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97-664: There is no designated type locality for this formation. It was named for a development at the mouth of Moencopie Wash in the Grand Canyon area by Ward in 1901. In 1917 a 'substitute' type locality was located by Gregory in the wall of the Little Colorado Canyon , about 5 miles below Tanner Crossing in Coconino County, Arizona . While in the Great Basin, Bassler and Reeside characterized and named

194-458: A lithological sequence as being on or very close to the Permian–Triassic boundary in rocks that are unsuitable for radiometric dating or have a lack of suitable index fossils . However, 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 during the earliest Triassic. The very idea of

291-620: A brief period of domination in the early Spathian, probably related to a transient oxygenation of deep waters. Neospathodid conodonts survived the crisis but underwent proteromorphosis. In the PTME's aftermath, disaster taxa of benthic foraminifera filled many of their vacant niches. The recovery of benthic foraminifera was very slow and frequently interrupted until the Spathian. In the Tethys, foraminiferal communities remained low in diversity into

388-632: A ceiling limiting the maximum ecological complexity of marine ecosystems until the Spathian. Recovery biotas appear to have been ecologically uneven and unstable into the Anisian , making them vulnerable to environmental stresses. Whereas most marine communities were fully recovered by the Middle Triassic, global marine diversity reached pre-extinction values no earlier than the Middle Jurassic, approximately 75 million years after

485-449: A duration of 60 ± 48 thousand years. A large, abrupt global decrease in δ C , the ratio of the stable isotope carbon-13 to that of carbon-12 , coincides with this extinction, and is sometimes used to identify the Permian–Triassic boundary and PTME in rocks that are unsuitable for radiometric dating . The negative carbon isotope excursion's magnitude was 4-7% and lasted for approximately 500 kyr, though estimating its exact value

582-513: A family of large-size fusuline foraminifera . 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. 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

679-420: A fungal origin for Reduviasporonites , diluting these critiques. 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 or that the extinction was long and spread out over a few million years, with a sharp peak in the last million years of

776-491: A fungal spike has been criticized on several grounds, including: Reduviasporonites , the most common supposed fungal spore, may be a fossilized alga ; the spike did not appear worldwide; and in many places it did not fall on the Permian–Triassic boundary. 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. Newer chemical evidence agrees better with

873-581: 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. In South China, the subtropical Cathaysian gigantopterid dominated rainforests abruptly collapsed. The floral extinction in South China is associated with bacterial blooms in soil and nearby lacustrine ecosystems, with soil erosion resulting from

970-420: A high background extinction rate (by implication, taxa with a high turnover ). The extinction rate of marine organisms was catastrophic. Bioturbators were extremely severely affected, as evidenced by the loss of the sedimentary mixed layer in many marine facies during the end-Permian extinction. Surviving marine invertebrate groups included articulate brachiopods (those with a hinge), which had undergone

1067-487: 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 first appearance of the conodont Hindeodus parvus has been used to delineate the Permian-Triassic boundary. The extinction occurred between 251.941 ± 0.037 and 251.880 ± 0.031 million years ago,

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1164-566: A massive rearrangement of ecosystems does occur, with plant abundances and distributions changing profoundly and all the forests virtually disappearing. The dominant floral groups changed, with many groups of land plants entering abrupt decline, such as Cordaites ( gymnosperms ) and Glossopteris ( seed ferns ). The severity of plant extinction has been disputed. The Glossopteris -dominated flora that characterised high-latitude Gondwana collapsed in Australia around 370,000 years before

1261-553: A severe bottleneck in diversity. Evidence from South China indicates the foraminiferal extinction had two pulses. Foraminiferal biodiversity hotspots shifted into deeper waters during the PTME. 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. Foraminifera that were calcaerous suffered an extinction rate of 91%. The reason why lagenides survived while fusulinoidean fusulinides went completely extinct may have been due to

1358-481: A slow decline in numbers since the P–Tr extinction; the Ceratitida order of ammonites ; and crinoids ("sea lilies"), 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

1455-406: A specific region were more likely to go extinct than cosmopolitan taxa. There was little latitudinal difference in the survival rates of taxa. Organisms that inhabited refugia less affected by global warming experienced lesser or delayed extinctions. Among benthic organisms the extinction event multiplied background extinction rates , and therefore caused maximum species loss to taxa that had

