Procolophonidae

{{Short description|Extinct family of reptiles}} {{Automatic taxobox | taxon = Procolophonidae | fossil_range = {{fossilrange|Late Permian|Late Triassic}} | image = Kapes_full_skeleton.jpg | image_caption = | image2 = Kapes bentoni life restoration.jpg | image2_caption = Skeleton (top) and life restoration (bottom) of ''[[Kapes bentoni]]'' (Procolophoninae) scale bar = 1cm | authority = [[Harry Seeley|Seeley]], 1888 | subdivision_ranks = Subgroups and Genera | subdivision = * {{extinct}}''[[Coletta (reptile)|Coletta]]'' * {{extinct}}''[[Gomphiosauridion]]''? * {{extinct}}''[[Kinelia]]''? * {{extinct}}''[[Kitchingnathus]]'' * {{extinct}}''[[Lasasaurus]]'' * {{extinct}}''[[Oryporan]]'' * {{extinct}}''[[Phaanthosaurus]]'' * {{extinct}}''[[Pintosaurus]]'' * {{extinct}}''[[Sauropareion]]'' * {{extinct}}''[[Sauropia]]''? * {{extinct}}''[[Smilodonterpeton]]'' * {{extinct}}''[[Sphenosaurus]]'' * {{extinct}}''[[Suchonosaurus]]'' * {{extinct}}''[[Spondylolestes]]''? * {{extinct}}''[[Tichvinskia]]'' * {{extinct}}''[[Xenodiphyodon]]''? * {{extinct}}''[[Youngetta]]'' * {{extinct}}'''[[Leptopleuroninae]]''' * {{extinct}}'''[[Procolophoninae]]''' * {{extinct}}'''[[Theledectinae]]''' | synonyms = * Sclerosauridae <small>Nopcsa, 1923</small><ref name=CJC08>{{Cite journal | last1 = Cisneros | first1 = J. C. | title = Phylogenetic relationships of procolophonid parareptiles with remarks on their geological record | doi = 10.1017/S1477201907002350 | journal = Journal of Systematic Palaeontology | volume = 6 | issue = 3 | pages = 345–366 | year = 2008 | bibcode = 2008JSPal...6..345C | s2cid = 84468714 }}</ref><ref name=Sclerosaurus08>{{Cite journal | author = Hans-Dieter Sues and Robert R. Reisz | title = Anatomy and Phylogenetic Relationships of ''Sclerosaurus armatus'' (Amniota: Parareptilia) from the Buntsandstein (Triassic) of Europe | journal = Journal of Vertebrate Paleontology | volume = 28 | issue = 4 | pages = 1031–1042 | year = 2008 | doi=10.1671/0272-4634-28.4.1031| bibcode = 2008JVPal..28.1031S | s2cid = 53967912 }}</ref> }} [[File:Kapes skull.jpg|left|thumb|Skull of ''[[Kapes bentoni]]'']] '''Procolophonidae''' is an [[Extinction|extinct]] [[Family (taxonomy)|family]] of small, lizard-like [[Procolophonia|procolophonian]] "[[parareptile]]s" known from the [[Late Permian]] to [[Late Triassic]] that were distributed across [[Pangaea]], having been reported from Europe, North America, China, South Africa, South America, [[Antarctica]], and Australia.

