{{Short description|Extinct clade of reptiles}} {{Automatic taxobox | name = Pterosauromorphs | fossil_range = <br />Late Triassic – Late Cretaceous, {{Fossil range|236|earliest=244|66|ref=<ref>{{Cite journal|author1=Claudia A. Marsicano|author2=Randall B. Irmis|author3=Adriana C. Mancuso|author4=Roland Mundil|author5=Farid Chemale|year=2016|title=The precise temporal calibration of dinosaur origins|journal=Proceedings of the National Academy of Sciences of the United States of America|volume=113|issue=3|pages=509–513|bibcode=2016PNAS..113..509M|doi=10.1073/pnas.1512541112|pmc=4725541|pmid=26644579|doi-access=free}}</ref>}} | image = Lagerpeton skeleton.png | image_caption = Skeletal restoration of ''{{nobr|Lagerpeton chanarensis}}'' (known elements represented in white) | image2 = Dimorphodon skeleton.jpg | image2_caption = Skeletal restoration of ''{{nobr|Dimorphodon macronyx}}'' | taxon = Pterosauromorpha | authority = Kuhn-Schnyder & Rieber, 1986 | subdivision_ranks = Subgroups | subdivision = *{{extinct}}''Maehary''? *{{extinct}}''Scleromochlus'' *{{extinct}}Lagerpetidae *{{extinct}}'''Pterosauria''' }} '''Pterosauromorpha''' (meaning "pterosaur-like forms") is one of the two basic divisions of Ornithodira that includes pterosaurs and all taxa that are closer to them than to dinosaurs and their close relatives (i.e. Dinosauromorpha).<ref name="Padian1997">{{cite book |author=Padian |first=K. |title=The Encyclopedia of Dinosaurs |publisher=Academic Press |year=1997 |isbn=978-0-12-226810-6 |editor1=Currie |editor-first=P. J. |pages=617–618 |contribution=Pterosauromorpha |editor2=Padian |editor-first2=K.}}</ref><ref name="Phylonyms">{{cite book |title=Phylonyms: A Companion to the PhyloCode |publisher=CRC Press |year=2020 |isbn=978-1-138-33293-5 |editor-last=de Queiroz |editor-first=K. |location=Boca Raton |pages=1195–1199 |language= |chapter=''Pterosauromorpha'' E. Kuhn-Schnyder and H. Rieber 1986 [B. Andres and K. Padian], converted clade name |editor-last2=Cantino |editor-first2=P. D. |editor-last3=Gauthier |editor-first3=J. A.}}</ref> In addition to pterosaurs, Pterosauromorpha also includes the basal clade Lagerpetidae and some other Late Triassic ornithodirans (such as ''Scleromochlus'').<ref name="Kammerer2020">{{Cite journal |last1=Kammerer |first1=C. F. |last2=Nesbitt |first2=S. J. |last3=Flynn |first3=J. J. |last4=Ranivoharimanana |first4=L. |last5=Wyss |first5=A. R. |date=2020 |title=A tiny ornithodiran archosaur from the Triassic of Madagascar and the role of miniaturization in dinosaur and pterosaur ancestry |journal=Proceedings of the National Academy of Sciences |language=en |volume=117 |issue=30 |pages=17932–17936 |doi=10.1073/pnas.1916631117 |issn=0027-8424 |pmc=7395432 |pmid=32631980|doi-access=free |bibcode=2020PNAS..11717932K }}</ref><ref name="Ezcurra2020">{{Cite journal |last1=Ezcurra |first1=M. D. |last2=Nesbitt |first2=S. J. |last3=Bronzati |first3=M. |last4=Dalla Vecchia |first4=F. M. |last5=Agnolin |first5=F. L. |last6=Benson |first6=R. B. J. |last7=Brissón Egli |first7=F. |last8=Cabreira |first8=S. F. |last9=Evers |first9=S. W. |last10=Gentil |first10=A. R. |last11=Irmis |first11=R. B. |display-authors=10 |date=2020 |title=Enigmatic dinosaur precursors bridge the gap to the origin of Pterosauria |journal=Nature |language=en |volume=588 |issue=7838 |pages=445–449 |doi=10.1038/s41586-020-3011-4 |issn=0028-0836 |pmid=33299179 |bibcode=2020Natur.588..445E |s2cid=228077525|url=http://doc.rero.ch/record/329918/files/ser_edp.pdf }}</ref><ref name="Baron2021">{{Cite journal |last1=Baron |first1=M. G. |date=2021 |title=The origin of Pterosaurs |journal=Earth-Science Reviews |language=en |volume=221 |article-number=103777 |doi=10.1016/j.earscirev.2021.103777 |bibcode=2021ESRv..22103777B |issn=0012-8252}}</ref><ref name="Kellner2022">{{cite journal |last1=Kellner |first1=A.W.A. |last2=Holgado |first2=B. |last3=Grillo |first3=O. |last4=Pretto |first4=F.A. |last5=Kerber |first5=L. |last6=Pinheiro |first6=F.L. |last7=Soares |first7=M.B. |last8=Schultz |first8=C.L. |last9=Lopes |first9=R.T. |last10=Araújo |first10=A. |last11=Müller |first11=R.T. |year=2022 |title=Reassessment of ''Faxinalipterus minimus'', a purported Triassic pterosaur from southern Brazil with the description of a new taxon |journal=PeerJ |volume=10 |article-number=e13276 |doi=10.7717/peerj.13276 |pmc=9074864 |pmid=35529502 |doi-access=free}}</ref>
== Classification == [[File:Scleromochlus taylori.jpg|left|thumb|Skeletal restoration of ''Scleromochlus'']] The name Pterosauromorpha was originally coined by Emil Kuhn-Schnyder and Hans Rieber (1986) for a reptilian subclass distinct from Archosauria which includes pterosaurs.<ref>{{Cite book |last1=Kuhn-Schnyder |first1=E. |title=Handbook of Paleozoology |last2=Rieber |first2=H. |publisher=Johns Hopkins University Press |year=1986 |location=Baltimore, MD}}</ref> In 1997, Kevin Padian classified Pterosauromorpha as a clade of archosaurs and proposed phylogenetic definition for this group: "Pterosauria and all ornithodiran archosaurs closer to them than to dinosaurs".<ref name="Padian1997" /> Brian Andres and Kevin Padian redefined Pterosauromorpha as: "The clade consisting of ''Pterodactylus'' (originally ''Ornithocephalus'') ''antiquus'' (Sömmerring 1812) (Pterosauria) and all organisms or species that share a more recent common ancestor with it than with ''Alligator'' (originally ''Crocodilus'') ''mississippiensis'' (Daudin 1802) (Suchia) and ''Compsognathus longipes'' Wagner 1859 (Dinosauromorpha)".<ref name="Phylonyms" /><ref>{{Cite web|title=Pterosauromorpha|url=https://www.phyloregnum.org/?term=Pterosauromorpha|access-date=30 May 2022|website=RegNum}}</ref>
Lagerpetidae was traditionally considered the earliest diverging group of dinosauromorphs. This caused no other reptiles besides the true pterosaurs to be placed in Pterosauromorpha.<ref name="Nesbitt2011">{{cite journal |last=Nesbitt |first=S.J. |year=2011 |title=The Early Evolution of Archosaurs: Relationships and the Origin of Major Clades |journal=Bulletin of the American Museum of Natural History |volume=352 |page=189 |doi=10.1206/352.1 |issn=0003-0090 |hdl=2246/6112 |s2cid=83493714|doi-access=free |hdl-access=free }}</ref><ref name="Nesbitt2017">{{cite journal |last1=Nesbitt |first1=S. J. |last2=Butler |first2=R. J. |last3=Ezcurra |first3=M. D. |last4=Barrett |first4=P. M. |last5=Stocker |first5=M. R. |last6=Angielczyk |first6=K. D. |last7=Smith |first7=R. M. H. |last8=Sidor |first8=C. A. |last9=Niedźwiedzki |first9=G. |last10=Sennikov |first10=A. G. |last11=Charig |first11=A. J. |date=2017 |title=The earliest bird-line archosaurs and the assembly of the dinosaur body plan |url=http://pure-oai.bham.ac.uk/ws/files/39495711/Nesbitt_et_al._in_press.pdf |journal=Nature |volume=544 |issue=7651 |pages=484–487 |bibcode=2017Natur.544..484N |doi=10.1038/nature22037 |pmid=28405026|s2cid=9095072 }}</ref> The only notable exception was a small reptile named ''Scleromochlus'', whose placement within the clade itself remained controversial due to the poor preservation of its otherwise complete remains. Different phylogenetic analyses found it as a basal pterosauromorph,<ref name="Kammerer2020" /><ref name="Ezcurra2020" /> a non-aphanosaurian, non-pterosaur basal avemetatarsalian, a basal dinosauromorph,<ref name="Nesbitt2017" /> or a basal archosauriform.