Permian–Triassic extinction event

The Permian–Triassic (P–T, P–Tr)[3][4] extinction event, also known as the End-Permian Extinction[5] and colloquially as the Great Dying,[6] formed the boundary between the Permian and Triassic geologic periods, as well as between the Paleozoic and Mesozoic eras, approximately 251.9 million years ago.[7] It is the Earth's most severe known extinction event,[8] with the extinction of 57% of biological families, 83% of genera, 81% of marine species[9][10][11] and 70% of terrestrial vertebrate species.[12] It was the largest known mass extinction of insects.

Marine extinction intensity during the Phanerozoic
%
Millions of years ago
(H)
Cap
Plot of extinction intensity (percentage of marine genera that are present in each interval of time but do not exist in the following interval) vs time in the past.[1] Geological periods are annotated (by abbreviation and colour) above. The Permian–Triassic extinction event is the most significant event for marine genera, with just over 50% (according to this source) perishing. (source and image info)
Permian–Triassic boundary at Frazer Beach in New South Wales, with the End Permian extinction event located just above the coal layer.[2]

There is evidence for one to three distinct pulses, or phases, of extinction.[13][12][14][15][16]

The scientific consensus is that the main cause of extinction was the large amount of carbon dioxide emitted by the volcanic eruptions that created the Siberian Traps, which elevated global temperatures, and in the oceans led to widespread anoxia and acidification.[17] Proposed contributing factors include: the emission of much additional carbon dioxide from the thermal decomposition of hydrocarbon deposits, including oil and coal, triggered by the eruptions; and emissions of methane by novel methanogenic microorganisms, perhaps nourished by minerals dispersed in the eruptions.[18][19]

The speed of recovery from the extinction is disputed. Some scientists estimate that it took 10 million years (until the Middle Triassic), due both to the severity of the extinction and because grim conditions returned periodically for another 5 million years, causing further extinction events, such as the Smithian-Spathian boundary extinction.[8][20] However, studies in Bear Lake County, near Paris, Idaho,[21] and nearby sites in Idaho and Nevada[22] showed a relatively quick rebound in a localized Early Triassic marine ecosystem, taking around 3 million years to recover, suggesting that the impact of the extinction may have been felt less severely in some areas than in others.

Dating

Previously, it was thought that rock sequences spanning the Permian–Triassic boundary were too few and contained too many gaps for scientists to reliably determine its details.[26] However, it is now possible to date the extinction with millennial precision. U–Pb zircon dates from five volcanic ash beds from the Global Stratotype Section and Point for the Permian–Triassic boundary at Meishan, China, establish a high-resolution age model for the extinction – allowing exploration of the links between global environmental perturbation, carbon cycle disruption, mass extinction, and recovery at millennial timescales. The extinction occurred between 251.941 ± 0.037 and 251.880 ± 0.031 million years ago, a duration of 60 ± 48 thousand years.[27] A large (approximately 0.9%), abrupt global decrease in the ratio of the stable isotope carbon-13 to that of carbon-12 coincides with this extinction,[24][28][29][30][31] and is sometimes used to identify the Permian–Triassic boundary in rocks that are unsuitable for radiometric dating.[32] Further evidence for environmental change around the P–Tr boundary suggests an 8 °C (14 °F) rise in temperature,[24] and an increase in CO
2 levels by 2000 ppm (for comparison, the concentration immediately before the Industrial Revolution was 280 ppm,[24] and the amount today is about 415 ppm[33]). There is also evidence of increased ultraviolet radiation reaching the earth, causing the mutation of plant spores.[24][34]

It has been suggested that the Permian–Triassic boundary is associated with a sharp increase in the abundance of marine and terrestrial fungi, caused by the sharp increase in the amount of dead plants and animals fed upon by the fungi.[35] For a while this "fungal spike" was used by some paleontologists to identify the Permian–Triassic boundary in rocks that are unsuitable for radiometric dating or lack suitable index fossils, but even the proposers of the fungal spike hypothesis pointed out that "fungal spikes" may have been a repeating phenomenon created by the post-extinction ecosystem in the earliest Triassic.[35] The very idea of a fungal spike has been criticized on several grounds, including: Reduviasporonites, the most common supposed fungal spore, may be a fossilized alga;[24][36] the spike did not appear worldwide;[37][38] and in many places it did not fall on the Permian–Triassic boundary.[39] The reduviasporonites may even represent a transition to a lake-dominated Triassic world rather than an earliest Triassic zone of death and decay in some terrestrial fossil beds.[40] Newer chemical evidence agrees better with a fungal origin for Reduviasporonites, diluting these critiques.[41]

