Editing Permian–Triassic extinction event (section)

=== Terrestrial vertebrates ===
The tempo of the terrestrial vertebrate extinction is disputed. Some evidence from the Karoo Basin indicates a protracted extinction lasting a million years.<ref>{{Cite journal |last1=Viglietti |first1=Pia A. |last2=Benson |first2=Roger B. J. |last3=Smith |first3=Roger M. H. |last4=Botha |first4=Jennifer |last5=Kammerer |first5=Christian F. |last6=Skosan |first6=Zaituna |last7=Butler |first7=Elize |last8=Crean |first8=Annelise |last9=Eloff |first9=Bobby |last10=Kaal |first10=Sheena |last11=Mohoi |first11=Joël |last12=Molehe |first12=William |last13=Mtalana |first13=Nolusindiso |last14=Mtungata |first14=Sibusiso |last15=Ntheri |first15=Nthaopa |last16=Ntsala |first16=Thabang |last17=Nyaphuli |first17=John |last18=October |first18=Paul |last19=Skinner |first19=Georgina |last20=Strong |first20=Mike |last21=Stummer |first21=Hedi |last22=Wolvaart |first22=Frederik P. |last23=Angielczyk |first23=Kenneth D. |date=27 April 2021 |title=Evidence from South Africa for a protracted end-Permian extinction on land |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |language=en |volume=118 |issue=17 |doi=10.1073/pnas.2017045118 |doi-access=free |issn=0027-8424 |pmc=8092562 |pmid=33875588|bibcode=2021PNAS..11817045V }}</ref> Other evidence from the Karoo deposits suggest it took 50,000 years or less,<ref>{{cite journal |last1=Smith |first1=Roger M. H. |last2=Ward |first2=Peter D. |date=1 December 2001 |title=Pattern of vertebrate extinctions across an event bed at the Permian-Triassic boundary in the Karoo Basin of South Africa |url=https://pubs.geoscienceworld.org/gsa/geology/article-abstract/29/12/1147/192006/Pattern-of-vertebrate-extinctions-across-an-event |journal=[[Geology (journal)|Geology]] |volume=29 |issue=12 |pages=1147–1150 |doi=10.1130/0091-7613(2001)029<1147:POVEAA>2.0.CO;2 |bibcode=2001Geo....29.1147S |access-date=23 May 2023|url-access=subscription }}</ref> while a study of coprolites in the Vyazniki fossil beds in Russia suggests it took only a few thousand years.<ref>{{Cite journal |last1=Niedźwiedzki |first1=Grzegorz |last2=Bajdek |first2=Piotr |last3=Qvarnström |first3=Martin |last4=Sulej |first4=Tomasz |last5=Sennikov |first5=Andrey G. |last6=Golubev |first6=Valeriy K. |date=15 May 2016 |title=Reduction of vertebrate coprolite diversity associated with the end-Permian extinction event in Vyazniki region, European Russia |url=https://linkinghub.elsevier.com/retrieve/pii/S0031018216001590 |journal=[[Palaeogeography, Palaeoclimatology, Palaeoecology]] |language=en |volume=450 |pages=77–90 |doi=10.1016/j.palaeo.2016.02.057 |bibcode=2016PPP...450...77N |access-date=18 June 2024 |via=Elsevier Science Direct|url-access=subscription }}</ref> Aridification induced by global warming was the chief culprit behind terrestrial vertebrate extinctions.<ref name="SmithBotha2014">{{cite journal |last1=Smith |first1=Roger M. H. |last2=Botha-Brink |first2=Jennifer |date=15 February 2014 |title=Anatomy of a mass extinction: Sedimentological and taphonomic evidence for drought-induced die-offs at the Permo-Triassic boundary in the main Karoo Basin, South Africa |url=https://www.sciencedirect.com/science/article/abs/pii/S0031018214000030 |journal=[[Palaeogeography, Palaeoclimatology, Palaeoecology]] |volume=396 |pages=99–118 |doi=10.1016/j.palaeo.2014.01.002 |bibcode=2014PPP...396...99S |access-date=23 May 2023|url-access=subscription }}</ref><ref>{{cite journal |last1=Gastaldo |first1=Robert A. |last2=Neveling |first2=Johann |date=1 April 2016 |title=Comment on: "Anatomy of a mass extinction: Sedimentological and taphonomic evidence for drought-induced die-offs at the Permo–Triassic boundary in the main Karoo Basin, South Africa" by R.M.H. Smith and J. Botha-Brink, Palaeogeography, Palaeoclimatology, Palaeoecology 396:99-118 |journal=[[Palaeogeography, Palaeoclimatology, Palaeoecology]] |volume=447 |pages=88–91 |doi=10.1016/j.palaeo.2014.06.027 |bibcode=2016PPP...447...88G |doi-access=free }}</ref> There is enough evidence to indicate that over two thirds of terrestrial [[labyrinthodont]] [[amphibian]]s, [[sauropsid]] ("reptile") and [[therapsid]] ("proto-mammal") [[taxa]] became extinct. Large [[herbivore]]s suffered the heaviest losses.

