Editing Lizard (section)

==Evolution==

===Fossil history===
[[File:DalinghesaurusLongidigitus-PaleozoologicalMuseumOfChina-May23-08 (cropped).jpg|thumb|Fossil lizard ''[[Dalinghosaurus|Dalinghosaurus longidigitus]]'', [[Early Cretaceous]], China]]

The closest living relatives of lizards are [[rhynchocephalia]]ns, a once diverse order of reptiles, of which is there is now only one living species, the [[tuatara]] of New Zealand. Some reptiles from the Early and Middle [[Triassic]], like ''[[Sophineta]]'' and ''[[Megachirella]]'', are suggested to be [[stem-group]] squamates, more closely related to modern lizards than rhynchocephalians, however, their position is disputed, with some studies recovering them as less closely related to squamates than rhynchocephalians are.<ref>{{Cite journal |last1=Tałanda |first1=Mateusz |last2=Fernandez |first2=Vincent |last3=Panciroli |first3=Elsa |last4=Evans |first4=Susan E. |last5=Benson |first5=Roger J. |date=2022-11-03 |title=Synchrotron tomography of a stem lizard elucidates early squamate anatomy |url=https://www.nature.com/articles/s41586-022-05332-6 |journal=Nature |language=en |volume=611 |issue=7934 |pages=99–104 |doi=10.1038/s41586-022-05332-6 |pmid=36289329 |bibcode=2022Natur.611...99T |s2cid=253160713 |issn=0028-0836 |access-date=2022-12-31 |archive-date=2023-12-28 |archive-url=https://web.archive.org/web/20231228173131/https://www.nature.com/articles/s41586-022-05332-6 |url-status=live }}</ref> The oldest undisputed lizards date to the Middle Jurassic, from remains found In Europe, Asia and North Africa.<ref>{{Citation |last=Evans |first=Susan E. |title=The Origin and Early Diversification of Squamates |date=2022-08-11 |url=https://www.cambridge.org/core/product/identifier/9781108938891%23CN-bp-2/type/book_part |work=The Origin and Early Evolutionary History of Snakes |pages=7–25 |editor-last=Gower |editor-first=David J. |access-date=2024-01-10 |edition=1 |publisher=Cambridge University Press |doi=10.1017/9781108938891.004 |isbn=978-1-108-93889-1 |editor2-last=Zaher |editor2-first=Hussam|url-access=subscription }}</ref> Lizard morphological and ecological diversity substantially increased over the course of the [[Cretaceous]].<ref>{{Cite journal |last1=Herrera-Flores |first1=Jorge A. |last2=Stubbs |first2=Thomas L. |last3=Benton |first3=Michael J. |date=March 2021 |title=Ecomorphological diversification of squamates in the Cretaceous |journal=Royal Society Open Science |language=en |volume=8 |issue=3 |pages=rsos.201961, 201961 |doi=10.1098/rsos.201961 |issn=2054-5703 |pmc=8074880 |pmid=33959350|bibcode=2021RSOS....801961H }}</ref> In the [[Paleogene|Palaeogene]], lizard body sizes in North America peaked during the middle of the period.<ref>{{Cite journal |last=ElShafie |first=Sara J. |date=5 January 2024 |editor-last=Meloro |editor-first=Carlo |title=Body size estimation from isolated fossil bones reveals deep time evolutionary trends in North American lizards |journal=[[PLOS ONE]] |language=en |volume=19 |issue=1 |pages=e0296318 |doi=10.1371/journal.pone.0296318 |doi-access=free |issn=1932-6203 |pmc=10769094 |pmid=38180961 |bibcode=2024PLoSO..1996318E }}</ref>

