Peristiwa kepupusan

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.
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".
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".

Patterns in frequency

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.
Others have argued that there is strong evidence supporting periodicity in a variety of records,
and additional evidence in the form of coincident periodic variation in nonbiological geochemical variables.
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.

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. 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.

Identifying causes of particular 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.
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.

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. 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.

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. 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.

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.
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.

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

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.
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.

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.

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.
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. Gamma ray bursts are fairly rare, occurring only a few times in a given galaxy per million years.
It has been suggested that a supernova or gamma ray burst caused the End-Ordovician extinction.

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.

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.

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.

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. 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.
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; and some writers estimate that the recovery was not complete until 30M years after the P-T extinction, i.e. in the late Triassic.  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.
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.