Editing Permian–Triassic extinction event (section)

== Extinction patterns ==
{| style="margin: 0 0 0.5em 1em; float:right; width:350px; text-align:left; font-size:90%;" class="wikitable"
|-
! style="background:#aec; text-align:center;"| Marine extinctions
| style="background:#aec; text-align:center;"| [[genus|Genera]] extinct
| style="background:#aec; text-align:center;"| Notes
|-
| colspan="5" style="background:#bfd" | '''[[Arthropoda]]'''
|-
| [[Eurypterid]]s
| style="text-align:center;"| '''100%''' || May have become extinct shortly before the P–Tr boundary
|-
| [[Ostracod]]s
| style="text-align:center;"| 74% || &nbsp;
|-
| [[Trilobite]]s
| style="text-align:center;"| '''100%''' || In decline since the Devonian; only 5 genera living before the extinction
|-
| colspan="5" style="background:#bfd" | '''[[Brachiopoda]]'''
|-
| [[Brachiopod]]s
| style="text-align:center;"| 96% || [[orthida|Orthids]], [[Orthotetida|Orthotetids]] and [[Productida|Productids]] died out
|-
| colspan="5" style="background:#bfd" | '''[[Bryozoa]]'''
|-
| [[Bryozoa]]ns
| style="text-align:center;"| 79% || Fenestrates, trepostomes, and cryptostomes died out
|-
| colspan="5" style="background:#bfd" | '''[[Chordata]]'''
|-
| [[Acanthodii|Acanthodians]]
| style="text-align:center;"| '''100%''' || In decline since the [[Devonian]], with only one living family
|-
| colspan="5" style="background:#bfd" | '''[[Cnidaria]]'''
|-
| [[Anthozoa]]ns
| style="text-align:center;"| 96% || [[tabulate coral|Tabulate]] and [[rugosa|rugose]] corals died out
|-
| colspan="5" style="background:#bfd" | '''[[Echinodermata]]'''
|-
| [[Blastoid]]s
| style="text-align:center;"| '''100%''' || 
|-
| [[Crinoid]]s
| style="text-align:center;"| 98% || Inadunates and camerates died out
|-
| colspan="5" style="background:#bfd" | '''[[Mollusca]]'''
|-
| [[Ammonite]]s
| style="text-align:center;"| 97% || [[Goniatite]]s and [[Prolecanitida|Prolecantids]] died out
|-
| [[Bivalvia|Bivalves]]
| style="text-align:center;"| 59% || &nbsp;
|-
| [[Gastropod]]s
| style="text-align:center;"| 98% || &nbsp;
|-
| colspan="3" style="background:#bfd" | '''[[Retaria]]'''
|-
| [[Foraminifera]]ns
| style="text-align:center;"| 97% || [[Fusulinid]]s died out, but were almost extinct before the catastrophe
|-
| [[Radiolaria]]ns
| style="text-align:center;"| 99%<ref>{{cite journal|vauthors=Racki G|year=1999|title=Silica-secreting biota and mass extinctions: survival processes and patterns|volume=154|pages=107–132|doi=10.1016/S0031-0182(99)00089-9|journal=[[Palaeogeography, Palaeoclimatology, Palaeoecology]]|issue=1–2|bibcode=1999PPP...154..107R}}</ref> ||
|}

=== Marine organisms ===
[[Marine invertebrates]] suffered the greatest losses during the P–Tr extinction. Evidence of this was found in samples from south China sections at the P–Tr boundary. Here, 286 out of 329 marine invertebrate genera disappear within the final two sedimentary zones containing [[conodont]]s from the Permian.<ref name="Jin2000" /> The decrease in [[Biodiversity|diversity]] was probably caused by a sharp increase in extinctions, rather than a decrease in [[speciation]].<ref>{{Cite journal |last1=Bambach |first1=R. K. |last2=Knoll |first2=A. H. |last3=Wang |first3=S. C. |title=Origination, extinction, and mass depletions of marine diversity |journal=[[Paleobiology (journal)|Paleobiology]] |volume=30 |issue=4 |pages=522–542 |date=December 2004 |url=http://www.bioone.org/perlserv/?request=get-document&issn=0094-8373&volume=30&page=522 |doi=10.1666/0094-8373(2004)030<0522:OEAMDO>2.0.CO;2 |bibcode=2004Pbio...30..522B |s2cid=17279135 |issn=0094-8373}}</ref>

The extinction primarily affected organisms with [[calcium carbonate]] skeletons, especially those reliant on stable CO<sub>2</sub> levels to produce their skeletons. These organisms were susceptible to the effects of the [[ocean acidification]] that resulted from increased atmospheric CO<sub>2</sub>.<ref name="Knoll2004">{{cite book |url=http://www.geochem.geos.vt.edu/bgep/pubs/Chapter_11_Knoll.pdf |title=Reviews in Mineralogy and Geochemistry |vauthors=Knoll AH |year=2004 |veditors=Dove PM, DeYoreo JJ, Weiner S |article=Biomineralization and evolutionary history |archive-url=https://web.archive.org/web/20100620222809/http://www.geochem.geos.vt.edu/bgep/pubs/Chapter_11_Knoll.pdf |archive-date=2010-06-20 |url-status=dead}}</ref> Organisms that relied on [[haemocyanin]] or [[haemoglobin]] for transporting oxygen were more resistant to extinction than those utilizing [[hemerythrin]] or oxygen diffusion.<ref>{{Cite journal |last1=Song |first1=Haijun |last2=Wu |first2=Yuyang |last3=Dai |first3=Xu |last4=Dal Corso |first4=Jacopo |last5=Wang |first5=Fengyu |last6=Feng |first6=Yan |last7=Chu |first7=Daoliang |last8=Tian |first8=Li |last9=Song |first9=Huyue |last10=Foster |first10=William J. |date=6 May 2024 |title=Respiratory protein-driven selectivity during the Permian-Triassic mass extinction |journal=The Innovation |language=en |volume=5 |issue=3 |pages=100618 |doi=10.1016/j.xinn.2024.100618 |pmid=38638583 |pmc=11025005 |bibcode=2024Innov...500618S }}</ref> There is also evidence that endemism was a strong risk factor influencing a taxon's likelihood of extinction. Bivalve taxa that were endemic and localized to a specific region were more likely to go extinct than cosmopolitan taxa.<ref>{{cite journal |last1=Yan |first1=Jia |last2=Song |first2=Haijun |last3=Dai |first3=Xu |date=1 February 2023 |title=Increased bivalve cosmopolitanism during the mid-Phanerozoic mass extinctions |url=https://www.sciencedirect.com/science/article/abs/pii/S0031018222005338 |journal=[[Palaeogeography, Palaeoclimatology, Palaeoecology]] |volume=611 |page=111362 |doi=10.1016/j.palaeo.2022.111362 |bibcode=2023PPP...61111362Y |access-date=20 February 2023|url-access=subscription }}</ref> There was little latitudinal difference in the survival rates of taxa.<ref>{{cite journal |last1=Allen |first1=Bethany J. |last2=Clapham |first2=Matthew E. |last3=Saupe |first3=Erin E. |last4=Wignall |first4=Paul Barry |last5=Hill |first5=Daniel J. |last6=Dunhill |first6=Alexander M. |date=10 February 2023 |title=Estimating spatial variation in origination and extinction in deep time: a case study using the Permian–Triassic marine invertebrate fossil record |journal=[[Paleobiology (journal)|Paleobiology]] |volume=49 |issue=3 |pages=509–526 |doi=10.1017/pab.2023.1 |bibcode=2023Pbio...49..509A |s2cid=256801383 |doi-access=free |hdl=20.500.11850/598054 |hdl-access=free }}</ref> Organisms that inhabited refugia less affected by global warming experienced lesser or delayed extinctions.<ref>{{Cite journal |last1=Jiang |first1=Haishui |last2=Joachimski |first2=Michael M. |last3=Wignall |first3=Paul Barry |last4=Zhang |first4=Muhui |last5=Lai |first5=Xulong |date=15 December 2015 |title=A delayed end-Permian extinction in deep-water locations and its relationship to temperature trends (Bianyang, Guizhou Province, South China) |url=https://linkinghub.elsevier.com/retrieve/pii/S0031018215005647 |journal=[[Palaeogeography, Palaeoclimatology, Palaeoecology]] |language=en |volume=440 |pages=690–695 |doi=10.1016/j.palaeo.2015.10.002 |bibcode=2015PPP...440..690J |access-date=13 October 2024 |via=Elsevier Science Direct|url-access=subscription }}</ref>