1552-521: Is challenging due to diagenetic alteration of many sedimentary facies spanning the boundary. Further evidence for environmental change around the Permian-Triassic boundary suggests an 8 °C (14 °F) rise in temperature, and an increase in CO 2 levels to 2,500  ppm (for comparison, the concentration immediately before the Industrial Revolution was 280 ppm , and

1649-468: Is difficult to know whether the end-Capitanian had finished, depending on the factor considered. Many of the extinctions once dated to the Permian-Triassic boundary have more 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

1746-600: Is likely attributable to their ability to thrive in a wide range of environmental conditions. Conodonts saw a rapid recovery during the Induan, with anchignathodontids experiencing a diversity peak in the earliest Induan. Gondolellids diversified at the end of the Griesbachian; this diversity spike was most responsible for the overall conodont diversity peak in the Smithian. Segminiplanate conodonts again experienced

1843-451: Is likely that post-extinction microbial mats played a vital, indispensable role in the survival and recovery of various bioturbating organisms. The microbialite refuge hypothesis has been criticised as reflecting a taphonomic bias due to the greater preservation potential of microbialite deposits, however, rather than a genuine phenomenon. Ichnocoenoses show that marine ecosystems recovered to pre-extinction levels of ecological complexity by

1940-702: Is the type locality. The concept is similar to type site in archaeology . End-Permian mass extinction Approximately 251.9 million years ago, the Permian–Triassic ( P–T , P–Tr ) extinction event ( PTME ; also known as the Late Permian extinction event , the Latest Permian extinction event , the End-Permian extinction event , and colloquially as the Great Dying ) forms

2037-525: The Bowen Basin of Queensland indicates numerous intermittent periods of marine environmental stress from the middle to late Lopingian leading up to the end-Permian extinction proper, supporting aspects of the gradualist hypothesis. Additionally, the decline in marine species richness and the structural collapse of marine ecosystems may have been decoupled as well, with the former preceding the latter by about 61,000 years according to one study. Whether

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2134-555: The Cedar Mesa area to deltaic sandstones and shallow marine limestones at Capitol Reef. In eastern Nevada and northwestern Utah, it thickens dramatically, then transitions to the Woodside , Thaynes , and Mahogany formations . The general deposition setting was sluggish rivers traversing a flat, featureless coastal plain to the sea. The low relief meant that the shoreline moved great distances with changes of sea level or even with

2231-643: The Chinle Formation in turn rests unconformably on the Moenkopi. Both unconformities are locally angular unconformities . The lower unconformity corresponds to the regional Tr-1 unconformity and the upper to the regional Tr-3 unconformity. The Tr-1 unconformity represents a hiatus of at least 20 million years while Tr-2 represents a hiatus of about 10 million years. Members differ considerably from east to west, in part because sandstone beds corresponding to marine transgressions are used to define members to

2328-598: The Middle Triassic ) due to the severity of the extinction. However, studies in Bear Lake County , near Paris, Idaho , and nearby sites in Idaho and Nevada showed a relatively quick rebound in a localized Early Triassic marine ecosystem ( Paris biota ), taking around 1.3 million years to recover, while an unusually diverse and complex ichnobiota is known from Italy less than a million years after

2425-565: The Olenekian Virgin Limestone Member of the Moenkopi Formation, in south-western Utah. The discovery of 27 species from 18 genera of two subclasses in these sites in 2013 cast doubt on previous claims that the bivalve fauna only recovered in the Middle Triassic after the end-Permian mass extinction . The first subclass, Pteriomorphia , includes mainly genera that survived the mass extinction, while

2522-516: 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)

2619-489: The Wupatki Member . [REDACTED] Media related to Moenkopi Formation at Wikimedia Commons Type locality (geology) Type locality , also called type area , is the locality where a particular rock type , stratigraphic unit or mineral species is first identified. If the stratigraphic unit in a locality is layered, it is called a stratotype , whereas the standard of reference for unlayered rocks

2716-406: The ocean acidification that resulted from increased atmospheric CO 2 . Organisms that relied on haemocyanin or haemoglobin for transporting oxygen were more resistant to extinction than those utilising haemerythrin or oxygen diffusion. There is also evidence that endemism was a strong risk factor influencing a taxon's likelihood of extinction. Bivalve taxa that were endemic and localised to