All Permian [[anapsid]] reptiles died out except the [[Procolophonidae|procolophonids]] (although [[testudines]] have ''morphologically''-anapsid skulls, they are now thought to have separately evolved from diapsid ancestors). [[Pelycosaurs]] died out before the end of the Permian. Too few Permian diapsid fossils have been found to support any conclusion about the effect of the Permian extinction on [[diapsid]]s (the "reptile" group from which lizards, snakes, crocodilians, and [[dinosaur]]s (including birds) evolved).<ref>{{cite journal | author=Maxwell, W.D. | year=1992 | title=Permian and Early Triassic extinction of non-marine tetrapods | journal=[[Palaeontology (journal)|Palaeontology]] | volume=35 | pages=571–583}}</ref><ref name="Erwin1990">{{cite journal | author=Erwin, D.H. | title=The End-Permian Mass Extinction | journal=[[Annual Review of Ecology, Evolution, and Systematics]] | volume=21 | pages=69–91 | year=1990 | issue=1 | doi=10.1146/annurev.es.21.110190.000441 | bibcode=1990AnRES..21...69E }}</ref> [[Tangasauridae|Tangasaurids]] were largely unaffected.<ref>{{Cite journal |last1=Ketchum |first1=Hilary F |last2=Barrett |first2=Paul M |date=21 January 2004 |title=New reptile material from the Lower Triassic of Madagascar: implications for the PermianTriassic extinction event |url=https://pubs.geoscienceworld.org/csp/cjes/article-abstract/41/1/1/53678/New-reptile-material-from-the-Lower-Triassic-of?redirectedFrom=fulltext |journal=[[Canadian Journal of Earth Sciences]] |language=en |volume=41 |issue=1 |pages=1–8 |doi=10.1139/e03-084 |bibcode=2004CaJES..41....1K |issn=0008-4077 |access-date=13 October 2024 |via=GeoScienceWorld|url-access=subscription }}</ref> Gorgonopsians are traditionally thought to have gone extinct during the PTME, but some tentative evidence suggests they may have survived into the Triassic.<ref>{{Cite journal |last1=Benoit |first1=Julien |last2=Kammerer |first2=Christian F. |last3=Dollman |first3=Kathleen |last4=Groenewald |first4=David P. |last5=Smith |first5=Roger M.H. |date=15 March 2024 |title=Did gorgonopsians survive the end-Permian "Great Dying"? A re-appraisal of three gorgonopsian specimens (Therapsida, Theriodontia) reported from the Triassic Lystrosaurus declivis Assemblage Zone, Karoo Basin, South Africa |url=https://linkinghub.elsevier.com/retrieve/pii/S0031018224000336 |journal=[[Palaeogeography, Palaeoclimatology, Palaeoecology]] |language=en |volume=638 |pages=112044 |doi=10.1016/j.palaeo.2024.112044 |bibcode=2024PPP...63812044B |access-date=21 May 2024 |via=Elsevier Science Direct|url-access=subscription }}</ref> Freshwater and euryhaline fishes, having experienced minimal diversity losses before the PTME, were unaffected during the PTME and actually appear to have increased in diversity across the Permian-Triassic boundary.<ref>{{Cite journal |last=Pitrat |first=Charles W. |date=December 1973 |title=Vertebrates and the Permo-Triassic extinction |url=https://www.sciencedirect.com/science/article/abs/pii/0031018273900114 |journal=[[Palaeogeography, Palaeoclimatology, Palaeoecology]] |language=en |volume=14 |issue=4 |pages=249–264 |doi=10.1016/0031-0182(73)90011-4 |bibcode=1973PPP....14..249P |access-date=13 October 2024 |via=Elsevier Science Direct|url-access=subscription }}</ref> However, faunal turnovers in freshwater fish communities occurred in areas like the Kuznetsk Basin.