Uncertainty exists regarding the duration of the overall extinction and about the timing and duration of various groups' extinctions within the greater process. Some evidence suggests that there were multiple extinction pulses[12] or that the extinction was spread out over a few million years, with a sharp peak in the last million years of the Permian.[39][42] Statistical analyses of some highly fossiliferous strata in Meishan, Zhejiang Province in southeastern China, suggest that the main extinction was clustered around one peak.[14] Recent research shows that different groups became extinct at different times; for example, while difficult to date absolutely, ostracod and brachiopod extinctions were separated by 670,000 to 1.17 million years.[43] In a well-preserved sequence in east Greenland, the decline of animals is concentrated in a period 10,000 to 60,000 years long, with plants taking an additional several hundred thousand years to show the full impact of the event.[44]

An older theory, still supported in some recent papers,[12][45] is that there were two major extinction pulses 9.4 million years apart, separated by a period of extinctions well above the background level, and that the final extinction killed off only about 80% of marine species alive at that time while the other losses occurred during the first pulse or the interval between pulses. According to this theory one of these extinction pulses occurred at the end of the Guadalupian epoch of the Permian.[12][46] For example, all dinocephalian genera died out at the end of the Guadalupian,[45] as did the Verbeekinidae, a family of large-size fusuline foraminifera.[47] The impact of the end-Guadalupian extinction on marine organisms appears to have varied between locations and between taxonomic groups – brachiopods and corals had severe losses.[48][49]

Studies of the timing and causes of the Permian-Triassic extinction are complicated by the often-overlooked Capitanian extinction (also called the Guadalupian extinction), just one of perhaps two mass extinctions in the late Permian that closely preceded the Permian-Triassic event. In short, when the Permian-Triassic starts it is difficult to know whether the end-Capitanian had finished, depending on the factor considered.[50][51] Some of the extinctions dated to the Permian-Triassic boundary have recently been redated to the end-Capitanian. Further, it is unclear whether some species who survived the prior extinction(s) had recovered well enough for their final demise in the Permian-Triassic event to be considered separate from Capitanian event. A minority point of view considers the sequence of environmental disasters to have effectively constituted a single, prolonged extinction event, perhaps depending on which species is considered.

Extinction patterns
Marine extinctions Genera extinct Notes
Arthropoda
Eurypterids 100%May have become extinct shortly before the P–Tr boundary
Ostracods 59% 
Trilobites 100%In decline since the Devonian; only 2 genera living before the extinction
Brachiopoda
Brachiopods 96%Orthids and productids died out
Bryozoa
Bryozoans 79%Fenestrates, trepostomes, and cryptostomes died out
Chordata
Acanthodians 100%In decline since the Devonian, with only one living family
Cnidaria
Anthozoans 96%Tabulate and rugose corals died out
Echinodermata
Blastoids 100%May have become extinct shortly before the P–Tr boundary
Crinoids 98%Inadunates and camerates died out
Mollusca
Ammonites 97%Goniatites died out
Bivalves 59% 
Gastropods 98% 
Retaria
Foraminiferans 97%Fusulinids died out, but were almost extinct before the catastrophe
Radiolarians 99%[52]

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 conodonts from the Permian.[14] The decrease in diversity was probably caused by a sharp increase in extinctions, rather than a decrease in speciation.[53]

The extinction primarily affected organisms with calcium carbonate skeletons, especially those reliant on stable CO2 levels to produce their skeletons.[54] These organisms were susceptible to the effects of the ocean acidification that resulted from increased atmospheric CO2.

Among benthic organisms the extinction event multiplied background extinction rates, and therefore caused maximum species loss to taxa that had a high background extinction rate (by implication, taxa with a high turnover).[55][56] The extinction rate of marine organisms was catastrophic.[14][57][58][59]

Surviving marine invertebrate groups included articulate brachiopods (those with a hinge),[60] which had undergone a slow decline in numbers since the P–Tr extinction; the Ceratitida order of ammonites;[61] and crinoids ("sea lilies"),[61] which very nearly became extinct but later became abundant and diverse.

The groups with the highest survival rates generally had active control of circulation, elaborate gas exchange mechanisms, and light calcification; more heavily calcified organisms with simpler breathing apparatuses suffered the greatest loss of species diversity.[23][62] In the case of the brachiopods, at least, surviving taxa were generally small, rare members of a formerly diverse community.[63]

The ammonoids, which had been in a long-term decline for the 30 million years since the Roadian (middle Permian), suffered a selective extinction pulse 10 million years before the main event, at the end of the Capitanian stage. In this preliminary extinction, which greatly reduced 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.[64]

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.[65]

Terrestrial invertebrates

The Permian had great diversity in insect and other invertebrate species, including the largest insects ever to have existed. The end-Permian is the largest known mass extinction of insects;[66] according to some sources, it may well be the only mass extinction to significantly affect insect diversity.[67][68] Eight or nine insect orders became extinct and ten more were greatly reduced in diversity. 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.[57]

Most fossil insect groups found after the Permian–Triassic boundary differ significantly from those before: Of Paleozoic insect groups, only the Glosselytrodea, Miomoptera, and Protorthoptera have been discovered in deposits from after the extinction. The caloneurodeans, monurans, paleodictyopteroids, protelytropterans, and protodonates became extinct by the end of the Permian. Though Triassic insects are very different from those of the Permian, a gap in the insect fossil record spans approximately 15 million years from the late Permian to early Triassic. In well-documented Late Triassic deposits, fossils overwhelmingly consist of modern fossil insect groups.[67]

