Extinction Event
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An extinction event (also known as a mass extinction or biotic crisis) is a widespread and rapid decrease in the
biodiversity Biodiversity is the variability of life, life on Earth. It can be measured on various levels. There is for example genetic variability, species diversity, ecosystem diversity and Phylogenetics, phylogenetic diversity. Diversity is not distribut ...
on
Earth Earth is the third planet from the Sun and the only astronomical object known to Planetary habitability, harbor life. This is enabled by Earth being an ocean world, the only one in the Solar System sustaining liquid surface water. Almost all ...
. Such an event is identified by a sharp fall in the diversity and abundance of
multicellular organism A multicellular organism is an organism that consists of more than one cell (biology), cell, unlike unicellular organisms. All species of animals, Embryophyte, land plants and most fungi are multicellular, as are many algae, whereas a few organism ...
s. It occurs when the rate of
extinction Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
increases with respect to the
background extinction rate Background extinction rate (BER), also known as the normal extinction rate, refers to the standard rate of extinction in Earth's geological and biological history, excluding major extinction events, including the current human-induced Holocene ex ...
and the rate of
speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...
. 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 disagreement as to what constitutes a "major" extinction event, and the data chosen to measure past diversity.


The "Big Five" mass extinctions

In a landmark paper published in 1982,
Jack Sepkoski Joseph John Sepkoski Jr. (July 26, 1948 – May 1, 1999) was a University of Chicago paleontologist. Sepkoski studied the fossil record and the diversity of life on Earth. Sepkoski and David Raup produced a new understanding of extinction events ...
and David M. Raup identified five particular geological intervals with excessive diversity loss. They were originally identified as outliers on a general trend of decreasing extinction rates during the
Phanerozoic The Phanerozoic is the current and the latest of the four eon (geology), geologic eons in the Earth's geologic time scale, covering the time period from 538.8 million years ago to the present. It is the eon during which abundant animal and ...
, but as more stringent statistical tests have been applied to the accumulating data, it has been established that in the current, Phanerozoic Eon, multicellular animal life has experienced at least five major and many minor mass extinctions. 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. All of the five in the Phanerozoic Eon were anciently preceded by the presumed far more extensive mass extinction of microbial life during the
Great Oxidation Event The Great Oxidation Event (GOE) or Great Oxygenation Event, also called the Oxygen Catastrophe, Oxygen Revolution, Oxygen Crisis or Oxygen Holocaust, was a time interval during the Earth's Paleoproterozoic era when the Earth's atmosphere an ...
(a.k.a. Oxygen Catastrophe) early in the
Proterozoic Eon The Proterozoic ( ) is the third of the four geologic eons of Earth's history, spanning the time interval from 2500 to 538.8 Mya, and is the longest eon of Earth's geologic time scale. It is preceded by the Archean and followed by the Phanerozoic ...
. At the end of the
Ediacaran The Ediacaran ( ) is a geological period of the Neoproterozoic geologic era, Era that spans 96 million years from the end of the Cryogenian Period at 635 Million years ago, Mya to the beginning of the Cambrian Period at 538.8 Mya. It is the last ...
and just before the Cambrian explosion, yet another Proterozoic extinction event (of unknown magnitude) is speculated to have ushered in the Phanerozoic.
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Triassic–Jurassic extinction event The Triassic–Jurassic (Tr-J) extinction event (TJME), often called the end-Triassic extinction, marks the boundary between the Triassic and Jurassic periods, . It represents one of five major extinction events during the Phanerozoic, profoundly ...
, style="vertical-align:top;text-align:left;", 201.3  Ma , - , style="vertical-align:top;text-align:left;" colspan="3", The End Triassic extinction marks the
Triassic The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is t ...
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...
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. Most non-dinosaurian
archosaur Archosauria () or archosaurs () is a clade of diapsid sauropsid tetrapods, with birds and crocodilians being the only extant taxon, extant representatives. Although broadly classified as reptiles, which traditionally exclude birds, the cladistics ...
s, most therapsids, and most of the large
amphibian Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
s were eliminated, leaving
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
s with little terrestrial competition. Non-dinosaurian archosaurs continued to dominate aquatic environments, while non-archosaurian diapsids continued to dominate marine environments. The
Temnospondyl Temnospondyli (from Greek language, Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order (biology), order of small to giant tetrapods—often considered Labyrinth ...
lineage of large amphibians also survived until the Cretaceous in Australia (such as ''
Koolasuchus ''Koolasuchus'' is an extinct genus of brachyopoid temnospondyl in the family Chigutisauridae. Fossils have been found from Victoria, Australia and date back 125-120 million years ago to Barremian-Aptian stages of the Early Cretaceous. ''Koola ...
''). , - , style="vertical-align:top;text-align:center;" rowspan="2", , style="vertical-align:top;text-align:left;",
Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event, also known as the K–T extinction, was the extinction event, mass extinction of three-quarters of the plant and animal species on Earth approximately 66 million years ago. The event cau ...
, style="vertical-align:top;text-align:left;",   Ma , - , style="vertical-align:top;text-align:left;" colspan="3", The End Cretaceous extinction, or the K–Pg extinction (formerly K–T extinction) occurred at the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
(
Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
) –
Paleogene The Paleogene Period ( ; also spelled Palaeogene or Palæogene) is a geologic period and system that spans 43 million years from the end of the Cretaceous Period Ma (million years ago) to the beginning of the Neogene Period Ma. It is the fir ...
(
Danian The Danian is the oldest age or lowest stage of the Paleocene Epoch or Series, of the Paleogene Period or System, and of the Cenozoic Era or Erathem. The beginning of the Danian (and the end of the preceding Maastrichtian) is at the Cretac ...
) transition. The event was formerly called the Cretaceous-Tertiary or K–T extinction or K–T boundary; it is now officially named the Cretaceous–Paleogene (or K–Pg) extinction event. : About 17% of all families, 50% of all
genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
and 75% of all species became extinct. In the seas all the
ammonites Ammonoids are extinct, (typically) coiled-shelled cephalopods comprising the subclass Ammonoidea. They are more closely related to living octopuses, squid, and cuttlefish (which comprise the clade Coleoidea) than they are to nautiluses (family N ...
,
plesiosaurs The Plesiosauria or plesiosaurs are an Order (biology), order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia. Plesiosaurs first appeared in the latest Triassic Period (geology), Period, possibly in the Rhaetian st ...
and
mosasaur Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Ancient Greek, Greek ' meaning 'lizard') are an extinct group of large aquatic reptiles within the family Mosasauridae that lived during the Late Cretaceous. Their first fossil remains wer ...
s disappeared and the percentage of
sessile Sessility, or sessile, may refer to: * Sessility (motility), organisms which are not able to move about * Sessility (botany), flowers or leaves that grow directly from the stem or peduncle of a plant * Sessility (medicine), tumors and polyps that ...
animals was reduced to about 33%. All known non-avian
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
s became extinct during that time. The boundary event was severe with a significant amount of variability in the rate of extinction between and among different
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
s.
Mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s, descended from the
synapsid Synapsida is a diverse group of tetrapod vertebrates that includes all mammals and their extinct relatives. It is one of the two major clades of the group Amniota, the other being the more diverse group Sauropsida (which includes all extant rept ...
s, and
bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
s, a side-branch of the
theropod Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
dinosaurs, emerged as the two predominant clades of terrestrial tetrapods. Despite the common presentation focusing only on these five events, no measure of extinction shows any definite line separating them from the many other
Phanerozoic The Phanerozoic is the current and the latest of the four eon (geology), geologic eons in the Earth's geologic time scale, covering the time period from 538.8 million years ago to the present. It is the eon during which abundant animal and ...
extinction events that appear only slightly lesser catastrophes; further, using different methods of calculating an extinction's impact can lead to other events featuring in the top five. Fossil records of older events are more difficult to interpret. This is because: * Older fossils are more difficult to find, as they are usually buried at a considerable depth. * Dating of 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. * Marine fossils tend to be better preserved than their more sought-after land-based counterparts, but the deposition and preservation of fossils on land is more erratic. 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 (geologically rapid increase in
fungal A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as one of the tradit ...
abundance) 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 In statistics, sampling bias is a bias (statistics), bias in which a sample is collected in such a way that some members of the intended statistical population, population have a lower or higher sampling probability than others. It results in a b ...
.


