Extinction event

An extinction event (also known as a mass extinction or biotic crisis) is a widespread and rapid decrease in the biodiversity on Earth. Such an event is identified by a sharp change in the diversity and abundance of multicellular organisms. It occurs when the rate of extinction increases with respect to the rate of speciation. Estimates of the number of major mass extinctions in the last 540 million years range from as few as five to more than twenty. These differences stem from the threshold chosen for describing an extinction event as "major", and the data chosen to measure past diversity.

Because most diversity and biomass on Earth is microbial, and thus difficult to measure, recorded extinction events affect the easily observed, biologically complex component of the biosphere rather than the total diversity and abundance of life.[1] Extinction occurs at an uneven rate. Based on the fossil record, the background rate of extinctions on Earth is about two to five taxonomic families of marine animals every million years. Marine fossils are mostly used to measure extinction rates because of their superior fossil record and stratigraphic range compared to land animals.

The Great Oxygenation Event, which occurred around 2.45 billion years ago, was probably the first major extinction event. Since the Cambrian explosion five further major mass extinctions have significantly exceeded the background extinction rate. The most recent and arguably best-known, the Cretaceous–Paleogene extinction event, which occurred approximately 66 million years ago (Ma), was a large-scale mass extinction of animal and plant species in a geologically short period of time.[2] In addition to the five major mass extinctions, there are numerous minor ones as well, and the ongoing mass extinction caused by human activity is sometimes called the sixth extinction.[3] Mass extinctions seem to be a mainly Phanerozoic phenomenon, with extinction rates low before large complex organisms arose.[4]

CambrianOrdovicianSilurianDevonianCarboniferousPermianTriassicJurassicCretaceousPaleogeneNeogene
Marine extinction intensity during the Phanerozoic
%
Millions of years ago
CambrianOrdovicianSilurianDevonianCarboniferousPermianTriassicJurassicCretaceousPaleogeneNeogene
The blue graph shows the apparent percentage (not the absolute number) of marine animal genera becoming extinct during any given time interval. It does not represent all marine species, just those that are readily fossilized. The labels of the traditional "Big Five" extinction events and the more recently recognised End-Capitanian extinction event are clickable hyperlinks. (source and image info)

Major extinction events

KT boundary 054
Badlands near Drumheller, Alberta, where erosion has exposed the K–Pg boundary
Kainops invius lateral and ventral
Trilobites were highly successful marine animals until the Permian–Triassic extinction event wiped them all out.

In a landmark paper published in 1982, Jack Sepkoski and David M. Raup identified five mass extinctions. They were originally identified as outliers to a general trend of decreasing extinction rates during the Phanerozoic,[5] but as more stringent statistical tests have been applied to the accumulating data, it has been established that multicellular animal life has experienced five major and many minor mass extinctions.[6] The "Big Five" cannot be so clearly defined, but rather appear to represent the largest (or some of the largest) of a relatively smooth continuum of extinction events.[5]

  1. Ordovician–Silurian extinction events (End Ordovician or O–S): 450–440 Ma (million years ago) at the OrdovicianSilurian transition. Two events occurred that killed off 27% of all families, 57% of all genera and 60% to 70% of all species.[7] Together they are ranked by many scientists as the second largest of the five major extinctions in Earth's history in terms of percentage of genera that became extinct.
  2. Late Devonian extinction: 375–360 Ma near the DevonianCarboniferous transition. At the end of the Frasnian Age in the later part(s) of the Devonian Period, a prolonged series of extinctions eliminated about 19% of all families, 50% of all genera[7] and at least 70% of all species.[8] This extinction event lasted perhaps as long as 20 million years, and there is evidence for a series of extinction pulses within this period.
  3. Permian–Triassic extinction event (End Permian): 252 Ma at the PermianTriassic transition.[9] Earth's largest extinction killed 57% of all families, 83% of all genera and 90% to 96% of all species[7] (53% of marine families, 84% of marine genera, about 96% of all marine species and an estimated 70% of land species,[2] including insects).[10] The highly successful marine arthropod, the trilobite, became extinct. The evidence regarding plants is less clear, but new taxa became dominant after the extinction.[11] The "Great Dying" had enormous evolutionary significance: on land, it ended the primacy of mammal-like reptiles. The recovery of vertebrates took 30 million years,[12] but the vacant niches created the opportunity for archosaurs to become ascendant. In the seas, the percentage of animals that were sessile dropped from 67% to 50%. The whole late Permian was a difficult time for at least marine life, even before the "Great Dying".
  4. Triassic–Jurassic extinction event (End Triassic): 201.3 Ma at the TriassicJurassic transition. About 23% of all families, 48% of all genera (20% of marine families and 55% of marine genera) and 70% to 75% of all species became extinct.[7] Most non-dinosaurian archosaurs, most therapsids, and most of the large amphibians were eliminated, leaving dinosaurs with little terrestrial competition. Non-dinosaurian archosaurs continued to dominate aquatic environments, while non-archosaurian diapsids continued to dominate marine environments. The Temnospondyl lineage of large amphibians also survived until the Cretaceous in Australia (e.g., Koolasuchus).
  5. Cretaceous–Paleogene extinction event (End Cretaceous, K–Pg extinction, or formerly K–T extinction): 66 Ma at the Cretaceous (Maastrichtian) – Paleogene (Danian) transition interval.[13] The event formerly called the Cretaceous-Tertiary or K–T extinction or K–T boundary is now officially named the Cretaceous–Paleogene (or K–Pg) extinction event. About 17% of all families, 50% of all genera[7] and 75% of all species became extinct.[14] In the seas all the ammonites, plesiosaurs and mosasaurs disappeared and the percentage of sessile animals (those unable to move about) was reduced to about 33%. All non-avian dinosaurs became extinct during that time.[15] The boundary event was severe with a significant amount of variability in the rate of extinction between and among different clades. Mammals and birds, the latter descended from theropod dinosaurs, emerged as dominant large land animals.