2813-568: The "Big Five" mass extinctions of the Phanerozoic . There is evidence for one to three distinct pulses, or phases, of extinction. The scientific consensus is that the main cause of the extinction was the flood basalt volcanic eruptions that created the Siberian Traps , which released sulfur dioxide and carbon dioxide , resulting in euxinia (oxygen-starved, sulfurous oceans), elevating global temperatures, and acidifying

2910-609: The Anisian. Metazoan reefs became common again during the Anisian because the oceans cooled down then from their overheated state during the Early Triassic. Biodiversity amongst metazoan reefs did not recover until well into the Anisian, millions of years after non-reef ecosystems recovered their diversity. Microbially induced sedimentary structures (MISS) from the earliest Triassic have been found to be associated with abundant opportunistic bivalves and vertical burrows, and it

3007-634: The Black Dragon, Torrey, and Moody Canyon members in the Paradox Basin and Plateau Sedimentary Province in 1974. Contacts were revised yet again by Welsh and others in 1979. Kietzke modified the age to Early and Middle Triassic using biostratigraphic dating in 1988. The Anton Chico Member was assigned in the Palo Duro Basin and areal limits set by Lucas and Hunt in 1989. In 1991 areal limits were set again by Lucas and Hayden. An overview

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3104-712: The Early Triassic; and they dominated many surviving communities across the recovery from the mass extinction. Microbialite deposits appear to have declined in the early Griesbachian synchronously with a significant sea level drop that occurred then. Metazoan-built reefs reemerged during the Olenekian, mainly being composed of sponge biostrome and bivalve builups. Keratose sponges were particularly noteworthy in their integral importance to Early Triassic microbial-metazoan reef communities, and they helped to create stability in heavily damaged ecosystems during early phases of biotic recovery. " Tubiphytes "-dominated reefs appeared at

3201-804: The Karoo Basin indicates a protracted extinction lasting a million years. Other evidence from the Karoo deposits suggest it took 50,000 years or less, while a study of coprolites in the Vyazniki fossil beds in Russia suggests it took only a few thousand years. Aridification induced by global warming was the chief culprit behind terrestrial vertebrate extinctions. 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

3298-484: The Late Cretaceous to recover their full diversity. Crinoids ("sea lilies") suffered a selective extinction, resulting in a decrease in the variety of their forms. Though cladistic analyses suggest the beginning of their recovery to have taken place in the Induan, the recovery of their diversity as measured by fossil evidence was far less brisk, showing up in the late Ladinian. Their adaptive radiation after

3395-514: The Middle Triassic, with the exception of a notable Ladinian fauna from the Catalonian Basin. Microbial reefs were common across shallow seas for a short time during the earliest Triassic, predominating in low latitudes while being rarer in higher latitudes, occurring both in anoxic and oxic waters. Polybessurus -like microfossils often dominated these earliest Triassic microbialites . Microbial-metazoan reefs appeared very early in

3492-420: The PTME, being the most severely affected clade among the lophophorates. 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. Foraminifera suffered

3589-666: The PTME, but some tentative evidence suggests they may have survived into the Triassic. 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. However, faunal turnovers in freshwater fish communities occurred in areas like the Kuznetsk Basin. The groups that survived suffered extremely heavy losses of species and some terrestrial vertebrate groups very nearly became extinct at

3686-517: The PTME. The Cordaites flora, which dominated the Angaran floristic realm corresponding to Siberia, collapsed over the course of the extinction. In the Kuznetsk Basin , the aridity-induced extinction of the regions's 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

3783-566: 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, and the increase in predation pressure and durophagy led to the Mesozoic Marine Revolution . Marine vertebrates recovered relatively quickly, with complex predator-prey interactions with vertebrates at

3880-447: The Permian-Triassic boundary are highly variable depending on the location and preservation quality of any given site. Plants are relatively immune to mass extinction, with the impact of all the major mass extinctions "insignificant" at a family level. Floral diversity losses were more superficial than those of marine animals. Even the reduction observed in species diversity (of 50%) may be mostly due to taphonomic processes. However,

3977-462: 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. 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. Palynological or pollen studies from East Greenland of sedimentary rock strata laid down during