Sixth mass extinction

Research completed after the seminal 1982 paper (Sepkoski and Raup) has concluded that a sixth mass extinction event due to human activities is currently underway:


Extinctions by severity

Extinction events can be tracked by several methods, including geological change, ecological impact, extinction vs. origination (
speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...
) rates, and most commonly diversity loss among
taxonomic 280px, Generalized scheme of taxonomy Taxonomy is a practice and science concerned with classification or categorization. Typically, there are two parts to it: the development of an underlying scheme of classes (a taxonomy) and the allocation ...
units. Most early papers used
families Family (from ) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictability, structure, and safety as ...
as the unit of taxonomy, based on compendiums of marine animal families by Sepkoski (1982, 1992). Later papers by Sepkoski and other authors switched to
genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
, which are more precise than families and less prone to taxonomic bias or incomplete sampling relative to species. These are several major papers estimating loss or ecological impact from fifteen commonly-discussed extinction events. Different methods used by these papers are described in the following section. The "Big Five" mass extinctions are bolded. Graphed but not discussed by Sepkoski (1996), considered continuous with the Late Devonian mass extinction
At the time considered continuous with the end-Permian mass extinction
Includes late
Norian The Norian is a division of the Triassic geological period, Period. It has the rank of an age (geology), age (geochronology) or stage (stratigraphy), stage (chronostratigraphy). It lasted from ~227.3 to Mya (unit), million years ago. It was prec ...
time slices
Diversity loss of both pulses calculated together
Pulses extend over adjacent time slices, calculated separately
Considered ecologically significant, but not analyzed directly
Excluded due to a lack of consensus on Late Triassic chronology