Despite the popularization of these five events, there is no definite line separating them from other extinction events; using different methods of calculating an extinction's impact can lead to other events featuring in the top five.[16]

Older fossil records are more difficult to interpret. This is because:

  • Older fossils are harder to find as they are usually buried at a considerable depth.
  • Dating older fossils is more difficult.
  • Productive fossil beds are researched more than unproductive ones, therefore leaving certain periods unresearched.
  • Prehistoric environmental events can disturb the deposition process.
  • The preservation of fossils varies on land, but marine fossils tend to be better preserved than their sought after land-based counterparts.[17]

It has been suggested that the apparent variations in marine biodiversity may actually be an artifact, with abundance estimates directly related to quantity of rock available for sampling from different time periods.[18] However, statistical analysis shows that this can only account for 50% of the observed pattern, and other evidence (such as fungal spikes) provides reassurance that most widely accepted extinction events are real. A quantification of the rock exposure of Western Europe indicates that many of the minor events for which a biological explanation has been sought are most readily explained by sampling bias.[19]

Research completed after the seminal 1982 paper has concluded that a sixth mass extinction event is ongoing:

6. Holocene extinction: Currently ongoing. Extinctions have occurred at over 1000 times the background extinction rate since 1900.[20][21] The mass extinction is a result of human activity.[22][23][24]

More recent research has indicated that the End-Capitanian extinction event likely constitutes a separate extinction event from the Permian–Triassic extinction event; if so, it would be larger than many of the "Big Five" extinction events.

Evolutionary importance

Mass extinctions have sometimes accelerated the evolution of life on Earth. When dominance of particular ecological niches passes from one group of organisms to another, it is rarely because the new dominant group is "superior" to the old and usually because an extinction event eliminates the old dominant group and makes way for the new one.[25][26]

For example, mammaliformes ("almost mammals") and then mammals existed throughout the reign of the dinosaurs, but could not compete for the large terrestrial vertebrate niches which dinosaurs monopolized. The end-Cretaceous mass extinction removed the non-avian dinosaurs and made it possible for mammals to expand into the large terrestrial vertebrate niches. Ironically, the dinosaurs themselves had been beneficiaries of a previous mass extinction, the end-Triassic, which eliminated most of their chief rivals, the crurotarsans.

Another point of view put forward in the Escalation hypothesis predicts that species in ecological niches with more organism-to-organism conflict will be less likely to survive extinctions. This is because the very traits that keep a species numerous and viable under fairly static conditions become a burden once population levels fall among competing organisms during the dynamics of an extinction event.

Furthermore, many groups which survive mass extinctions do not recover in numbers or diversity, and many of these go into long-term decline, and these are often referred to as "Dead Clades Walking".[27] However, clades that survive for a considerable period of time after a mass extinction, and which were reduced to only a few species, are likely to have experienced a rebound effect called the "push of the past".[28]

Darwin was firmly of the opinion that biotic interactions, such as competition for food and space—the ‘struggle for existence’—were of considerably greater importance in promoting evolution and extinction than changes in the physical environment. He expressed this in The Origin of Species: "Species are produced and exterminated by slowly acting causes…and the most import of all causes of organic change is one which is almost independent of altered…physical conditions, namely the mutual relation of organism to organism-the improvement of one organism entailing the improvement or extermination of others".[29]

Patterns in frequency

It has been suggested variously that extinction events occurred periodically, every 26 to 30 million years,[30][31] or that diversity fluctuates episodically every ~62 million years.[32] Various ideas attempt to explain the supposed pattern, including the presence of a hypothetical companion star to the sun,[33][34] oscillations in the galactic plane, or passage through the Milky Way's spiral arms.[35] However, other authors have concluded that the data on marine mass extinctions do not fit with the idea that mass extinctions are periodic, or that ecosystems gradually build up to a point at which a mass extinction is inevitable.[5] Many of the proposed correlations have been argued to be spurious.[36][37] Others have argued that there is strong evidence supporting periodicity in a variety of records,[38] and additional evidence in the form of coincident periodic variation in nonbiological geochemical variables.[39]

Phanerozoic biodiversity as shown by the fossil record
All genera
"Well-defined" genera
Trend line
"Big Five" mass extinctions
Other mass extinctions
Million years ago
Thousands of genera
Phanerozoic biodiversity blank 01
Phanerozoic biodiversity as shown by the fossil record
Phanerozoic biodiversity blank 01

Mass extinctions are thought to result when a long-term stress is compounded by a short term shock.[40] Over the course of the Phanerozoic, individual taxa appear to be less likely to become extinct at any time,[41] which may reflect more robust food webs as well as less extinction-prone species and other factors such as continental distribution.[41] However, even after accounting for sampling bias, there does appear to be a gradual decrease in extinction and origination rates during the Phanerozoic.[5] This may represent the fact that groups with higher turnover rates are more likely to become extinct by chance; or it may be an artefact of taxonomy: families tend to become more speciose, therefore less prone to extinction, over time;[5] and larger taxonomic groups (by definition) appear earlier in geological time.[42]

It has also been suggested that the oceans have gradually become more hospitable to life over the last 500 million years, and thus less vulnerable to mass extinctions,[note 1][43][44] but susceptibility to extinction at a taxonomic level does not appear to make mass extinctions more or less probable.[41]

Causes

There is still debate about the causes of all mass extinctions. In general, large extinctions may result when a biosphere under long-term stress undergoes a short-term shock.[40] An underlying mechanism appears to be present in the correlation of extinction and origination rates to diversity. High diversity leads to a persistent increase in extinction rate; low diversity to a persistent increase in origination rate. These presumably ecologically controlled relationships likely amplify smaller perturbations (asteroid impacts, etc.) to produce the global effects observed.[5]

Identifying causes of specific mass extinctions

A good theory for a particular mass extinction should: (i) explain all of the losses, not just focus on a few groups (such as dinosaurs); (ii) explain why particular groups of organisms died out and why others survived; (iii) provide mechanisms which are strong enough to cause a mass extinction but not a total extinction; (iv) be based on events or processes that can be shown to have happened, not just inferred from the extinction.

