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Archosauromorpha: Difference between revisions

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| subdivision = *{{extinct}}''[[Aenigmastropheus]]''
*{{extinct}}''[[Elachistosuchus]]''?
*{{extinct}}''[[Lusitanosaurus]]''?
*{{extinct}}''[[Prolacertoides]]''
*{{extinct}}[[Choristodera]]?
*{{extinct}}[[Kuehneosauridae]]?
*{{extinct}}[[Drepanosauridae]]?
*{{extinct}}[[Helveticosauridae]]?
*{{extinct}}[[Ichthyosauromorpha]]?
*{{extinct}}[[Protorosauria]] (likely [[Polyphyly|polyphyleticparaphyletic]])
*{{extinct}}[[Sauropterygia]]?
*{{extinct}}[[Thalattosauria]]?
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}}
 
'''Archosauromorpha''' ([[Greek language|Greek]] for "ruling lizard forms") is a [[clade]] of [[diapsid]] [[reptile]]s containing all reptiles more closely related to [[archosaur]]s (such as [[crocodilia]]ns and [[dinosaur]]s, including [[bird]]s) rather than [[Lepidosauria|lepidosaurs]] (such as [[tuatara]]s, [[lizard]]s, and [[snake]]s).<ref name=":1" /> Archosauromorphs first appeared during the late [[Middle Permian]] or [[Late Permian]],<ref>{{Cite journal |last1=Martinelli |first1=Agustín G. |last2=Francischini |first2=Heitor |last3=Dentzien-Dias |first3=Paula C. |last4=Soares |first4=Marina B. |last5=Schultz |first5=Cesar L. |date=2017-01-02 |title=The oldest archosauromorph from South America: postcranial remains from the Guadalupian (mid-Permian) Rio do Rasto Formation (Paraná Basin), southern Brazil |url=https://www.tandfonline.com/doi/full/10.1080/08912963.2015.1125897 |journal=Historical Biology |language=en |volume=29 |issue=1 |pages=76–84 |doi=10.1080/08912963.2015.1125897 |bibcode=2017HBio...29...76M |s2cid=86151472 |issn=0891-2963}}</ref> though they became much more common and diverse during the [[Triassic|Triassic period]].<ref name=":0">{{Cite journal|last1=Butler|first1=Richard J.|last2=Scheyer|first2=Torsten M.|last3=Ezcurra|first3=Martín D.|date=2014-02-27|title=The Origin and Early Evolution of Sauria: Reassessing the Permian Saurian Fossil Record and the Timing of the Crocodile-Lizard Divergence|journal=PLOS ONE|language=en|volume=9|issue=2|pages=e89165|doi=10.1371/journal.pone.0089165|issn=1932-6203|pmc=3937355|pmid=24586565|bibcode=2014PLoSO...989165E|doi-access=free}}</ref>
 
