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122-621: The Tākaka Terrane is a Paleozoic terrane that outcrops in the South Island of New Zealand. It is most extensively exposed within the Kahurangi National Park in the Tasman District . The terrane is mostly made up of marble and volcanic rocks but is highly variable in composition. It ranges in age from mid- Cambrian to Devonian time (510–400 Ma), including New Zealand's oldest rocks, which are found in

244-458: 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

366-620: 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

488-622: A cataclysm known as " The Great Dying ", the third and most severe Phanerozoic mass extinction. The early Cambrian climate was probably moderate at first, becoming warmer over the course of the Cambrian, as the second-greatest sustained sea level rise in the Phanerozoic got underway. However, as if to offset this trend, Gondwana moved south, so that, in Ordovician time, most of West Gondwana (Africa and South America) lay directly over

610-632: 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

732-449: 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

854-513: 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

976-445: A foothold on land. These early plants were the forerunners of all plant life on land. During this time, there were four continents: Gondwana (Africa, South America, Australia, Antarctica, Siberia), Laurentia (North America), Baltica (Northern Europe), and Avalonia (Western Europe). The recent rise in sea levels allowed many new species to thrive in water. The Devonian spanned from 419–359 million years ago. Also known as "The Age of

1098-420: 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

1220-491: 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

1342-581: 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

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1464-420: 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

1586-487: 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,

1708-566: 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

1830-553: 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

1952-481: 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

2074-406: 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

2196-521: 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

2318-663: Is considered the first Phanerozoic mass extinction event, and the second deadliest. The Silurian spanned from 444–419 million years ago. The Silurian saw the rejuvenation of life as the Earth recovered from the previous glaciation. This period saw the mass evolution of fish, as jawless fish became more numerous, jawed fish evolved, and the first freshwater fish evolved, though arthropods, such as sea scorpions , were still apex predators . Fully terrestrial life evolved, including early arachnids, fungi, and centipedes. The evolution of vascular plants ( Cooksonia ) allowed plants to gain

2440-468: 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

2562-600: 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

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2684-451: 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

2806-575: Is the sudden appearance of nearly all of the invertebrate animal phyla in great abundance at the beginning of the Cambrian. The first vertebrates appeared in the form of primitive fish, which greatly diversified in the Silurian and Devonian Periods. The first animals to venture onto dry land were the arthropods. Some fish had lungs, and powerful bony fins that in the late Devonian, 367.5 million years ago, allowed them to crawl onto land. The bones in their fins eventually evolved into legs and they became

2928-605: The Appalachians , Caledonides , Ural Mountains , and mountains of Tasmania . The Cambrian spanned from 539–485 million years ago and is the first period of the Paleozoic Era of the Phanerozoic. The Cambrian marked a boom in evolution in an event known as the Cambrian explosion in which the largest number of creatures evolved in any single period of the history of the Earth. Creatures like algae evolved, but

3050-525: 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

3172-531: The Carboniferous Rainforest Collapse . Gondwana was glaciated as much of it was situated around the south pole. The Permian spanned from 299–252 million years ago and was the last period of the Paleozoic Era. At the beginning of this period, all continents joined together to form the supercontinent Pangaea, which was encircled by one ocean called Panthalassa . The land mass was very dry during this time, with harsh seasons, as

3294-645: The Early Palaeozoic Icehouse , culminating in the Hirnantian glaciation, 445  million years ago at the end of the Ordovician. The middle Paleozoic was a time of considerable stability. Sea levels had dropped coincident with the ice age, but slowly recovered over the course of the Silurian and Devonian. The slow merger of Baltica and Laurentia, and the northward movement of bits and pieces of Gondwana created numerous new regions of relatively warm, shallow sea floor. As plants took hold on

3416-598: 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

3538-531: 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 the "Big Five" mass extinctions of

3660-533: 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

3782-516: 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)

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3904-571: The South Pole . The early Paleozoic climate was strongly zonal, with the result that the "climate", in an abstract sense, became warmer, but the living space of most organisms of the time – the continental shelf marine environment – became steadily colder. However, Baltica (Northern Europe and Russia) and Laurentia (eastern North America and Greenland) remained in the tropical zone, while China and Australia lay in waters which were at least temperate. The early Paleozoic ended, rather abruptly, with