It may be necessary to consider combinations of causes. For example, the marine aspect of the end-Cretaceous extinction appears to have been caused by several processes which partially overlapped in time and may have had different levels of significance in different parts of the world.[45]

Arens and West (2006) proposed a "press / pulse" model in which mass extinctions generally require two types of cause: long-term pressure on the eco-system ("press") and a sudden catastrophe ("pulse") towards the end of the period of pressure.[46] Their statistical analysis of marine extinction rates throughout the Phanerozoic suggested that neither long-term pressure alone nor a catastrophe alone was sufficient to cause a significant increase in the extinction rate.

Most widely supported explanations

Macleod (2001)[47] summarized the relationship between mass extinctions and events which are most often cited as causes of mass extinctions, using data from Courtillot et al. (1996),[48] Hallam (1992)[49] and Grieve et al. (1996):[50]

  • Flood basalt events: 11 occurrences, all associated with significant extinctions[51][52] But Wignall (2001) concluded that only five of the major extinctions coincided with flood basalt eruptions and that the main phase of extinctions started before the eruptions.[53]
  • Sea-level falls: 12, of which seven were associated with significant extinctions.[52]
  • Asteroid impacts: one large impact is associated with a mass extinction, i.e. the Cretaceous–Paleogene extinction event; there have been many smaller impacts but they are not associated with significant extinctions.[54]

The most commonly suggested causes of mass extinctions are listed below.

Flood basalt events

The formation of large igneous provinces by flood basalt events could have:

  • produced dust and particulate aerosols which inhibited photosynthesis and thus caused food chains to collapse both on land and at sea[55]
  • emitted sulfur oxides which were precipitated as acid rain and poisoned many organisms, contributing further to the collapse of food chains
  • emitted carbon dioxide and thus possibly causing sustained global warming once the dust and particulate aerosols dissipated.

Flood basalt events occur as pulses of activity punctuated by dormant periods. As a result, they are likely to cause the climate to oscillate between cooling and warming, but with an overall trend towards warming as the carbon dioxide they emit can stay in the atmosphere for hundreds of years.

It is speculated that massive volcanism caused or contributed to the End-Permian, End-Triassic and End-Cretaceous extinctions.[56] The correlation between gigantic volcanic events expressed in the large igneous provinces and mass extinctions was shown for the last 260 Myr.[57][58] Recently such possible correlation was extended for the whole Phanerozoic Eon.[59]

Sea-level falls

These are often clearly marked by worldwide sequences of contemporaneous sediments which show all or part of a transition from sea-bed to tidal zone to beach to dry land – and where there is no evidence that the rocks in the relevant areas were raised by geological processes such as orogeny. Sea-level falls could reduce the continental shelf area (the most productive part of the oceans) sufficiently to cause a marine mass extinction, and could disrupt weather patterns enough to cause extinctions on land. But sea-level falls are very probably the result of other events, such as sustained global cooling or the sinking of the mid-ocean ridges.

Sea-level falls are associated with most of the mass extinctions, including all of the "Big Five"—End-Ordovician, Late Devonian, End-Permian, End-Triassic, and End-Cretaceous.

A study, published in the journal Nature (online June 15, 2008) established a relationship between the speed of mass extinction events and changes in sea level and sediment.[60] The study suggests changes in ocean environments related to sea level exert a driving influence on rates of extinction, and generally determine the composition of life in the oceans.[61]

Impact events

The impact of a sufficiently large asteroid or comet could have caused food chains to collapse both on land and at sea by producing dust and particulate aerosols and thus inhibiting photosynthesis.[62] Impacts on sulfur-rich rocks could have emitted sulfur oxides precipitating as poisonous acid rain, contributing further to the collapse of food chains. Such impacts could also have caused megatsunamis and/or global forest fires.

Most paleontologists now agree that an asteroid did hit the Earth about 66 Ma ago, but there is an ongoing dispute whether the impact was the sole cause of the Cretaceous–Paleogene extinction event.[63][64]

Global cooling

Sustained and significant global cooling could kill many polar and temperate species and force others to migrate towards the equator; reduce the area available for tropical species; often make the Earth's climate more arid on average, mainly by locking up more of the planet's water in ice and snow. The glaciation cycles of the current ice age are believed to have had only a very mild impact on biodiversity, so the mere existence of a significant cooling is not sufficient on its own to explain a mass extinction.

It has been suggested that global cooling caused or contributed to the End-Ordovician, Permian–Triassic, Late Devonian extinctions, and possibly others. Sustained global cooling is distinguished from the temporary climatic effects of flood basalt events or impacts.

Global warming

This would have the opposite effects: expand the area available for tropical species; kill temperate species or force them to migrate towards the poles; possibly cause severe extinctions of polar species; often make the Earth's climate wetter on average, mainly by melting ice and snow and thus increasing the volume of the water cycle. It might also cause anoxic events in the oceans (see below).

Global warming as a cause of mass extinction is supported by several recent studies.[65]

The most dramatic example of sustained warming is the Paleocene–Eocene Thermal Maximum, which was associated with one of the smaller mass extinctions. It has also been suggested to have caused the Triassic–Jurassic extinction event, during which 20% of all marine families became extinct. Furthermore, the Permian–Triassic extinction event has been suggested to have been caused by warming.[66][67][68]

Clathrate gun hypothesis

Clathrates are composites in which a lattice of one substance forms a cage around another. Methane clathrates (in which water molecules are the cage) form on continental shelves. These clathrates are likely to break up rapidly and release the methane if the temperature rises quickly or the pressure on them drops quickly—for example in response to sudden global warming or a sudden drop in sea level or even earthquakes. Methane is a much more powerful greenhouse gas than carbon dioxide, so a methane eruption ("clathrate gun") could cause rapid global warming or make it much more severe if the eruption was itself caused by global warming.

The most likely signature of such a methane eruption would be a sudden decrease in the ratio of carbon-13 to carbon-12 in sediments, since methane clathrates are low in carbon-13; but the change would have to be very large, as other events can also reduce the percentage of carbon-13.[69]

It has been suggested that "clathrate gun" methane eruptions were involved in the end-Permian extinction ("the Great Dying") and in the Paleocene–Eocene Thermal Maximum, which was associated with one of the smaller mass extinctions.