Plant ecosystem response

The geological record of terrestrial plants is sparse and based mostly on pollen and spore studies. Plants are relatively immune to mass extinction, with the impact of all the major mass extinctions "insignificant" at a family level.[24] Even the reduction observed in species diversity (of 50%) may be mostly due to taphonomic processes.[24] However, a massive rearrangement of ecosystems does occur, with plant abundances and distributions changing profoundly and all the forests virtually disappearing;[24][69] the Palaeozoic flora scarcely survived this extinction.[70]

At the P–Tr boundary, the dominant floral groups changed, with many groups of land plants entering abrupt decline, such as Cordaites (gymnosperms) and Glossopteris (seed ferns).[71] Dominant gymnosperm genera were replaced post-boundary by lycophytes – extant lycophytes are recolonizers of disturbed areas.[72]

Palynological or pollen studies from East Greenland of sedimentary rock strata laid down during the extinction period indicate dense gymnosperm woodlands 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. Later, other groups of gymnosperms again become dominant but again suffered major die-offs. These cyclical flora shifts occurred a few times over the course of the extinction period and afterward. These fluctuations of the dominant flora between woody and herbaceous taxa indicate chronic environmental stress resulting in a loss of most large woodland plant species. The successions and extinctions of plant communities do not coincide with the shift in δ13C values but occurred many years after.[38] The recovery of gymnosperm forests took 4–5 million years.[24]

In China, the subtropical Cathaysian gigantopterid dominated rainforests abruptly collapsed, and were replaced by low-lying herbaceous vegetation dominated by the isoetalean Tomiostrobus.[73]

Coal gap

No coal deposits are known from the Early Triassic, and those in the Middle Triassic are thin and low-grade.[25] This "coal gap" has been explained in many ways. It has been suggested that new, more aggressive fungi, insects, and vertebrates evolved and killed vast numbers of trees. These decomposers themselves suffered heavy losses of species during the extinction and are not considered a likely cause of the coal gap.[25] It could simply be that all coal-forming plants were rendered extinct by the P–Tr extinction and that it took 10 million years for a new suite of plants to adapt to the moist, acid conditions of peat bogs.[25] Abiotic factors (factors not caused by organisms), such as decreased rainfall or increased input of clastic sediments, may also be to blame.[24]

On the other hand, the lack of coal may simply reflect the scarcity of all known sediments from the Early Triassic. Coal-producing ecosystems, rather than disappearing, may have moved to areas where we have no sedimentary record for the Early Triassic.[24] For example, in eastern Australia a cold climate had been the norm for a long period, with a peat mire ecosystem adapted to these conditions. Approximately 95% of these peat-producing plants went locally extinct at the P–Tr boundary;[74] coal deposits in Australia and Antarctica disappear significantly before the P–Tr boundary.[24]

Terrestrial vertebrates

There is enough evidence to indicate that over two thirds of terrestrial labyrinthodont amphibians, sauropsid ("reptile") and therapsid ("proto-mammal") taxa became extinct. Large herbivores suffered the heaviest losses.

All Permian anapsid reptiles died out except the 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 diapsids (the "reptile" group from which lizards, snakes, crocodilians, and dinosaurs (including birds) evolved).[75][76]

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 million years for the terrestrial vertebrate fauna to fully recover both numerically and ecologically.[77]

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.[78]

Biotic recovery

In the wake of the extinction event, the ecological structure of present-day biosphere evolved from the stock of surviving taxa. In the sea, the "Modern Evolutionary Fauna" became dominant over elements of the "Palaeozoic Evolutionary Fauna".[79] Typical taxa of shelly benthic faunas were now bivalves, snails, sea urchins and Malacostraca, whereas bony fishes[80] and marine reptiles[81] diversified in the pelagic zone. On land, dinosaurs and mammals arose in the course of the Triassic. The profound change in the taxonomic composition was partly a result of the selectivity of the extinction event, which affected some taxa (e.g., brachiopods) more severely than others (e.g., bivalves).[82] However, recovery was also differential between taxa. Some survivors became extinct some million years after the extinction event without having rediversified (dead clade walking,[83] e.g. the snail family Bellerophontidae),[84] whereas others rose to dominance over geologic times (e.g., bivalves).[85][86]

Changes in marine ecosystems
Shell bed with the bivalve Claraia clarai, a common early Triassic disaster taxon.