4026-550: The coal beds of Europe and eastern North America . Towards the end of the era, large, sophisticated synapsids and diapsids were dominant and the first modern plants ( conifers ) appeared. The Paleozoic Era ended with the largest extinction event of the Phanerozoic Eon , the Permian–Triassic extinction event . The effects of this catastrophe were so devastating that it took life on land 30 million years into

4148-406: 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

4270-609: 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

4392-593: The Cobb Valley in north-west Nelson. The Cobb Valley is also the location of " Trilobite Rock" a glacial dropstone made from the moulted exoskeletons of trilobites. Asbestos was mined in the Cobb Valley from the Tākaka Terrene between the late 1880s and 1917. The Tākaka Terrane is highly deformed and has been intruded by many batholiths . The Tākaka Terrane has two main igneous units, the arc-related Devil River Volcanics Group ( Middle to Late Cambrian ) and

4514-712: 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

4636-648: The Fish", the Devonian featured a huge diversification of fish, including armored fish like Dunkleosteus and lobe-finned fish which eventually evolved into the first tetrapods. On land, plant groups diversified rapidly in an event known as the Devonian explosion when plants made lignin , leading to taller growth and vascular tissue; the first trees and seeds evolved. These new habitats led to greater arthropod diversification. The first amphibians appeared and fish occupied

4758-804: 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

4880-484: 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

5002-658: The Mesozoic Era to recover. Recovery of life in the sea may have been much faster. The base of the Paleozoic is one of the major divisions in geological time representing the divide between the Proterozoic and Phanerozoic eons, the Paleozoic and Neoproterozoic eras and the Ediacaran and Cambrian periods. When Adam Sedgwick named the Paleozoic in 1835, he defined the base as the first appearance of complex life in

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5124-610: The Middle Carboniferous). An important evolutionary development of the time was the evolution of amniotic eggs , which allowed amphibians to move farther inland and remain the dominant vertebrates for the duration of this period. Also, the first reptiles and synapsids evolved in the swamps. Throughout the Carboniferous, there was a cooling trend, which led to the Permo-Carboniferous glaciation or

5246-514: 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

5368-420: 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

5490-666: 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

5612-517: 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

5734-629: The Palaeozoic had very few facultatively motile animals that could easily adjust to disturbance, with such creatures composing 1% of its assemblages in contrast to 50% in Cenozoic faunal assemblages. Non-motile animals untethered to the substrate, extremely rare in the Cenozoic, were abundant in the Palaeozoic. Palaeozoic phytoplankton overall were both nutrient-poor themselves and adapted to nutrient-poor environmental conditions. This phytoplankton nutrient poverty has been cited as an explanation for

5856-644: The Palaeozoic's relatively low biodiversity. Permian%E2%80%93Triassic extinction event 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 the boundary between

5978-459: The Paleozoic and Mesozoic eras and the Permian and Triassic periods is marked by the first occurrence of the conodont Hindeodus parvus . This is the first biostratigraphic event found worldwide that is associated with the beginning of the recovery following the end- Permian mass extinctions and environmental changes. In non-marine strata, the equivalent level is marked by the disappearance of

6100-530: The Permian Dicynodon tetrapods . This means events previously considered to mark the Permian-Triassic boundary, such as the eruption of the Siberian Traps flood basalts , the onset of greenhouse climate, ocean anoxia and acidification and the resulting mass extinction are now regarded as being of latest Permian in age. The GSSP is near Meishan , Zhejiang Province, southern China. Radiometric dating of volcanic clay layers just above and below

6222-511: 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

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6344-447: 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,

6466-462: 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

6588-526: 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

6710-864: 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

6832-520: 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

6954-529: The Silurian Period, about 420 million years ago, when they began to transition onto dry land. Terrestrial flora reached its climax in the Carboniferous, when towering lycopsid rainforests dominated the tropical belt of Euramerica . Climate change caused the Carboniferous Rainforest Collapse which fragmented this habitat, diminishing the diversity of plant life in the late Carboniferous and Permian periods. A noteworthy feature of Paleozoic life

7076-432: 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

7198-545: 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

7320-446: 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

7442-677: The Tākaka Terrane is equivalent to rocks in Tasmania , Australia and was separated from them with the opening of the Tasman Sea . The Arthur Marble has been chemically weathered by rain and groundwater due to its high calcium carbonate content forming a karst geomorphology . This has led to the formation of extensive cave systems like Harwoods Hole and the Riuwaka Resurgence . Paleozoic The name Paleozoic