Anoxic events

Anoxic events are situations in which the middle and even the upper layers of the ocean become deficient or totally lacking in oxygen. Their causes are complex and controversial, but all known instances are associated with severe and sustained global warming, mostly caused by sustained massive volcanism.[70]

It has been suggested that anoxic events caused or contributed to the Ordovician–Silurian, late Devonian, Permian–Triassic and Triassic–Jurassic extinctions, as well as a number of lesser extinctions (such as the Ireviken, Mulde, Lau, Toarcian and Cenomanian–Turonian events). On the other hand, there are widespread black shale beds from the mid-Cretaceous which indicate anoxic events but are not associated with mass extinctions.

The bio-availability of essential trace elements (in particular selenium) to potentially lethal lows has been shown to coincide with, and likely have contributed to, at least three mass extinction events in the oceans, i.e. at the end of the Ordovician, during the Middle and Late Devonian, and at the end of the Triassic. During periods of low oxygen concentrations very soluble selenate (Se6+) is converted into much less soluble selenide (Se2-), elemental Se and organo-selenium complexes. Bio-availability of selenium during these extinction events dropped to about 1% of the current oceanic concentration, a level that has been proven lethal to many extant organisms.[71]

Hydrogen sulfide emissions from the seas

Kump, Pavlov and Arthur (2005) have proposed that during the Permian–Triassic extinction event the warming also upset the oceanic balance between photosynthesising plankton and deep-water sulfate-reducing bacteria, causing massive emissions of hydrogen sulfide which poisoned life on both land and sea and severely weakened the ozone layer, exposing much of the life that still remained to fatal levels of UV radiation.[72][73][74]

Oceanic overturn

Oceanic overturn is a disruption of thermo-haline circulation which lets surface water (which is more saline than deep water because of evaporation) sink straight down, bringing anoxic deep water to the surface and therefore killing most of the oxygen-breathing organisms which inhabit the surface and middle depths. It may occur either at the beginning or the end of a glaciation, although an overturn at the start of a glaciation is more dangerous because the preceding warm period will have created a larger volume of anoxic water.[75]

Unlike other oceanic catastrophes such as regressions (sea-level falls) and anoxic events, overturns do not leave easily identified "signatures" in rocks and are theoretical consequences of researchers' conclusions about other climatic and marine events.

It has been suggested that oceanic overturn caused or contributed to the late Devonian and Permian–Triassic extinctions.

A nearby nova, supernova or gamma ray burst

A nearby gamma-ray burst (less than 6000 light-years away) would be powerful enough to destroy the Earth's ozone layer, leaving organisms vulnerable to ultraviolet radiation from the Sun.[76] Gamma ray bursts are fairly rare, occurring only a few times in a given galaxy per million years.[77] It has been suggested that a supernova or gamma ray burst caused the End-Ordovician extinction.[78]

Geomagnetic reversal

One theory is that periods of increased geomagnetic reversals will weaken Earth's magnetic field long enough to expose the atmosphere to the solar winds, causing oxygen ions to escape the atmosphere in a rate increased by 3–4 orders, resulting in a disastrous decrease in oxygen.[79]

Plate tectonics

Movement of the continents into some configurations can cause or contribute to extinctions in several ways: by initiating or ending ice ages; by changing ocean and wind currents and thus altering climate; by opening seaways or land bridges which expose previously isolated species to competition for which they are poorly adapted (for example, the extinction of most of South America's native ungulates and all of its large metatherians after the creation of a land bridge between North and South America). Occasionally continental drift creates a super-continent which includes the vast majority of Earth's land area, which in addition to the effects listed above is likely to reduce the total area of continental shelf (the most species-rich part of the ocean) and produce a vast, arid continental interior which may have extreme seasonal variations.

Another theory is that the creation of the super-continent Pangaea contributed to the End-Permian mass extinction. Pangaea was almost fully formed at the transition from mid-Permian to late-Permian, and the "Marine genus diversity" diagram at the top of this article shows a level of extinction starting at that time which might have qualified for inclusion in the "Big Five" if it were not overshadowed by the "Great Dying" at the end of the Permian.[80]

Other hypotheses

Many other hypotheses have been proposed, such as the spread of a new disease, or simple out-competition following an especially successful biological innovation. But all have been rejected, usually for one of the following reasons: they require events or processes for which there is no evidence; they assume mechanisms which are contrary to the available evidence; they are based on other theories which have been rejected or superseded.

Scientists have been concerned that human activities could cause more plants and animals to become extinct than any point in the past. Along with human-made changes in climate (see above), some of these extinctions could be caused by overhunting, overfishing, invasive species, or habitat loss. A study published in May 2017 in Proceedings of the National Academy of Sciences argued that a “biological annihilation” akin to a sixth mass extinction event is underway as a result of anthropogenic causes, such as over-population and over-consumption. The study suggested that as much as 50% of the number of animal individuals that once lived on Earth were already extinct, threatening the basis for human existence too.[81][24]

Future biosphere extinction/sterilization

The eventual warming and expanding of the Sun, combined with the eventual decline of atmospheric carbon dioxide could actually cause an even greater mass extinction, having the potential to wipe out even microbes (in other words, the Earth is completely sterilized), where rising global temperatures caused by the expanding Sun will gradually increase the rate of weathering, which in turn removes more and more carbon dioxide from the atmosphere. When carbon dioxide levels get too low (perhaps at 50 ppm), all plant life will die out, although simpler plants like grasses and mosses can survive much longer, until CO2 levels drop to 10 ppm.[82][83]

With all photosynthetic organisms gone, atmospheric oxygen can no longer be replenished, and is eventually removed by chemical reactions in the atmosphere, perhaps from volcanic eruptions. Eventually the loss of oxygen will cause all remaining aerobic life to die out via asphyxiation, leaving behind only simple anaerobic prokaryotes. When the Sun becomes 10% brighter in about a billion years,[82] Earth will suffer a moist greenhouse effect resulting in its oceans boiling away, while the Earth's liquid outer core cools due to the inner core's expansion and causes the Earth's magnetic field to shut down. In the absence of a magnetic field, charged particles from the Sun will deplete the atmosphere and further increase the Earth's temperature to an average of ~420 K (147 °C, 296 °F) in 2.8 billion years, causing the last remaining life on Earth to die out. This is the most extreme instance of a climate-caused extinction event. Since this will only happen late in the Sun's life, such will cause the final mass extinction in Earth's history (albeit a very long extinction event).[82][83]