The study of major extinction events


Breakthrough studies in the 1980s–1990s

For much of the 20th century, the study of mass extinctions was hampered by insufficient data. Mass extinctions, though acknowledged, were considered mysterious exceptions to the prevailing gradualistic view of prehistory, where slow evolutionary trends define faunal changes. The first breakthrough was published in 1980 by a team led by Luis Alvarez, who discovered trace metal evidence for an
asteroid impact An impact event is a collision between astronomical objects causing measurable effects. Impact events have been found to regularly occur in planetary systems, though the most frequent involve asteroids, comets or meteoroids and have minimal effe ...
at the end of the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
period. The
Alvarez hypothesis The Alvarez hypothesis posits that the Extinction event, mass extinction of the non-avian dinosaurs and many other living things during the Cretaceous–Paleogene extinction event was caused by the Impact event, impact of a large asteroid on the Ea ...
for the end-Cretaceous extinction gave mass extinctions, and catastrophic explanations, newfound popular and scientific attention. Another landmark study came in 1982, when a paper written by David M. Raup and
Jack Sepkoski Joseph John Sepkoski Jr. (July 26, 1948 – May 1, 1999) was a University of Chicago paleontologist. Sepkoski studied the fossil record and the diversity of life on Earth. Sepkoski and David Raup produced a new understanding of extinction events ...
was published in the journal ''
Science Science is a systematic discipline that builds and organises knowledge in the form of testable hypotheses and predictions about the universe. Modern science is typically divided into twoor threemajor branches: the natural sciences, which stu ...
''. This paper, originating from a compendium of extinct marine animal
families Family (from ) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictability, structure, and safety as ...
developed by Sepkoski, identified five peaks of marine family extinctions that stand out among a backdrop of decreasing extinction rates through time. Four of these peaks were statistically significant: the
Ashgillian The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era, and the second of twelve periods of the Phanerozoic Eon. The Ordovician spans 41.6 million years from the end of the Cambrian Period Ma (millio ...
( end-Ordovician),
Late Permian Late or LATE may refer to: Everyday usage * Tardy, or late, not being on time * Late (or the late) may refer to a person who is dead Music * ''Late'' (The 77s album), 2000 * Late (Alvin Batiste album), 1993 * Late!, a pseudonym used by Dave Groh ...
,
Norian The Norian is a division of the Triassic geological period, Period. It has the rank of an age (geology), age (geochronology) or stage (stratigraphy), stage (chronostratigraphy). It lasted from ~227.3 to Mya (unit), million years ago. It was prec ...
( end-Triassic), and
Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
(end-Cretaceous). The remaining peak was a broad interval of high extinction smeared over the later half of the
Devonian The Devonian ( ) is a period (geology), geologic period and system (stratigraphy), system of the Paleozoic era (geology), era during the Phanerozoic eon (geology), eon, spanning 60.3 million years from the end of the preceding Silurian per ...
, with its apex in the
Frasnian The Frasnian is one of two faunal stages in the Late Devonian Period. It lasted from million years ago to million years ago. It was preceded by the Givetian Stage and followed by the Famennian Stage. Major reef-building was under way during ...
stage. Through the 1980s, Raup and Sepkoski continued to elaborate and build upon their extinction and origination data, defining a high-resolution
biodiversity Biodiversity is the variability of life, life on Earth. It can be measured on various levels. There is for example genetic variability, species diversity, ecosystem diversity and Phylogenetics, phylogenetic diversity. Diversity is not distribut ...
curve (the "Sepkoski curve") and successive evolutionary faunas with their own patterns of diversification and extinction. Though these interpretations formed a strong basis for subsequent studies of mass extinctions, Raup and Sepkoski also proposed a more controversial idea in 1984: a 26-million-year periodic pattern to mass extinctions. Two teams of
astronomer An astronomer is a scientist in the field of astronomy who focuses on a specific question or field outside the scope of Earth. Astronomers observe astronomical objects, such as stars, planets, natural satellite, moons, comets and galaxy, galax ...
s linked this to a hypothetical
brown dwarf Brown dwarfs are substellar objects that have more mass than the biggest gas giant planets, but less than the least massive main sequence, main-sequence stars. Their mass is approximately 13 to 80 Jupiter mass, times that of Jupiter ()not big en ...
in the distant reaches of the
Solar System The Solar SystemCapitalization of the name varies. The International Astronomical Union, the authoritative body regarding astronomical nomenclature, specifies capitalizing the names of all individual astronomical objects but uses mixed "Sola ...
, inventing the "
Nemesis hypothesis Nemesis is a hypothetical red dwarf or brown dwarf, originally postulated in 1984 to be orbiting the Sun at a distance of about 95,000 AU (1.5 light-years), somewhat beyond the Oort cloud, to explain a perceived cycle of mass extinctions in the ...
", which has been strongly disputed by other astronomers. Around the same time, Sepkoski began to devise a compendium of marine animal
genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
, which would allow researchers to explore extinction at a finer taxonomic resolution. He began to publish preliminary results of this in-progress study as early as 1986, in a paper that identified 29 extinction intervals of note. By 1992, he also updated his 1982 family compendium, finding minimal changes to the diversity curve despite a decade of new data. In 1996, Sepkoski published another paper that tracked marine genera extinction (in terms of net diversity loss) by stage, similar to his previous work on family extinctions. The paper filtered its sample in three ways: all genera (the entire unfiltered sample size), multiple-interval genera (only those found in more than one stage), and "well-preserved" genera (excluding those from groups with poor or understudied fossil records). Diversity trends in marine animal families were also revised based on his 1992 update. Revived interest in mass extinctions led many other authors to re-evaluate geological events in the context of their effects on life. A 1995 paper by
Michael Benton Michael James Benton (born 8 April 1956) is a British palaeontologist, and professor of vertebrate paleontology, vertebrate palaeontology in the School of Earth Sciences at the University of Bristol. His published work has mostly concentrated on ...
tracked extinction and origination rates among both marine and continental (freshwater & terrestrial) families, identifying 22 extinction intervals and no periodic pattern. Overview books by O.H. Walliser (1996) and A. Hallam and P.B. Wignall (1997) summarized the new extinction research of the previous two decades. One chapter in the former source lists over 60 geological events that could conceivably be considered global extinctions of varying sizes. These texts, and other widely circulated publications in the 1990s, helped to establish the popular image of mass extinctions as a "big five" alongside many smaller extinctions through prehistory.