Although Archosauromorpha was first named in 1946, its membership did not become well-established until the 1980s. Currently Archosauromorpha encompasses four main groups of reptiles: the stocky, herbivorous [[Allokotosauria|allokotosaurs]] and [[rhynchosaur]]s, the hugely diverse [[Archosauriformes]], and a [[Polyphyly|polyphyletic]] grouping of various long-necked reptiles including ''[[Protorosaurus]]'', [[Tanystropheidae|tanystropheids]], and ''[[Prolacerta]]''. Other groups including [[pantestudines]] ([[turtle]]s and their extinct relatives) and the semiaquatic [[Choristodera|choristoderes]] have also been placed in Archosauromorpha by some authors.
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The advent of [[cladistics]] helped to sort out at least some of the relationships within Reptilia, and it became clear that there was a split between the archosaur lineage and the lepidosaur lineage somewhere within the Permian, with certain reptiles clearly closer to archosaurs and others allied with lepidosaurs. [[Jacques Gauthier]] reused the term Archosauromorpha for the archosaur lineage at the 1982 annual meeting of the [[Society for Integrative and Comparative Biology|American Society of Zoologists]], and later used it within his 1984 Ph.D. thesis.<ref name=":2">{{Cite journal|author1=Jacques Gauthier |author2=Arnold G. Kluge |author3=Timothy Rowe | title = Amniote phylogeny and the importance of fossils | journal = [[Cladistics (journal)|Cladistics]] | volume = 4 | issue = 2 | pages = 105–209 | date = 1988 | doi = 10.1111/j.1096-0031.1988.tb00514.x |pmid=34949076 |hdl=2027.42/73857 |s2cid=83502693 |url=https://deepblue.lib.umich.edu/bitstream/2027.42/73857/1/j.1096-0031.1988.tb00514.x.pdf | hdl-access = free }}</ref> Archosauromorpha, as formulated by Gauthier, included four main groups of reptiles: Rhynchosauria, "Prolacertiformes", "Trilophosauria", and [[Archosauria]] (now equivalent to the group [[Archosauriformes]]). Cladistic analyses created during the 1980s by Gauthier, [[Michael Benton|Michael J. Benton]], and [[Susan E. Evans]] implemented Gauthier's classification scheme within large studies of reptile relations.<ref>{{Cite journal|last=Benton|first=Michael J.|date=1985-06-01|title=Classification and phylogeny of the diapsid reptiles|url=https://www.academia.edu/10694425|journal=Zoological Journal of the Linnean Society|language=en|volume=84|issue=2|pages=97–164|doi=10.1111/j.1096-3642.1985.tb01796.x|issn=0024-4082}}</ref><ref name=":2" /><ref name=":3">{{Cite book|title=The Phylogeny and Classification of the Tetrapods, Volume 1: Amphibians, Reptiles, Birds|last=Evans|first=Susan E.|publisher=Clarendon Press|year=1988|editor-last=Benton|editor-first=M. J.|location=Oxford|pages=221–260|chapter=The early history and relationships of the Diapsida|chapter-url=https://www.researchgate.net/publication/228117622}}</ref>
 
[[Michel Laurin]] (1991) defined Archosauromorpha as thea [[Node-based taxon|node-based]] clade containing the [[most recent common ancestor]] of ''[[Prolacerta]]'', ''[[Trilophosaurus]]'', ''[[Hyperodapedon]]'' and all of its descendants.<ref name="Apsisaurus">{{Cite journal|author=Michel Laurin |year=1991 |title=The osteology of a Lower Permian eosuchian from Texas and a review of diapsid phylogeny |journal=Zoological Journal of the Linnean Society |volume=101 |issue=1 |pages=59–95 |doi=10.1111/j.1096-3642.1991.tb00886.x }}</ref> David Dilkes (1998) formulated a more inclusive (and currently more popularcommon) definition of Archosauromorpha, defining it as thea [[Branch-based taxon|branch-based]] [[total group]] clade containing ''[[Protorosaurus]]'' and all other [[sauria]]ns that are more closely related to ''Protorosaurus'' than to Lepidosauria.<ref name="Dilkes98">{{cite journal|author=David M. Dilkes|year=1998|title=The Early Triassic rhynchosaur ''Mesosuchus browni'' and the interrelationships of basal archosauromorph reptiles|journal=Philosophical Transactions of the Royal Society of London, Series B|volume=353|issue=1368|pages=501–541|doi=10.1098/rstb.1998.0225|pmc=1692244}}</ref> Gauthier, as an author for ''[[Phylonyms]]'' (2020), redefined Archosauromorpha as a node-based clade containing ''[[Chicken|Gallus]], [[Alligator]], [[Mesosuchus]], Trilophosaurus, Prolacerta,'' and ''Protorosaurus''. The new name '''Pan-Archosauria''' was established for the broader total group of Archosauromorpha, similar to the definition of Dilkes (1998).<ref name=":12">{{Cite book |last1=Gauthier |first1=Jacques A. |url=https://www.taylorfrancis.com/books/mono/10.1201/9780429446276/phylonyms-kevin-de-queiroz-philip-cantino-jacques-gauthier |title=Phylonyms: A Companion to the PhyloCode |publisher=CRC Press |year=2020 |isbn=9780429446276 |editor-last1=De Queiroz |editor-first1=Kevin |edition=1st |location=Boca Raton |chapter=Archosauromorpha F. von Huene 1946 [J. A. Gauthier], converted clade name |doi=10.1201/9780429446276 |editor-last2=Cantino |editor-first2=Philip |editor-last3=Gauthier |editor-first3=Jacques |s2cid=242704712}}</ref>
 