Effects and recovery

The impact of mass extinction events varied widely. After a major extinction event, usually only weedy species survive due to their ability to live in diverse habitats.[84] Later, species diversify and occupy empty niches. Generally, biodiversity recovers 5 to 10 million years after the extinction event. In the most severe mass extinctions it may take 15 to 30 million years.[84]

The worst event, the Permian–Triassic extinction, devastated life on earth, killing over 90% of species. Life seemed to recover quickly after the P-T extinction, but this was mostly in the form of disaster taxa, such as the hardy Lystrosaurus. The most recent research indicates that the specialized animals that formed complex ecosystems, with high biodiversity, complex food webs and a variety of niches, took much longer to recover. It is thought that this long recovery was due to successive waves of extinction which inhibited recovery, as well as prolonged environmental stress which continued into the Early Triassic. Recent research indicates that recovery did not begin until the start of the mid-Triassic, 4M to 6M years after the extinction;[85] and some writers estimate that the recovery was not complete until 30M years after the P-T extinction, i.e. in the late Triassic.[86] Subsequent to the P-T extinction, there was an increase in provincialization, with species occupying smaller ranges – perhaps removing incumbents from niches and setting the stage for an eventual rediversification.[87]

The effects of mass extinctions on plants are somewhat harder to quantify, given the biases inherent in the plant fossil record. Some mass extinctions (such as the end-Permian) were equally catastrophic for plants, whereas others, such as the end-Devonian, did not affect the flora.[88]

See also

Notes

  1. ^ Dissolved oxygen became more widespread and penetrated to greater depths; the development of life on land reduced the run-off of nutrients and hence the risk of eutrophication and anoxic events; and marine ecosystems became more diversified so that food chains were less likely to be disrupted.

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External links

Araguainha crater

The Araguainha crater or Araguainha dome is an impact crater on the border of Mato Grosso and Goiás states, Brazil, between the villages of Araguainha and Ponte Branca. With a diameter of 40 kilometres (25 mi), it is the second largest known impact crater in South America, and possibly the oldest one.

The crater has most recently been dated to 254.7 ± 2.5 million years ago, when the region was probably a shallow sea. The margins of error of this date overlap the time of the Permian–Triassic extinction event, one of the largest mass extinction events in Earth's history. The impact punched through Paleozoic sedimentary units belonging to the Paraná Basin formations, and exposed the underlying Ordovician granite basement rocks. It is estimated that the crater was initially 24 kilometres (15 mi) wide and 2.4 kilometres (1.5 mi) deep, which then widened to 40 kilometres (25 mi) as its walls subsided inwards.

Averostra

Averostra, or "bird snouts", is a clade that includes most theropod dinosaurs that have a promaxillary fenestra (fenestra promaxillaris), an extra opening in the front outer side of the maxilla, the bone that makes up the upper jaw. Two groups of averostrans, the Ceratosauria and the Orionides, survived into the Cretaceous period. When the Cretaceous–Paleogene extinction event occurred, ceratosaurians and two groups of orionideans within the clade Coelurosauria, the Tyrannosauroidea and Maniraptoriformes, were still extant. Only one subgroup of maniraptoriformes, Aves, survived the extinction event and persisted to the present day.

Cambrian–Ordovician extinction event

The Cambrian–Ordovician extinction event occurred approximately 488 million years ago (m.y.a.). This early Phanerozoic Eon extinction event eliminated many brachiopods and conodonts, and severely reduced the number of trilobite species.

It was preceded by the less-documented (but probably worse) End-Botomian extinction event around 517 million years ago and the Dresbachian extinction event about 502 million years ago.

The Cambrian–Ordovician event ended the Cambrian Period, and led into the Ordovician Period in the Paleozoic Era.

Cretaceous–Paleogene extinction event

The Cretaceous–Paleogene (K–Pg) extinction event, also known as the Cretaceous–Tertiary (K–T) extinction, was a sudden mass extinction of some three-quarters of the plant and animal species on Earth, approximately 66 million years ago. With the exception of some ectothermic species such as the leatherback sea turtle and crocodiles, no tetrapods weighing more than 25 kilograms (55 lb) survived. It marked the end of the Cretaceous period and with it, the entire Mesozoic Era, opening the Cenozoic Era that continues today.

In the geologic record, the K–Pg event is marked by a thin layer of sediment called the K–Pg boundary, which can be found throughout the world in marine and terrestrial rocks. The boundary clay shows high levels of the metal iridium, which is rare in the Earth's crust, but abundant in asteroids.As originally proposed in 1980 by a team of scientists led by Luis Alvarez and his son Walter Alvarez, it is now generally thought that the K–Pg extinction was caused by the impact of a massive comet or asteroid 10 to 15 km (6 to 9 mi) wide, 66 million years ago, which devastated the global environment, mainly through a lingering impact winter which halted photosynthesis in plants and plankton. The impact hypothesis, also known as the Alvarez hypothesis, was bolstered by the discovery of the 180-kilometer-wide (112 mi) Chicxulub crater in the Gulf of Mexico in the early 1990s, which provided conclusive evidence that the K–Pg boundary clay represented debris from an asteroid impact. The fact that the extinctions occurred simultaneously provides strong evidence that they were caused by the asteroid. A 2016 drilling project into the Chicxulub peak ring, confirmed that the peak ring comprised granite ejected within minutes from deep in the earth, but contained hardly any gypsum, the usual sulfate-containing sea floor rock in the region: the gypsum would have vaporized and dispersed as an aerosol into the atmosphere, causing longer-term effects on the climate and food chain.

Other causal or contributing factors to the extinction may have been the Deccan Traps and other volcanic eruptions, climate change, and sea level change.