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7564-476: 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

7686-549: The basal Cambrian Global Stratotype Section and Point (GSSP) at the base of the Treptichnus pedum assemblage of trace fossils and immediately above the last occurrence of the Ediacaran problematica fossils Harlaniella podolica and Palaeopsacichnus . The base of the Phanerozoic, Paleozoic and Cambrian is dated at 538.8+/-0.2 Ma and now lies below both the first appearance of trilobites and SSF. The boundary between

7808-516: The boundary confine its age to a narrow range of 251.902+/-0.024 Ma. The beginning of the Paleozoic Era witnessed the breakup of the supercontinent of Pannotia and ended while the supercontinent Pangaea was assembling. The breakup of Pannotia began with the opening of the Iapetus Ocean and other Cambrian seas and coincided with a dramatic rise in sea level. Paleoclimatic studies and evidence of glaciers indicate that Central Africa

7930-423: 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

8052-511: The climate of the interior of Pangaea was not regulated by large bodies of water. Diapsids and synapsids flourished in the new dry climate. Creatures such as Dimetrodon and Edaphosaurus ruled the new continent. The first conifers evolved, and dominated the terrestrial landscape. Near the end of the Permian, however, Pangaea grew drier. The interior was desert, and new taxa such as Scutosaurus and Gorgonopsids filled it. Eventually they disappeared, along with 95% of all life on Earth, in

8174-561: The continental margins, oxygen levels increased and carbon dioxide dropped, although much less dramatically. The north–south temperature gradient also seems to have moderated, or metazoan life simply became hardier, or both. At any event, the far southern continental margins of Antarctica and West Gondwana became increasingly less barren. The Devonian ended with a series of turnover pulses which killed off much of middle Paleozoic vertebrate life, without noticeably reducing species diversity overall. There are many unanswered questions about

8296-480: 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.

8418-635: 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

8540-465: 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

8662-423: 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

8784-507: The empty continent of Gondwana. By the end of the Ordovician, Gondwana was at the south pole, early North America had collided with Europe, closing the intervening ocean. Glaciation of Africa resulted in a major drop in sea level, killing off all life that had established along coastal Gondwana. Glaciation may have caused the Ordovician–Silurian extinction events , in which 60% of marine invertebrates and 25% of families became extinct, and

8906-492: 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

9028-560: The end of the Permian period. In late middle Permian the pareiasaurs originated, successful herbivores and the only sauropsids that could reach sizes comparable to some of the largest synapsids. The Palaeozoic marine fauna was notably lacking in predators relative to the present day. Predators made up about 4% of the fauna in Palaeozoic assemblages while making up 17% of temperate Cenozoic assemblages and 31% of tropical ones. Infaunal animals made up 4% of soft substrate Palaeozoic communities but about 47% of Cenozoic communities. Additionally,

9150-440: 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

9272-504: 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

9394-655: 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

9516-498: 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

9638-415: 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

9760-552: 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

9882-409: 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

10004-588: 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 ,

10126-409: 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

10248-461: The first tetrapods, 390  million years ago , and began to develop lungs. Amphibians were the dominant tetrapods until the mid-Carboniferous, when climate change greatly reduced their diversity, allowing amniotes to take over. Amniotes would split into two clades shortly after their origin in the Carboniferous; the synapsids, which was the dominant group, and the sauropsids . The synapsids continued to prosper and increase in number and variety till

10370-619: 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

10492-469: 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

10614-441: 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 ,

10736-421: 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

10858-628: 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

10980-423: 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

11102-548: The late Paleozoic. The Mississippian (early Carboniferous Period) began with a spike in atmospheric oxygen, while carbon dioxide plummeted to new lows. This destabilized the climate and led to one, and perhaps two, ice ages during the Carboniferous. These were far more severe than the brief Late Ordovician ice age; but, this time, the effects on world biota were inconsequential. By the Cisuralian Epoch, both oxygen and carbon dioxide had recovered to more normal levels. On

11224-504: The majority of Ediacaran to Cambrian rock sequences are composed of siliciclastic rocks where skeletal fossils are rarely preserved. This led the International Commission on Stratigraphy (ICS) to use trace fossils as an indicator of complex life. Unlike later in the fossil record, Cambrian trace fossils are preserved in a wide range of sediments and environments, which aids correlation between different sites around