In 2016, [[Martin Ezcurra]] named a subgroup of Archosauromorpha, '''Crocopoda''' ("crocodile feet"). Crocopoda is defined as all archosauromorphs more closely related to allokotosaurs (specifically ''[[Azendohsaurus]]'' and ''[[Trilophosaurus]]''), rhynchosaurs (specifically ''[[Rhynchosaurus]]''), or archosauriforms (specifically ''Proterosuchus'') rather than ''Protorosaurus'' or tanystropheids (specifically ''[[Tanystropheus]]''). This group roughly corresponds to Laurin's definition of Archosauromorpha.<ref name=":1">{{Cite journal|last=Ezcurra|first=Martín D.|date=2016-04-28|title=The phylogenetic relationships of basal archosauromorphs, with an emphasis on the systematics of proterosuchian archosauriforms|journal=PeerJ|language=en|volume=4|pages=e1778|doi=10.7717/peerj.1778|pmid=27162705|pmc=4860341|issn=2167-8359 |doi-access=free }}</ref>
 
==Members==
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Apart from these four groups, Archosauromorpha is sometimes considered to encompass several additional groups of reptiles. One of the most common additions is [[Choristodera]], a group of semiaquatic reptiles with mysterious origins. Although choristodere fossils are only known from the [[Jurassic]] through the [[Miocene]], it is theorized that they first appeared during the Permian alongside the earliest archosauromorphs. Choristoderes share numerous otherwise unique traits with archosauromorphs, but they share an equal or greater number of unique traits with lepidosauromorphs as well, so there is still some debate over their inclusion within either group.<ref name=":3" /><ref name=":1" /> The [[chameleon]]- or [[tamandua]]-like [[drepanosaur]]s are also semi-regularly placed within Archosauromorpha,<ref name="Dilkes98" /> although some studies have considered them to be part of a much more basal lineage of reptiles.<ref name=":02">{{Cite journal|last1=Pritchard|first1=Adam C.|last2=Nesbitt|first2=Sterling J.|date=2017-10-01|title=A bird-like skull in a Triassic diapsid reptile increases heterogeneity of the morphological and phylogenetic radiation of Diapsida|journal=Royal Society Open Science|language=en|volume=4|issue=10|pages=170499|doi=10.1098/rsos.170499|pmid=29134065|pmc=5666248|issn=2054-5703|bibcode=2017RSOS....470499P}}</ref> The aquatic [[thalattosaur]]s<ref name=":3" /> and gliding [[Kuehneosauridae|kuehneosaurids]]<ref name=":3" /><ref name=":02" /> are also irregularly considered archosauromorphs.
 