[[Mosasaurs]] likely evolved from an extinct group of aquatic lizards<ref>{{cite AV media |last= Dash |first= Sean |title= Prehistoric Monsters Revealed |location= United States |publisher= Workaholic Productions / History Channel |date= 2008 |url= https://www.youtube.com/watch?v=lEEhm4rzxEg | archive-url=https://web.archive.org/web/20160127070909/https://www.youtube.com/watch?v=lEEhm4rzxEg| archive-date=2016-01-27|access-date= December 18, 2015}}</ref> known as [[Aigialosauridae|aigialosaur]]s in the [[Early Cretaceous]]. [[Dolichosauridae]] is a family of [[Late Cretaceous]] aquatic varanoid lizards closely related to the mosasaurs.<ref name=Primitivus>{{Cite journal|author1=Ilaria Paparella |author2=Alessandro Palci |author3=Umberto Nicosia |author4=Michael W. Caldwell |year=2018 |title=A new fossil marine lizard with soft tissues from the Late Cretaceous of southern Italy |journal=Royal Society Open Science |volume=5 |issue=6 |pages=172411 |doi=10.1098/rsos.172411 |pmid=30110414 |pmc=6030324 |bibcode=2018RSOS....572411P }}</ref><ref name="moremosa">{{Cite journal|date=1999-01-01|title=Squamate phylogeny and the relationships of snakes and mosasauroids|journal=Zoological Journal of the Linnean Society|volume=125|issue=1|pages=115–147|doi=10.1006/zjls.1997.0144|issn=0024-4082|last1=Caldwell|first1=M.|doi-access=free}}</ref>

===Phylogeny===

====External====

The position of the lizards and other [[Squamata]] among the reptiles was studied using fossil evidence by Rainer Schoch and Hans-Dieter Sues in 2015. Lizards form about 60% of the extant non-avian reptiles.<ref name=pappochelys2015>{{Cite journal | last1=Schoch | first1=Rainer R. | last2=Sues | first2=Hans-Dieter | title=A Middle Triassic stem-turtle and the evolution of the turtle body plan | journal=[[Nature (journal)|Nature]] | volume=523 | issue=7562 | pages=584–587 | doi=10.1038/nature14472 | date=24 June 2015 | pmid=26106865 | bibcode=2015Natur.523..584S | s2cid=205243837 }}</ref>
{{clade
|label1=[[Archelosauria]]
|1={{clade
   |1=[[Archosauromorpha]]<span style="{{MirrorH}}">[[File:Deinosuchus riograndensis.png|100px]]</span>[[File:Meyers grosses Konversations-Lexikon - ein Nachschlagewerk des allgemeinen Wissens (1908) (Antwerpener Breiftaube).jpg|40 px]]
   |label2=[[Lepidosauromorpha]]
   |2={{clade
      |1={{clade
         |1=[[Pantestudines]] [[File:Erpétologie générale, ou, Histoire naturelle complète des reptiles (Centrochelys sulcata).jpg|70px]]
         |2={{clade
            |1={{extinct}}[[Kuehneosauridae]][[File:Icarosaurus white background.jpg|80px]]
            |label2=[[Lepidosauria]]
            |2={{clade
               |1=[[Rhynchocephalia]][[File:Hatteria white background.jpg|80px]]
               |2='''[[Squamata]]'''[[File:British reptiles, amphibians, and fresh-water fishes (1920) (Lacerta agilis).jpg|80px]]
               }}
            }}
         }}
      }}
   }} 
}}