Among [[Benthos|benthic organisms]] the extinction event multiplied [[background extinction rate]]s, and therefore caused maximum species loss to [[Taxon|taxa]] that had a high background extinction rate (by implication, taxa with a high [[Population turnover|turnover]]).<ref name=Stanley2008>{{cite journal | last = Stanley |first=S. M. | year = 2008 | title = Predation defeats competition on the seafloor | journal = [[Paleobiology (journal)|Paleobiology]] | volume = 34 | issue = 1 | pages = 1–21 | url = http://paleobiol.geoscienceworld.org/cgi/content/short/34/1/1 | access-date = 2008-05-13 | doi = 10.1666/07026.1|bibcode=2008Pbio...34....1S | s2cid = 83713101 | url-access = subscription }}</ref><ref name=Stanley2007>{{cite journal | last = Stanley | first = S. M. | year = 2007 | title = An Analysis of the History of Marine Animal Diversity | journal = [[Paleobiology (journal)|Paleobiology]] | volume = 33 | issue = sp6 | pages = 1–55 | url=http://paleobiol.geoscienceworld.org/cgi/content/abstract/33/4_Suppl/1 | doi = 10.1666/06020.1| bibcode = 2007Pbio...33R...1S | s2cid = 86014119 | url-access = subscription }}</ref> The extinction rate of marine organisms was catastrophic.<ref name="Jin2000" /><ref>{{cite journal|last=McKinney |first=M. L.|year=1987|title=Taxonomic selectivity and continuous variation in mass and background extinctions of marine taxa|journal=[[Nature (journal)|Nature]]|volume=325|issue=6100|pages=143–145|doi=10.1038/325143a0|bibcode = 1987Natur.325..143M |s2cid=13473769}}</ref><ref name="Twitchett">{{cite journal |title=Rapid and synchronous collapse of marine and terrestrial ecosystems during the end-Permian biotic crisis |vauthors=Twitchett RJ, Looy CV, Morante R, Visscher H, Wignall PB |journal=[[Geology (journal)|Geology]] |year=2001 |volume=29 |issue=4 |pages=351–354 |doi=10.1130/0091-7613(2001)029<0351:RASCOM>2.0.CO;2 |issn=0091-7613 |bibcode = 2001Geo....29..351T }}</ref> Bioturbators were extremely severely affected, as evidenced by the loss of the sedimentary mixed layer in many marine facies during the end-Permian extinction.<ref>{{cite journal |last1=Hofmann |first1=R. |last2=Buatois |first2=L. A. |last3=MacNaughton |first3=R. B. |last4=Mángano |first4=M. G. |date=15 June 2015 |title=Loss of the sedimentary mixed layer as a result of the end-Permian extinction |url=https://www.sciencedirect.com/science/article/abs/pii/S0031018215001674 |journal=[[Palaeogeography, Palaeoclimatology, Palaeoecology]] |volume=428 |pages=1–11 |doi=10.1016/j.palaeo.2015.03.036 |bibcode=2015PPP...428....1H |access-date=19 December 2022|url-access=subscription }}</ref>

Surviving marine invertebrate groups included articulate [[brachiopods]] (those with a hinge),<ref>{{Cite web|title = Permian: The Marine Realm and The End-Permian Extinction|url = http://paleobiology.si.edu/geotime/main/htmlversion/permian4.html|website = paleobiology.si.edu|access-date = 2016-01-26}}</ref> which had undergone a slow decline in numbers since the P–Tr extinction; the [[Ceratitida]] order of [[ammonite]]s;<ref name="Encyclopædia Britannica">{{Cite web|title = Permian extinction|url = https://www.britannica.com/science/Permian-extinction|website = Encyclopædia Britannica|access-date = 2016-01-26}}</ref> and [[crinoid]]s ("sea lilies"),<ref name="Encyclopædia Britannica" /> which very nearly became extinct but later became abundant and diverse. The groups with the highest survival rates generally had active control of [[circulatory system|circulation]], elaborate gas exchange mechanisms, and light calcification; more heavily calcified organisms with simpler breathing apparatuses suffered the greatest loss of species diversity.<ref name="Payne2004Local">{{cite journal |author=Payne, J.L. |author2=Lehrmann, D.J. |author3=Wei, J. |author4=Orchard, M.J. |author5=Schrag, D.P. |author6=Knoll, A.H. |year=2004 |title=Large Perturbations of the Carbon Cycle During Recovery from the End-Permian Extinction |url=http://pangea.stanford.edu/~jlpayne/Payne_et_al_2004.pdf |journal=[[Science (journal)|Science]] |volume=305 |issue=5683 |pages=506–9 |bibcode=2004Sci...305..506P |citeseerx=10.1.1.582.9406 |doi=10.1126/science.1097023 |pmid=15273391 |s2cid=35498132}}</ref><ref name=Knoll1996>{{cite journal | last1 = Knoll |first1=A. H. |last2=Bambach |first2=R. K. |last3=Canfield |first3=D. E. |last4=Grotzinger |first4=J. P. | year = 1996 | title = Comparative Earth history and Late Permian mass extinction | journal = [[Science (journal)|Science]] | volume = 273| issue = 5274 | pages = 452–457 | doi = 10.1126/science.273.5274.452 | pmid = 8662528 |bibcode = 1996Sci...273..452K |s2cid=35958753 }}</ref> In the case of the brachiopods, at least, surviving [[Taxon|taxa]] were generally small, rare members of a formerly diverse community.<ref name=Leighton2008>{{cite journal | last1 = Leighton |first1=L. R. |last2=Schneider |first2=C. L.| year = 2008 | title = Taxon characteristics that promote survivorship through the Permian–Triassic interval: transition from the Paleozoic to the Mesozoic brachiopod fauna | journal = [[Paleobiology (journal)|Paleobiology]]| volume = 34 | issue = 1 | pages = 65–79 | doi = 10.1666/06082.1|s2cid=86843206
 }}</ref>