Genetic studies have found evidence that modern testudines ([[turtle]]s and [[tortoise]]s) are more closely related to crocodilians than to lizards.<ref>{{cite journal |author=Bhart-Anjan S. Bhullar |author2=Gabe S. Bever |year=2009 |title=An archosaur-like laterosphenoid in early turtles (Reptilia: Pantestudines) |journal=Breviora |volume=518 |pages=1–11 |url=http://www.mcz.harvard.edu/Publications/pubs/Breviora_2009_0518_2476.pdf |doi=10.3099/0006-9698-518.1.1|s2cid=42333056 }}</ref><ref>{{Cite journal|author1=Sean Modesto |author2=Robert Reisz |author3=Diane Scott |year=2011 |title=A neodiapsid reptile from the Lower Permian of Oklahoma |journal=Society of Vertebrate Paleontology 71st Annual Meeting Program and Abstracts |page=160}}</ref> If this evidence is accurate, then turtles are part of basal Archosauromorpha. Likewise, extinct turtle relatives known as [[Pantestudines]] would also fall within Archosauromorpha. Some geneticists have proposed a name to refer to reptiles within the group formed by relatives of turtles and archosaurs. This name is the clade '''[[Archelosauria]]'''. Since Pantestudines may encompass the entire aquatic reptile order [[Sauropterygia]], this means that Archosauromorpha (as Archelosauria) may be a much wider group than commonly believed.<ref>{{Cite journal|author=John W. Merck |year=1997 |title=A phylogenetic analysis of the euryapsid reptiles |journal=Journal of Vertebrate Paleontology |volume=17 |issue=Supplement to 3 |pages=1–93 | doi = 10.1080/02724634.1997.10011028}}</ref> However, anatomical data disagrees with this genetic evidence, instead placing Pantestudines within Lepidosauromorpha<ref>{{Cite journal|last=Rieppel|first=Olivier|date=1999-02-12|title=Turtle Origins|journal=Science|language=en|volume=283|issue=5404|pages=945–946|doi=10.1126/science.283.5404.945|pmid=10075558|s2cid=42475241|issn=1095-9203}}</ref> but many modern studies have supported Archelosauria. Several recent studies place sauropterygians within Archosauromorpha group, forming a large clade including [[Ichthyosauromorpha]] and [[Thalattosauria]] as opposed to the Pantestudine relations.<ref>{{Cite journal | last1 = Neenan | first1 = J. M. | last2 = Klein | first2 = N. | last3 = Scheyer | first3 = T. M. | title = European origin of placodont marine reptiles and the evolution of crushing dentition in Placodontia | doi = 10.1038/ncomms2633 | journal = Nature Communications | volume = 4 | pages = 1621 | year = 2013 | pmid = 23535642| bibcode = 2013NatCo...4.1621N | doi-access = free }}</ref><ref>{{cite journal |last1=Simões |first1=T. |last2=Kammerer |first2=C. |date=August 2022 |title=Successive climate crises in the deep past drove the early evolution and radiation of reptiles |urljournal=https://www.science.org/doi/10.1126/sciadv.abq1898Science |journal=ScienceAdvancesAdvances |volume=08 |issue=33 |pages= eabq1898|doi=10.1126/sciadv.abq1898 |pmid=35984885 |pmc=9390993 |bibcode=2022SciA....8.1898S |s2cid=251694019 |access-date=October 27, 2023|doi-access=free }}</ref><ref>{{cite journal |last1=Wang |first1=W. |last2=Shang |first2=Q. |date=December 2022 |title=Ancestral body plan and adaptive radiation of sauropterygian marine reptiles |journal= iScience|volume=25 |issue=12 |pages= |doi=10.1016/j.isci.2022.105635 |pmid=36483013 |pmc=9722468 |bibcode=2022iSci...25j5635W }}</ref>
 
== Anatomy ==
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=== Forelimbs ===
The [[humerus]] (forearmupper arm bone) is solid in archosauromorphs, completely lacking a hole near the elbow known as the [[entepicondylar foramen]]. This hole, present in most other tetrapods, is also absent in choristoderes yet not fully enclosed in some proterosuchids. In many advanced archosauromorphs, the [[Capitulum of the humerus|capitulum]] and [[Trochlea of humerus|trochlea]] (elbow joints) of the humerus are poorly developed. Early archosauromorphs retain well-developed elbow joints, but all archosauromorphs apart from ''Aenigmastropheus'' have a trochlea ([[ulna]] joint) which is shifted towards the outer surface of the humerus, rather than the midpoint of the elbow as in other reptiles. In conjunction with this shift, the [[Olecranon|olecranon process]] of the ulna is poorly developed in archosauromorphs apart from ''Aenigmastropheus'' and ''Protorosaurus''.<ref name=":1" />
 