====Internal====
Both the snakes and the [[Amphisbaenia]] (worm lizards) are [[clade]]s deep within the [[Squamata]] (the smallest clade that contains all the lizards), so "lizard" is [[paraphyletic]].<ref>{{cite journal |last1=Reeder |first1=Tod W. |last2=Townsend |first2=Ted M. |last3=Mulcahy |first3=Daniel G. |last4=Noonan |first4=Brice P.|last5=Wood |first5=Perry L. |last6=Sites |first6=Jack W. |last7=Wiens |first7=John J.|title=Integrated Analyses Resolve Conflicts over Squamate Reptile Phylogeny and Reveal Unexpected Placements for Fossil Taxa |journal=PLOS ONE |date=2015 |volume=10 |issue=3 |pages=e0118199 |doi=10.1371/journal.pone.0118199 |pmid=25803280|pmc=4372529|bibcode=2015PLoSO..1018199R |doi-access=free }}</ref>
The cladogram is based on genomic analysis by Wiens and colleagues in 2012 and 2016.<ref name=wiensetal2012>{{cite journal | last1=Wiens | first1=J. J. | last2=Hutter | first2=C. R. | last3=Mulcahy | first3=D. G. | last4=Noonan | first4=B. P. | last5=Townsend | first5=T. M. | last6=Sites | first6=J. W. | last7=Reeder | first7=T. W. | year=2012 | title=Resolving the phylogeny of lizards and snakes (Squamata) with extensive sampling of genes and species | journal=Biology Letters | volume=8 | issue=6| pages=1043–1046 | doi=10.1098/rsbl.2012.0703 | pmid=22993238 | pmc=3497141}}</ref><ref name="zheng & wiens 2016">{{cite journal|last1=Zheng |first1=Yuchi|last2=Wiens |first2=John J. |title=Combining phylogenomic and supermatrix approaches, and a time-calibrated phylogeny for squamate reptiles (lizards and snakes) based on 52 genes and 4162 species |journal=Molecular Phylogenetics and Evolution |date=2016 |volume=94 |issue=Pt B|pages=537–547 |doi=10.1016/j.ympev.2015.10.009|pmid=26475614|bibcode=2016MolPE..94..537Z }}</ref> Excluded taxa are shown in upper case on the cladogram.