New data on genera: Sepkoski's compendium

Though Sepkoski died in 1999, his marine genera compendium was formally published in 2002. This prompted a new wave of studies into the dynamics of mass extinctions. These papers utilized the compendium to track origination rates (the rate that new species appear or
speciate Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...
) parallel to extinction rates in the context of geological stages or substages. A review and re-analysis of Sepkoski's data by Bambach (2006) identified 18 distinct mass extinction intervals, including 4 large extinctions in the
Cambrian The Cambrian ( ) is the first geological period of the Paleozoic Era, and the Phanerozoic Eon. The Cambrian lasted 51.95 million years from the end of the preceding Ediacaran period 538.8 Ma (million years ago) to the beginning of the Ordov ...
. These fit Sepkoski's definition of extinction, as short substages with large diversity loss and overall high extinction rates relative to their surroundings. Bambach et al. (2004) considered each of the "Big Five" extinction intervals to have a different pattern in the relationship between origination and extinction trends. Moreover, background extinction rates were broadly variable and could be separated into more severe and less severe time intervals. Background extinctions were least severe relative to the origination rate in the middle Ordovician-early Silurian, late Carboniferous-Permian, and Jurassic-recent. This argues that the Late Ordovician, end-Permian, and end-Cretaceous extinctions were statistically significant outliers in biodiversity trends, while the Late Devonian and end-Triassic extinctions occurred in time periods that were already stressed by relatively high extinction and low origination. Computer models run by Foote (2005) determined that abrupt pulses of extinction fit the pattern of prehistoric biodiversity much better than a gradual and continuous background extinction rate with smooth peaks and troughs. This strongly supports the utility of rapid, frequent mass extinctions as a major driver of diversity changes. Pulsed origination events are also supported, though to a lesser degree that is largely dependent on pulsed extinctions. Similarly, Stanley (2007) used extinction and origination data to investigate turnover rates and extinction responses among different evolutionary faunas and taxonomic groups. In contrast to previous authors, his diversity simulations show support for an overall exponential rate of biodiversity growth through the entire Phanerozoic.


Tackling biases in the fossil record

As data continued to accumulate, some authors began to re-evaluate Sepkoski's sample using methods meant to account for
sampling bias In statistics, sampling bias is a bias (statistics), bias in which a sample is collected in such a way that some members of the intended statistical population, population have a lower or higher sampling probability than others. It results in a b ...
es. As early as 1982, a paper by Phillip W. Signor and Jere H. Lipps noted that the true sharpness of extinctions was diluted by the incompleteness of the fossil record. This phenomenon, later called the Signor-Lipps effect, notes that a species' true extinction must occur after its last fossil, and that origination must occur before its first fossil. Thus, species that appear to die out just prior to an abrupt extinction event may instead be a victim of the event, despite an apparent gradual decline looking at the fossil record alone. A model by Foote (2007) found that many geological stages had artificially inflated extinction rates due to Signor-Lipps "backsmearing" from later stages with extinction events. Other biases include the difficulty in assessing taxa with high turnover rates or restricted occurrences, which cannot be directly assessed due to a lack of fine-scale temporal resolution. Many paleontologists opt to assess diversity trends by randomized sampling and
rarefaction Rarefaction is the reduction of an item's density, the opposite of compression. Like compression, which can travel in waves (sound waves, for instance), rarefaction waves also exist in nature. A common rarefaction wave is the area of low relati ...
of fossil abundances rather than raw temporal range data, in order to account for all of these biases. But that solution is influenced by biases related to sample size. One major bias in particular is the "
Pull of the recent The Pull of the Recent (POR) describes a phenomenon in which a combination of factors causes palaeontologists to overestimate diversity towards the present day. Biased preservation and sampling in the fossil record results in past biodiversity estim ...
", the fact that the fossil record (and thus known diversity) generally improves closer to the modern day. This means that biodiversity and abundance for older geological periods may be underestimated from raw data alone. Alroy (2010) attempted to circumvent sample size-related biases in diversity estimates using a method he called "
shareholder A shareholder (in the United States often referred to as stockholder) of corporate stock refers to an individual or legal entity (such as another corporation, a body politic, a trust or partnership) that is registered by the corporation as the ...
quorum subsampling" (SQS). In this method, fossils are sampled from a "collection" (such as a time interval) to assess the relative diversity of that collection. Every time a new species (or other
taxon In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
) enters the sample, it brings over all other fossils belonging to that species in the collection (its " share" of the collection). For example, a skewed collection with half its fossils from one species will immediately reach a sample share of 50% if that species is the first to be sampled. This continues, adding up the sample shares until a "coverage" or "
quorum A quorum is the minimum number of members of a group necessary to constitute the group at a meeting. In a deliberative assembly (a body that uses parliamentary procedure, such as a legislature), a quorum is necessary to conduct the business of ...
" is reached, referring to a pre-set desired sum of share percentages. At that point, the number of species in the sample are counted. A collection with more species is expected to reach a sample quorum with more species, thus accurately comparing the relative diversity change between two collections without relying on the biases inherent to sample size. Alroy also elaborated on three-timer algorithms, which are meant to counteract biases in estimates of extinction and origination rates. A given taxon is a "three-timer" if it can be found before, after, and within a given time interval, and a "two-timer" if it overlaps with a time interval on one side. Counting "three-timers" and "two-timers" on either end of a time interval, and sampling time intervals in sequence, can together be combined into equations to predict extinction and origination with less bias. In subsequent papers, Alroy continued to refine his equations to improve lingering issues with precision and unusual samples. McGhee et al. (2013), a paper that primarily focused on ecological effects of mass extinctions, also published new estimates of extinction severity based on Alroy's methods. Many extinctions were significantly more impactful under these new estimates, though some were less prominent. Stanley (2016) was another paper that attempted to remove two common errors in previous estimates of extinction severity. The first error was the unjustified removal of "singletons", genera unique to only a single time slice. Their removal would mask the influence of groups with high turnover rates or lineages cut short early in their diversification. The second error was the difficulty in distinguishing background extinctions from brief mass extinction events within the same short time interval. To circumvent this issue, background rates of diversity change (extinction/origination) were estimated for stages or substages without mass extinctions, and then assumed to apply to subsequent stages with mass extinctions. For example, the
Santonian The Santonian is an age in the geologic timescale or a chronostratigraphic stage. It is a subdivision of the Late Cretaceous Epoch or Upper Cretaceous Series. It spans the time between 86.3 ± 0.7 mya ( million years ago) and 83.6 ± 0.7 m ...
and
Campanian The Campanian is the fifth of six ages of the Late Cretaceous epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campa ...
stages were each used to estimate diversity changes in the
Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
prior to the K-Pg mass extinction. Subtracting background extinctions from extinction tallies had the effect of reducing the estimated severity of the six sampled mass extinction events. This effect was stronger for mass extinctions that occurred in periods with high rates of background extinction, like the Devonian.