<ref name="bennett2020">{{Cite journal |last=Bennett |first=S.C. |year=2020 |title=Reassessment of the Triassic archosauriform ''Scleromochlus taylori'': neither runner nor biped, but hopper |journal=PeerJ |volume=8 |article-number=e8418 |doi=10.7717/peerj.8418 |issn=2167-8359 |pmc=7035874 |pmid=32117608 |doi-access=free}}</ref> This has resulted in a large gap between the fully aerial pterosaurs and their terrestrial ancestors, as the earliest pterosaurs were already capable flyers.<ref name="Nesbitt2017" />
First iteration of phylogenetic analysis produced by Kammerer et al. (2020) restored lagerpetids as a basal dinosauromorphs, which corresponds to the traditional point of view. But the second iteration, in which were added ''Scleromochlus,'' found Lagerpetidae as the most basal pterosauromorphs, and ''Scleromochlus'' as the sister taxon of pterosaurs.<ref name="Kammerer2020" /> In a study that used micro-CT scans, Ezcurra et al. (2020) have found additional similarities, including large semicircular canals within the bones of some lagerpetids that resemble that of pterosaurs. It is assumed that large semicircular canals are related to arboreal, aerial or other agile forms of terrestrial locomotion as well as rapid movements. The flocculus, the part of the brain that aids in transmitting information, was also large in both pterosaurs and lagerpetids, though to a lesser extent. When Ezcurra et al. (2020) included ''Scleromochlus'' in their analysis, they found it to be the most basal pterosauromorph, sister to a clade including lagerpetids and pterosaurs.<ref name="Ezcurra2020" /> Baron (2021) conducted his own analysis, which confirmed the relationship between lagerpetids and pterosaurs.<ref name="Baron2021" />
{{Cquote|1=It has to do with the semicircular canals [in the ear], which orients you in 3D space. The shape of those canals correlates with ecology and how you move your head — basically, are you agile or not? And a lot of things that have flight have semicircular canals with a really large and characteristic [shape] because you're flying, you're in a lot more 3D space.|author=Sterling Nesbitt<ref>{{Cite web |date=December 16, 2020 |title="Missing Link" in Pterosaurs' Family Tree Has Been Identified |url=https://mysteriousuniverse.org/2020/12/missing-link-in-pterosaurs-family-tree-has-been-identified/ |website=Mysterious Universe |publisher=Jocelyne LeBlanc}}</ref>}}
Kellner et al. (2022) described ''Maehary,'' a small archosaur from the Late Triassic of Brazil. Along with lagerpetids, it was interpreted as a basal pterosauromorph. It is noteworthy that left maxilla of ''Maehary'' was previously considered to be a specimen of ''Faxinalipterus'' that was re-classified as a lagerpetid.<ref name="Kellner2022" /> Contrary to the initial classification, phylogenetic analyses conducted by Müller et al. (2023) and Müller (2024) recovered ''Maehary'' as a gracilisuchid pseudosuchian rather than a pterosauromorph.