{{clade| style=font-size:85%;line-height:80%
|label1='''Squamata'''
|1={{clade
  |label1=[[Dibamia]]
  |1=[[Dibamidae]]
  |label2=[[Bifurcata]]
  |2={{clade
    |label1=[[Gekkota]]
    |1={{clade
      |label1=[[Pygopodomorpha]]
      |1={{clade
        |1=[[Diplodactylidae]] [[File:Hoplodactylus pomarii white background.jpg|70 px]]
        |2={{clade
          |1=[[Pygopodidae]] [[File:The zoology of the voyage of the H.M.S. Erebus and Terror (Lialis burtonis).jpg|70 px]]
          |2=[[Carphodactylidae]]
           }}
         }}
      |label2=[[Gekkomorpha]]
      |2={{clade
        |1=[[Eublepharidae]]
        |label2=[[Gekkonoidea]]
        |2={{clade
          |1=[[Sphaerodactylidae]] 
          |2={{clade
            |1=[[Phyllodactylidae]] [[File:Phyllodactylus gerrhopygus 1847 - white background.jpg|70 px]]
            |2=[[Gekkonidae]] [[File:G gecko 060517 6167 trij (washout).jpg|70px]]
             }}
           }}
         }}
       }}
    |label2=[[Unidentata]]
    |2={{clade
      |label1=[[Scincomorpha|Scinciformata]]
      |1={{clade
        |label1=[[Scincomorpha]]
        |1=[[Scincidae]] [[File:Bilder-Atlas zur wissenschaftlich-populären Naturgeschichte der Wirbelthiere (Plate (24)) Tribolonotus novaeguineae.jpg|70 px]]
        |label2=[[Cordylomorpha]]
        |2={{clade
          |1=[[Xantusiidae]]
          |2={{clade
            |1=[[Gerrhosauridae]] [[File:Gerrhosaurus ocellatus flipped.jpg|70 px]]
            |2=[[Cordylidae]] [[File:Illustrations of the zoology of South Africa (Smaug giganteus).jpg|70 px]]
             }}
           }}
         }}
      |label2=[[Episquamata]]
      |2={{clade
        |label1=[[Lacertoidea|Laterata]]
        |1={{clade
          |label1=[[Teiformata]]
          |1={{clade
            |1=[[Gymnophthalmidae]] [[File:PZSL1851PlateReptilia06 Cercosaura ocellata.png|70 px]]
            |2=[[Teiidae]] [[File:Bilder-Atlas zur wissenschaftlich-populären Naturgeschichte der Wirbelthiere (Tupinambis teguixin).jpg|70 px]]
             }}
          |label2=[[Lacertibaenia]]
          |2={{clade
            |label1=[[Lacertiformata]]
            |1=[[Lacertidae]] [[File:Brockhaus' Konversations-Lexikon (1892) (Lacerta agilis).jpg|70 px]]
            |2='''[[Amphisbaenia|AMPHISBAENIA]]''' (worm lizards, not usually considered "true lizards") [[File:Amphisbaena microcephalum 1847 - white background.jpg|70 px]]   
             }}
           }}
        |label2=[[Toxicofera]]
        |2={{clade
          |1={{clade
            |label1=[[Anguimorpha]]
            |1={{clade
              |label1=[[Palaeoanguimorpha]]
              |1={{clade
                |label1=[[Shinisauria]]
                |1=[[Shinisauridae]] 
                |label2=[[Varanoidea]]
                |2={{clade
                  |1=[[Lanthanotidae]]
                  |2=[[Varanidae]] [[File:Zoology of Egypt (1898) (Varanus griseus).png|90 px]]
                   }}
                 }}
              |label2=[[Neoanguimorpha]]
              |2={{clade
                |label1=[[Helodermatoidea]]
                |1=[[Helodermatidae]] [[File:Gila monster ncd 2012 white background.jpg|70 px]]
                |2={{clade
                  |label1=[[Xenosauroidea]]
                  |1=[[Xenosauridae]]
                  |label2=[[Anguioidea]]
                  |2={{clade
                    |1=[[Diploglossidae]]
                    |2={{clade
                      |1=[[Anniellidae]]
                      |2=[[Anguidae]] [[File:Anguis fragilis (cropped).jpg|60px]]
                       }}
                     }}
                   }}
                 }}
               }}
            |label2=[[Iguanomorpha]]
            |2={{clade
              |label1=[[Acrodonta (lizard)|Acrodonta]]
              |1={{clade
                |1=[[Chamaeleonidae]] [[File:Zoology of Egypt (1898) (Chamaeleo calyptratus).jpg|70 px]]
                |2=[[Agamidae]] [[File:Haeckel Lacertilia (Chlamydosaurus kingii).jpg|55 px]]
                 }}
              |label2=[[Pleurodonta]]
              |2={{clade
                |1=[[Leiocephalidae]]
                |2={{clade
                  |1=[[Iguanidae]] [[File:Stamps of Germany (Berlin) 1977, Cyclura cornuta.jpg|70 px]]
                  |2={{clade
                    |1={{clade
                      |1=[[Hoplocercidae]] 
                      |2={{clade
                        |1=[[Crotaphytidae]]
                        |2=[[Corytophanidae]] [[File:SDC10934 - Basiliscus plumifrons (extracted).JPG|70px]]
                         }}
                       }}
                    |2={{clade
                      |1=[[Tropiduridae]]
                      |2={{clade
                        |1={{clade
                          |1=[[Phrynosomatidae]]
                          |2={{clade
                            |1=[[Dactyloidae]]
                            |2=[[Polychrotidae]]
                             }}
                           }}
                        |2={{clade
                          |1=[[Liolaemidae]]
                          |2={{clade
                            |1=[[Leiosauridae]]
                            |2=[[Opluridae]]
                             }}
                           }}
                         }}
                       }}
                     }}
                   }}
                 }}
               }}
             }}
          |2='''[[Serpentes|SERPENTES]]''' (snakes, not considered to be lizards) [[File:Python natalensis Smith 1840 white background.jpg|120 px]]