=== Hindlimbs ===
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|label2=[[Archelosauria]]/ |sublabel2='''Archosauromorpha ''[[sensu lato|s. l.]]'''''
|2={{clade
|label1='''[[Pan-Archosauria'''Testudines]]/ |sublabel1=[[Pantestudines]]
|1={{clade
|1={{extinct}}[[Choristodera]][[File:Champsosaurus BW flipped.jpg|80px]]
|label2='''Archosauromorpha ''[[sensu stricto|s. s.]]'''''
|2={{clade
|1={{extinct}}[[Prolacertiformes]] [[File:Prolacerta_broomi.jpg|50px]]
|2={{clade
|1={{clade
|1={{extinct}}''[[Trilophosaurus]]''[[File:Trilophosaurus buettneri (flipped).jpg|80px]]
|2={{extinct}}[[Rhynchosauria]][[File:Hyperodapedon BW2 white background.jpg|80px]] }}
|2=[[Archosauriformes]]<span style="{{MirrorH}}">[[File:Deinosuchus riograndensis.png|80px]]</span> }} }} }}
|label2=[[Pan-Testudines]]/ |sublabel2=[[Pantestudines]]
|2={{clade
|1={{extinct}}[[Eosauropterygia]][[File:Dolichorhynchops BW flipped.jpg|80px]]
|2={{clade
Line 111 ⟶ 103:
|2={{clade
|1={{extinct}}''[[Odontochelys]]''
|label2=&nbsp;[[Testudinata]]&nbsp;
|2={{clade
|1={{extinct}}''[[Proganochelys]]''
|2=[[Testudines]][[File:Erpétologie générale, ou, Histoire naturelle complète des reptiles (Centrochelys sulcata).jpg|70px]]
}} }} }} }} }} }} }} }}
|label2=[['''Pan-Testudines]]/ |sublabel2=[[Pantestudines]]Archosauria'''
|2={{clade
|1={{extinct}}[[Choristodera]][[File:Champsosaurus BW flipped.jpg|80px]]
|label2='''Archosauromorpha ''[[sensu stricto|s. s.]]'''''
|2={{clade
|1={{extinct}}[[Prolacertiformes]] [[File:Prolacerta_broomi.jpg|50px]]
|2={{clade
|1={{clade
|1={{extinct}}''[[Trilophosaurus]]''[[File:Trilophosaurus buettneri (flipped).jpg|80px]]
|2={{extinct}}[[Rhynchosauria]][[File:Hyperodapedon BW2 white background.jpg|80px]] }}
|2=[[Archosauriformes]]<span style="{{MirrorH}}">[[File:Deinosuchus riograndensis.png|80px]]</span> }} }} }}
}} }} }}
 
The following cladogram is based on a large analysis of [[Archosauriformes|archosauriform]]s published by M.D. Ezcurra in 2016.<ref name=":1" />
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|1=[[Lepidosauromorpha]][[File:British reptiles, amphibians, and fresh-water fishes (1920) (Lacerta agilis).jpg|80px]]
|2={{extinct}}[[Choristodera]][[File:Champsosaurus BW flipped.jpg|80px]]
|label3=&nbsp;'''Archosauromorpha'''&nbsp;
|3={{clade
|1={{clade
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|2={{clade
|1={{extinct}}[[Tanystropheidae]] [[File:Macrocnemus BW.jpg|80px]]
|label2=&nbsp;'''Crocopoda'''&nbsp;
|2={{clade
|label1={{extinct}}[[Allokotosauria]]
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[[Category:Guadalupian first appearances]]
[[Category:Taxa named by Friedrich von Huene]]
[[Category:Archelosauria]]