Marine post-extinction faunas were mostly species-poor and dominated by few disaster species such as the bivalves Claraia and Unionites. Seafloor communities maintained a comparatively low diversity until the end of the Early Triassic, approximately 4 million years after the extinction event.[87] This slow recovery stands in remarkable contrast with the quick recovery seen in nektonic organisms such as ammonoids, which exceeded pre-extinction diversities already two million years after the crisis.[88] The relative delay in the recovery of benthic organisms has been attributed to widespread anoxia,[89] but high abundances of benthic species contradict this explanation.[90] More recent work suggests that the pace of recovery was intrinsically driven by the intensity of competition among species, which drives rates of niche differentiation and speciation.[91] Accordingly, low levels of interspecific competition in seafloor communities that are dominated by primary consumers correspond to slow rates of diversification and high levels of interspecific competition among nektonic secondary and tertiary consumers to high diversification rates. Whereas most marine communities were fully recovered by the Middle Triassic,[92][93] global marine diversity reached pre-extinction values no earlier than the Middle Jurassic, approximately 75 million years after the extinction event.[94]

Sessile filter feeders like this Carboniferous crinoid, the mushroom crinoid (Agaricocrinus americanus), were significantly less abundant after the P–Tr extinction.

Prior to the extinction, about two-thirds of marine animals were sessile and attached to the seafloor. During the Mesozoic, only about half of the marine animals were sessile while the rest were free-living. Analysis of marine fossils from the period indicated a decrease in the abundance of sessile epifaunal suspension feeders such as brachiopods and sea lilies and an increase in more complex mobile species such as snails, sea urchins and crabs.[95]

Before the Permian mass extinction event, both complex and simple marine ecosystems were equally common. After the recovery from the mass extinction, the complex communities outnumbered the simple communities by nearly three to one,[95] and the increase in predation pressure led to the Mesozoic Marine Revolution.

Bivalves were fairly rare before the P–Tr extinction but became numerous and diverse in the Triassic, and one group, the rudist clams, became the Mesozoic's main reef-builders. Some researchers think much of the change happened in the 5 million years between the two major extinction pulses.[96]

Crinoids ("sea lilies") suffered a selective extinction, resulting in a decrease in the variety of their forms.[97] Their ensuing adaptive radiation was brisk, and resulted in forms possessing flexible arms becoming widespread; motility, predominantly a response to predation pressure, also became far more prevalent.[98]

Land vertebrates
Lystrosaurus was by far the most abundant early Triassic land vertebrate.

Lystrosaurus, a pig-sized herbivorous dicynodont therapsid, constituted as much as 90% of some earliest Triassic land vertebrate fauna. Smaller carnivorous cynodont therapsids also survived, including the ancestors of mammals. In the Karoo region of southern Africa, the therocephalians Tetracynodon, Moschorhinus and Ictidosuchoides survived, but do not appear to have been abundant in the Triassic.[99]

Archosaurs (which included the ancestors of dinosaurs and crocodilians) were initially rarer than therapsids, but they began to displace therapsids in the mid-Triassic. In the mid to late Triassic, the dinosaurs evolved from one group of archosaurs, and went on to dominate terrestrial ecosystems during the Jurassic and Cretaceous.[100] This "Triassic Takeover" may have contributed to the evolution of mammals by forcing the surviving therapsids and their mammaliform successors to live as small, mainly nocturnal insectivores; nocturnal life probably forced at least the mammaliforms to develop fur, better hearing and higher metabolic rates,[101] while losing part of the differential color-sensitive retinal receptors reptilians and birds preserved. The archosaur dominance would end again due to the K-Pg extinction event, after which both birds (only extant dinosaurs) and mammals (only extant synapsids) would diversify and share the world.

Some temnospondyl amphibians made a relatively quick recovery, in spite of nearly becoming extinct. Mastodonsaurus and trematosaurians were the main aquatic and semiaquatic predators during most of the Triassic, some preying on tetrapods and others on fish.[102]

Land vertebrates took an unusually long time to recover from the P–Tr extinction; Palaeontologist Michael Benton estimated the recovery was not complete until 30 million years after the extinction, i.e. not until the Late Triassic, when the first dinosaurs had risen from bipedal archosaurian ancestors and the first mammals from small cynodont ancestors.[10]

Hypotheses about cause

Pinpointing the exact causes of the Permian–Triassic extinction event is difficult, mostly because it occurred over 250 million years ago, and since then much of the evidence that would have pointed to the cause has been destroyed or is concealed deep within the Earth under many layers of rock. The sea floor is completely recycled over around 200 million years by the ongoing process of plate tectonics and seafloor spreading, leaving no useful indications beneath the ocean.

Yet, scientists have gathered significant evidence for causes, and several mechanisms have been proposed. The proposals include both catastrophic and gradual processes (similar to those theorized for the Cretaceous–Paleogene extinction event).

  • The catastrophic group includes one or more large bolide impact events, increased volcanism, and sudden release of methane from the seafloor, either due to dissociation of methane hydrate deposits or metabolism of organic carbon deposits by methanogenic microbes.
  • The gradual group includes sea level change, increasing hypoxia, and increasing aridity.

