae and Anhangueridae (Pterosauria, Pterodactyloidea) and discussion of pterosaur phylogeny|journal=Columbia University}}</ref> | Least-inclusive clade containing ''Quetzalcoatlus'', ''Tapejara'', and ''Dsungaripterus'' | May be synonymous with Azhdarchoidea if ''Dsungaripterus'' is more closely related to ''Quetzalcoatlus'' than either are to ''Tapejara''; alternatively used for a clade containing Tapejaridae and Thalassodromidae<ref name="Banguela"/> |- |}

===Taxonomy=== [[File:Mimodactylus_in_life.png|thumb|Reconstruction of ''Mimodactylus'', a marine pteranodontoid]] There are competing theories of pterodactyloid phylogeny. Below is cladogram following a topology recovered by Brian Andres, using the most recent iteration of his data set (Andres, 2021). This study found the two traditional groupings of ctenochasmatoids and kin as an early branching group, with all other pterodactyloids grouped into the Eupterodactyloidea. A simplified version of the cladogram included in that publication is shown below.<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='''Pterodactyloidea''' |1={{clade |1=''Kryptodrakon'' |2={{clade |1=''Propterodactylus'' |label2=Lophocratia |2={{clade |label1=Archaeopterodactyloidea |1={{clade |1=Germanodactylidae |label2=Euctenochasmatia |2={{clade |1=''Pterodactylus'' |2=Ctenochasmatoidea 50 px}} }} |label2=Eupterodactyloidea |2={{clade |1=''Haopterus'' |label2=Ornithocheiroidea |2={{clade |label1=Pteranodontoidea |1={{clade |2=Pteranodontia 50 px |1=Ornithocheiromorpha 40 px }} |label2=Azhdarchoidea |2={{clade |1=Tapejaridae50 px |2=Neoazhdarchia 50 px}} }} }} }} }} }} }}

Some studies based on a different type of analysis have found that this basic division into primitive (archaeopterodactyloid) and advanced (eupterodactyloid) species may not be correct. Beginning in 2014, Steven Vidovic and David Martill constructed an analysis in which several pterosaurs traditionally thought of as archaeopterodactyloids closely related to the ctenochasmatoids may have been more closely related to ornithocheiroids, or in some cases, fall outside both groups. The results of their updated 2017 analysis are shown below.<ref name=Altmuehlopterus>Vidovic, S.U. and Martill, D.M. (2017). The taxonomy and phylogeny of ''Diopecephalus kochi'' (Wagner, 1837) and "''Germanodactylus rhamphastinus''" (Wagner, 1851). In Hone, D. W. E., Witton, M. P. &Martill, D. M. (eds) ''New Perspectives on Pterosaur Palaeobiology''. Geological Society, London, Special Publications, 455 {{doi|10.1144/SP455.12}}</ref> {{clade| style=font-size:80%;line-height:80% |label1='''Pterodactyloidea''' |1={{clade |1=''Eosipterus yangi'' |2={{clade |1=''Pterodactylus antiquus'' |label2=Lophocratia |2={{clade |label1=Euctenochasmatia |1={{clade |1=''Diopecephalus kochi'' |label2=Ctenochasmatoidea |2={{clade |label1=Aurorazhdarchia |1={{clade |1=''Aerodactylus scolopaciceps'' |2={{clade |1=Aurorazhdarchidae |2=Gallodactylidae }} }} |2=Ctenochasmatidae }} }} |label2=Eupterodactyloidea |2={{clade |1=''Altmuehlopterus ramphastinus'' |label2=Dsungaripteroidea |2={{clade |1=''Germanodactylus cristatus'' |2={{clade |1=''Elanodactylus prolatus'' |label2=Ornithocheiroidea |2={{clade |label1=Dsungaripteromorpha |1={{clade |1={{clade |1=''Lonchodectes compressirostris'' |2={{clade |1=''Prejanopterus curvirostra'' |2={{clade |1=''Kepodactylus insperatus'' |2=Dsungaripteridae}} }} }} |label2=Pteranodontoidea |2={{clade |1=Ornithocheiromorpha |2={{clade |1={{clade |1=''Hamipterus tianshanensis'' |2=''Ikrandraco avatar'' }} |2={{clade |1={{clade |1=''Istiodactylus'' |label2=Pteranodontia |2={{clade |1=Nyctosauridae |2=''Pteranodon'' }} }} }} }} }} }} |label2=Azhdarchoidea |2={{clade |1=''Sinopterus dongi'' |2={{clade |1=''Eoazhdarcho liaoxiensis'' |label2=Neoazhdarchia |2={{clade |1=Tapejaromorpha |2=''Bennettazhia oregonensis'' |3=''Huaxiapterus jii'' |4=''Thalassodromeus sethi'' |5=''Tupuxuara leonardii'' |6=Azhdarchidae }} }} }} }} }} }} }} }} }} }} }}

==References== {{Reflist}} {{Pterosauria|M.}} {{Taxonbar|from=Q746655}}

Category:Pterodactyloidea Category:Reptile clades Category:Jurassic reptiles Category:Cretaceous reptiles