A wide range of species perished in the K–Pg extinction, the best-known being the non-avian dinosaurs. It also destroyed a plethora of other terrestrial organisms, including certain mammals, pterosaurs, birds, lizards, insects, and plants. In the oceans, the K–Pg extinction killed off plesiosaurs and the giant marine lizards (Mosasauridae) and devastated fish, sharks, mollusks (especially ammonites, which became extinct), and many species of plankton. It is estimated that 75% or more of all species on Earth vanished. Yet the extinction also provided evolutionary opportunities: in its wake, many groups underwent remarkable adaptive radiation—sudden and prolific divergence into new forms and species within the disrupted and emptied ecological niches. Mammals in particular diversified in the Paleogene, evolving new forms such as horses, whales, bats, and primates. Birds, fish, and perhaps lizards also radiated.

Dresbachian

The Dresbachian is a Maentwrogian regional stage of North America, lasting from 501 to 497 million years ago. It is part of the Upper Cambrian and is defined by four trilobite zones. It overlaps with the ICS-stages Guzhangian, Paibian and the lowest Jiangshanian.The Dresbachian overlies the Middle Cambrian Albertan series, and is the lowest stage of the Upper Cambrian Croixian series, followed by the Franconian stage. The Dresbachian extinction event, about 502 million years ago, was followed by the Cambrian–Ordovician extinction event about 485.4 million years ago.

Emeishan Traps

The Emeishan Traps constitute a flood basalt volcanic province, or large igneous province, in south-western China, centred in Sichuan province. It is sometimes referred to as the Permian Emeishan Large Igneous Province or Emeishan Flood Basalts. Like other volcanic provinces or "traps," the Emeishan Traps are multiple layers of igneous rock laid down by large mantle plume volcanic eruptions. The Emeishan Traps eruptions were serious enough to have global ecological and paleontological impact.It is named for Emeishan, a mountain in Sichuan.

Eocene–Oligocene extinction event

The transition between the end of the Eocene (33.9 Ma) and the beginning of the Oligocene is marked by large-scale extinction and floral and faunal turnover (although minor in comparison to the largest mass extinctions). Most of the affected organisms were marine or aquatic in nature. They included the last of the ancient cetaceans, the Archaeoceti.

This was a time of major climatic change, especially cooling, not obviously linked with any single major impact or any catastrophic volcanic event. One cause of the extinction event is speculated to be extended volcanic activity. Another speculation is that the extinctions are related to several large meteorite impacts that occurred about this time. One such event caused the Chesapeake Bay impact crater 40 km (25 mi), and another at the Popigai crater 100 km (62 mi) of central Siberia, scattering debris perhaps as far as Europe. New dating of the Popigai meteor suggests it may be a cause of the mass extinction.A leading scientific theory on climate cooling at this time predicts a decrease in atmospheric carbon dioxide, which slowly declined in the mid to late Eocene and possibly reached some threshold approximately 34 million years ago. This boundary is closely linked with the Oligocene Oi-1 event, an oxygen isotope excursion that marks the beginning of ice sheet coverage on Antarctica.

Famennian

The Famennian is the latter of two faunal stages in the Late Devonian epoch. It lasted from 372.2 million years ago to 358.9 million years ago. It was preceded by the Frasnian stage and followed by the Tournaisian stage.

It was during this age that tetrapods first appeared. In the seas, a novel major group of ammonoid cephalopods called clymeniids appeared, underwent tremendous diversification and spread worldwide, then just as suddenly went extinct.

The beginning of the Famennian is marked by a major extinction event, the Kellwasser Event, and the end with a smaller but still quite severe extinction event, the Hangenberg Event.

North American subdivisions of the Famennian include the Chautauquan, Canadaway, Conneaut, Conneautan, Conewango and Conewangan.

Haplogroup

A haplotype is a group of alleles in an organism that are inherited together from a single parent, and a haplogroup (haploid from the Greek: ἁπλούς, haploûs, "onefold, simple" and English: group) is a group of similar haplotypes that share a common ancestor with a single-nucleotide polymorphism mutation. More specifically, a haplogroup is a combination of alleles at different chromosomes regions that are closely linked and that tend to be inherited together. As a haplogroup consists of similar haplotypes, it is usually possible to predict a haplogroup from haplotypes. Haplogroups pertain to a single line of descent. As such, membership of a haplogroup, by any individual, relies on a relatively small proportion of the genetic material possessed by that individual.

Each haplogroup originates from, and remains part of, a preceding single haplogroup (or paragroup). As such, any related group of haplogroups may be precisely modelled as a nested hierarchy, in which each set (haplogroup) is also a subset of a single broader set (as opposed, that is, to biparental models, such as human family trees).

Haplogroups are normally identified by an initial letter of the alphabet, and refinements consist of additional number and letter combinations, such as (for example) A → A1 → A1a.

In human genetics, the haplogroups most commonly studied are Y-chromosome (Y-DNA) haplogroups and mitochondrial DNA (mtDNA) haplogroups, each of which can be used to define genetic populations. Y-DNA is passed solely along the patrilineal line, from father to son, while mtDNA is passed down the matrilineal line, from mother to offspring of both sexes. Neither recombines, and thus Y-DNA and mtDNA change only by chance mutation at each generation with no intermixture between parents' genetic material.

Holocene extinction

The Holocene extinction, otherwise referred to as the Sixth extinction or Anthropocene extinction, is a current event, and is one of the most significant extinction events in the history of the Earth. This ongoing extinction of species coincides with the present Holocene epoch (approx. 11,700 years), and is mainly a result of human activity. This large number of extinctions spans numerous families of plants and animals, including mammals, birds, amphibians, reptiles and arthropods. With widespread degradation of highly biodiverse habitats such as coral reefs and rainforests, as well as other areas, the vast majority of these extinctions are thought to be undocumented, as no one is even aware of the existence of the species before they go extinct, or no one has yet discovered their extinction. The current rate of extinction of species is estimated at 100 to 1,000 times higher than natural background rates.The Holocene extinction includes the disappearance of large land animals known as megafauna, starting at the end of the last Ice Age. Megafauna outside of the African continent, which did not evolve alongside humans, proved highly sensitive to the introduction of new predation, and many died out shortly after early humans began spreading and hunting across the Earth (additionally, many African species have also gone extinct in the Holocene). These extinctions, occurring near the Pleistocene–Holocene boundary, are sometimes referred to as the Quaternary extinction event.