== Ecology == The most primitive procolophonids were likely [[Insectivore|insectivorous]] or [[Omnivore|omnivorous]], while more [[Derived (phylogenetics)|derived]] members of the [[clade]] developed [[Cusp (anatomy)|bicusped]] [[Molar (tooth)|molars]], likely either [[Herbivore|herbivores]] feeding on high fiber vegetation or [[Durophagy|durophagous]] omnivores.<ref>{{Cite journal |last1=Pinheiro |first1=Felipe L. |last2=Silva-Neves |first2=Eduardo |last3=Da-Rosa |first3=Átila A. S. |date=August 2021 |editor-last=Ruta |editor-first=Marcello |title=An early-diverging procolophonid from the lowermost Triassic of South America and the origins of herbivory in Procolophonoidea |url=https://onlinelibrary.wiley.com/doi/10.1002/spp2.1355 |journal=[[Papers in Palaeontology]] |language=en |volume=7 |issue=3 |pages=1601–1612 |doi=10.1002/spp2.1355 |bibcode=2021PPal....7.1601P |s2cid=233797716 |issn=2056-2799 |url-access=subscription |archive-date=10 July 2021 |access-date=20 June 2022 |archive-url=https://web.archive.org/web/20210710001355/https://onlinelibrary.wiley.com/doi/10.1002/spp2.1355 |url-status=live }}</ref> Many members of the group are noted for spines projecting from the [[quadratojugal bone]] of the skull, which likely served a defensive purpose as well as possibly also for display.<ref>{{Cite journal |last1=Zaher |first1=Marta |last2=Coram |first2=Robert A. |last3=Benton |first3=Michael J. |date=February 2019 |editor-last=Angielczyk |editor-first=Kenneth |title=The Middle Triassic procolophonid Kapes bentoni: computed tomography of the skull and skeleton |journal=[[Papers in Palaeontology]] |language=en |volume=5 |issue=1 |pages=111–138 |doi=10.1002/spp2.1232|s2cid=134058607 |doi-access=free |bibcode=2019PPal....5..111Z |hdl=1983/3dd2d71d-a439-404a-997f-758063f40678 |hdl-access=free }}</ref> At least some taxa were likely [[fossorial]] burrowers.<ref>{{Cite journal |last1=Botha-Brink |first1=Jennifer |last2=Smith |first2=Roger Malcolm Harris |date=September 2012 |title=Palaeobiology of Triassic procolophonids, inferred from bone microstructure |url=https://linkinghub.elsevier.com/retrieve/pii/S1631068312000735 |journal=[[Comptes Rendus Palevol]] |language=en |volume=11 |issue=6 |pages=419–433 |doi=10.1016/j.crpv.2012.03.002|bibcode=2012CRPal..11..419B |url-access=subscription }}</ref>

== Evolution == Procolophonids radiated in the aftermath of the [[Permian-Triassic extinction event]], adapting to changes in the availability of different food resources.<ref>{{Cite journal |last1=Burgardt |first1=Caroline D’Ávila |last2=Pohlmann |first2=Karine |last3=Fabrício Machado |first3=Arielli |last4=Santos |first4=Mateus A. Costa |last5=Meira |first5=João L. |last6=Pinheiro |first6=Felipe L. |date=12 January 2026 |title=A peculiar procolophonid lower jaw from the Sanga do Cabral Formation (Lower Triassic, Brazil) |url=https://www.tandfonline.com/doi/full/10.1080/08912963.2025.2608698#d1e642 |journal=[[Historical Biology]] |language=en |pages=1–10 |doi=10.1080/08912963.2025.2608698 |issn=0891-2963 |access-date=18 March 2026 |via=Taylor and Francis Online|url-access=subscription }}</ref> While diverse during the [[Early Triassic|Early]] and [[Middle Triassic]], they had very low diversity during the Late Triassic, and were extinct by the beginning of the [[Jurassic]].<ref>{{Cite journal |last1=MacDougall |first1=Mark J. |last2=Brocklehurst |first2=Neil |last3=Fröbisch |first3=Jörg |date=20 March 2019 |title=Species richness and disparity of parareptiles across the end-Permian mass extinction |journal=[[Proceedings of the Royal Society B: Biological Sciences]] |volume=286 |issue=1899 |article-number=20182572 |doi=10.1098/rspb.2018.2572 |pmid=30890099 |pmc=6452079 |issn=0962-8452}}</ref>

==Phylogeny== The family is defined as all taxa more closely related to ''[[Procolophon trigoniceps]]'' than to ''[[Owenetta rubidgei]]''.<ref name=CJC08/> Below is a [[cladogram]] from Ruta ''et al.'' (2011):<ref name=RCLTM11>{{Cite journal | last1 = Ruta | first1 = M. | last2 = Cisneros | first2 = J. C. | last3 = Liebrecht | first3 = T. | last4 = Tsuji | first4 = L. A. | last5 = Müller | first5 = J. | title = Amniotes through major biological crises: Faunal turnover among Parareptiles and the end-Permian mass extinction | doi = 10.1111/j.1475-4983.2011.01051.x | journal = [[Palaeontology (journal)|Palaeontology]] | volume = 54 | issue = 5 | pages = 1117–1137 | year = 2011 | s2cid = 83693335 | doi-access = free | bibcode = 2011Palgy..54.1117R }}</ref>

