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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 or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic ('' genetic variability''), species ('' species diversity''), and ecosystem ('' ecosystem diversity'') ...
on
Earth Earth is the third planet from the Sun and the only astronomical object known to harbor life. While large volumes of water can be found throughout the Solar System, only Earth sustains liquid surface water. About 71% of Earth's sur ...
. Such an event is identified by a sharp change in the diversity and abundance of multicellular organisms. It occurs when the rate of
extinction Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the Endling, last individual of the species, although the Functional ext ...
increases with respect to the background extinction rate and the rate of speciation. Estimates of the number of major mass extinctions in the last 540 million years range from as few as five to more than twenty. These differences stem from 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 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, but as more stringent statistical tests have been applied to the accumulating data, it has been established that 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. An earlier (first) event at the end of the Ediacaran is speculated. # Ordovician–Silurian extinction events (End Ordovician or O–S): 445–444  Ma, just prior to and at the Ordovician
Silurian The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleoz ...
transition. Two events occurred that killed off 27% of all families, 57% of all genera and 85% of all
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
. Together they are ranked by many scientists as the second-largest of the five major extinctions in Earth's history in terms of percentage of genera that became extinct. In May 2020, studies suggested the cause of the mass extinction was due to
global warming In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to ...
, related to volcanism, and anoxia, and not due, as considered earlier, to cooling and
glaciation A glacial period (alternatively glacial or glaciation) is an interval of time (thousands of years) within an ice age that is marked by colder temperatures and glacier advances. Interglacials, on the other hand, are periods of warmer climate bet ...
. However, this is at odds with numerous previous studies, which have indicated global cooling as the primary driver. Most recently, the deposition of volcanic ash has been suggested to be the trigger for reductions in atmospheric carbon dioxide leading to the glaciation and anoxia observed in the geological record. # Late Devonian extinctions: 372–359  Ma, occupying much of the Late Devonian up to the
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wh ...
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carboniferou ...
transition. The Late Devonian was an interval of high diversity loss, concentrated into two extinction events. The largest extinction was the ''
Kellwasser Event The Late Devonian extinction consisted of several extinction events in the Late Devonian Epoch, which collectively represent one of the five largest mass extinction events in the history of life on Earth. The term primarily refers to a major ex ...
'' (
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 th ...
-
Famennian The Famennian is the latter of two faunal stages in the Late Devonian Epoch. The most recent estimate for its duration estimates that it lasted from around 371.1 million years ago to 359.3 million years ago. An earlier 2012 estimate, still used ...
, or F-F, 372 Ma), an extinction event at the end of the Frasnian, about midway through the Late Devonian. This extinction annihilated
coral reefs A coral reef is an underwater ecosystem characterized by reef-building corals. Reefs are formed of colonies of coral polyps held together by calcium carbonate. Most coral reefs are built from stony corals, whose polyps cluster in groups. ...
and numerous tropical
benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from ancient Greek, βένθος (bénthos), meaning " ...
(seabed-living) animals such as jawless fish, brachiopods, and
trilobites Trilobites (; meaning "three lobes") are extinct marine arthropods that form the class Trilobita. Trilobites form one of the earliest-known groups of arthropods. The first appearance of trilobites in the fossil record defines the base of the A ...
. Another major extinction was the '' Hangenberg Event'' (Devonian-Carboniferous, or D-C, 359 Ma), which brought an end to the Devonian as a whole. This extinction wiped out the armored placoderm fish and nearly led to the extinction of the newly-evolved ammonoids. These two closely-spaced extinction events collectively eliminated about 19% of all families, 50% of all genera and at least 70% of all species. Sepkoski and Raup (1982) did not initially consider the Late Devonian extinction interval ( Givetian, Frasnian, and Famennian stages) to be statistically significant. Regardless, later studies have affirmed the strong ecological impacts of the Kellwasser and Hangenberg Events. #
Permian–Triassic extinction event The Permian–Triassic (P–T, P–Tr) extinction event, also known as the Latest Permian extinction event, the End-Permian Extinction and colloquially as the Great Dying, formed the boundary between the Permian and Triassic geologic periods, ...
(End Permian): 252  Ma, at the
Permian The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleo ...
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest per ...
transition. Earth's largest extinction killed 57% of all families, 83% of all genera and 90% to 96% of all species (53% of marine families, 84% of marine genera, about 81% of all marine species and an estimated 70% of land species, including
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pa ...
s). The highly successful marine arthropod, the
trilobite Trilobites (; meaning "three lobes") are extinct marine arthropods that form the class Trilobita. Trilobites form one of the earliest-known groups of arthropods. The first appearance of trilobites in the fossil record defines the base of the ...