Conodonts were severely affected both in terms of taxonomic and morphological diversity, although not as severely as during the Capitanian mass extinction.<ref>{{Cite journal |last1=Xue |first1=Chunling |last2=Yuan |first2=Dong-xun |last3=Chen |first3=Yanlong |last4=Stubbs |first4=Thomas L. |last5=Zhao |first5=Yueli |last6=Zhang |first6=Zhifei |date=15 May 2024 |title=Morphological innovation after mass extinction events in Permian and Early Triassic conodonts based on Polygnathacea |url=https://linkinghub.elsevier.com/retrieve/pii/S003101822400138X |journal=[[Palaeogeography, Palaeoclimatology, Palaeoecology]] |language=en |volume=642 |pages=112149 |doi=10.1016/j.palaeo.2024.112149 |bibcode=2024PPP...64212149X |access-date=21 May 2024 |via=Elsevier Science Direct|url-access=subscription }}</ref>

The [[Ammonoidea|ammonoids]], which had been in a long-term decline for the 30&nbsp;million years since the [[Roadian]] (middle Permian), suffered a selective extinction pulse 10&nbsp;million years before the main event, at the end of the [[Capitanian]] stage. In this preliminary extinction, which greatly reduced [[Guild (ecology)|disparity]], or the range of different ecological guilds, environmental factors were apparently responsible. Diversity and disparity fell further until the P–Tr boundary; the extinction here (P–Tr) was non-selective, consistent with a catastrophic initiator. During the Triassic, diversity rose rapidly, but disparity remained low.<ref>{{Cite journal| last1 = Villier | first1 = L. | last2 = Korn | first2 = D. | title = Morphological Disparity of Ammonoids and the Mark of Permian Mass Extinctions | journal = [[Science (journal)|Science]] | volume = 306 | issue =5694 | pages = 264–266 | date=October 2004 | issn = 0036-8075 | pmid = 15472073 | doi = 10.1126/science.1102127 |bibcode = 2004Sci...306..264V| s2cid = 17304091 }}</ref> The range of morphospace occupied by the ammonoids, that is, their range of possible forms, shapes or structures, became more restricted as the Permian progressed. A few million years into the Triassic, the original range of ammonoid structures was once again reoccupied, but the parameters were now shared differently among [[clades]].<ref>{{Cite journal| first1 = W. B.| last1 = Saunders| first2 = E.| last2 = Greenfest-Allen| first3 = D. M.| last3 = Work| first4 = S. V.| last4 = Nikolaeva| title = Morphologic and taxonomic history of Paleozoic ammonoids in time and morphospace| journal = [[Paleobiology (journal)|Paleobiology]]| volume = 34| issue = 1| pages = 128–154| year = 2008| doi = 10.1666/07053.1| bibcode = 2008Pbio...34..128S| s2cid = 83650272}}</ref>

Ostracods experienced prolonged diversity perturbations during the Changhsingian before the PTME proper, when immense proportions of them abruptly vanished.<ref>{{cite journal |last1=Crasquin |first1=Sylvie |last2=Forel |first2=Marie-Béatrice |last3=Qinglai |first3=Feng |last4=Aihua |first4=Yuan |last5=Baudin |first5=François |last6=Collin |first6=Pierre-Yves |date=30 July 2010 |title=Ostracods (Crustacea) through the Permian-Triassic boundary in South China: the Meishan stratotype (Zhejiang Province) |url=https://www.tandfonline.com/doi/abs/10.1080/14772011003784992 |journal=[[Journal of Systematic Palaeontology]] |volume=8 |issue=3 |pages=331–370 |doi=10.1080/14772011003784992 |bibcode=2010JSPal...8..331C |s2cid=85986762 |access-date=3 July 2023|url-access=subscription }}</ref> At least 74% of ostracods died out during the PTME itself.<ref name="Forel2012">{{cite journal |last1=Forel |first1=Marie-Béatrice |date=1 August 2012 |title=Ostracods (Crustacea) associated with microbialites across the Permian-Triassic boundary in Dajiang (Guizhou Province, South China) |url=https://europeanjournaloftaxonomy.eu/index.php/ejt/article/view/117 |journal=[[European Journal of Taxonomy]] |issue=19 |pages=1–34 |doi=10.5852/ejt.2012.19 |access-date=3 July 2023|doi-access=free }}</ref>

Bryozoans had been on a long-term decline throughout the Late Permian epoch before they suffered even more catastrophic losses during the PTME,<ref>{{Cite journal |last1=Powers |first1=Catherine M. |last2=Bottjer |first2=David J. |date=1 November 2007 |title=Bryozoan paleoecology indicates mid-Phanerozoic extinctions were the product of long-term environmental stress |url=https://pubs.geoscienceworld.org/geology/article/35/11/995-998/129752 |journal=[[Geology (journal)|Geology]] |language=en |volume=35 |issue=11 |pages=995 |doi=10.1130/G23858A.1 |bibcode=2007Geo....35..995P |issn=0091-7613 |access-date=30 December 2023|url-access=subscription }}</ref> being the most severely affected clade among the lophophorates.<ref name="PowersAndBottjer2009" />

Deep water sponges suffered a significant diversity loss and exhibited a decrease in spicule size over the course of the PTME. Shallow water sponges were affected much less strongly; they experienced an increase in spicule size and much lower loss of morphological diversity compared to their deep water counterparts.<ref name="DeclineOfSiliceousSponges">{{cite journal |last1=Liu |first1=Guichun |last2=Feng |first2=Qinglai |last3=Shen |first3=Jun |last4=Yu |first4=Jianxin |last5=He |first5=Weihong |last6=Algeo |first6=Thomas J. |title=Decline of Siliceous Sponges and Spicule Miniaturization Induced by Marine Productivity Collapse and Expanding Anoxia During the Permian-Triassic Crisis in South China |date=1 August 2013 |url=https://pubs.geoscienceworld.org/sepm/palaios/article-abstract/28/8/664/146368/DECLINE-OF-SILICEOUS-SPONGES-AND-SPICULE |journal=[[PALAIOS]] |volume=28 |issue=8 |pages=664–679 |doi=10.2110/palo.2013.p13-035r |s2cid=128751510 |access-date=31 May 2023|url-access=subscription }}</ref>