Any hypothesis about the cause must explain the selectivity of the event, which affected organisms with calcium carbonate skeletons most severely; the long period (4 to 6 million years) before recovery started, and the minimal extent of biological mineralization (despite inorganic carbonates being deposited) once the recovery began.[54]

Volcanism

The final stages of the Permian had two flood basalt events. A smaller one, the Emeishan Traps in China, occurred at the same time as the end-Guadalupian extinction pulse, in an area close to the equator at the time.[103][104] The flood basalt eruptions that produced the Siberian Traps constituted one of the largest known volcanic events on Earth and covered over 2,000,000 square kilometres (770,000 sq mi) with lava.[105][106][107] The date of the Siberian Traps eruptions and the extinction event are in good agreement.[27][108] The Siberian Traps are underlain by thick sequences of Early-Mid Paleozoic aged carbonate and evaporite deposits, as well as Carboniferous-Permian aged coal bearing clastic rocks. When heated, such as by igneous intrusions, these rocks are capable of emitting large amounts of greenhouse and toxic gases. The unique setting of the Siberian Traps over these deposits is likely the reason for the severity of the extinction.[109]

The Emeishan and Siberian Traps eruptions may have caused dust clouds and acid aerosols, which would have blocked out sunlight and thus disrupted photosynthesis both on land and in the photic zone of the ocean, causing food chains to collapse. The eruptions may also have caused acid rain as the aerosols washed out of the atmosphere. That may have killed land plants and mollusks and planktonic organisms which had calcium carbonate shells. The eruptions would also have emitted carbon dioxide, causing global warming. When all of the dust clouds and aerosols washed out of the atmosphere, the excess carbon dioxide would have remained and the warming would have proceeded without any mitigating effects.[110]

The Siberian Traps had unusual features that made them even more dangerous. Pure flood basalts produce fluid, low-viscosity lava, and do not hurl debris into the atmosphere. It appears, however, that 20% of the output of the Siberian Traps eruptions was pyroclastic (consisted of ash and other debris thrown high into the atmosphere), increasing the short-term cooling effect.[111] The basalt lava erupted or intruded into carbonate rocks and into sediments that were in the process of forming large coal beds, both of which would have emitted large amounts of carbon dioxide, leading to stronger global warming after the dust and aerosols settled.[110] In addition, these volcanic eruptions released significant quantities of toxic mercury into the atmosphere and ocean, further contributing to large scale die-offs of terrestrial and marine life.[112]

In January 2011, a team, led by Stephen Grasby of the Geological Survey of Canada – Calgary, reported evidence that volcanism caused massive coal beds to ignite, possibly releasing more than 3 trillion tons of carbon. The team found ash deposits in deep rock layers near what is now the Buchanan Lake Formation. According to their article, "coal ash dispersed by the explosive Siberian Trap eruption would be expected to have an associated release of toxic elements in impacted water bodies where fly ash slurries developed. ... Mafic megascale eruptions are long-lived events that would allow significant build-up of global ash clouds."[113][114] In a statement, Grasby said, "In addition to these volcanoes causing fires through coal, the ash it spewed was highly toxic and was released in the land and water, potentially contributing to the worst extinction event in earth history."[115] In 2013, a team led by Q.Y. Yang reported the total amounts of important volatiles emitted from the Siberian Traps are

  • 8.5 × 107 Tg CO2,
  • 4.4 × 106 Tg CO,
  • 7.0 × 106 Tg H2S, and
  • 6.8 × 107 Tg SO2.

The data support a popular notion that the end-Permian mass extinction on the Earth was caused by the emission of enormous amounts of volatiles from the Siberian Traps into the atmosphere.[116]

In 2015, evidence and a timeline indicated the extinction was caused by events in the large igneous province of the Siberian Traps.[117][118][119][120] Carbon dioxide levels prior to and after the eruptions are poorly constrained, but may have jumped from between 500 and 4000 PPM prior to the extinction event to around 8000 PPM after the extinction.[121]

In 2020 scientists reconstructed the mechanisms that led to the extinction event in a biogeochemical model, showed the consequences of the greenhouse effect on the marine environment and reported that the mass extinction can be traced back to volcanic CO2 emissions.[122][123] Further evidence – based on paired coronene-mercury spikes – for a volcanic combustion cause of the mass extinction was published in 2020.[124][125]

Methane clathrate gasification

Methane clathrates, also known as methane hydrates, consist of methane molecules trapped in cages of water molecules. The methane, produced by methanogens (microscopic single-celled organisms), has a 13C ⁄ 12C ratio about 6.0% below normal (δ13C −6.0%). At the right combination of pressure and temperature, the methane is trapped in clathrates fairly close to the surface of permafrost and, in much larger quantities, on continental shelves and the deeper seabed close to them. Oceanic methane hydrates are usually found buried in sediments where the seawater is at least 300 m (980 ft) deep. They can be found up to about 2,000 m (6,600 ft) below the sea floor, but usually only about 1,100 m (3,600 ft) below the sea floor.[126]