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4074-526: The Permian-Triassic mass extinction marked a key turning point in this ecological shift that began after the Capitanian mass extinction and culminated in the Late Jurassic . Typical taxa of shelly benthic faunas were now bivalves , snails , sea urchins and Malacostraca , whereas bony fishes and marine reptiles diversified in the pelagic zone . On land, dinosaurs and mammals arose in

4171-864: The Permian. Statistical analyses of some highly fossiliferous strata in Meishan, Zhejiang Province in southeastern China, suggest that the main extinction was clustered around one peak, while a study of the Liangfengya section found evidence of two extinction waves, MEH-1 and MEH-2, which varied in their causes, and a study of the Shangsi section showed two extinction pulses with different causes too. 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 around 670,000 to 1.17 million years. Palaeoenvironmental analysis of Lopingian strata in

4268-520: 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. 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

4365-815: The Rock Canyon Conglomerate, Virgin Limestone, and Shnabkaib Shale members in 1921. Salt Creek (later replaced by Wupatki and Moqui Members) and the Holbrook Member were found and named in the Black Mesa basin by Hager in 1922. The Sinbad Limestone Member was named in the Paradox Basin by Gilluly and Reeside in 1928. Williams and Gregory named the Timpoweap Member in the Plateau sedimentary province in 1947. The Wupatki Member

4462-432: The Spathian and Anisian. 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. Other explanations state that life was delayed in its recovery because grim conditions returned periodically over

4559-545: The Spathian. Despite high taxonomic turnover, the ecological life modes of Early Triassic ostracods remained rather similar to those of pre-PTME ostracods. Bryozoans in the Early Triassic were restricted to the Boreal realm. They were also not diverse, represented mainly by members of Trepostomatida . During the Middle Triassic, there was a rise in bryozoan diversity, which peaked in the Carnian. However, bryozoans took until

4656-446: The Triassic, taking over niches that were filled primarily by brachiopods before the mass extinction event. Bivalves were once thought to have outcompeted brachiopods, but this outdated hypothesis about the brachiopod-bivalve transition has been disproven by Bayesian analysis . The success of bivalves in the aftermath of the extinction event may have been a function of them possessing greater resilience to environmental stress compared to

4753-476: The amount today is about 422 ppm ). There is also evidence of increased ultraviolet radiation reaching the earth, causing the mutation of plant spores. 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. This "fungal spike" has been used by some paleontologists to identify

4850-517: The boundary between the Permian and Triassic geologic periods , and with them the Paleozoic and Mesozoic eras. It is Earth 's most severe known extinction event , with the extinction of 57% of biological families , 83% of genera, 81% of marine species and 70% of terrestrial vertebrate species. It is also the greatest known mass extinction of insects . It is the greatest of

4947-423: The brachiopods that they coexisted with, whilst other studies have emphasised the greater niche breadth of the former. The rise of bivalves to taxonomic and ecological dominance over brachiopods was not synchronous, however, and brachiopods retained an outsized ecological dominance into the Middle Triassic even as bivalves eclipsed them in taxonomic diversity. Some researchers think the brachiopod-bivalve transition

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5044-480: 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 ). However, recovery was also differential between taxa. Some survivors became extinct some million years after the extinction event without having rediversified ( dead clade walking , e.g.

5141-635: The course of the Early Triassic, causing further extinction events, such as the Smithian-Spathian boundary extinction . Continual episodes of extremely hot climatic conditions during the Early Triassic have been held responsible for the delayed recovery of oceanic life, in particular skeletonised taxa that are most vulnerable to high carbon dioxide concentrations. The relative delay in the recovery of benthic organisms has been attributed to widespread anoxia, but high abundances of benthic species contradict this explanation. A 2019 study attributed

5238-465: The die-off of plants being their likely cause. Wildfires too likely played a role in the fall of Gigantopteris . 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. The tempo of the terrestrial vertebrate extinction is disputed. Some evidence from

5335-423: The dissimilarity of recovery times between different ecological communities to differences in local environmental stress during the biotic recovery interval, with regions experiencing persistent environmental stress post-extinction recovering more slowly, supporting the view that recurrent environmental calamities were culpable for retarded biotic recovery. Recurrent Early Triassic environmental stresses also acted as

