{{Short description|Paraphyletic grouping of reptiles}} {{For|the extinct fish|Anaspid}} thumb|Generic diagram of an anapsid reptile skull An '''anapsid''' is an amniote whose skull lacks one or more skull openings (fenestra, or fossae) near the temples.<ref>Pough, F. H. et al. (2002) ''Vertebrate Life'', 6th ed. Upper Saddle River, New Jersey: Prentice Hall. {{ISBN|0-13-041248-1}}</ref> Traditionally, the Anapsida are considered the most primitive subclass of amniotes, the ancestral stock from which Synapsida and Diapsida evolved, making anapsids paraphyletic. It is, however, doubtful that all anapsids lack temporal fenestra as a primitive trait, and that all the groups traditionally seen as anapsids truly lacked fenestra.
==Anapsids and the turtles== [[File:Caretta carettaZZ.jpg|thumb|left|Anapsid skull of ''Caretta caretta'' (loggerhead sea turtle), a testudine]]While "anapsid reptiles" or "Anapsida" were traditionally spoken of as if they were a monophyletic group, it has been suggested that several groups of reptiles that had anapsid skulls might be only distantly related. Scientists still debate the exact relationship between the basal (original) reptiles that first appeared in the late Carboniferous, the various Permian reptiles that had anapsid skulls, and the Testudines (turtles, tortoises, and terrapins). However, it was later suggested that the anapsid-like turtle skull is due to reversion rather than to anapsid descent. The majority of modern paleontologists believe that the Testudines are descended from diapsid reptiles that lost their temporal fenestrae. More recent morphological phylogenetic studies with this in mind placed turtles firmly within diapsids,<ref name=debraga&rieppel1997>{{cite journal | last1 = deBraga | first1 = M. | last2 = Rieppel | first2 = O. | year = 1997 | title = Reptile phylogeny and the interrelationships of turtles | url = https://academic.oup.com/zoolinnean/article-pdf/120/3/281/16872152/j.1096-3642.1997.tb01280.x.pdf| 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><ref name=Parareptilia1>{{cite journal |first1=Linda A. |last1=Tsuji |first2=Johannes |last2=Muller |year=2009 |title=Assembling the history of the Parareptilia: phylogeny, diversification, and a new definition of the clade |journal=Fossil Record |volume=12 |issue=1 |pages=71–81 |doi=10.1002/mmng.200800011|doi-access=free |bibcode=2009FossR..12...71T }}</ref><ref name=Parareptilia2>{{cite journal |first1=Marcello |last1=Ruta |first2=Juan C. |last2=Cisneros |first3=Torsten |last3=Liebrect |first4=Linda A. |last4=Tsuji |first5=Johannes |last5=Muller |year=2011 |title=Amniotes through major biological crises: faunal turnover among Parareptiles and the end-Permian mass extinction |journal=Palaeontology |volume=54 |issue=5 |pages=1117–1137 |doi=10.1111/j.1475-4983.2011.01051.x |s2cid=83693335 |doi-access=free |bibcode=2011Palgy..54.1117R }}</ref><ref name="Evans2009">{{Cite journal|first=Susan E. |last=Evans|year=2009|title=An early kuehneosaurid reptile (Reptilia: Diapsida) from the Early Triassic of Poland|journal=Paleontologica Polonica|volume=65|pages=145–178|url=http://palaeontologia.pan.pl/PP65/PP65_145-178.pdf}}.</ref><ref name="Borsuk−Białynicka2009">{{Cite journal|first1=Magdalena |last1=Borsuk−Białynicka |first2=Susan E. |last2=Evans|year=2009|title=A long−necked archosauromorph from the Early Triassic of Poland|journal=Paleontologica Polonica|volume=65|pages=203–234|url=http://palaeontologia.pan.pl/PP65/PP65_203-234.pdf}}</ref> most commonly within Archelosauria.<ref name="ReferenceA">{{Cite journal|last1=Field|first1=Daniel J.|last2=Gauthier|first2=Jacques A.|last3=King|first3=Benjamin L.|last4=Pisani|first4=Davide|last5=Lyson|first5=Tyler R.|last6=Peterson|first6=Kevin J.|date=July 2014|title=Toward consilience in reptile phylogeny: miRNAs support an archosaur, not lepidosaur, affinity for turtles: Reptile phylogeny from miRNAs|journal=Evolution & Development|language=en|volume=16|issue=4|pages=189–196|doi=10.1111/ede.12081|pmc=4215941|pmid=24798503}}</ref>