The most popular theory is that human overhunting of species added to existing stress conditions as the extinction coincides with human emergence. Although there is debate regarding how much human predation affected their decline, certain population declines have been directly correlated with human activity, such as the extinction events of New Zealand and Hawaii. Aside from humans, climate change may have been a driving factor in the megafaunal extinctions, especially at the end of the Pleistocene.

Ecologically, humanity has been noted as an unprecedented "global superpredator" that consistently preys on the adults of other apex predators, and has worldwide effects on food webs. There have been extinctions of species on every land mass and in every ocean: there are many famous examples within Africa, Asia, Europe, Australia, North and South America, and on smaller islands. Overall, the Holocene extinction can be linked to the human impact on the environment. The Holocene extinction continues into the 21st century, with meat consumption, overfishing, ocean acidification and the decline in amphibian populations being a few broader examples of an almost universal, cosmopolitan decline in biodiversity. Human overpopulation (and continued population growth) along with profligate consumption are considered to be the primary drivers of this rapid decline.

Late Devonian extinction

The Late Devonian extinction was one of five major extinction events in the history of life on Earth. A major extinction, the Kellwasser event, occurred at the boundary that marks the beginning of the last phase of the Devonian period, the Famennian faunal stage (the Frasnian–Famennian boundary), about 376–360 million years ago. Overall, 19% of all families and 50% of all genera became extinct. A second, distinct mass extinction, the Hangenberg event, closed the Devonian period.Although it is clear that there was a massive loss of biodiversity in the Late Devonian, the timespan of this event is uncertain, with estimates ranging from 500,000 to 25 million years, extending from the mid-Givetian to the end-Famennian. Nor is it clear whether there were two sharp mass extinctions or a series of smaller extinctions, though the latest research suggests multiple causes and a series of distinct extinction pulses during an interval of some three million years. Some consider the extinction to be as many as seven distinct events, spread over about 25 million years, with notable extinctions at the ends of the Givetian, Frasnian, and Famennian stages.By the Late Devonian, the land had been colonized by plants and insects. In the oceans were massive reefs built by corals and stromatoporoids. Euramerica and Gondwana were beginning to converge into what would become Pangaea. The extinction seems to have only affected marine life. Hard-hit groups include brachiopods, trilobites, and reef-building organisms; the reef-building organisms almost completely disappeared. The causes of these extinctions are unclear. Leading hypotheses include changes in sea level and ocean anoxia, possibly triggered by global cooling or oceanic volcanism. The impact of a comet or another extraterrestrial body has also been suggested, such as the Siljan Ring event in Sweden. Some statistical analysis suggests that the decrease in diversity was caused more by a decrease in speciation than by an increase in extinctions. This might have been caused by invasions of cosmopolitan species, rather than by any single event. Surprisingly, jawed vertebrates seem to have been unaffected by the loss of reefs or other aspects of the Kellwasser event, while agnathans were in decline long before the end of the Frasnian.

Maastrichtian

The Maastrichtian ( ) is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous epoch or Upper Cretaceous series, the Cretaceous period or system, and of the Mesozoic era or erathem. It spanned the interval from 72.1 to 66 million years ago. The Maastrichtian was preceded by the Campanian and succeeded by the Danian (part of the Paleogene and Paleocene).At the end of this period, there was a mass extinction known as the Cretaceous–Paleogene extinction event, (formerly known as the Cretaceous–Tertiary extinction event). At this extinction event, many commonly recognized groups such as non-avian dinosaurs, plesiosaurs and mosasaurs, as well as many other lesser-known groups, died out. The cause of the extinction is most commonly linked to an asteroid about 10 to 15 kilometres (6.2 to 9.3 mi) wide colliding with Earth at the end of the Cretaceous.

Mesozoic

The Mesozoic Era ( or ) is an interval of geological time from about 252 to 66 million years ago. It is also called the Age of Reptiles, a phrase introduced by the 19th century paleontologist Gideon Mantell who viewed it as dominated by diapsids such as Iguanodon, Megalosaurus, Plesiosaurus and Pterodactylus. To paleobotanists, this Era is also called the Age of Conifers.Mesozoic means "middle life", deriving from the Greek prefix meso-/μεσο- for "between" and zōon/ζῷον meaning "animal" or "living being". The name "Mesozoic" was proposed in 1840 by the British geologist John Phillips (1800–1874). It is one of three geologic eras of the Phanerozoic Eon, preceded by the Paleozoic ("ancient life") and succeeded by the Cenozoic ("new life"). The era is subdivided into three major periods: the Triassic, Jurassic, and Cretaceous, which are further subdivided into a number of epochs and stages.

The era began in the wake of the Permian–Triassic extinction event, the largest well-documented mass extinction in Earth's history, and ended with the Cretaceous–Paleogene extinction event, another mass extinction whose victims included the non-avian dinosaurs. The Mesozoic was a time of significant tectonic, climate and evolutionary activity. The era witnessed the gradual rifting of the supercontinent Pangaea into separate landmasses that would move into their current positions during the next era. The climate of the Mesozoic was varied, alternating between warming and cooling periods. Overall, however, the Earth was hotter than it is today. Dinosaurs first appeared in the Mid-Triassic, and became the dominant terrestrial vertebrates in the Late Triassic or Early Jurassic, occupying this position for about 150 or 135 million years until their demise at the end of the Cretaceous. Birds first appeared in the Jurassic (however, true toothless birds appeared first in the Cretaceous), having evolved from a branch of theropod dinosaurs. The first mammals also appeared during the Mesozoic, but would remain small—less than 15 kg (33 lb)—until the Cenozoic.

Neoaves

Neoaves is a clade that consists of all modern birds (Neornithes or Aves) with the exception of Paleognathae (ratites and kin) and Galloanserae (ducks, chickens and kin). Almost 95% of the roughly 10,000 known species of modern birds belong to the Neoaves.