5432-492: The end of the Olenekian, representing the earliest platform-margin reefs of the Triassic, though they did not become abundant until the late Anisian, when reefs' species richness increased. The first scleractinian corals appear in the late Anisian as well, although they would not become the dominant reef builders until the end of the Triassic period. Bryozoans, after sponges, were the most numerous organisms in Tethyan reefs during

5529-440: 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. It is difficult to analyze extinction and survival rates of land organisms in detail because few terrestrial fossil beds span

5626-504: 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. 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

5723-655: The end-Permian extinction. Additionally, the complex Guiyang biota found near Guiyang , China also indicates life thrived in some places just a million years after the mass extinction, as does a fossil assemblage known as the Shanggan fauna found in Shanggan, China, the Wangmo biota from the Luolou Formation of Guizhou, and a gastropod fauna from the Al Jil Formation of Oman. Regional differences in

5820-498: The expansion of more habitable climatic zones. Brachiopod taxa during the Anisian recovery interval were only phylogenetically related to Late Permian brachiopods at a familial taxonomic level or higher; the ecology of brachiopods had radically changed from before in the mass extinction's aftermath. Ostracods were extremely rare during the basalmost Early Triassic. Taxa associated with microbialites were disproportionately represented among ostracod survivors. Ostracod recovery began in

5917-415: The extinction event resulted in forms possessing flexible arms becoming widespread; motility , predominantly a response to predation pressure, also became far more prevalent. Though their taxonomic diversity remained relatively low, crinoids regained much of their ecological dominance by the Middle Triassic epoch. Stem-group echinoids survived the PTME. The survival of miocidarid echinoids such as Eotiaris

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6014-552: The extinction event. 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 . Before

6111-409: 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 . Data from Kap Stosch suggest that floral species richness was not significantly affected during

6208-570: The fauna. Temnospondyli include Eocyclotosaurus , Quasicyclotosaurus , Wellesaurus , Vigilius , and Cosgriffius . The rhynchosaur Ammorhynchus is known, but rare. Anisodontosaurus is an enigmatic reptile only known from a few tooth-bearing jaws. The poposauroid archosaur Arizonasaurus is known from one relatively complete skeleton and a significant amount of other isolated material. Footprints and several fragmentary body fossils are known from dicynodonts . The footprints of Cheirotherium and Rhynchosauroides are common in

6305-588: The final extinction killed off only about 80% of marine species alive at that time, whereas 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. For example, all dinocephalian genera died out at the end of the Guadalupian, as did the Verbeekinidae ,

6402-409: The final two sedimentary zones containing conodonts from the Permian. The decrease in diversity was probably caused by a sharp increase in extinctions, rather than a decrease in speciation . The extinction primarily affected organisms with calcium carbonate skeletons, especially those reliant on stable CO 2 levels to produce their skeletons. These organisms were susceptible to the effects of

6499-619: The full impact of the event. Many sedimentary sequences from South China show synchronous terrestrial and marine extinctions. Research in the Sydney Basin of the PTME's duration and course also supports a synchronous occurrence of the terrestrial and marine biotic collapses. Other scientists believe the terrestrial mass extinction began between 60,000 and 370,000 years before the onset of the marine mass extinction. Chemostratigraphic analysis from sections in Finnmark and Trøndelag shows

6596-469: The gasification of methane clathrates ; emissions of methane by novel methanogenic microorganisms nourished by minerals dispersed in the eruptions; longer and more intense El Niño events; and an extraterrestrial impact which created the Araguainha crater and caused seismic release of methane and the destruction of the ozone layer with increased exposure to solar radiation. Previously, it

6693-441: The greater range of environmental tolerance and greater geographic distribution of the former compared to the latter. 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. Ichthyosaurs , which evolved immediately before the PTME, were also PTME survivors. The Lilliput effect ,

6790-421: The greatest loss of species diversity. In the case of the brachiopods, at least, surviving taxa were generally small, rare members of a formerly diverse community. Conodonts were severely affected both in terms of taxonomic and morphological diversity, although not as severely as during the Capitanian mass extinction. The ammonoids , which had been in a long-term decline for the 30 million years since

6887-628: 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). Tangasaurids were largely unaffected. Gorgonopsians are traditionally thought to have gone extinct during