==Phylogenetic position of turtles== All molecular studies have strongly upheld the placement of turtles within diapsids; some place turtles within Archosauria,<ref name = "Mannen">{{Cite journal|last1 = Mannen|first1 = Hideyuki|last2=Li |first2=Steven S.-L. |title = Molecular evidence for a clade of turtles|journal = Molecular Phylogenetics and Evolution|volume = 13|issue = 1|pages = 144–148| date = October 1999|doi = 10.1006/mpev.1999.0640 | pmid = 10508547| bibcode=1999MolPE..13..144M }}</ref> or, more commonly, as a sister group to extant archosaurs.<ref name = "Zardoya">{{cite journal| last1=Zardoya| first1=R.| last2=Meyer |first2=A.| year=1998|title=Complete mitochondrial genome suggests diapsid affinities of turtles| journal=Proceedings of the National Academy of Sciences| issn=0027-8424| volume=95| issue=24| pages=14226–14231| doi=10.1073/pnas.95.24.14226| pmid=9826682| pmc=24355| bibcode=1998PNAS...9514226Z| doi-access=free}}</ref><ref name = "Iwabe">{{Cite journal | last1 = Iwabe | first1 = N. |last2=Hara |first2=Y.|last3=Kumazawa |first3=Y.|last4=Shibamoto |first4=K.|last5=Saito |first5=Y.|last6=Miyata |first6=T. |last7=Katoh |first7=K. | title = Sister group relationship of turtles to the bird-crocodilian clade revealed by nuclear DNA-coded proteins | journal = Molecular Biology and Evolution | volume = 22 | issue = 4 | pages = 810–813 | date = 2004-12-29 | doi = 10.1093/molbev/msi075 | pmid = 15625185 | doi-access = free }}</ref><ref name = "Roos">{{Cite journal | last1 = Roos | first1 = Jonas |last2= Aggarwal |first2=Ramesh K. |last3=Janke |first3=Axel | title = Extended mitogenomic phylogenetic analyses yield new insight into crocodylian evolution and their survival of the Cretaceous–Tertiary boundary | journal = Molecular Phylogenetics and Evolution | volume = 45 | issue = 2 | pages = 663–673 | date = November 2007 | doi = 10.1016/j.ympev.2007.06.018 | pmid = 17719245| bibcode = 2007MolPE..45..663R }}</ref><ref name = "Katsu">{{Cite journal | last1 = Katsu | first1 = Y. |last2= Braun |first2=E. L. |last3= Guillette |first3=L. J. Jr. |last4= Iguchi |first4=T. |title = From reptilian phylogenomics to reptilian genomes: analyses of c-Jun and DJ-1 proto-oncogenes|journal = Cytogenetic and Genome Research|volume = 127|issue = 2–4|pages = 79–93|date = 2010-03-17|doi = 10.1159/000297715 | pmid = 20234127| s2cid = 12116018 }}</ref><ref name=Crawford12>{{Cite journal|last1 = Crawford|first1 = N. G.|last2 = Faircloth|first2 = B. C.|last3 = McCormack| first3 = J. E.|last4 = Brumfield|first4 = R. T.|last5 = Winker|first5 = K.|last6 = Glenn|first6 = T. C.|title = More than 1000 ultraconserved elements provide evidence that turtles are the sister group of archosaurs|doi = 10.1098/rsbl.2012.0331|journal = Biology Letters|year = 2012|pmid = 22593086| pmc = 3440978 | volume=8 | issue=5 | pages=783–786}}</ref> One molecular study, published in 2012, suggests that turtles are lepidosauromorph diapsids, most closely related to the lepidosaurs (lizards, snakes, and tuataras).<ref name ="turtlelizard">{{Cite journal |first1=Tyler R. |last1=Lyson |first2=Erik A. |last2=Sperling |first3=Alysha M. |last3=Heimberg |first4=Jacques A. |last4=Gauthier |first5=Benjamin L. |last5=King |first6=Kevin J. |last6=Peterson | title = MicroRNAs support a turtle + lizard clade | journal = Biology Letters | volume = 8 | issue = 1 | pages = 104–107 | doi = 10.1098/rsbl.2011.0477 | pmid=21775315 | pmc=3259949|date=2012-02-23 |bibcode=2012BiLet...8..104L }}</ref> However, in a later paper from the same authors, published in 2014, based on more extensive data, the archosauromorph hypothesis is supported.<ref name="ReferenceA"/>
Reanalysis of prior phylogenies suggests that they classified turtles as anapsids both because they assumed this classification (most of them were studying what sort of anapsid turtles are) and because they did not sample fossil and extant taxa broadly enough for constructing the cladogram. Testudines is suggested to have diverged from other diapsids between 200 and 279 million years ago, though the debate is far from settled.<ref name = "Rieppel">{{cite journal |vauthors=Rieppel O, DeBraga M |title=Turtles as diapsid reptiles |journal=Nature |volume=384 |issue= 6608|pages=453–455 |year=1996 |doi=10.1038/384453a0 |bibcode=1996Natur.384..453R |s2cid=4264378 |url=https://doc.rero.ch/record/16242/files/PAL_E3477.pdf }}</ref><ref name = "Zardoya"/><ref>{{cite book|last=Benton|first=M. J.|title=Vertebrate Paleontology|edition=2nd|publisher=Blackwell Science|location=London|year=2000|isbn=978-0-632-05614-9|title-link=Vertebrate Paleontology (Benton)}} 3rd edition (2004) {{ISBN|0-632-05637-1}}.</ref> Although procolophonids managed to survive into the Triassic, most of the other reptiles with anapsid skulls, including the millerettids, nycteroleterids, and pareiasaurs, became extinct in the Late Permian period by the Permian-Triassic extinction event.
