An extinction event (also known as: mass extinction; extinction-level event (ELE), or biotic crisis) is a sharp decrease in the diversity and abundance of macroscopic life. They occur when the rate of extinction increases with respect to the rate of speciation. Because the majority of 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.
Over 98% of documented species are now extinct, but 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 invertebrates and vertebrates every million years.Marine fossils are mostly used to measure extinction rates because of their superior fossil record and stratigraphic range compared to land organisms.
Since life began on Earth, several major mass extinctions have significantly exceeded the background extinction rate. The most recent, the Cretaceous–Paleogene extinction event, which occurred approximately 65.5 million years ago (Ma), was a large-scale mass extinction of animal and plant species in a geologically short period of time. In the past 540 million years there have been five major events when over 50% of animal species died. Mass extinctions seem to be a Phanerozoic phenomenon, with extinction rates low before large complex organisms arose.
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.
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, but as more stringent statistical tests have been applied to the accumulating data, 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.
Despite the popularization of these five events, there is no fine line separating them from other extinction events; indeed, using different methods of calculating an extinction's impact can lead to other events featuring in the top five.
The older the fossil record gets, the more difficult it is to read. This is because:
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. 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 indeed real. A quantification of the rock exposure of Western Europe does indicate that many of the minor events for which a biological explanation has been sought are most readily explained by sampling bias.
Lesser extinction events include:
|Quaternary extinction event||50 ka to now||Anthropogenic|
|Neogene||Middle Miocene disruption||14.5 Ma||Nördlinger Ries bolide impact? Volcanoes in African Rift Valley|
|Eocene–Oligocene extinction event||33.9 Ma||Volcanoes? Chesapeake Bay and Popigai crater bolide impacts?|
|Cretaceous||Aptian extinction||117 Ma||Rahjamal Traps volcanism episode in Bengal?|
|End-Jurassic extinction||145.5 Ma|
|Jurassic||Toarcian turnover||183 Ma|
|Permian||Olson's Extinction||270 Ma|
|Carboniferous||Carboniferous Rainforest Collapse||318 Ma||Climate change|
|End Silurian||416 Ma|
|Lau event||420 Ma|
|Mulde event||424 Ma||Global drop in sea level?|
|Silurian||Ireviken event||428 Ma||Deep-ocean anoxia?|
|Cambrian–Ordovician extinction event||488 Ma||Glaciation? Depletion of oxygen in marine waters?|
|Cambrian||End Botomian extinction event||517 Ma|
|Precambrian||End-Ediacaran extinction||542 Ma||Ocean anoxia?|
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. 
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". So analysing extinctions in terms of "what died and what survived" often fails to tell the full story.
It has been suggested variously that extinction events occurred periodically, every 26 to 30 million years, or that diversity fluctuates episodically every ~62 million years. Various ideas attempt to explain the supposed pattern, including the presence of a hypothetical companion star to the sun, oscillations in the galactic plane, or passage through the Milky Way's spiral arms. However, other authors have concluded 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. Many of the proposed correlations have been argued to be spurious.Mass extinctions are thought to result when a long-term stress is compounded by a short term shock. Over the course of the Phanerozoic, individual taxa appear to be less likely to become extinct at any time, which may reflect more robust food webs as well as less extinction-prone species and other factors such as continental distribution.However, even after accounting for sampling bias, there does appear to be a gradual decrease in extinction and origination rates during the Phanerozoic. 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; and larger taxonomic groups (by definition) appear earlier in geological time.
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,  but susceptibility to extinction at a taxonomic level does not appear to make mass extinctions more or less probable.
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. 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.
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.
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. 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.
Macleod (2001) 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), Hallam (1992) and Grieve et al. (1996):
The most commonly suggested causes of mass extinctions are listed below.
The formation of large igneous provinces by flood basalt events could have:
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.
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.
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. 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.
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. 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 65 Ma, but there is an ongoing dispute whether the impact was the sole cause of the Cretaceous–Tertiary extinction event. There is evidence that there was an interval of about 300 ka from the impact to the mass extinction. In 1997, paleontologist Sankar Chatterjee drew attention to the proposed and much larger 600 km (370 mi) Shiva crater and the possibility of a multiple-impact scenario.
In 2007, a hypothesis was put forth that argued the impactor that killed the dinosaurs 65 Ma years ago belonged to the Baptistina family of asteroids. Concerns have been raised regarding the reputed link, in part because very few solid observational constraints exist of the asteroid or family. Indeed, it was recently discovered that 298 Baptistina does not share the same chemical signature as the source of the K-T impact. Although this finding may make the link between the Baptistina family and K-T impactor more difficult to substantiate, it does not preclude the possibility.
Sustained 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.
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.
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 went extinct. Furthermore, the Permian–Triassic extinction event has been suggested to have been caused by warming.  
See main article: 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.
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 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.
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.
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.  
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.
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.
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. Gamma ray bursts are fairly rare, occurring only a few times in a given galaxy per million years. A proposal that a supernova or gamma ray burst had caused a mass extinction would also have to be backed up by astronomical evidence of such an explosion at the right place and time.
It has been suggested that a supernova or gamma ray burst caused the End-Ordovician extinction.
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.
It is widely thought 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.
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.
The impact of mass extinction events varied widely. The worst event, the Permian–Triassic extinction event, devastated life on earth and is estimated to have killed off over 90% of species. Life on Earth seemed to recover quickly after this 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 the successive waves of extinction which inhibited recovery, as well as to prolonged environmental stress to organisms 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; and some writers estimate that the recovery was not complete until 30M years after the P-Tr extinction, i.e. in the late Triassic.
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.
After a major extinction event, species diversify and occupy empty niches.