6984-423: The inarticulate brachiopod Lingularia , and the foraminifera Earlandia and Rectocornuspira kalhori , the latter of which is sometimes classified under the genus Ammodiscus . Their guild diversity was also low. Post-PTME faunas had a flat, insignificant latitudinal diversity gradient. The speed of recovery from the extinction is disputed. Some scientists estimate that it took 10 million years (until

7081-724: The mass extinction, exemplifying the Lilliput effect's opposite, which has been dubbed the Brobdingnag effect. 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; according to some sources, it may well be the only mass extinction to significantly affect insect diversity. 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

7178-512: 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. However, some observed entomofaunal declines in the PTME were biogeographic changes rather than outright extinctions. The geological record of terrestrial plants is sparse and based mostly on pollen and spore studies. Floral changes across

7275-411: The oceans . The level of atmospheric carbon dioxide rose from around 400 ppm to 2,500 ppm with approximately 3,900 to 12,000 gigatonnes of carbon being added to the ocean-atmosphere system during this period. Several other contributing factors have been proposed, including the emission of carbon dioxide from the burning of oil and coal deposits ignited by the eruptions; emissions of methane from

7372-475: The pace of biotic recovery existed, which suggests that the impact of the extinction may have been felt less severely in some areas than others, with differential environmental stress and instability being the source of the variance. In addition, it has been proposed that although overall taxonomic diversity rebounded rapidly, functional ecological diversity took much longer to return to its pre-extinction levels; one study concluded that marine ecological recovery

7469-478: The parameters were now shared differently among clades . Ostracods experienced prolonged diversity perturbations during the Changhsingian before the PTME proper, when immense proportions of them abruptly vanished. At least 74% of ostracods died out during the PTME itself. Bryozoans had been on a long-term decline throughout the Late Permian epoch before they suffered even more catastrophic losses during

7566-516: The phenomenon of dwarfing of species during and immediately following a mass extinction event, has been observed across the Permian-Triassic boundary, notably occurring in foraminifera, brachiopods, bivalves, and ostracods. Though gastropods that survived the cataclysm were smaller in size than those that did not, it remains debated whether the Lilliput effect truly took hold among gastropods. Some gastropod taxa, termed "Gulliver gastropods", ballooned in size during and immediately following

7663-405: The previous extinction interval. Another study of latest Permian vertebrates in the Karoo Basin found that 54% of them went extinct due to the PTME. 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 "Palaeozoic evolutionary fauna" declined while the "modern evolutionary fauna" achieved greater dominance;

7760-456: The quick recovery seen in nektonic organisms such as ammonoids , which exceeded pre-extinction diversities already two million years after the crisis, and conodonts, which diversified considerably over the first two million years of the Early Triassic. 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 . That recovery

7857-479: The second, Heteroconchia , is represented mainly by genera that evolved in the Early Triassic. A diverse fossil vertebrate fauna has been described from the Moenkopi Formation , mainly from the Wupatki Member and Holbrook Member of northern Arizona. Described basal vertebrates include freshwater hybodont sharks , coelacanths , and lungfish . Temnospondyl amphibians are a common component of

7954-412: The sequence of environmental disasters to have effectively constituted a single, prolonged extinction event, perhaps depending on which species is considered. This older theory, still supported in some recent papers, proposes that there were two major extinction pulses 9.4 million years apart, separated by a period of extinctions that were less extensive, but still well above the background level, and that

8051-417: The snail family Bellerophontidae ), whereas others rose to dominance over geologic times (e.g., bivalves). A cosmopolitanism event began immediately after the end-Permian extinction event. Marine post-extinction faunas were mostly species-poor and were dominated by few disaster taxa such as the bivalves Claraia , Unionites , Eumorphotis , and Promyalina , the conodonts Clarkina and Hindeodus ,

8148-427: The terrestrial and marine extinctions were synchronous or asynchronous is another point of controversy. Evidence from a well-preserved sequence in east Greenland suggests that the terrestrial and marine extinctions began simultaneously. In this sequence, the decline of animal life is concentrated in a period approximately 10,000 to 60,000 years long, with plants taking an additional several hundred thousand years to show

8245-415: The terrestrial extinction occurred after the marine extinction in the tropics. 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