==Anapsida in modern taxonomy== Anapsida is still sporadically recognized as a valid group, but is not favoured by current workers.<ref name=modestoanderson2004>{{cite journal | last1 = Modesto | first1 = S.P. | last2 = Anderson | first2 = J.S. | year = 2004 | title = The phylogenetic definition of Reptilia | journal = Systematic Biology | volume = 53 | issue = 5| pages = 815–821 | doi = 10.1080/10635150490503026 | doi-access = free | pmid = 15545258 }}</ref><ref name=TsujiMuller2009FR>{{cite journal |first1=Linda A. |last1=Tsuji |first2=Johannes |last2=Müller |year=2009 |title=Assembling the history of the Parareptilia: Phylogeny, diversification, and a new definition of the clade |journal=Fossil Record |volume=12 |issue=1 |pages=71–81 |doi=10.1002/mmng.200800011 |doi-access=free |bibcode=2009FossR..12...71T }}</ref> Anapsids in the traditional meaning of the word are not a clade, but rather a paraphyletic group composed of all the early reptiles retaining the primitive skull morphology, grouped together by the absence of temporal openings.<ref name=modestoanderson2004/><ref name=TsujiMuller2009FR/> {{harvp|Gauthier|Kluge|Rowe|1988}} attempted to redefine 'Anapsida' so it would be monophyletic, defining it as the clade containing "extant turtles and all other extinct taxa that are more closely related to them than they are to other reptiles".<ref name=GKT88>{{cite book |last1=Gauthier |first1=J.A. |last2=Kluge |first2=A.G. |last3=Rowe |first3=T. |year=1988 |chapter=The early evolution of the Amniota |title=The Phylogeny and Classification of the Tetrapods |volume=1 |editor-last=Benton |editor-first=M.J. |publisher=Clarendon Press |location=Oxford, UK |isbn=978-0198577058 |pages=103–155 }}</ref>
This definition explicitly includes turtles in Anapsida; because the phylogenetic placement of turtles within Amniota is very uncertain, it is unclear what taxa, other than turtles themselves, would be included in such defined Anapsida, and whether its content would be similar to the Anapsida of tradition. Indeed, {{harvp|Gauthier|Kluge|Rowe|1988}} themselves included only turtles and Captorhinidae in their Anapsida, while excluding the majority of clades from it that had been traditionally included under the older definition of anapsids.<ref name=GKT88/>
==Temporal openings in traditional anapsids== Tsuji and Müller (2009) noted that the name Anapsida implies a morphology (lack of temporal openings) that is in fact absent in the skeletons of a number of taxa traditionally included in the group.<ref name=TsujiMuller2009FR/> A temporal opening in the skull roof behind each eye, similar to that present in the skulls of synapsids, has been discovered in the skulls of a number of members of Parareptilia (the group containing most of reptiles traditionally referred to as anapsids), including lanthanosuchoids, millerettids, bolosaurids, some nycteroleterids, some procolophonoids and at least some mesosaurs.<ref name=TsujiMuller2009FR /><ref>{{cite journal |first1=Juan C. |last1=Cisneros |first2=Ross |last2=Damiani |first3=Cesar |last3=Schultz |first4=Átila |last4=da Rosa |first5=Cibele |last5=Schwanke |first6=Leopoldo W. |last6=Neto |first7=Pedro L. P. |last7=Aurélio |year=2004 |title=A procolophonoid reptile with temporal fenestration from the Middle Triassic of Brazil |journal=Proceedings of the Royal Society B: Biological Sciences |volume=271 |issue=1547 |pages=1541–1546 |doi=10.1098/rspb.2004.2748 |pmid=15306328 |pmc=1691751}}</ref><ref name=PineiroetalCRP2012>{{cite journal |first1=Graciela |last1=Piñeiro |first2=Jorge |last2=Ferigolo |first3=Alejandro |last3=Ramos |first4=Michel |last4=Laurin |year=2012 |title=Cranial morphology of the Early Permian mesosaurid ''Mesosaurus tenuidens'' and the evolution of the lower temporal fenestration reassessed |journal=Comptes Rendus Palevol |volume=11 |issue=5 |pages=379–391 |doi=10.1016/j.crpv.2012.02.001 |bibcode=2012CRPal..11..379P }}</ref> The presence of temporal openings in the skulls of these taxa makes it uncertain whether the ancestral reptiles had an anapsid-like skull as traditionally assumed or a synapsid-like skull instead.<ref name=PineiroetalCRP2012/>
==See also== * Euryapsida
== References == {{Reflist}}
==External links== {{Wikispecies|Anapsida}} *[https://www.ucmp.berkeley.edu/anapsids/anapsida.html Introduction to Anapsida] from UCMP
{{Sauropsida|E.}} {{Taxonbar|from=Q331974}}
Category:Reptile taxonomy Category:Paraphyletic groups