Foraminifera suffered a severe bottleneck in diversity.<ref name="NatureThroughTime">{{cite book |last1=Delfini |first1=Massimo |last2=Kustatscher |first2=Evelyn |last3=Lavezzi |first3=Fabrizio |last4=Bernardi |first4=Massimo |title=Nature through Time |chapter=The End-Permian Mass Extinction: Nature's Revolution |series=Springer Textbooks in Earth Sciences, Geography and Environment |editor-last1=Martinetto |editor-first1=Edoardo |editor-last2=Tschopp |editor-first2=Emanuel |editor-last3=Gastaldo |editor-first3=Robert A. |date=29 July 2021 |url=https://link.springer.com/book/10.1007/978-3-030-35058-1 |chapter-url=https://link.springer.com/chapter/10.1007/978-3-030-35058-1_10?error=cookies_not_supported&code=aba29752-f20d-4748-90c7-88fdd0fc23ed |publisher=Springer Cham |pages=253–267 |doi=10.1007/978-3-030-35058-1_10 |isbn=978-3-030-35060-4|s2cid=226405085 }}</ref> Evidence from South China indicates the foraminiferal extinction had two pulses.<ref>{{Cite journal |last1=Song |first1=Haijun |last2=Tong |first2=Jinnan |last3=Chen |first3=Zhong-Qiang Chen |date=13 July 2008 |title=Two episodes of foraminiferal extinction near the Permian–Triassic boundary at the Meishan section, South China |url=http://www.tandfonline.com/doi/abs/10.1080/08120090903002599 |journal=[[Australian Journal of Earth Sciences]] |language=en |volume=56 |issue=6 |pages=765–773 |doi=10.1080/08120090903002599 |issn=0812-0099 |access-date=28 October 2024 |via=Taylor and Francis Online|url-access=subscription }}</ref> Foraminiferal biodiversity hotspots shifted into deeper waters during the PTME.<ref name="LiuEtAl2020Foraminifera">{{Cite journal |last1=Liu |first1=Xiaokang |last2=Song |first2=Haijun |last3=Bond |first3=David P.G. |last4=Tong |first4=Jinnan |last5=Benton |first5=Michael James |date=October 2020 |title=Migration controls extinction and survival patterns of foraminifers during the Permian-Triassic crisis in South China |url=https://linkinghub.elsevier.com/retrieve/pii/S0012825220303755 |journal=[[Earth-Science Reviews]] |language=en |volume=209 |pages=103329 |doi=10.1016/j.earscirev.2020.103329 |bibcode=2020ESRv..20903329L |access-date=28 October 2024 |via=Elsevier Science Direct}}</ref> Approximately 93% of latest Permian foraminifera became extinct, with 50% of the clade Textulariina, 92% of Lagenida, 96% of Fusulinida, and 100% of Miliolida disappearing.<ref>{{cite journal |last1=Song |first1=Haijun |last2=Tong |first2=Jinnan |last3=Chen |first3=Zhong-Qiang |last4=Yang |first4=Hao |last5=Wang |first5=Yongbiao |date=14 July 2015 |title=End-Permian mass extinction of foraminifers in the Nanpanjiang basin, South China |url=https://www.cambridge.org/core/journals/journal-of-paleontology/article/abs/endpermian-mass-extinction-of-foraminifers-in-the-nanpanjiang-basin-south-china/627B9121604EBB9E6020DEFF730206BA |journal=[[Journal of Paleontology]] |volume=83 |issue=5 |pages=718–738 |doi=10.1666/08-175.1 |s2cid=130765890 |access-date=23 March 2023|url-access=subscription }}</ref> Foraminifera that were calcareous suffered an extinction rate of 91%.<ref>{{Cite journal |last1=Groves |first1=John R. |last2=Altiner |first2=Demír |date=September–October 2005 |title=Survival and recovery of calcareous foraminifera pursuant to the end-Permian mass extinction |url=https://www.sciencedirect.com/science/article/pii/S1631068305000199 |journal=[[Comptes Rendus Palevol]] |language=en |volume=4 |issue=6–7 |pages=487–500 |doi=10.1016/j.crpv.2004.12.007 |bibcode=2005CRPal...4..487G |access-date=28 October 2024 |via=Elsevier Science Direct|url-access=subscription }}</ref> The reason why lagenides survived while fusulinoidean fusulinides went completely extinct may have been due to the greater range of environmental tolerance and greater geographic distribution of the former compared to the latter.<ref name="GrovesEtAl2017">{{cite journal |last1=Groves |first1=John R. |last2=Altiner |first2=Demír |last3=Rettori |first3=Roberto |date=11 August 2017 |title=Extinction, Survival, and Recovery of Lagenide Foraminifers in the Permian–Triassic Boundary Interval, Central Taurides, Turkey |url=https://www.cambridge.org/core/journals/journal-of-paleontology/article/abs/extinction-survival-and-recovery-of-lagenide-foraminifers-in-the-permiantriassic-boundary-interval-central-taurides-turkey/5DFB53BB450B1C47FF194171D5EAEA69 |journal=[[Journal of Paleontology]] |volume=79 |issue=S62 |pages=1–38 |doi=10.1666/0022-3360(2005)79[1:ESAROL]2.0.CO;2 |hdl=11511/41696 |s2cid=130620805 |access-date=23 March 2023|url-access=subscription }}</ref>

Cladodontomorph sharks likely survived the extinction by surviving in refugia in the deep oceans, a hypothesis based on the discovery of [[Early Cretaceous]] cladodontomorphs in deep, outer shelf environments.<ref name="Cladodontomorphs">{{cite journal |last1=Guinot |first1=Guillaume |last2=Adnet |first2=Sylvain |last3=Cavin |first3=Lionel |last4=Cappetta |first4=Henri |date=29 October 2013 |title=Cretaceous stem chondrichthyans survived the end-Permian mass extinction |journal=[[Nature Communications]] |volume=4 |page=2669 |doi=10.1038/ncomms3669 |pmid=24169620 |bibcode=2013NatCo...4.2669G |s2cid=205320689 |doi-access=free }}</ref> [[Ichthyosaurs]], which evolved immediately before the PTME, were also PTME survivors.<ref name="KearEtAl2023Ichthyosaurs">{{cite journal |last1=Kear |first1=Benjamin P. |last2=Engelschiøn |first2=Victoria S. |last3=Hammer |first3=Øyvind |last4=Roberts |first4=Aubrey J. |last5=Hurum |first5=Jørn H. |date=13 March 2023 |title=Earliest Triassic ichthyosaur fossils push back oceanic reptile origins |journal=[[Current Biology]] |volume=33 |issue=5 |pages=R178–R179 |doi=10.1016/j.cub.2022.12.053 |pmid=36917937 |s2cid=257498390 |doi-access=free |bibcode=2023CBio...33R.178K }}</ref>