{{clade| style=font-size:85%;line-height:85% |label1=Procolophonidae |1={{clade |1=''[[Coletta (reptile)|Coletta seca]]'' |2=''[[Pintosaurus magnidentis]]'' |3=''[[Sauropareion anoplus]]'' |4=''[[Kitchingnathus untabeni]]'' |5={{clade |1=''[[Phaanthosaurus ignatjevi]]'' |2=''[[Phaanthosaurus simus]]''}} |6={{clade |label1=[[Theledectinae]] |1={{clade |1=''[[Eumetabolodon dongshengensis]]'' |2=''[[Theledectes perforatus]]'' }} |2={{clade |1=''[[Tichvinskia vjatkensis]]'' |2={{clade |label1=[[Leptopleuroninae]] |1={{clade |1=''[[Pentaedrusaurus ordosianus]]'' |2={{clade |1=''[[Neoprocolophon asiaticus]]'' |2={{clade |1={{clade |1=''[[Sclerosaurus armatus]]'' |2=''[[Scoloparia glyphanodon]]''}} |2={{clade |1=''[[Leptopleuron lacertinum]]'' |2={{clade |1=''[[Soturnia caliodon]]'' |2=''[[Hypsognathus fenneri]]''}} }} }} }} }} |label2=[[Procolophoninae]] |2={{clade |1={{clade |1=''[[Eumetabolodon bathycephalus]]'' |2={{clade |1=''[[Procolophon trigoniceps]]'' |2=''[[Teratophon spinigenis]]'' |3=''[[Thelerpeton oppressus]]'' }} }} |2={{clade |1=''[[Timanophon raridentatus]]'' |2={{clade |1=''[[Thelephon contritus]]'' |2={{clade |1=''[[Anomoiodon liliensterni]]'' |2={{clade |1=''[[Kapes amaenus]]'' |2=''[[Kapes bentoni]]'' |3=''[[Kapes komiensis]]'' |4=''[[Kapes majmesculae]]'' }} }} }} }} }} }} }} }} }} }}

Below are three cladograms that follow [[phylogenetic analyses]] by Butler ''et al''. (2023). '''Analysis 1:''' Strict consensus of 760 [[maximum parsimony (phylogenetics)|most parsimonious trees]] (MPTs):<ref name=Butler2023>{{Cite journal |last1=Butler |first1=R. J. |last2=Meade |first2=L. E. |last3=Cleary |first3=T. J. |last4=McWhirter |first4=K. T. |last5=Brown |first5=E. E. |last6=Kemp |first6=T. S. |last7=Benito |first7=J. |last8=Fraser |first8=N. C. |year=2023 |title=''Hwiccewyrm trispiculum'' gen. et sp. nov., a new leptopleuronine procolophonid from the Late Triassic of southwest England |journal=[[The Anatomical Record]] |doi=10.1002/ar.25316 |pmid=37735997 |doi-access=free}}</ref>

{{clade| style=font-size:85%;line-height:85% |label1='''Procolophonidae''' |1={{clade |1=''[[Eomurruna]]'' |2=''[[Procolina]]'' |3=''[[Anomoiodon]]'' |4=''[[Lasasaurus]]'' |5=''[[Kitchingnathus]]'' |6=''[[Phonodus]]'' |7=''[[Thelephon]]'' |8=''[[Kapes bentoni]]'' |9=''[[Kapes majmesculae]]'' |10=''[[Timanophon]]'' |11=''[[Tichvinskia]]'' |12=''[[Theledectes]]'' |13=''[[Eumetabolodon dongshengensis]]'' |14=''[[Phaantosaurus]]'' ([[sic]]) |15=''[[Sauropareion]]'' |16=''[[Pintosaurus]]'' |17=''[[Coletta (reptile)|Coletta]]'' |18={{clade |1=''[[Eumetabolodon bathycephalus]]'' |label2=[[Procolophoninae]] |2={{clade |1=''[[Teratophon]]'' |2=''[[Thelerpeton]]'' |3=''[[Procolophon]]'' }} |label3=[[Leptopleuroninae]] |3={{clade |1=''[[Neoprocolophon]]'' |2=''[[Pentaedrusaurus]]'' |3={{clade |1=''[[Mandaphon]]'' |2={{clade |1={{clade |1=''[[Scoloparia]]'' |2=''[[Sclerosaurus]]'' }} |2={{clade |1=''[[Hwiccewyrm]]'' |2=''[[Hypsognathus]]'' |3=''[[Soturnia]]'' |4=''[[Leptopleuron]]'' }} }} }} }} }} }} }}