           }}
         }}
       }}
     }}
   }}
}}

===Taxonomy===
{{Main|List of Lacertilia families}}
[[Image:Prognathodon3.jpg|thumb|Artistic restoration of a mosasaur, ''[[Prognathodon]]'']]
In the 13th century, lizards were recognized in Europe as part of a broad category of ''reptiles'' that consisted of a miscellany of egg-laying creatures, including "snakes, various fantastic monsters, […], assorted amphibians, and worms", as recorded by [[Vincent of Beauvais]] in his ''Mirror of Nature''.<ref>{{cite book | last = Franklin-Brown | first = Mary | title = Reading the world : encyclopedic writing in the scholastic age | publisher = The University of Chicago Press | location = Chicago London | year = 2012 | isbn = 9780226260709|page=223;377}}</ref> The seventeenth century saw changes in this loose description. The name [[Sauria]] was coined by [[James Macartney (anatomist)|James Macartney]] (1802);<ref>James Macartney: Table III ''in'': George Cuvier (1802) "Lectures on Comparative Anatomy" (translated by William Ross under the inspection of James Macartney). Vol I. London, Oriental Press, Wilson and Co.</ref> it was the Latinisation of the French name ''Sauriens'', coined by [[Alexandre Brongniart]] (1800) for an order of reptiles in the classification proposed by the author, containing lizards and [[crocodilia]]ns,<ref>Alexandre Brongniart (1800) "Essai d'une classification naturelle des reptiles. 1ère partie: Etablissement des ordres." Bulletin de la Science. Société Philomathique de Paris 2 (35): 81–82</ref> later discovered not to be each other's closest relatives. Later authors used the term "Sauria" in a more restricted sense, i.e. as a synonym of Lacertilia, a suborder of [[Squamata]] that includes all lizards but excludes [[snake]]s. This classification is rarely used today because Sauria so-defined is a [[paraphyletic]] group. It was defined as a [[clade]] by [[Jacques Gauthier]], Arnold G. Kluge and Timothy Rowe (1988) as the group containing the most recent common ancestor of [[archosaur]]s and [[Lepidosauria|lepidosaurs]] (the groups containing crocodiles and lizards, as per Mcartney's original definition) and all its descendants.<ref>{{Cite journal| last=Gauthier | first=J. A. | author-link=Jacques Gauthier |author2=Kluge, A. G. |author3=Rowe, T. | title=Amniote phylogeny and the importance of fossils | journal=[[Cladistics (journal)|Cladistics]] | volume=4 | issue=2 | pages=105–209 | date=June 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> A different definition was formulated by Michael deBraga and Olivier Rieppel (1997), who defined Sauria as the clade containing the most recent common ancestor of [[Choristodera]], [[Archosauromorpha]], [[Lepidosauromorpha]] and all their descendants.<ref>{{cite journal |author1=Debraga, M. |author2=Rieppel, O. |name-list-style=amp | year=1997 | title=Reptile phylogeny and the interrelationships of turtles | journal=Zoological Journal of the Linnean Society | volume=120 | issue=3 | pages=281–354 | doi=10.1111/j.1096-3642.1997.tb01280.x | doi-access=free }}</ref> However, these uses have not gained wide acceptance among specialists.