The [[Lilliput effect]], the phenomenon of dwarfing of species during and immediately following a mass extinction event, has been observed across the Permian-Triassic boundary,<ref name="LilliputEffectAftermathEndPermianExtinctionEvent">{{cite journal |last1=Twitchett |first1=Richard J. |date=20 August 2007 |title=The Lilliput effect in the aftermath of the end-Permian extinction event |url=https://www.sciencedirect.com/science/article/abs/pii/S0031018207001137 |journal=[[Palaeogeography, Palaeoclimatology, Palaeoecology]] |volume=252 |issue=1–2 |pages=132–144 |doi=10.1016/j.palaeo.2006.11.038 |bibcode=2007PPP...252..132T |access-date=13 January 2023|url-access=subscription }}</ref><ref>{{cite journal |last1=Schaal |first1=Ellen K. |last2=Clapham |first2=Matthew E. |last3=Rego |first3=Brianna L. |last4=Wang |first4=Steve C. |last5=Payne |first5=Jonathan L. |date=6 November 2015 |title=Comparative size evolution of marine clades from the Late Permian through Middle Triassic |url=https://pubs.geoscienceworld.org/paleobiol/article/42/1/127/140555/Comparative-size-evolution-of-marine-clades-from |journal=[[Paleobiology (journal)|Paleobiology]] |volume=42 |issue=1 |pages=127–142 |doi=10.1017/pab.2015.36 |s2cid=18552375 |access-date=13 January 2023}}</ref> notably occurring in foraminifera,<ref>{{cite journal |last1=Feng |first1=Yan |last2=Song |first2=Haijun |last3=Bond |first3=David P. G. |date=1 November 2020 |title=Size variations in foraminifers from the early Permian to the Late Triassic: implications for the Guadalupian–Lopingian and the Permian–Triassic mass extinctions |journal=[[Paleobiology (journal)|Paleobiology]] |volume=46 |issue=4 |pages=511–532 |doi=10.1017/pab.2020.37 |bibcode=2020Pbio...46..511F |s2cid=224855811 |doi-access=free }}</ref><ref>{{cite journal |last1=Song |first1=Haijun |last2=Tong |first2=Jinnan |last3=Chen |first3=Zhong-Qiang |date=15 July 2011 |title=Evolutionary dynamics of the Permian–Triassic foraminifer size: Evidence for Lilliput effect in the end-Permian mass extinction and its aftermath |url=https://www.sciencedirect.com/science/article/abs/pii/S0031018210006528 |journal=[[Palaeogeography, Palaeoclimatology, Palaeoecology]] |volume=308 |issue=1–2 |pages=98–110 |doi=10.1016/j.palaeo.2010.10.036 |bibcode=2011PPP...308...98S |access-date=13 January 2023|url-access=subscription }}</ref><ref>{{cite journal |last1=Rego |first1=Brianna L. |last2=Wang |first2=Steve C. |last3=Altiner |first3=Demír |last4=Payne |first4=Jonathan L. |date=8 February 2016 |title=Within- and among-genus components of size evolution during mass extinction, recovery, and background intervals: a case study of Late Permian through Late Triassic foraminifera |url=https://www.cambridge.org/core/journals/paleobiology/article/abs/within-and-amonggenus-components-of-size-evolution-during-mass-extinction-recovery-and-background-intervals-a-case-study-of-late-permian-through-late-triassic-foraminifera/DDF33E40E62ACA1918481D99F495FA4D |journal=[[Paleobiology (journal)|Paleobiology]] |volume=38 |issue=4 |pages=627–643 |doi=10.1666/11040.1 |hdl=11511/34619 |s2cid=17284750 |access-date=23 March 2023|url-access=subscription }}</ref> brachiopods,<ref>{{cite journal |last1=He |first1=Weihong |last2=Twitchett |first2=Richard J. |last3=Zhang |first3=Y. |last4=Shi |first4=G. R. |last5=Feng |first5=Q.-L. |last6=Yu |first6=J.-X. |last7=Wu |first7=S.-B. |last8=Peng |first8=X.-F. |date=9 November 2010 |title=Controls on body size during the Late Permian mass extinction event |url=https://onlinelibrary.wiley.com/doi/abs/10.1111/j.1472-4669.2010.00248.x |journal=[[Geobiology (journal)|Geobiology]] |volume=8 |issue=5 |pages=391–402 |doi=10.1111/j.1472-4669.2010.00248.x |pmid=20550584 |bibcode=2010Gbio....8..391H |s2cid=23096333 |access-date=13 January 2023|url-access=subscription }}</ref><ref>{{cite journal |last1=Chen |first1=Jing |last2=Song |first2=Haijun |last3=He |first3=Weihong |last4=Tong |first4=Jinnan |last5=Wang |first5=Fengyu |last6=Wu |first6=Shunbao |date=1 April 2019 |title=Size variation of brachiopods from the Late Permian through the Middle Triassic in South China: Evidence for the Lilliput Effect following the Permian-Triassic extinction |url=https://www.sciencedirect.com/science/article/abs/pii/S0031018218304929 |journal=[[Palaeogeography, Palaeoclimatology, Palaeoecology]] |volume=519 |pages=248–257 |doi=10.1016/j.palaeo.2018.07.013 |bibcode=2019PPP...519..248C |s2cid=134806479 |access-date=13 January 2023|url-access=subscription }}</ref><ref>{{cite journal |last1=Shi |first1=Guang R. |last2=Zhang |first2=Yi-chun |last3=Shen |first3=Shu-zhong |last4=He |first4=Wei-hong |date=15 April 2016 |title=Nearshore–offshore–basin species diversity and body size variation patterns in Late Permian (Changhsingian) brachiopods |journal=[[Palaeogeography, Palaeoclimatology, Palaeoecology]] |volume=448 |pages=96–107 |doi=10.1016/j.palaeo.2015.07.046 |bibcode=2016PPP...448...96S |doi-access=free |hdl=10536/DRO/DU:30080099 |hdl-access=free }}</ref> bivalves,<ref>{{cite journal |last1=Huang |first1=Yunfei |last2=Tong |first2=Jinnan |last3=Tian |first3=Li |last4=Song |first4=Haijun |last5=Chu |first5=Daoliang |last6=Miao |first6=Xue |last7=Song |first7=Ting |date=1 January 2023 |title=Temporal shell-size variations of bivalves in South China from the Late Permian to the early Middle Triassic |url=https://www.sciencedirect.com/science/article/abs/pii/S0031018222004783 |journal=[[Palaeogeography, Palaeoclimatology, Palaeoecology]] |volume=609 |page=111307 |doi=10.1016/j.palaeo.2022.111307 |bibcode=2023PPP...60911307H |s2cid=253368808 |access-date=13 January 2023|url-access=subscription }}</ref><ref>{{cite journal |last1=Yates |first1=Adam M. |last2=Neumann |first2=Frank H. |last3=Hancox |first3=P. John |date=2 February 2012 |title=The Earliest Post-Paleozoic Freshwater Bivalves Preserved in Coprolites from the Karoo Basin, South Africa |journal=[[PLOS ONE]] |volume=7 |issue=2 |pages=e30228 |doi=10.1371/journal.pone.0030228 |pmid=22319562 |pmc=3271088 |bibcode=2012PLoSO...730228Y |doi-access=free }}</ref><ref>{{cite journal |last1=Foster |first1=W. J. |last2=Gliwa |first2=J. |last3=Lembke |first3=C. |last4=Pugh |first4=A. C. |last5=Hofmann |first5=R. |last6=Tietje |first6=M. |last7=Varela |first7=S. |last8=Foster |first8=L. C. |last9=Korn |first9=D. |last10=Aberhan |first10=M. |date=January 2020 |title=Evolutionary and ecophenotypic controls on bivalve body size distributions following the end-Permian mass extinction |url=https://www.sciencedirect.com/science/article/abs/pii/S0921818119305739 |journal=[[Global and Planetary Change]] |volume=185 |page=103088 |doi=10.1016/j.gloplacha.2019.103088 |bibcode=2020GPC...18503088F |s2cid=213294384 |access-date=13 January 2023|url-access=subscription }}</ref> and ostracods.<ref>{{cite journal |last1=Chu |first1=Daoliang |last2=Tong |first2=Jinnan |last3=Song |first3=Haijun |last4=Benton |first4=Michael James |last5=Song |first5=Huyue |last6=Yu |first6=Jianxin |last7=Qiu |first7=Xincheng |last8=Huang |first8=Yunfei |last9=Tian |first9=Li |date=1 October 2015 |title=Lilliput effect in freshwater ostracods during the Permian–Triassic extinction |url=https://www.sciencedirect.com/science/article/abs/pii/S0031018215003004 |journal=[[Palaeogeography, Palaeoclimatology, Palaeoecology]] |volume=435 |pages=38–52 |doi=10.1016/j.palaeo.2015.06.003 |bibcode=2015PPP...435...38C |access-date=13 January 2023|url-access=subscription }}</ref><ref>{{cite journal |last1=Forel |first1=Marie-Béatrice |last2=Crasquin |first2=Sylvie |last3=Chitnarin |first3=Anisong |last4=Angiolini |first4=Lucia |last5=Gaetani |first5=Maurizio |date=25 February 2015 |title=Precocious sexual dimorphism and the Lilliput effect in Neo-Tethyan Ostracoda (Crustacea) through the Permian–Triassic boundary |url=https://onlinelibrary.wiley.com/doi/full/10.1111/pala.12151 |journal=[[Palaeontology (journal)|Palaeontology]] |volume=58 |issue=3 |pages=409–454 |doi=10.1111/pala.12151 |bibcode=2015Palgy..58..409F |hdl=2434/354270 |s2cid=140198533 |access-date=23 May 2023|hdl-access=free }}</ref> Though gastropods that survived the cataclysm were smaller in size than those that did not,<ref>{{cite journal |last1=Payne |first1=Jonathan L. |date=8 April 2016 |title=Evolutionary dynamics of gastropod size across the end-Permian extinction and through the Triassic recovery interval |url=https://www.cambridge.org/core/journals/paleobiology/article/abs/evolutionary-dynamics-of-gastropod-size-across-the-endpermian-extinction-and-through-the-triassic-recovery-interval/5E66427F02CD63A731971EBD09F12ACF |journal=[[Paleobiology (journal)|Paleobiology]] |volume=31 |issue=2 |pages=269–290 |doi=10.1666/0094-8373(2005)031[0269:EDOGSA]2.0.CO;2 |s2cid=7367192 |access-date=23 March 2023|url-access=subscription }}</ref> it remains debated whether the Lilliput effect truly took hold among gastropods.<ref>{{cite journal |last1=Brayard |first1=Arnaud |last2=Nützel |first2=Alexander |last3=Stephen |first3=Daniel A. |last4=Bylund |first4=Kevin G. |last5=Jenks |first5=Jim |last6=Bucher |first6=Hugo |date=1 February 2010 |title=Gastropod evidence against the Early Triassic Lilliput effect |url=https://pubs.geoscienceworld.org/gsa/geology/article-abstract/38/2/147/130196/Gastropod-evidence-against-the-Early-Triassic?redirectedFrom=fulltext&casa_token=60Fb3K0zMJwAAAAA:DhYezXj8KGzLNoRTZrBE_j0wA7r4dukCPgwOrHMetVKPp6jP0m5KqM1cWquxpdbOu5QbCM3f |journal=[[Geology (journal)|Geology]] |volume=37 |issue=2 |pages=147–150 |doi=10.1130/G30553.1 |bibcode=2010Geo....38..147B |access-date=13 January 2023|url-access=subscription }}</ref><ref>{{cite journal |last1=Fraiser |first1=M. L. |last2=Twitchett |first2=Richard J. |last3=Frederickson |first3=J. A. |last4=Metcalfe |first4=B. |last5=Bottjer |first5=D. J. |date=1 January 2011 |title=Gastropod evidence against the Early Triassic Lilliput effect: COMMENT |journal=[[Geology (journal)|Geology]] |volume=39 |issue=1 |pages=e232 |doi=10.1130/G31614C.1 |bibcode=2011Geo....39E.232F |doi-access=free }}</ref><ref>{{cite journal |last1=Brayard |first1=Arnaud |last2=Nützel |first2=Alexander |last3=Kajm |first3=Andrzej |last4=Escarguel |first4=Gilles |last5=Hautmann |first5=Michael |last6=Stephen |first6=Daniel A. |last7=Bylund |first7=Kevin G. |last8=Jenks |first8=Jim |last9=Bucher |first9=Hugo |date=1 January 2011 |title=Gastropod evidence against the Early Triassic Lilliput effect: REPLY |journal=[[Geology (journal)|Geology]] |volume=39 |issue=1 |pages=e233 |doi=10.1130/G31765Y.1 |bibcode=2011Geo....39E.233B |doi-access=free }}</ref> Some gastropod taxa, termed "Gulliver gastropods", ballooned in size during and immediately following the mass extinction,<ref>{{cite journal |last1=Brayard |first1=Arnaud |last2=Meier |first2=Maximiliano |last3=Escarguel |first3=Gilles |last4=Fara |first4=Emmanuel |last5=Nützel |first5=Alexander |last6=Olivier |first6=Nicolas |last7=Bylund |first7=Kevin G. |last8=Jenks |first8=James F. |last9=Stephen |first9=Daniel A. |last10=Hautmann |first10=Michael |last11=Vennin |first11=Emmanuelle |last12=Bucher |first12=Hugo |date=July 2015 |title=Early Triassic Gulliver gastropods: Spatio-temporal distribution and significance for biotic recovery after the end-Permian mass extinction |url=https://www.sciencedirect.com/science/article/abs/pii/S0012825215000574 |journal=[[Earth-Science Reviews]] |volume=146 |pages=31–64 |doi=10.1016/j.earscirev.2015.03.005 |bibcode=2015ESRv..146...31B |access-date=20 January 2023|url-access=subscription }}</ref> exemplifying the Lilliput effect's opposite, which has been dubbed the Brobdingnag effect.<ref>{{cite journal |last1=Atkinson |first1=Jed W. |last2=Wignall |first2=Paul Barry |last3=Morton |first3=Jacob D. |last4=Aze |first4=Tracy |date=9 January 2019 |title=Body size changes in bivalves of the family Limidae in the aftermath of the end-Triassic mass extinction: the Brobdingnag effect |url=https://onlinelibrary.wiley.com/doi/10.1111/pala.12415 |journal=[[Palaeontology (journal)|Palaeontology]] |volume=62 |issue=4 |pages=561–582 |doi=10.1111/pala.12415 |bibcode=2019Palgy..62..561A |s2cid=134070316 |access-date=20 January 2023}}</ref>