The early diversification of the various neoavian groups occurred very rapidly around the Cretaceous–Paleogene extinction event, and attempts to resolve their relationships with each other have resulted initially in much controversy.

Ordovician–Silurian extinction events

The Ordovician–Silurian extinction events, when combined, are the second-largest of the five major extinction events in Earth's history in terms of percentage of genera that became extinct. This event greatly affected marine communities, which caused the disappearance of one third of all brachiopod and bryozoan families, as well as numerous groups of conodonts, trilobites, and graptolites. The Ordovician–Silurian extinction occurred during the Hirnantian stage of the Ordovician Period and the subsequent Rhuddanian stage of the Silurian Period. The last event is dated in the interval of 455–430 Ma ago, i.e., lasting from the Middle Ordovician to Early Silurian, thus including the extinction period. This event was the first of the big five Phanerozoic events and was the first to significantly affect animal-based communities.Almost all major taxonomic groups were affected during this extinction event. Extinction was global during this period, eliminating 49-60% of marine genera and nearly 85% of marine species.Brachiopods, bivalves, echinoderms, bryozoans and corals were particularly affected. Before the late Ordovician cooling, temperatures were relatively warm and it is the suddenness of the climate changes and the elimination of habitats due to sea-level fall that are believed to have precipitated the extinctions. The falling sea level disrupted or eliminated habitats along the continental shelves. Evidence for the glaciation was found through deposits in the Sahara Desert. A combination of lowering of sea level and glacially driven cooling were likely driving agents for the Ordovician mass extinction.

Permian

The Permian is a geologic period and system which spans 47 million years from the end of the Carboniferous Period 298.9 million years ago (Mya), to the beginning of the Triassic period 251.902 Mya. It is the last period of the Paleozoic era; the following Triassic period belongs to the Mesozoic era. The concept of the Permian was introduced in 1841 by geologist Sir Roderick Murchison, who named it after the city of Perm.

The Permian witnessed the diversification of the early amniotes into the ancestral groups of the mammals, turtles, lepidosaurs, and archosaurs. The world at the time was dominated by two continents known as Pangaea and Siberia, surrounded by a global ocean called Panthalassa. The Carboniferous rainforest collapse left behind vast regions of desert within the continental interior. Amniotes, who could better cope with these drier conditions, rose to dominance in place of their amphibian ancestors.

The Permian (along with the Paleozoic) ended with the Permian–Triassic extinction event, the largest mass extinction in Earth's history, in which nearly 96% of marine species and 70% of terrestrial species died out. It would take well into the Triassic for life to recover from this catastrophe. Recovery from the Permian–Triassic extinction event was protracted; on land, ecosystems took 30 million years to recover.

Permian–Triassic extinction event

The Permian–Triassic (P–Tr or P–T) extinction event, colloquially known as the Great Dying, the End-Permian Extinction or the Great Permian Extinction, occurred about 252 Ma (million years) ago, forming the boundary between the Permian and Triassic geologic periods, as well as between the Paleozoic and Mesozoic eras. It is the Earth's most severe known extinction event, with up to 96% of all marine species and 70% of terrestrial vertebrate species becoming extinct. It is the only known mass extinction of insects. Some 57% of all biological families and 83% of all genera became extinct. Because so much biodiversity was lost, the recovery of land-dwelling life took significantly longer than after any other extinction event, possibly up to 10 million years. Studies in Bear Lake County, near Paris, Idaho, showed a relatively quick rebound in a localized marine ecosystem, taking around 2 million years to recover, suggesting that the impact of the extinction may have been felt less severely in some areas than others.

There is evidence for one to three distinct pulses, or phases, of extinction. Suggested mechanisms for the latter include one or more large meteor impact events, massive volcanism such as that of the Siberian Traps, and the ensuing coal or gas fires and explosions, and a runaway greenhouse effect triggered by sudden release of methane from the sea floor due to methane clathrate dissociation according to the clathrate gun hypothesis or methane-producing microbes known as methanogens. Possible contributing gradual changes include sea-level change, increasing anoxia, increasing aridity, and a shift in ocean circulation driven by climate change.

Triassic

The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.9 million years ago (Mya), to the beginning of the Jurassic Period 201.3 Mya. The Triassic is the first period of the Mesozoic Era. Both the start and end of the period are marked by major extinction events.Triassic began in the wake of the Permian–Triassic extinction event, which left the Earth's biosphere impoverished; it was well into the middle of the Triassic before life recovered its former diversity. Therapsids and archosaurs were the chief terrestrial vertebrates during this time. A specialized subgroup of archosaurs, called dinosaurs, first appeared in the Late Triassic but did not become dominant until the succeeding Jurassic Period.The first true mammals, themselves a specialized subgroup of therapsids, also evolved during this period, as well as the first flying vertebrates, the pterosaurs, who, like the dinosaurs, were a specialized subgroup of archosaurs. The vast supercontinent of Pangaea existed until the mid-Triassic, after which it began to gradually rift into two separate landmasses, Laurasia to the north and Gondwana to the south.

The global climate during the Triassic was mostly hot and dry, with deserts spanning much of Pangaea's interior. However, the climate shifted and became more humid as Pangaea began to drift apart. The end of the period was marked by yet another major mass extinction, the Triassic–Jurassic extinction event, that wiped out many groups and allowed dinosaurs to assume dominance in the Jurassic.

The Triassic was named in 1834 by Friedrich von Alberti, after the three distinct rock layers (tri meaning "three") that are found throughout Germany and northwestern Europe—red beds, capped by marine limestone, followed by a series of terrestrial mud- and sandstones—called the "Trias".

Triassic–Jurassic extinction event

The Triassic–Jurassic extinction event marks the boundary between the Triassic and Jurassic periods, 201.3 million years ago, and is one of the major extinction events of the Phanerozoic eon, profoundly affecting life on land and in the oceans. In the seas, a whole class (conodonts) and 23-34% of marine genera disappeared. On land, all archosaurs other than crocodylomorphs (Sphenosuchia and Crocodyliformes) and Avemetatarsalia (pterosaurs and dinosaurs), some remaining therapsids, and many of the large amphibians became extinct.

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