{{colbegin}}
Suborder Lacertilia (Sauria) – (lizards) 
* Family †[[Bavarisauridae]]
* Family †[[Eichstaettisauridae]]
* Infraorder [[Iguanomorpha]]
** Family †[[Arretosauridae]]
** Family †[[Euposauridae]]
** Family [[Corytophanidae]] (casquehead lizards)
** Family [[Iguanidae]] ([[iguana]]s and [[spinytail iguanas]])
** Family [[Phrynosomatidae]] ([[Earless lizard|earless]], [[spiny lizard|spiny]], [[Urosaurus|tree]], [[side-blotched lizard|side-blotched]] and [[horned lizard|horned]] lizards)
** Family [[Polychrotidae]] ([[anole]]s)
*** Family [[Leiosauridae]] (see Polychrotinae)
** Family [[Tropiduridae]] (neotropical ground lizards)
*** Family [[Liolaemidae]] (see Tropidurinae)
*** Family [[Leiocephalidae]] (see Tropidurinae)
** Family [[Crotaphytidae]] ([[crotaphytus|collared]] and [[gambelia|leopard]] lizards)
** Family [[Opluridae]] (Madagascar iguanids)
** Family [[Hoplocercidae]] (wood lizards, clubtails)
** Family †[[Priscagamidae]]
** Family †[[Isodontosauridae]]
** Family [[Agamidae]] ([[Agama (lizard)|agamas]], [[Chlamydosaurus|frilled lizard]]s)
** Family [[Chamaeleonidae]] ([[chameleon]]s)
* Infraorder [[Gekkota]]
** Family [[Gekkonidae]] ([[gecko]]s)
** Family [[Pygopodidae]] (legless geckos)
** Family [[Dibamidae]] (blind lizards)

<!--gap is to allow columns to break here naturally, col-break only works on some browsers-->
* Infraorder [[Scincomorpha]]
** Family †[[Paramacellodidae]]
** Family †[[Slavoiidae]]
** Family [[Scincidae]] (skinks)
** Family [[Cordylidae]] (spinytail lizards)
** Family [[Gerrhosauridae]] (plated lizards)
** Family [[Xantusiidae]] (night lizards)
** Family [[Lacertidae]] (wall lizards or true lizards)
** Family †[[Mongolochamopidae]]
** Family †[[Adamisauridae]]
** Family [[Teiidae]] ([[tegu]]s and whiptails)
** Family [[Gymnophthalmidae]] (spectacled lizards)
* Infraorder [[Diploglossa]]
** Family [[Anguidae]] (slowworms, glass lizards)
** Family [[Anniellidae]] (American legless lizards)
** Family [[Xenosauridae]] (knob-scaled lizards)
* Infraorder [[Platynota]] ([[Varanoidea]])
** Family [[Varanidae]] (monitor lizards)
** Family [[Lanthanotidae]] (earless monitor lizards)
** Family [[Helodermatidae]] ([[Gila monster]]s and [[Mexican beaded lizard|beaded lizards]])
** Family †[[Mosasaur]]idae (marine lizards)
{{colend}}

[[File:Anguidae.jpg|thumb|The slowworms, ''[[Anguis]]'', are among over twenty groups of lizards that have [[convergent evolution|convergently evolved]] a legless [[body plan]].<ref name=Brandley/>]]

===Convergence===

Lizards have frequently [[convergent evolution|evolved convergently]], with multiple groups independently developing similar morphology and [[ecological niche]]s. [[Anolis ecomorphs|''Anolis'' ecomorphs]] have become a model system in evolutionary biology for studying convergence.<ref>{{cite journal |last1=Losos |first1=Jonathan B. |title=The Evolution of Convergent Structure in Caribbean Anolis Communities |journal=Systematic Biology |date=1992 |volume=41 |issue=4 |pages=403–420 |doi=10.1093/sysbio/41.4.403}}</ref> Limbs have been lost or reduced independently [[legless lizard|over two dozen times across lizard evolution]], including in the [[Anniellidae]], [[Anguidae]], [[Cordylidae]], [[Dibamidae]], [[Gymnophthalmidae]], [[Pygopodidae]], and [[Scincidae]]; snakes are just the most famous and species-rich group of Squamata to have followed this path.<ref name=Brandley>{{cite journal |last1=Brandley |first1=Matthew C.|display-authors=etal|title=Rates And Patterns In The Evolution Of Snake-Like Body Form In Squamate Reptiles: Evidence For Repeated Re-Evolution Of Lost Digits And Long-Term Persistence Of Intermediate Body Forms |journal=Evolution |date=August 2008 |volume=62 |issue=8 |pages=2042–2064 |doi=10.1111/j.1558-5646.2008.00430.x|pmid=18507743 |s2cid=518045|doi-access=free }}</ref>