=== Terrestrial invertebrates ===
The Permian had great diversity in insect and other invertebrate species, including the [[List of largest insects|largest insects]] ever to have existed. The end-Permian is the largest known mass extinction of insects;<ref name="ConradLabandeira">{{citation |journal=American Entomologist |date=1 January 2005 |volume=51 |pages=14–29 |title=The fossil record of insect extinction: New approaches and future directions |first=Conrad |last=Labandeira |doi=10.1093/ae/51.1.14|doi-access=free }}</ref><ref>{{Cite journal |last=Ponomarenko |first=A. G. |date=13 May 2016 |title=Insects during the time around the Permian—Triassic crisis |url=http://link.springer.com/10.1134/S0031030116020052 |journal=[[Paleontological Journal]] |language=en |volume=50 |issue=2 |pages=174–186 |doi=10.1134/S0031030116020052 |bibcode=2016PalJ...50..174P |issn=0031-0301 |access-date=13 October 2024 |via=Springer Link|url-access=subscription }}</ref> according to some sources, it may well be the only mass extinction to significantly affect insect diversity.<ref name="Labandeira">{{cite journal |vauthors=Labandeira CC, Sepkoski JJ |title=Insect diversity in the fossil record |journal=[[Science (journal)|Science]] |volume=261 |issue=5119 |pages=310–315 |year=1993 |pmid=11536548 |doi= 10.1126/science.11536548
|bibcode = 1993Sci...261..310L |citeseerx=10.1.1.496.1576 }}</ref><ref name="sole">{{cite encyclopedia |author1=Sole, R. V. |author2=Newman, M. |editor1=Canadell, J. G. |editor2=Mooney, H. A. |article=Extinctions and Biodiversity in the Fossil Record |title=Encyclopedia of Global Environmental Change, The Earth System |series=Biological and Ecological Dimensions of Global Environmental Change |volume=2 |publisher=Wiley |location=New York |year=2003 |pages=297–391 |isbn=978-0-470-85361-0}}</ref> Eight or nine insect [[Order (biology)|orders]] became extinct and ten more were greatly reduced in diversity. [[Palaeodictyopteroidea|Palaeodictyopteroids]] (insects with piercing and sucking mouthparts) began to decline during the mid-Permian; these extinctions have been linked to a change in flora. The greatest decline occurred in the Late Permian and was probably not directly caused by weather-related floral transitions.<ref name="Erwin1993" /> However, some observed entomofaunal declines in the PTME were biogeographic changes rather than outright extinctions.<ref>{{Cite journal |last=Shcherbakov |first=D. E. |date=30 January 2008 |title=On Permian and Triassic insect faunas in relation to biogeography and the Permian-Triassic crisis |url=https://link.springer.com/10.1134/S0031030108010036 |journal=[[Paleontological Journal]] |language=en |volume=42 |issue=1 |pages=15–31 |doi=10.1134/S0031030108010036 |bibcode=2008PalJ...42...15S |issn=0031-0301 |access-date=18 June 2024 |via=Springer Link|url-access=subscription }}</ref>