11346-724: 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

11468-512: 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

11590-434: The most rapid and widespread diversification of life in Earth's history, known as the Cambrian explosion , in which most modern phyla first appeared. Arthropods , molluscs , fish , amphibians , reptiles , and synapsids all evolved during the Paleozoic. Life began in the ocean but eventually transitioned onto land, and by the late Paleozoic, great forests of primitive plants covered the continents, many of which formed

11712-603: The most ubiquitous of that period were the armored arthropods, like trilobites. Almost all marine phyla evolved in this period. During this time, the supercontinent Pannotia begins to break up, most of which later became the supercontinent Gondwana. The Ordovician spanned from 485–444 million years ago. The Ordovician was a time in Earth's history in which many of the biological classes still prevalent today evolved, such as primitive fish, cephalopods, and coral. The most common forms of life, however, were trilobites, snails and shellfish. The first arthropods went ashore to colonize

11834-411: 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

11956-520: The other hand, the assembly of Pangaea created huge arid inland areas subject to temperature extremes. The Lopingian Epoch is associated with falling sea levels, increased carbon dioxide and general climatic deterioration, culminating in the devastation of the Permian extinction. While macroscopic plant life appeared early in the Paleozoic Era and possibly late in the Neoproterozoic Era of the earlier eon, plants mostly remained aquatic until

12078-475: 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

12200-478: 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

12322-516: 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

12444-405: 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;

12566-456: 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

12688-424: The rift‐related Gendarme Dolerite (latest Cambrian to Early Ordovician age). The Devil River Volcanics Group sediments contain trilobites , brachiopods and conodonts . Sedimentary units in the Tākaka Terrane ( Haupiri Group and Junction Formation rocks) likely formed in a back-arc basin . The distinctive Arthur Marble from Tākaka Hill and Mount Arthur is of Ordovician age. It has been speculated that

12810-401: The rock record as shown by the presence of trilobite -dominated fauna. Since then evidence of complex life in older rock sequences has increased and by the second half of the 20th century, the first appearance of small shelly fauna (SSF), also known as early skeletal fossils, were considered markers for the base of the Paleozoic. However, whilst SSF are well preserved in carbonate sediments,

12932-412: 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

13054-479: The short, but apparently severe, late Ordovician ice age. This cold spell caused the second-greatest mass extinction of the Phanerozoic Eon. Over time, the warmer weather moved into the Paleozoic Era. The Ordovician and Silurian were warm greenhouse periods, with the highest sea levels of the Paleozoic (200 m above today's); the warm climate was interrupted only by a 30 million year cool period,

13176-417: 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 ,

13298-427: 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

13420-415: 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

13542-502: 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

13664-554: The top of the food chain. Earth's second Phanerozoic mass extinction event (a group of several smaller extinction events), the Late Devonian extinction , ended 70% of existing species. The Carboniferous is named after the large coal deposits laid down during the period. It spanned from 359–299 million years ago. During this time, average global temperatures were exceedingly high; the early Carboniferous averaged at about 20 degrees Celsius (but cooled to 10 °C during

13786-420: 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

13908-521: The world. Trace fossils reflect the complexity of the body plan of the organism that made them. Ediacaran trace fossils are simple, sub-horizontal feeding traces. As more complex organisms evolved, their more complex behaviour was reflected in greater diversity and complexity of the trace fossils they left behind. After two decades of deliberation, the ICS chose Fortune Head , Burin Peninsula, Newfoundland as

14030-588: 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

14152-469: Was first used by Adam Sedgwick (1785–1873) in 1838 to describe the Cambrian and Ordovician periods. It was redefined by John Phillips (1800–1874) in 1840 to cover the Cambrian to Permian periods. It is derived from the Greek palaiós (παλαιός, "old") and zōḗ (ζωή, "life") meaning "ancient life". The Paleozoic was a time of dramatic geological, climatic, and evolutionary change. The Cambrian witnessed

14274-539: Was most likely in the polar regions during the early Paleozoic. The breakup of Pannotia was followed by the assembly of the huge continent Gondwana ( 510  million years ago ). By the mid-Paleozoic, the collision of North America and Europe produced the Acadian-Caledonian uplifts, and a subducting plate uplifted eastern Australia . By the late Paleozoic, continental collisions formed the supercontinent of Pangaea and created great mountain chains, including

14396-410: 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

14518-425: 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

14640-519: 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

14762-482: 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

14884-435: 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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