Uncertainty in the Proterozoic and earlier eons

Because most diversity and
biomass Biomass is a term used in several contexts: in the context of ecology it means living organisms, and in the context of bioenergy it means matter from recently living (but now dead) organisms. In the latter context, there are variations in how ...
on Earth is
microbial A microorganism, or microbe, is an organism of microscopic size, which may exist in its single-celled form or as a colony of cells. The possible existence of unseen microbial life was suspected from antiquity, with an early attestation in ...
, and thus difficult to measure via fossils, extinction events placed on-record are those that affect the easily observed, biologically complex component of the
biosphere The biosphere (), also called the ecosphere (), is the worldwide sum of all ecosystems. It can also be termed the zone of life on the Earth. The biosphere (which is technically a spherical shell) is virtually a closed system with regard to mat ...
rather than the total diversity and abundance of life. For this reason, well-documented extinction events are confined to the
Phanerozoic eon The Phanerozoic is the current and the latest of the four geologic eons in the Earth's geologic time scale, covering the time period from 538.8 million years ago to the present. It is the eon during which abundant animal and plant life has ...
– with the sole exception of the Oxygen Catastrophe in the
Proterozoic The Proterozoic ( ) is the third of the four geologic eons of Earth's history, spanning the time interval from 2500 to 538.8 Mya, and is the longest eon of Earth's geologic time scale. It is preceded by the Archean and followed by the Phanerozo ...
– since before the Phanerozoic, all living organisms were either microbial, or if multicellular then soft-bodied. Perhaps due to the absence of a robust microbial fossil record, mass extinctions might only ''seem'' to be mainly a Phanerozoic phenomenon, with merely the ''observable'' extinction rates appearing low before large complex organisms with hard body parts arose. Extinction occurs at an uneven rate. Based on the
fossil record A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
, the background rate of extinctions on Earth is about two to five
taxonomic 280px, Generalized scheme of taxonomy Taxonomy is a practice and science concerned with classification or categorization. Typically, there are two parts to it: the development of an underlying scheme of classes (a taxonomy) and the allocation ...
families Family (from ) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictability, structure, and safety as ...
of
marine animal Marine life, sea life or ocean life is the collective ecological communities that encompass all aquatic animals, aquatic plant, plants, algae, marine fungi, fungi, marine protists, protists, single-celled marine microorganisms, microorganisms ...
s every million years. The Oxygen Catastrophe, which occurred around 2.45 billion years ago in the
Paleoproterozoic The Paleoproterozoic Era (also spelled Palaeoproterozoic) is the first of the three sub-divisions ( eras) of the Proterozoic eon, and also the longest era of the Earth's geological history, spanning from (2.5–1.6  Ga). It is further sub ...
, is plausible as the first-ever major extinction event. It was perhaps also the worst-ever, in some sense, but with the Earth's ecology just before that time so poorly understood, and the concept of
prokaryote A prokaryote (; less commonly spelled procaryote) is a unicellular organism, single-celled organism whose cell (biology), cell lacks a cell nucleus, nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Ancient Gree ...
genera so different from genera of complex life, that it would be difficult to meaningfully compare it to any of the "Big Five" even if Paleoproterozoic life were better known. Since the Cambrian explosion, five further major mass extinctions have significantly exceeded the background extinction rate. The most recent and best-known, the
Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event, also known as the K–T extinction, was the extinction event, mass extinction of three-quarters of the plant and animal species on Earth approximately 66 million years ago. The event cau ...
, which occurred approximately  Ma (million years ago), was a large-scale mass extinction of animal and plant species in a geologically short period of time. In addition to the five major Phanerozoic mass extinctions, there are numerous lesser ones, and the ongoing mass extinction caused by human activity is sometimes called the
sixth mass extinction The Holocene extinction, also referred to as the Anthropocene extinction or the sixth mass extinction, is an ongoing extinction event caused exclusively by human activities during the Holocene epoch. This extinction event spans numerous families ...
.