8342-502: The terrestrial floral turnover occurred before the large negative δ C shift during the marine extinction. Dating of the boundary between the Dicynodon and Lystrosaurus assemblage zones in the Karoo Basin indicates that the terrestrial extinction occurred earlier than the marine extinction. The Sunjiagou Formation of South China also records a terrestrial ecosystem demise predating the marine crisis. Other research still has found that

8439-534: The tides. Thickness varies from a feather edge against the Uncompahgre highlands to the east to over 600 metres (2,000 ft) in southwestern Utah. The thickness varies greatly in the Paradox Basin , where the Moenkopi is thin to nonexistent on the crests of salt anticlines and over 400 meters (1,300 feet) thick in the corresponding synclines . The Moenkopi rests unconformably on Paleozoic beds and

8536-420: The top of the food web being known from coprolites five million years after the PTME. Post-PTME hybodonts exhibited extremely rapid tooth replacement. Ichthyopterygians appear to have ballooned in size extremely rapidly following the PTME. Bivalves rapidly recolonised many marine environments in the wake of the catastrophe. Bivalves were fairly rare before the P–Tr extinction but became numerous and diverse in

8633-488: The west but cannot be traced to the east. In different regions, by ascending stratigraphic order, the members are: Paradox Basin: Canyonlands and Glen Canyon area: San Juan Basin and Tucumcari: Other members listed in alphabetical order, with asterisks (*) indicating usage by the U.S. Geological Survey and other usages by state geological surveys: Found in these geologic locations: Found within these parks (incomplete list): Numerous fossils of bivalves were found in

8730-588: Was attributable not only to the end-Permian extinction but also the ecological restructuring that began as a result of the Capitanian extinction. Infaunal habits in bivalves became more common after the PTME. Linguliform brachiopods were commonplace immediately after the extinction event, their abundance having been essentially unaffected by the crisis. Adaptations for oxygen-poor and warm environments, such as increased lophophoral cavity surface, shell width/length ratio, and shell miniaturisation, are observed in post-extinction linguliforms. The surviving brachiopod fauna

8827-497: Was completed by Lucas in 1991, Sprinkel in 1994, Hintze and Axen in 1995 and later, Huntoon and others. The Moenkopi consists of thinly bedded sandstone , mudstone , and shale , with some limestone in the Capitol Reef area. It has a characteristic deep red color and tends to form slopes and benches. The depositional environment varies from fluvial channel and floodplain deposits in the eastern exposures to tidal mudflats in

8924-623: Was first used in the Plateau Sedimentary Province and its age was modified to Early and Middle(?) Triassic by McKee in 1951. Contacts were revised by Robeck in 1956 and Cooley in 1958. The Tenderfoot, Ali Baba, Sewemup, and Pariott Members were named in the Piceance and Uinta Basins by Shoemaker and Newman in 1959. The Hoskinnini Member was assigned in the Black Mesa and Paradox basins by Stewart in 1959. Contacts were revised again by Schell and Yochelson in 1966. Blakey named

9021-410: Was non-selective, consistent with a catastrophic initiator. During the Triassic, diversity rose rapidly, but disparity remained low. 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

9118-425: Was slow in the Early Triassic can be explained by low levels of biological competition due to the paucity of taxonomic diversity, and that biotic recovery explosively accelerated in the Anisian can be explained by niche crowding, a phenomenon that would have drastically increased competition, becoming prevalent by the Anisian. Biodiversity rise thus behaved as a positive feedback loop enhancing itself as it took off in

9215-519: Was still ongoing 50 million years after the extinction, during the latest Triassic, even though taxonomic diversity had rebounded in a tenth of that time. The pace and timing of recovery also differed based on clade and mode of life. Seafloor communities maintained a comparatively low diversity until the end of the Early Triassic, approximately 4 million years after the extinction event. Epifaunal benthos took longer to recover than infaunal benthos. This slow recovery stands in remarkable contrast with

9312-482: 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. 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

9409-435: Was very low in diversity and exhibited no provincialism whatsoever. Brachiopods began their recovery around 250.1 ± 0.3 Ma, as marked by the appearance of the genus Meishanorhynchia , believed to be the first of the progenitor brachiopods that evolved after the mass extinction. Major brachiopod rediversification only began in the late Spathian and Anisian in conjunction with the decline of widespread anoxia and extreme heat and

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