=== Terrestrial plants ===
The [[Geologic record|geological record]] of [[terrestrial plant|terrestrial]] plants is sparse and based mostly on [[pollen]] and [[spore]] studies. Floral changes across the Permian-Triassic boundary are highly variable depending on the location and preservation quality of any given site.<ref>{{cite journal |last1=Rees |first1=P. McAllister |date=1 September 2002 |title=Land-plant diversity and the end-Permian mass extinction |url=https://pubs.geoscienceworld.org/gsa/geology/article-abstract/30/9/827/192461/Land-plant-diversity-and-the-end-Permian-mass |journal=[[Geology (journal)|Geology]] |volume=30 |issue=9 |pages=827–830 |doi=10.1130/0091-7613(2002)030<0827:LPDATE>2.0.CO;2 |bibcode=2002Geo....30..827M |access-date=31 May 2023|url-access=subscription }}</ref> Plants are relatively immune to mass extinction, with the impact of all the major mass extinctions "insignificant" at a family level.<ref name=McElwain2007 />{{Dubious|date=May 2020}} Floral diversity losses were more superficial than those of marine animals.<ref>{{Cite journal |last=Ponomarenko |first=A. G. |date=August 2006 |title=Changes in terrestrial biota before the Permian-Triassic ecological crisis |url=https://link.springer.com/article/10.1134/S0031030106100066 |journal=[[Paleontological Journal]] |language=en |volume=40 |issue=4 |pages=S468–S474 |doi=10.1134/S0031030106100066 |bibcode=2006PalJ...40S.468P |issn=0031-0301 |access-date=13 October 2024 |via=Springer Link|url-access=subscription }}</ref> Even the reduction observed in species diversity (of 50%) may be mostly due to [[Taphonomy|taphonomic]] processes.<ref name=":0">{{cite journal |last1=Nowak |first1=Hendrik |last2=Schneebeli-Hermann |first2=Elke |last3=Kustatscher |first3=Evelyn |date=23 January 2019 |title=No mass extinction for land plants at the Permian–Triassic transition |journal=[[Nature Communications]] |volume=10 |issue=1 |page=384 |doi=10.1038/s41467-018-07945-w |pmid=30674875 |pmc=6344494 |bibcode=2019NatCo..10..384N }}</ref><ref name="McElwain2007" /> However, a massive rearrangement of ecosystems does occur, with plant abundances and distributions changing profoundly and all the forests virtually disappearing.<ref>{{cite web|url=http://www.nhm.ac.uk/nature-online/life/dinosaurs-other-extinct-creatures/mass-extinctions/end-permian-mass-extinction/index.html|title=The Dino Directory – Natural History Museum}}</ref><ref name="McElwain2007" /> The dominant floral groups changed, with many groups of land plants entering abrupt decline, such as ''[[Cordaites]]'' ([[gymnosperm]]s) and ''[[Glossopteris]]'' ([[Pteridospermatophyta|seed ferns]]).<ref name="GulbransonEtAl2022">{{cite journal |last1=Gulbranson |first1=Erik L. |last2=Mellum |first2=Morgan M. |last3=Corti |first3=Valentina |last4=Dahlseid |first4=Aidan |last5=Atkinson |first5=Brian A. |last6=Ryberg |first6=Patricia E. |last7=Cornamusini |first7=Giancula |date=24 May 2022 |title=Paleoclimate-induced stress on polar forested ecosystems prior to the Permian–Triassic mass extinction |journal=Scientific Reports |volume=12 |issue=1 |page=8702 |bibcode=2022NatSR..12.8702G |doi=10.1038/s41598-022-12842-w |pmc=9130125 |pmid=35610472}}</ref><ref name="Retallack1995">{{cite journal |last=Retallack |first=G. J. |year=1995 |title=Permian–Triassic life crisis on land |journal=[[Science (journal)|Science]] |volume=267 |issue=5194 |pages=77–80 |bibcode=1995Sci...267...77R |doi=10.1126/science.267.5194.77 |pmid=17840061 |s2cid=42308183}}</ref> The severity of plant extinction has been disputed.<ref name="Cascales-Miñana2011">{{Cite journal | last1 = Cascales-Miñana | first1 = B. | last2 = Cleal | first2 = C. J. | title = Plant fossil record and survival analyses | journal = [[Lethaia]] | pages = 71–82 | year = 2011 | doi = 10.1111/j.1502-3931.2011.00262.x | volume=45}}</ref><ref name=":0" />

The ''Glossopteris''-dominated flora that characterized high-latitude Gondwana collapsed in Australia around 370,000 years before the Permian-Triassic boundary, with this flora's collapse being less constrained in western Gondwana but still likely occurring a few hundred thousand years before the boundary.<ref name="JosefinaBodnar" /> The collapse of this flora is indirectly marked by an abrupt change in river morphology from meandering to braided river systems, signifying the widespread demise of rooted plants.<ref>{{Cite journal |last1=Ward |first1=Peter Douglas |last2=Montgomery |first2=David R. |last3=Smith |first3=Roger |date=8 September 2000 |title=Altered River Morphology in South Africa Related to the Permian-Triassic Extinction |url=https://www.science.org/doi/10.1126/science.289.5485.1740 |journal=[[Science (journal)|Science]] |language=en |volume=289 |issue=5485 |pages=1740–1743 |doi=10.1126/science.289.5485.1740 |pmid=10976065 |bibcode=2000Sci...289.1740W |issn=0036-8075 |access-date=13 October 2024|url-access=subscription }}</ref>