Evolutionary importance

Mass extinctions have sometimes accelerated the
evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
of life on Earth. When dominance of particular ecological niches passes from one group of organisms to another, it is rarely because the newly dominant group is "superior" to the old but usually because an extinction event eliminates the old, dominant group and makes way for the new one, a process known as
adaptive radiation In evolutionary biology, adaptive radiation is a process in which organisms diversify rapidly from an ancestral species into a multitude of new forms, particularly when a change in the environment makes new resources available, alters biotic int ...
. For example,
mammaliaformes Mammaliaformes ("mammalian forms") is a clade of synapsid tetrapods that includes the crown group mammals and their closest extinct relatives; the group radiated from earlier probainognathian cynodonts during the Late Triassic. It is defined as ...
("almost mammals") and then
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s existed throughout the reign of the
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
s, but could not compete in the large terrestrial vertebrate niches that 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. The dinosaurs themselves had been beneficiaries of a previous mass extinction, the end-Triassic, which eliminated most of their chief rivals, the
crurotarsans Crurotarsi is a clade of archosauriform reptiles that includes crocodilians and stem-crocodilians and possibly bird-line archosaurs too if the extinct, crocodile-like phytosaurs are more distantly related to crocodiles than traditionally thought ...
. Similarly, within
Synapsida Synapsida is a diverse group of tetrapod vertebrates that includes all mammals and their extinct relatives. It is one of the two major clades of the group Amniota, the other being the more diverse group Sauropsida (which includes all extant rep ...
, the replacement of taxa that originated in the earliest, Pennsylvanian and
Cisuralian The Cisuralian, also known as the Early Permian, is the first series/epoch of the Permian. The Cisuralian was preceded by the Pennsylvanian and followed by the Guadalupian. The Cisuralian Epoch is named after the western slopes of the Ural Mou ...
evolutionary radiation (often still called "
pelycosaur Pelycosaur ( ) is an older term for basal or primitive Late Paleozoic synapsids, excluding the therapsids and their descendants. Previously, the term mammal-like reptile was used, and Pelycosauria was considered an order, but this is now thoug ...
s", though this is a
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
group) by
therapsids Therapsida is a clade comprising a major group of eupelycosaurian synapsids that includes mammals and their ancestors and close relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including li ...
occurred around the
Kungurian In the geologic timescale, the Kungurian is an age or stage of the Permian. It is the latest or upper of four subdivisions of the Cisuralian Epoch or Series. The Kungurian lasted between and million years ago (Ma). It was preceded by the Arti ...
/
Roadian In the geologic timescale, the Roadian is an age or stage of the Permian. It is the earliest or lower of three subdivisions of the Guadalupian Epoch or Series. The Roadian lasted between and million years ago (Ma). It was preceded by the Kun ...
transition, which is often called
Olson's extinction Olson's Extinction was a mass extinction that occurred in the late Cisuralian or early Guadalupian epoch of the Permian period, predating the much larger Permian–Triassic extinction event. The event is named after American paleontologist E ...
(which may be a slow decline over 20 Ma rather than a dramatic, brief event). Another point of view put forward in the
Escalation hypothesis {{More citations needed, date=January 2019 The Escalation Hypothesis is an evolutionary theory in biology put forward by Geerat J. Vermeij. It states that organisms are in constant conflict with one another and therefore devote many resources t ...
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 that 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". However, clades that survive for a considerable period of time after a mass extinction, and that were reduced to only a few species, are likely to have experienced a rebound effect called the "
push of the past The push of the past is a type of survivorship bias associated with evolutionary diversification when extinction is possible. Groups that survive a long time are likely to have “got off to a flying start”, and this statistical bias creates an i ...
". 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 ''On the Origin of Species'' (or, more completely, ''On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life'')The book's full original title was ''On the Origin of Species by M ...
'': : "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

Various authors have suggested that extinction events occurred periodically, every 26 to 30 million years, or that diversity fluctuates episodically about every 62 million years. Different cycle lengths have been proposed; e.g. by Various ideas, mostly regarding
astronomical Astronomy is a natural science that studies celestial objects and the phenomena that occur in the cosmos. It uses mathematics, physics, and chemistry in order to explain their origin and their overall evolution. Objects of interest include ...
influences, 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 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. Many of the proposed correlations have been argued to be spurious or lacking statistical significance. 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 such as Strontium isotopes, flood basalts, anoxic events, orogenies, and evaporite deposition. One explanation for this proposed cycle is carbon storage and release by oceanic crust, which exchanges carbon between the atmosphere and mantle. Mass extinctions are thought to result when a long-term stress is compounded by a short-term shock. Over the course of the
Phanerozoic The Phanerozoic is the current and the latest of the four eon (geology), geologic eons in the Earth's geologic time scale, covering the time period from 538.8 million years ago to the present. It is the eon during which abundant animal and ...
, individual taxa appear to have become less likely to suffer extinction, which may reflect more robust food webs, as well as fewer 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 specific mass extinctions

A good theory for a particular mass extinction should: * explain all of the losses, not just focus on a few groups (such as dinosaurs); * explain why particular groups of organisms died out and why others survived; * provide mechanisms that are strong enough to cause a mass extinction but not a total extinction; * 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 that 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 The Phanerozoic is the current and the latest of the four eon (geology), geologic eons in the Earth's geologic time scale, covering the time period from 538.8 million years ago to the present. It is the eon during which abundant animal and ...
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) summarized the relationship between mass extinctions and events that are most often cited as causes of mass extinctions, using data from Courtillot, Jaeger & Yang ''et al.'' (1996), Hallam (1992) and Grieve & Pesonen (1992): *
Flood basalt A flood basalt (or plateau basalt) is the result of a giant volcanic eruption or series of eruptions that covers large stretches of land or the ocean floor with basalt lava. Many flood basalts have been attributed to the onset of a hotspot (geolo ...
events (giant volcanic eruptions): 11 occurrences, all associated with significant extinctions. 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. * Sea-level falls: 12, of which seven were associated with significant extinctions. * Asteroid impacts: one large impact is associated with a mass extinction, that is, the Cretaceous–Paleogene extinction event; there have been many smaller impacts but they are not associated with significant extinctions, or cannot be dated precisely enough. The impact that created the
Siljan Ring The Siljan Ring () is a prehistoric impact structure in Dalarna, central Sweden. It is one of the 15 largest known impact structures on Earth and the largest in Europe, with a diameter of about . The impact that created the Siljan Ring occurred ...
either was just before the Late Devonian Extinction or coincided with it. The most commonly suggested causes of mass extinctions are listed below.