'''Analysis 2:''' Single MPT:<ref name=Butler2023/> {{clade| style=font-size:85%;line-height:85% |label1='''Procolophonidae''' |1={{clade |1=''[[Coletta (reptile)|Coletta]]'' |2={{clade |1=''[[Lasasaurus]]'' |2={{clade |1=''[[Sauropareion]]'' |2={{clade |1={{clade |1=''[[Kitchingnathus]]'' |2=''[[Pintosaurus]]'' }} |2={{clade |1=''[[Phaantosaurus]]'' ([[sic]]) |2={{clade |1={{clade |1=''[[Eomurruna]]'' |2=''[[Eumetabolodon dongshengensis]]'' }} |2={{clade |1=''[[Timanophon]]'' |2={{clade |1=''[[Tichvinskia]]'' |2={{clade |label1=[[Procolophoninae]] |1={{clade |1=''[[Eumetabolodon|Eumetabolodon bathycephalus]]'' |2={{clade |1=''[[Thelerpeton]]'' |2={{clade |1=''[[Teratophon]]'' |2=''[[Procolophon]]'' }} }} }} |label2=[[Leptopleuroninae]] |2={{clade |1={{clade |1=''[[Kapes bentoni]]'' |2={{clade |1=''[[Thelephon]]'' |2=''[[Kapes majmesculae]]'' }} }} |2={{clade |1=''[[Pentaedrusaurus]]'' |2={{clade |1={{clade |1=''[[Anomoiodon]]'' |2={{clade |1=''[[Theledectes]]'' |2={{clade |1=''[[Procolina]]'' |2=''[[Phonodus]]'' }} }} }} |2={{clade |1=''[[Mandaphon]]'' |2={{clade |1=''[[Neoprocolophon]]'' |2={{clade |1={{clade |1=''[[Scoloparia]]'' |2=''[[Sclerosaurus]]'' }} |2={{clade |1=''[[Hwiccewyrm]]'' |2={{clade |1=''[[Leptopleuron]]'' |2={{clade |1=''[[Hypsognathus]]'' |2=''[[Soturnia]]'' }} }} }} }} }} }} }} }} }} }} }} }} }} }} }} }} }} }} }}

'''Analysis 3:''' Strict consensus of 18 MPTs:<ref name=Butler2023/> {{clade| style=font-size:85%;line-height:85% |label1='''Procolophonidae''' |1={{clade |1=''[[Coletta (reptile)|Coletta]]'' |2={{clade |1=''[[Sauropareion]]'' |2={{clade |1={{clade |1=''[[Kitchingnathus]]'' |2=''[[Phaantosaurus]]'' ([[sic]]) }} |2={{clade |1=''[[Eumetabolodon dongshengensis]]'' |2={{clade |1=''[[Eomurruna]]'' |2={{clade |1=''[[Theledectes]]'' |2={{clade |1={{clade |1=''[[Procolina]]'' |2=''[[Phonodus]]'' }} |2={{clade |1=''[[Lasasaurus]]'' |2={{clade |1=''[[Anomoiodon]]'' |2=''[[Thelephon]]'' |3=''[[Kapes bentoni]]'' |4=''[[Kapes majmesculae]]'' |5=''[[Timanophon]]'' |6={{clade |1=''[[Tichvinskia]]'' |2={{clade |1=''[[Eumetabolodon|Eumetabolodon bathycephalus]]'' |label2=[[Procolophoninae]] |2={{clade |1=''[[Teratophon]]'' |2=''[[Thelerpeton]]'' |3=''[[Procolophon]]'' }} |label3=[[Leptopleuroninae]] |3={{clade |1=''[[Neoprocolophon]]'' |2=''[[Pentaedrusaurus]]'' |3={{clade |1=''[[Mandaphon]]'' |2={{clade |1={{clade |1=''[[Scoloparia]]'' |2=''[[Sclerosaurus]]'' }} |2={{clade |1=''[[Hwiccewyrm]]'' |2=''[[Hypsognathus]]'' |3=''[[Soturnia]]'' |4=''[[Leptopleuron]]'' }} }} }} }} }} }} }} }} }} }} }} }} }} }} }} }}

==References== {{Reflist}}

==Sources== * {{cite book|author=Lambert, David|year=2001|title=Dinosaur Encyclopedia|url=https://archive.org/details/dinosaurencyclop00lamb|url-access=registration|publisher=Dorling Kindersley|location=New York|page=[https://archive.org/details/dinosaurencyclop00lamb/page/72 72]|isbn=978-0-7894-7935-8}}

==External links== * [http://www.ucmp.berkeley.edu/anapsids/procolophonoidea.html Introduction to Procolophonoidea]

[[Category:Procolophonidae]] [[Category:Prehistoric reptile families]] [[Category:Late Triassic extinctions]] [[Category:Lopingian first appearances]] [[Category:Long stubs with short prose]]