[[Palynological]] or pollen studies from East [[Greenland]] of sedimentary rock strata laid down during the extinction period indicate dense gymnosperm [[woodland]]s before the event. At the same time that marine [[invertebrate]] macrofauna declined, these large woodlands died out and were followed by a rise in diversity of smaller [[herbaceous]] plants including [[Lycopodiophyta]], both [[Selaginellales]] and [[Isoetales]].<ref name="LooyEtAl2005EndPermianDeadZone" /> Data from Kap Stosch suggest that floral species richness was not significantly affected during the PTME.<ref>{{Cite journal |last1=Schneebeli-Hermann |first1=Elke |last2=Hochuli |first2=Peter A. |last3=Bucher |first3=Hugo |date=August 2017 |title=Palynofloral associations before and after the Permian–Triassic mass extinction, Kap Stosch, East Greenland |url=https://linkinghub.elsevier.com/retrieve/pii/S0921818117301303 |journal=[[Global and Planetary Change]] |language=en |volume=155 |pages=178–195 |doi=10.1016/j.gloplacha.2017.06.009 |bibcode=2017GPC...155..178S |access-date=11 September 2024 |via=Elsevier Science Direct|url-access=subscription }}</ref>

The ''Cordaites'' flora, which dominated the Angaran floristic realm corresponding to Siberia, collapsed over the course of the extinction.<ref name="VajdaMcLoughlin2007" /> In the [[Kuznetsk Basin]], the aridity-induced extinction of the regional humid-adapted forest flora dominated by cordaitaleans occurred approximately 252.76 Ma, around 820,000 years before the end-Permian extinction in South China, suggesting that the end-Permian biotic catastrophe may have started earlier on land and that the ecological crisis may have been more gradual and asynchronous on land compared to its more abrupt onset in the marine realm.<ref name="DavydovEtAl2021PPP">{{cite journal |last1=Davydov |first1=V. I. |last2=Karasev |first2=E. V. |last3=Nurgalieva |first3=N. G. |last4=Schmitz |first4=M. D. |last5=Budnikov |first5=I. V. |last6=Biakov |first6=A. S. |last7=Kuzina |first7=D. M. |last8=Silantiev |first8=V. V. |last9=Urazaeva |first9=M. N. |last10=Zharinova |first10=V. V. |last11=Zorina |first11=S. O. |last12=Gareev |first12=B. |last13=Vasilenko |first13=D. V. |date=1 July 2021 |title=Climate and biotic evolution during the Permian-Triassic transition in the temperate Northern Hemisphere, Kuznetsk Basin, Siberia, Russia |url=https://www.sciencedirect.com/science/article/abs/pii/S0031018221002170 |journal=[[Palaeogeography, Palaeoclimatology, Palaeoecology]] |volume=573 |page=110432 |doi=10.1016/j.palaeo.2021.110432 |bibcode=2021PPP...57310432D |s2cid=235530804 |access-date=19 December 2022|url-access=subscription }}</ref>

In North China, the transition between the Upper Shihhotse and Sunjiagou Formations and their lateral equivalents marked a very large extinction of plants in the region. Those plant genera that did not go extinct still experienced a great reduction in their geographic range. Following this transition, coal swamps vanished. The North Chinese floral extinction correlates with the decline of the ''Gigantopteris'' flora of South China.<ref>{{cite journal |last1=Xiong |first1=Conghui |last2=Wang |first2=Jiashu |last3=Huang |first3=Pu |last4=Cascales-Miñana |first4=Borja |last5=Cleal |first5=Christopher J. |last6=Benton |first6=Michael James |last7=Xue |first7=Jinzhuang |date=December 2021 |title=Plant resilience and extinctions through the Permian to Middle Triassic on the North China Block: A multilevel diversity analysis of macrofossil records |url=https://www.sciencedirect.com/science/article/abs/pii/S0012825221003470 |journal=[[Earth-Science Reviews]] |volume=223 |page=103846 |doi=10.1016/j.earscirev.2021.103846 |bibcode=2021ESRv..22303846X |s2cid=240118558 |access-date=28 March 2023|hdl=20.500.12210/76649 |hdl-access=free }}</ref>

In South China, the subtropical [[Cathaysia]]n [[gigantopterid]] dominated rainforests abruptly collapsed.<ref>{{cite journal |last1=Zhang |first1=Hua |last2=Cao |first2=Chang-qun |last3=Liu |first3=Xiao-lei |last4=Mu |first4=Lin |last5=Zheng |first5=Quan-feng |last6=Liu |first6=Feng |last7=Xiang |first7=Lei |last8=Liu |first8=Lu-jun |last9=Shen |first9=Shu-zhong |date=15 April 2016 |title=The terrestrial end-Permian mass extinction in South China |url=https://www.sciencedirect.com/science/article/abs/pii/S0031018215003624 |journal=[[Palaeogeography, Palaeoclimatology, Palaeoecology]] |volume=448 |pages=108–124 |doi=10.1016/j.palaeo.2015.07.002 |bibcode=2016PPP...448..108Z |access-date=20 January 2023|url-access=subscription }}</ref><ref>{{cite journal |last1=Chu |first1=Daoliang |last2=Yu |first2=Jianxin |last3=Tong |first3=Jinnan |last4=Benton |first4=Michael James |last5=Song |first5=Haijun |last6=Huang |first6=Yunfei |last7=Song |first7=Ting |last8=Tian |first8=Li |date=November 2016 |title=Biostratigraphic correlation and mass extinction during the Permian-Triassic transition in terrestrial-marine siliciclastic settings of South China |url=https://www.sciencedirect.com/science/article/abs/pii/S0921818116300182#! |journal=[[Global and Planetary Change]] |volume=146 |pages=67–88 |doi=10.1016/j.gloplacha.2016.09.009 |bibcode=2016GPC...146...67C |hdl=1983/2d475883-42ea-4988-b01c-0a56912e6aec |access-date=12 November 2022|hdl-access=free }}</ref><ref name="FengEtAl2020EarthScience" /> The floral extinction in South China is associated with bacterial blooms in soil and nearby lacustrine ecosystems, with soil erosion resulting from the die-off of plants being their likely cause.<ref>{{cite journal |last1=Biswas |first1=Raman Kumar |last2=Kaiho |first2=Kunio |last3=Saito |first3=Ryosuke |last4=Tian |first4=Li |last5=Shi |first5=Zhiqiang |date=December 2020 |title=Terrestrial ecosystem collapse and soil erosion before the end-Permian marine extinction: Organic geochemical evidence from marine and non-marine records |url=https://www.sciencedirect.com/science/article/abs/pii/S0921818120302186 |journal=[[Global and Planetary Change]] |volume=195 |page=103327 |doi=10.1016/j.gloplacha.2020.103327 |bibcode=2020GPC...19503327B |s2cid=224900905 |access-date=21 December 2022|url-access=subscription }}</ref> Wildfires too likely played a role in the fall of ''Gigantopteris''.<ref name="EcologicalDisturbanceTropicalPeatlands" />

A conifer flora in what is now Jordan, known from fossils near the [[Dead Sea]], showed unusual stability over the Permian-Triassic transition, and appears to have been only minimally affected by the crisis.<ref>{{cite journal |last1=Blomenkemper |first1=Patrick |last2=Kerp |first2=Hans |last3=Abu Hamad |first3=Abdalla |last4=DiMichele |first4=William A. |last5=Bomfleur |first5=Benjamin |date=21 December 2018 |title=A hidden cradle of plant evolution in Permian tropical lowlands |journal=[[Science (journal)|Science]] |volume=362 |issue=6421 |pages=1414–1416 |doi=10.1126/science.aau4061 |pmid=30573628 |bibcode=2018Sci...362.1414B |s2cid=56582195 |doi-access=free }}</ref>

=== 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>