The release of methane from the clathrates has been considered as a cause because scientists have found worldwide evidence of a swift decrease of about 1% in the 13C ⁄ 12C isotope ratio in carbonate rocks from the end-Permian.[59][127] This is the first, largest, and most rapid of a series of negative and positive excursions (decreases and increases in 13C ⁄ 12C ratio) that continues until the isotope ratio abruptly stabilised in the middle Triassic, followed soon afterwards by the recovery of calcifying life forms (organisms that use calcium carbonate to build hard parts such as shells).[23] While a variety of factors may have contributed to this drop in the 13C ⁄ 12C ratio, , a 2002 review found most of them to be insufficient to account fully for the observed amount:[128]

  • Gases from volcanic eruptions have a 13C ⁄ 12C ratio about 0.5 to 0.8% below standard (δ13C about −0.5 to −0.8%), but an assessment made in 1995 concluded that the amount required to produce a reduction of about 1.0% worldwide requires eruptions greater by orders of magnitude than any for which evidence has been found.[129] (However, this analysis addressed only CO2 produced by the magma itself, not from interactions with carbon bearing sediments, as later proposed.)
  • A reduction in organic activity would extract 12C more slowly from the environment and leave more of it to be incorporated into sediments, thus reducing the 13C ⁄ 12C ratio. Biochemical processes preferentially use the lighter isotopes since chemical reactions are ultimately driven by electromagnetic forces between atoms and lighter isotopes respond more quickly to these forces, but a study of a smaller drop of 0.3 to 0.4% in 13C ⁄ 12C (δ13C −3 to −4 ‰) at the Paleocene-Eocene Thermal Maximum (PETM) concluded that even transferring all the organic carbon (in organisms, soils, and dissolved in the ocean) into sediments would be insufficient: Even such a large burial of material rich in 12C would not have produced the 'smaller' drop in the 13C ⁄ 12C ratio of the rocks around the PETM.[129]
  • Buried sedimentary organic matter has a 13C ⁄ 12C ratio 2.0 to 2.5% below normal (δ13C −2.0 to −2.5%). Theoretically, if the sea level fell sharply, shallow marine sediments would be exposed to oxidation. But 6500–8400 gigatons (1 gigaton = 109 metric tons) of organic carbon would have to be oxidized and returned to the ocean-atmosphere system within less than a few hundred thousand years to reduce the 13C ⁄ 12C ratio by 1.0%, which is not thought to be a realistic possibility.[57] Moreover, sea levels were rising rather than falling at the time of the extinction.[130]
  • Rather than a sudden decline in sea level, intermittent periods of ocean-bottom hyperoxia and anoxia (high-oxygen and low- or zero-oxygen conditions) may have caused the 13C ⁄ 12C ratio fluctuations in the Early Triassic;[23] and global anoxia may have been responsible for the end-Permian blip. The continents of the end-Permian and early Triassic were more clustered in the tropics than they are now, and large tropical rivers would have dumped sediment into smaller, partially enclosed ocean basins at low latitudes. Such conditions favor oxic and anoxic episodes; oxic/anoxic conditions would result in a rapid release/burial, respectively, of large amounts of organic carbon, which has a low 13C ⁄ 12C ratio because biochemical processes use the lighter isotopes more.[131] That or another organic-based reason may have been responsible for both that and a late Proterozoic/Cambrian pattern of fluctuating 13C ⁄ 12C ratios.[23]

Other hypotheses include mass oceanic poisoning, releasing vast amounts of CO2,[132] and a long-term reorganisation of the global carbon cycle.[128]

Prior to consideration of the inclusion of roasting carbonate sediments by volcanism, the only proposed mechanism sufficient to cause a global 1% reduction in the 13C ⁄ 12C ratio was the release of methane from methane clathrates.[57] Carbon-cycle models confirm that it would have had enough effect to produce the observed reduction.[128][132] It was also suggested that a large-scale release of methane and other greenhouse gases from the ocean into the atmosphere was connected to the anoxic events and euxinic (i.e. sulfidic) events at the time, with the exact mechanism compared to the 1986 Lake Nyos disaster [133]

The area covered by lava from the Siberian Traps eruptions is about twice as large as was originally thought, and most of the additional area was shallow sea at the time. The seabed probably contained methane hydrate deposits, and the lava caused the deposits to dissociate, releasing vast quantities of methane.[134] A vast release of methane might cause significant global warming since methane is a very powerful greenhouse gas. Strong evidence suggests the global temperatures increased by about 6 °C (10.8 °F) near the equator and therefore by more at higher latitudes: a sharp decrease in oxygen isotope ratios (18O ⁄ 16O);[135] the extinction of Glossopteris flora (Glossopteris and plants that grew in the same areas), which needed a cold climate, with its replacement by floras typical of lower paleolatitudes.[136]

However, the pattern of isotope shifts expected to result from a massive release of methane does not match the patterns seen throughout the Early Triassic. Not only would such a cause require the release of five times as much methane as postulated for the PETM,[23] but would it also have to be reburied at an unrealistically high rate to account for the rapid increases in the 13C ⁄ 12C ratio (episodes of high positive δ13C) throughout the early Triassic before it was released several times again.[23]