Flood basalt events

The formation of
large igneous province A large igneous province (LIP) is an extremely large accumulation of igneous rocks, including intrusive ( sills, dikes) and extrusive (lava flows, tephra deposits), arising when magma travels through the crust towards the surface. The format ...
s by flood basalt events could have: * produced dust and
particulate Particulate matter (PM) or particulates are microscopic particles of solid or liquid matter suspended in the air. An ''aerosol'' is a mixture of particulates and air, as opposed to the particulate matter alone, though it is sometimes define ...
aerosols, which inhibited photosynthesis and thus caused
food chain A food chain is a linear network of links in a food web, often starting with an autotroph (such as grass or algae), also called a producer, and typically ending at an apex predator (such as grizzly bears or killer whales), detritivore (such as ...
s to collapse both on land and at sea * emitted sulfur oxides that were precipitated as
acid rain Acid rain is rain or any other form of Precipitation (meteorology), precipitation that is unusually acidic, meaning that it has elevated levels of hydrogen ions (low pH). Most water, including drinking water, has a neutral pH that exists b ...
and poisoned many organisms, contributing further to the collapse of food chains * emitted
carbon dioxide Carbon dioxide is a chemical compound with the chemical formula . It is made up of molecules that each have one carbon atom covalent bond, covalently double bonded to two oxygen atoms. It is found in a gas state at room temperature and at norma ...
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. Flood basalt events have been implicated as the cause of many major extinction events. It is speculated that massive volcanism caused or contributed to the
Kellwasser Event The Late Devonian mass extinction, also known as the Kellwasser event, was a mass extinction event which occurred around 372 million years ago, at the boundary between the Frasnian and Famennian ages of the Late Devonian period.Racki, 2005McGhee, ...
, the End-Guadalupian Extinction Event, the End-Permian Extinction Event, the
Smithian-Spathian Extinction In the geologic timescale, the Olenekian is an age in the Early Triassic epoch; in chronostratigraphy, it is a stage in the Lower Triassic series. It spans the time between Ma and Ma (million years ago). The Olenekian is sometimes divided into ...
, the Triassic-Jurassic Extinction Event, the
Toarcian Oceanic Anoxic Event The Toarcian extinction event, also called the Pliensbachian-Toarcian extinction event, the Early Toarcian mass extinction, the Early Toarcian palaeoenvironmental crisis, or the Jenkyns Event, was an extinction event that occurred during the early ...
, the
Cenomanian-Turonian Oceanic Anoxic Event The Cenomanian-Turonian boundary event, also known as the Cenomanian-Turonian extinction, Cenomanian-Turonian Oceanic Anoxic Event ( OAE 2), and referred to also as the Bonarelli Event or Level, was an anoxic extinction event in the Cretaceous pe ...
, the Cretaceous-Palaeogene Extinction Event, and the Palaeocene-Eocene Thermal Maximum. The correlation between gigantic volcanic events expressed in the large igneous provinces and mass extinctions was shown for the last 260 million years. Recently such possible correlation was extended across the whole Phanerozoic Eon.


Sea-level fall

These are often clearly marked by worldwide sequences of contemporaneous sediments that 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 plate tectonics, mid-ocean ridges. Sea-level falls are associated with most of the mass extinctions, including all of the "Big Five"—Ordovician–Silurian extinction events, End-Ordovician, Late Devonian extinction, Late Devonian, Permian–Triassic extinction event, End-Permian, Triassic–Jurassic extinction event, End-Triassic, and Cretaceous–Paleogene extinction event, End-Cretaceous, along with the more recently recognised Capitanian mass extinction of comparable severity to the Big Five. A 2008 study, published in the journal ''Nature'', 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.


Extraterrestrial threats


= Impact events

= The impact of a sufficiently large asteroid or comet could have caused
food chain A food chain is a linear network of links in a food web, often starting with an autotroph (such as grass or algae), also called a producer, and typically ending at an apex predator (such as grizzly bears or killer whales), detritivore (such as ...
s to collapse both on land and at sea by producing dust and Atmospheric particulate matter, particulate aerosols and thus inhibiting photosynthesis. Impacts on sulfur-rich rocks could have emitted sulfur oxides precipitating as poisonous
acid rain Acid rain is rain or any other form of Precipitation (meteorology), precipitation that is unusually acidic, meaning that it has elevated levels of hydrogen ions (low pH). Most water, including drinking water, has a neutral pH that exists b ...
, 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, but there is lingering dispute whether the impact was the sole cause of the
Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event, also known as the K–T extinction, was the extinction event, mass extinction of three-quarters of the plant and animal species on Earth approximately 66 million years ago. The event cau ...
. Nonetheless, in October 2019, researchers reported that the Chicxulub impactor, Cretaceous Chicxulub asteroid impact that resulted in the Cretaceous–Paleogene extinction event, extinction of non-avian dinosaurs 66 Ma, also rapidly Ocean acidification, acidified the oceans, producing ecological collapse and long-lasting effects on the climate, and was a key reason for end-Cretaceous mass extinction. The Permian-Triassic extinction event has also been hypothesised to have been caused by an asteroid impact that formed the Araguainha crater due to the estimated date of the crater's formation overlapping with the end-Permian extinction event. However, this hypothesis has been widely challenged, with the impact hypothesis being rejected by most researchers. According to the Shiva hypothesis, the Earth is subject to increased asteroid impacts about once every 27 million years because of the Sun's passage through the plane of the Milky Way galaxy, thus causing extinction events at 27 million year intervals. Some evidence for this hypothesis has emerged in both marine and non-marine contexts. Alternatively, the Sun's passage through the higher density spiral arms of the galaxy could coincide with mass extinction on Earth, perhaps due to increased impact events. However, a reanalysis of the effects of the Sun's transit through the spiral structure based on maps of the spiral structure of the Milky Way in CO molecular line emission has failed to find a correlation.


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, 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 gamma ray burst caused the Ordovician–Silurian extinction events, End-Ordovician extinction, while a supernova has been proposed as the cause of the Hangenberg event. A supernova within 25 light-years would strip Earth of its atmosphere. Today there is in the Solar System's neighbourhood no critical star capable to produce a supernova dangerous to life on Earth.