<ref>{{Cite journal |last=Bakaev |first=Aleksandr S. |date=17 July 2024 |title=Actinopterygians from the continental Permian–Triassic boundary section at Babiy Kamen (Kuznetsk Basin, Siberia, Russia) |url=https://www.sciencedirect.com/science/article/abs/pii/S1871174X24000854 |journal=[[Palaeoworld]] |volume=34 |issue=2 |language=en |doi=10.1016/j.palwor.2024.05.007 |access-date=13 October 2024 |via=Elsevier Science Direct|url-access=subscription }}</ref>

The groups that survived suffered extremely heavy losses of species and some terrestrial vertebrate groups very nearly became extinct at the end of the Permian. Some of the surviving groups did not persist for long past this period, but others that barely survived went on to produce diverse and long-lasting lineages. However, it took 30{{nbsp}}million years for the terrestrial vertebrate fauna to fully recover both numerically and ecologically.<ref>{{cite web|url=http://www.bristol.ac.uk/news/2008/5785.html|title=Bristol University – News – 2008: Mass extinction}}</ref>

It is difficult to analyze extinction and survival rates of land organisms in detail because few terrestrial fossil beds span the Permian–Triassic boundary. The best-known record of [[vertebrate]] changes across the Permian–Triassic boundary occurs in the [[Karoo Supergroup]] of [[South Africa]], but statistical analyses have so far not produced clear conclusions.<ref name="KNollBambach2007Paleophysiology">{{cite journal |vauthors=Knoll AH, Bambach RK, Payne JL, Pruss S, Fischer WW |year=2007 |title=Paleophysiology and end-Permian mass extinction |url=https://earthscience.rice.edu/wp-content/uploads/2015/11/Knoll_P-T-extinction_07.pdf |journal=[[Earth and Planetary Science Letters]] |volume=256 |issue=3–4 |pages=295–313 |bibcode=2007E&PSL.256..295K |doi=10.1016/j.epsl.2007.02.018 |access-date=13 December 2021}}</ref> One study of the Karoo Basin found that 69% of terrestrial vertebrates went extinct over 300,000 years leading up to the Permian-Triassic boundary, followed by a minor extinction pulse involving four taxa that survived the previous extinction interval.<ref>{{cite journal |last1=Smith |first1=Roger M. H. |last2=Botha |first2=Jennifer |date=September–October 2005 |title=The recovery of terrestrial vertebrate diversity in the South African Karoo Basin after the end-Permian extinction |url=https://www.sciencedirect.com/science/article/pii/S1631068305000849 |journal=[[Comptes Rendus Palevol]] |volume=4 |issue=6–7 |pages=623–636 |doi=10.1016/j.crpv.2005.07.005 |bibcode=2005CRPal...4..623S |access-date=26 May 2023|url-access=subscription }}</ref> Another study of latest Permian vertebrates in the Karoo Basin found that 54% of them went extinct due to the PTME.<ref>{{cite journal |last1=Botha |first1=Jennifer |last2=Smith |first2=Roger M. H. |date=August 2006 |title=Rapid vertebrate recuperation in the Karoo Basin of South Africa following the End-Permian extinction |url=https://www.sciencedirect.com/science/article/abs/pii/S1464343X06001142 |journal=[[Journal of African Earth Sciences]] |volume=45 |issue=4–5 |pages=502–514 |doi=10.1016/j.jafrearsci.2006.04.006 |bibcode=2006JAfES..45..502B |access-date=26 May 2023|url-access=subscription }}</ref>