As of 2022, latest research suggests that greenhouse gas release during the extinction event was dominated by volcanic carbon dioxide:[137] while methane release had to have contributed, isotopic signatures show that thermogenic methane released from the Siberian Traps had consistently played a larger role than methane from clathrates and any other biogenic sources such as wetlands during the event.[138]

Hypercapnia and ocean acidification

Marine organisms are more sensitive to changes in CO2 (carbon dioxide) levels than terrestrial organisms for a variety of reasons. CO2 is 28 times more soluble in water than is oxygen. Marine animals normally function with lower concentrations of CO2 in their bodies than land animals, as the removal of CO2 in air-breathing animals is impeded by the need for the gas to pass through the respiratory system's membranes (lungs' alveolus, tracheae, and the like), even when CO2 diffuses more easily than oxygen. In marine organisms, relatively modest but sustained increases in CO2 concentrations hamper the synthesis of proteins, reduce fertilization rates, and produce deformities in calcareous hard parts. An analysis of marine fossils from the Permian's final Changhsingian stage found that marine organisms with a low tolerance for hypercapnia (high concentration of carbon dioxide) had high extinction rates, and the most tolerant organisms had very slight losses. The most vulnerable marine organisms were those that produced calcareous hard parts (from calcium carbonate) and had low metabolic rates and weak respiratory systems, notably calcareous sponges, rugose and tabulate corals, calcite-depositing brachiopods, bryozoans, and echinoderms; about 81% of such genera became extinct. Close relatives without calcareous hard parts suffered only minor losses, such as sea anemones, from which modern corals evolved. Animals with high metabolic rates, well-developed respiratory systems, and non-calcareous hard parts had negligible losses except for conodonts, in which 33% of genera died out. This pattern is also consistent with what is known about the effects of hypoxia, a shortage but not total absence of oxygen. However, hypoxia cannot have been the only killing mechanism for marine organisms. Nearly all of the continental shelf waters would have had to become severely hypoxic to account for the magnitude of the extinction, but such a catastrophe would make it difficult to explain the very selective pattern of the extinction. Mathematical models of the Late Permian and Early Triassic atmospheres show a significant but protracted decline in atmospheric oxygen levels, with no acceleration near the P–Tr boundary. Minimum atmospheric oxygen levels in the Early Triassic are never less than present-day levels and so the decline in oxygen levels does not match the temporal pattern of the extinction.[78]

In addition, an increase in CO2 concentration is inevitably linked to ocean acidification, consistent with the preferential extinction of heavily calcified taxa and other signals in the rock record that suggest a more acidic ocean.[139] The decrease in ocean pH is calculated to be up to 0.7 units.[140]

Anoxia and euxinia

Evidence for widespread ocean anoxia (severe deficiency of oxygen) and euxinia (presence of hydrogen sulfide) is found from the Late Permian to the Early Triassic. Throughout most of the Tethys and Panthalassic Oceans, evidence for anoxia, including fine laminations in sediments, small pyrite framboids, high uranium/thorium ratios, and biomarkers for green sulfur bacteria, appear at the extinction event.[141] However, in some sites, including Meishan, China, and eastern Greenland, evidence for anoxia precedes the extinction.[142][143] Biomarkers for green sulfur bacteria, such as isorenieratane, the diagenetic product of isorenieratene, are widely used as indicators of photic zone euxinia because green sulfur bacteria require both sunlight and hydrogen sulfide to survive. Their abundance in sediments from the P–T boundary indicates hydrogen sulfide was present even in shallow waters. The disproportionate extinction of high-latitude marine species provides further evidence for oxygen depletion as a killing mechanism; low-latitude species living in warmer, less oxygenated waters are naturally better adapted to lower levels of oxygen and are able to migrate to higher latitudes during periods of global warming, whereas high-latitude organisms are unable to escape from warming, hypoxic waters at the poles.[144]

This spread of toxic, oxygen-depleted water would have devastated marine life, causing widespread die-offs. Models of ocean chemistry suggest that anoxia and euxinia were closely associated with hypercapnia (high levels of carbon dioxide).[145] This suggests that poisoning from hydrogen sulfide, anoxia, and hypercapnia acted together as a killing mechanism. Hypercapnia best explains the selectivity of the extinction, but anoxia and euxinia probably contributed to the high mortality of the event. The persistence of anoxia through the Early Triassic may explain the slow recovery of marine life after the extinction. Models also show that anoxic events can cause catastrophic hydrogen sulfide emissions into the atmosphere (see below).[146]

The sequence of events leading to anoxic oceans may have been triggered by carbon dioxide emissions from the eruption of the Siberian Traps.[146] In that scenario, warming from the enhanced greenhouse effect would reduce the solubility of oxygen in seawater, causing the concentration of oxygen to decline. Increased weathering of the continents due to warming and the acceleration of the water cycle would increase the riverine flux of phosphate to the ocean. The phosphate would have supported greater primary productivity in the surface oceans. The increase in organic matter production would have caused more organic matter to sink into the deep ocean, where its respiration would further decrease oxygen concentrations. Once anoxia became established, it would have been sustained by a positive feedback loop because deep water anoxia tends to increase the recycling efficiency of phosphate, leading to even higher productivity.