Global cooling

Sustained and significant global cooling could kill many Polar circle, polar and temperate species and force others to migrate towards the equator; reduce the area available for Tropics, 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 Ordovician–Silurian extinction events, End-Ordovician, Permian–Triassic extinction event, Permian–Triassic, Late Devonian extinction, 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 Tropics, tropical species; kill temperate species or force them to migrate towards the Polar circle, 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 The Triassic–Jurassic (Tr-J) extinction event (TJME), often called the end-Triassic extinction, marks the boundary between the Triassic and Jurassic periods, . It represents one of five major extinction events during the Phanerozoic, profoundly ...
, 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 shelf, 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 effect, 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 Isotope analysis, 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 Permian–Triassic extinction event, 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 extinction events, Ordovician–Silurian, Late Devonian extinction, late Devonian, Capitanian mass extinction event, Capitanian, Permian–Triassic extinction event, Permian–Triassic, and Triassic–Jurassic extinction event, Triassic–Jurassic extinctions, as well as a number of lesser extinctions (such as the Ireviken event, Ireviken, Lundgreni Event, Lundgreni, Mulde event, Mulde, Lau event, Lau, Smithian-Spathian boundary extinction, Smithian-Spathian, Toarcian turnover, Toarcian, and Cenomanian-Turonian boundary event, Cenomanian–Turonian events). On the other hand, there are widespread black shale beds from the mid-Cretaceous that indicate anoxic events but are not associated with mass extinctions. The bio-availability of Essential trace element, 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, that is, 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 taxon, extant organisms. British oceanologist and atmospheric scientist, Andrew Watson (scientist), Andrew Watson, explained that, while the Holocene, Holocene epoch exhibits many processes reminiscent of those that have contributed to past anoxic events, full-scale ocean anoxia would take "thousands of years to develop".


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 that 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 that 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 extinction, late Devonian and Permian–Triassic extinction event, Permian–Triassic extinctions.


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 that expose previously isolated species to competition for which they are poorly adapted (for example, the extinction of most of South America's Meridiungulata, native ungulates and all of its Sparassodonta, large metatherians after the Great American Interchange, creation of a land bridge between North and South America). Occasionally continental drift creates a super-continent that 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 that may have extreme seasonal variations. Another theory is that the creation of the super-continent Pangaea contributed to the Permian-Triassic extinction event, 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.


Human activities

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 of the United States of America, Proceedings of the National Academy of Sciences'' argued that a "biological annihilation" akin to a Holocene extinction, sixth mass extinction event is underway as a result of anthropogenic causes, such as Human overpopulation, over-population and Overconsumption, 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.


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 that are contrary to the available evidence; they are based on other theories that have been rejected or superseded.


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 would be completely sterilized): rising global temperatures caused by the expanding Sun would gradually increase the rate of weathering, which would in turn remove more and more CO2 from the atmosphere. When CO2 levels get too low (perhaps at 50 ppm), most plant life will die out, although simpler plants like grasses and mosses can survive much longer, until levels drop to 10 ppm. With all photosynthetic organisms gone, atmospheric oxygen can no longer be replenished, and it 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
prokaryote A prokaryote (; less commonly spelled procaryote) is a unicellular organism, single-celled organism whose cell (biology), cell lacks a cell nucleus, nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Ancient Gree ...
s. When the Sun becomes 10% brighter in about a billion years, 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 around 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, it would represent the final mass extinction in Earth's history (albeit a very long extinction event).


Effects and recovery

The effects of mass extinction events varied widely. After a major extinction event, usually only weed#Weeds as adaptable species, weedy species survive due to their ability to live in diverse habitats. Later, species diversify and occupy empty niches. Generally, it takes millions of years for
biodiversity Biodiversity is the variability of life, life on Earth. It can be measured on various levels. There is for example genetic variability, species diversity, ecosystem diversity and Phylogenetics, phylogenetic diversity. Diversity is not distribut ...
to recover after extinction events. In the most severe mass extinctions it may take 15 to 30 million years. The worst
Phanerozoic The Phanerozoic is the current and the latest of the four eon (geology), geologic eons in the Earth's geologic time scale, covering the time period from 538.8 million years ago to the present. It is the eon during which abundant animal and ...
event, the Permian–Triassic extinction event, 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 pioneer organism, 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 that inhibited recovery, as well as prolonged environmental stress that continued into the Early Triassic. Recent research indicates that recovery did not begin until the start of the mid-Triassic, four to six million years after the extinction; and some writers estimate that the recovery was not complete until 30 million years after the P-T extinction, that is, 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.


In media

The term extinction level event (ELE) has been used in media. The 1998 film Deep Impact (film), Deep Impact describes a potential comet strike of earth as an E.L.E.Deep Impact
''Roger Ebert''. 8 May 1998. Retrieved 14 May 2024.


See also

* Bioevent * Elvis taxon * Endangered species * Geologic time scale * Global catastrophic risk * Holocene extinction * Human extinction * Kačák Event * Lazarus taxon * List of impact structures on Earth * List of largest volcanic eruptions * List of possible impact structures on Earth * Medea hypothesis * Rare species * Signor–Lipps effect * Snowball Earth * Speculative evolution * The Sixth Extinction: An Unnatural History, ''The Sixth Extinction: An Unnatural History'' (nonfiction book) * Timeline of extinctions in the Holocene * Late Pleistocene extinctions


Footnotes


References


Further reading

* * Edmeades B (2021) Megafauna: First victims of the human-caused extinction , Houndstooth Press , isbn 978-1-5445-2651-5


External links

* * – nonprofit organization producing a documentary about mass extinction titled ''"Call of Life: Facing the Mass Extinction"'' * * – Calculate extinction rates for yourself! {{DEFAULTSORT:Extinction Event Extinction events, Hypothetical impact events, * History of climate variability and change Evolutionary biology Meteorological hypotheses Natural disasters