<ref name="Venetoraptor">{{Cite journal |last1=Müller |first1=R. T. |last2=Ezcurra |first2=M. D. |last3=Garcia |first3=M. S. |last4=Agnolín |first4=F. L. |last5=Stocker |first5=M. R. |last6=Novas |first6=F. E. |last7=Soares |first7=M. B. |last8=Kellner |first8=A. W. A. |last9=Nesbitt |first9=S. J. |year=2023 |title=New reptile shows dinosaurs and pterosaurs evolved among diverse precursors |journal=Nature |volume=620 |issue=7974 |pages=589–594 |doi=10.1038/s41586-023-06359-z |doi-access=free}}</ref><ref name="Parvosuchus">{{Cite journal |last1=Müller |first1=Rodrigo T. |date=2024-06-20 |title=A new small-sized predatory pseudosuchian archosaur from the Middle-Late Triassic of Southern Brazil |journal=Scientific Reports |language=en |volume=14 |issue=1 |page=12706 |doi=10.1038/s41598-024-63313-3 |issn=2045-2322 |pmc=11189902 |doi-access=free}}</ref> In their study on ''Scleromochlus'', Foffa et al. (2023) noted that the phylogenetic position of ''Maehary'' remains uncertain due to the fragmentary nature of its material.<ref name="Foffa2023">{{Cite journal |last1=Foffa |first1=D. |last2=Nesbitt |first2=S. J. |last3=Butler |first3=R. J. |last4=Brusatte |first4=S. L. |last5=Walsh |first5=S. |last6=Fraser |first6=N. C. |last7=Barrett |first7=P. M. |year=2023 |title=The osteology of the Late Triassic reptile ''Scleromochlus taylori'' from μCT data |journal=The Anatomical Record |volume=307 |issue=4 |pages=1113–1146 |doi=10.1002/ar.25335 |pmid=37846180 |doi-access=free |hdl-access=free |hdl=20.500.11820/2411a888-1efb-42d1-a498-fc031145ed86}}</ref>
{{col-begin|width=100%}} {{col-break}} '''Nesbitt et al. (2011)'''<ref name="Nesbitt2011" /> {{clade|{{clade |label1=Dinosauromorpha |1={{clade |1=Lagerpetidae |2=Dinosauriformes }} |label2='''Pterosauromorpha''' |2=Pterosauria }}|style=font-size:90%;line-height:85%|label1=Ornithodira}} {{col-break}} '''Ezcurra et al. (2020)'''<ref name="Ezcurra2020 "/> {{clade|{{clade |1=Dinosauromorpha |label2='''Pterosauromorpha''' |2={{clade |1=Lagerpetidae |2=Pterosauria }} }}|style=font-size:90%;line-height:85%|label1=Ornithodira}} {{col-break}} '''Kellner et al. (2022)'''<ref name="Kellner2022" /> {{clade|{{clade |1=Dinosauromorpha |label2='''Pterosauromorpha''' |2={{clade |1={{clade |1=''Maehary'' |2={{clade |1=Lagerpetidae |2=Pterosauria }} }} }} }}|style=font-size:90%;line-height:85%|label1=Ornithodira}} {{col-end}}
An analysis by Garcia & Müller (2025) found lagerpetids as paraphyletic to pterosaurs, and ''Faxinalipterus'' not as a pterosauromorph but as a sister taxon to Ornithodira, more closely related to ornithodirans than to ''Mambachiton'' and Aphanosauria.<ref name="2025Review">{{Cite journal |last1=Garcia |first1=Maurício S. |last2=Müller |first2=Rodrigo T. |date=2025 |title=Triassic pterosaur precursors of Brazil: catalog, evolutionary context, and a new hypothesis for phylogenetic relationships of Pterosauromorpha |journal=Anais da Academia Brasileira de Ciências |volume=97 |issue=suppl 1 |article-number=e20240844 |doi=10.1590/0001-3765202520240844 |issn=1678-2690 |pmid=40008775 |doi-access=free |url=https://www.scielo.br/j/aabc/a/NJjRz5dD9PymgdfwfdpSXdJ/?lang=en}}</ref>{{clade|{{clade |1=''Faxinalipterus'' |label2=Ornithodira |2={{clade |1=Dinosauromorpha |label2='''Pterosauromorpha''' |2={{clade |grouplabel1='Lagerpetidae' |1=''Kongonaphon'' |barbegin1=gray |2={{clade |1=''Lagerpeton'' |bar1=gray |2={{clade |1=UFSM 11611 |bar1=gray |2={{clade |1=''Ixalerpeton'' |bar1=gray |2={{clade |1=PVSJ 883 |bar1=gray |2={{clade |1={{clade |1=''Dromomeron gigas'' |bar1=gray |2=''Dromomeron romeri'' |bar2=gray }} |3={{clade |1={{clade |1=''Venetoraptor'' |bar1=gray |2="''Dromomeron''" ''gregorii'' |barend2=gray }} |3={{clade |1=Pterosauria }} }} }} }} }} }} }} }} }} }}}}
== References == {{reflist}}
{{Avemetatarsalia}} {{Taxonbar|from=Q2189453}}
Category:Pterosauromorpha Category:Carnian first appearances Category:Maastrichtian extinctions