A severe anoxic event at the end of the Permian would have allowed sulfate-reducing bacteria to thrive, causing the production of large amounts of hydrogen sulfide in the anoxic ocean, turning it euxinic. Upwelling of this water may have released massive hydrogen sulfide emissions into the atmosphere and would poison terrestrial plants and animals and severely weaken the ozone layer, exposing much of the life that remained to fatal levels of UV radiation.[146] Indeed, biomarker evidence for anaerobic photosynthesis by Chlorobiaceae (green sulfur bacteria) from the Late-Permian into the Early Triassic indicates that hydrogen sulfide did upwell into shallow waters because these bacteria are restricted to the photic zone and use sulfide as an electron donor. The hypothesis has the advantage of explaining the mass extinction of plants, which would have added to the methane levels and should otherwise have thrived in an atmosphere with a high level of carbon dioxide. Fossil spores from the end-Permian further support the theory:[147] many show deformities that could have been caused by ultraviolet radiation, which would have been more intense after hydrogen sulfide emissions weakened the ozone layer.

Aridification

Analysis of the fossil river deposits of the floodplains indicate a shift from meandering to braided river patterns, indicating a very abrupt drying of the climate.[148] The climate change may have taken as little as 100,000 years, prompting the extinction of the unique Glossopteris flora and its associated herbivores, followed by the carnivorous guild.[149]

Evidence from the Sydney Basin of eastern Australia, on the other hand, suggests that the expansion of semi-arid and arid climatic belts across Pangaea was not immediate but was instead a gradual, prolonged process. Apart from the disappearance of peatlands, there was little evidence of significant sedimentological changes in depositional style across the Permian-Triassic boundary.[150] Instead, a modest shift to amplified seasonality and hotter summers is suggested by palaeoclimatological models based on weathering proxies from the region's Late Permian and Early Triassic deposits.[151]

Supercontinent Pangaea
Map of Pangaea showing where today's continents were at the Permian–Triassic boundary

In the mid-Permian (during the Kungurian age of the Permian's Cisuralian epoch), Earth's major continental plates joined, forming a supercontinent called Pangaea, which was surrounded by the superocean, Panthalassa.

Oceanic circulation and atmospheric weather patterns during the mid-Permian produced seasonal monsoons near the coasts and an arid climate in the vast continental interior.[152]

As the supercontinent formed, the ecologically diverse and productive coastal areas shrank. The shallow aquatic environments were eliminated and exposed formerly protected organisms of the rich continental shelves to increased environmental volatility.

Pangaea's formation depleted marine life at near catastrophic rates. However, Pangaea's effect on land extinctions is thought to have been smaller. In fact, the advance of the therapsids and increase in their diversity is attributed to the late Permian, when Pangaea's global effect was thought to have peaked.

While Pangaea's formation certainly initiated a long period of marine extinction, its impact on the "Great Dying" and the end of the Permian is uncertain.

Microbes

A hypothesis published in 2014 posits that a genus of anaerobic methanogenic archaea known as Methanosarcina was responsible for the event.[19] Three lines of evidence suggest that these microbes acquired a new metabolic pathway via gene transfer at about that time, enabling them to efficiently metabolize acetate into methane. That would have led to their exponential reproduction, allowing them to rapidly consume vast deposits of organic carbon that had accumulated in the marine sediment. The result would have been a sharp buildup of methane and carbon dioxide in the Earth's oceans and atmosphere, in a manner that may be consistent with the 13C/12C isotopic record. Massive volcanism facilitated this process by releasing large amounts of nickel, a scarce metal which is a cofactor for enzymes involved in producing methane.[19] On the other hand, in the canonical Meishan sections, the nickel concentration increases somewhat after the δ13C concentrations have begun to fall.[153]

Combination of causes

Possible causes supported by strong evidence appear to describe a sequence of catastrophes, each worse than the last: the Siberian Traps eruptions were bad enough alone, but because they occurred near coal beds and the continental shelf, they also triggered very large releases of carbon dioxide and methane.[78] The resultant global warming may have caused perhaps the most severe anoxic event in the oceans' history: according to this theory, the oceans became so anoxic, anaerobic sulfur-reducing organisms dominated the chemistry of the oceans and caused massive emissions of toxic hydrogen sulfide.[78]

However, there may be some weak links in this chain of events: the changes in the 13C/12C ratio expected to result from a massive release of methane do not match the patterns seen throughout the early Triassic;[23] and the types of oceanic thermohaline circulation that may have existed at the end of the Permian are not likely to have supported deep-sea anoxia.[154]

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