, became extinct. The evidence regarding
plant Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae excl ...
s is less clear, but new taxa became dominant after the extinction. The "Great Dying" had enormous evolutionary significance: On land, it ended the primacy of early synapsids. The recovery of vertebrates took 30 million years, but the vacant niches created the opportunity for
archosaur Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avia ...
s to become ascendant. In the seas, the percentage of animals that were sessile dropped from 67% to 50%. The whole late Permian was a difficult time, at least for marine life, even before the P–T boundary extinction. More recent research has indicated that the End-Capitanian extinction event that preceded the "Great Dying" likely constitutes a separate event from the P–T extinction; if so, it would be larger than some of the "Big Five" extinction events, and perhaps merit a separate place in this list immediately before this one. # Triassic–Jurassic extinction event (End Triassic): 201.3  Ma, at the
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest per ...
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
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 () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avia ...
s, most therapsids, and most of the large amphibians were eliminated, leaving
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
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 τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') is a diverse order of small to giant tetrapods—often considered primitive amphibians—that flourished worldwide during the Carb ...
lineage of large amphibians also survived until the Cretaceous in Australia (e.g., '' Koolasuchus''). #
Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event (also known as the Cretaceous–Tertiary extinction) was a sudden mass extinction of three-quarters of the plant and animal species on Earth, approximately 66 million years ago. With the ...
(''End Cretaceous'', ''K–Pg extinction'', or formerly ''K–T extinction''): Ma, at the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
(
Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the inte ...
) – Paleogene (
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 and 75% of all species became extinct. In the seas all the
ammonites Ammonoids are a group of extinct marine mollusc animals in the subclass Ammonoidea of the class Cephalopoda. These molluscs, commonly referred to as ammonites, are more closely related to living coleoids (i.e., octopuses, squid and cuttlefis ...
, plesiosaurs and
mosasaur Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Greek ' meaning 'lizard') comprise a group of extinct, large marine reptiles from the Late Cretaceous. Their first fossil remains were discovered in a limestone quarry at Maastricht on ...
s disappeared and the percentage of sessile animals (those unable to move about) was reduced to about 33%. All non-avian
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
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 A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...
s.
Mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur ...
s and
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
s, the former descended from the synapsids and the latter from theropod dinosaurs, emerged as dominant terrestrial animals. Despite the popularization of these five events, there is no definite line separating them from other extinction events; using different methods of calculating an extinction's impact can lead to other events featuring in the top five. Older fossil records are more difficult to interpret. This is because: * Older fossils are harder to find as they are usually buried at a considerable depth. * Dating 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 Deposition may refer to: * Deposition (law), taking testimony outside of court * Deposition (politics), the removal of a person of authority from political power * Deposition (university), a widespread initiation ritual for new students practiced f ...
process. * The preservation of fossils varies on land, but marine fossils tend to be better preserved than their sought after land-based counterparts. 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 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 in which a sample is collected in such a way that some members of the intended population have a lower or higher sampling probability than others. It results in a biased sample of a population (or non-human f ...
.


Sixth mass extinction

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


Extinctions by severity

Extinction events can be tracked by several methods, including geological change, ecological impact, extinction vs. origination ( speciation) rates, and most commonly diversity loss among taxonomic units. Most early papers used families 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, 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 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 at the end of the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
period. The
Alvarez hypothesis The Alvarez hypothesis posits that the mass extinction of the non-avian dinosaurs and many other living things during the Cretaceous–Paleogene extinction event was caused by the impact of a large asteroid on the Earth. Prior to 2013, it was ...
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 was published in the journal ''
Science Science is a systematic endeavor that builds and organizes knowledge in the form of testable explanations and predictions about the universe. Science may be as old as the human species, and some of the earliest archeological evidence ...
''. This paper, originating from a compendium of extinct marine animal families developed by Sepkoski, identified five peaks of marine family extinctions which 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. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya. The ...
( end-Ordovician), Late Permian, Norian ( end-Triassic), and
Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the inte ...
(end-Cretaceous). The remaining peak was a broad interval of high extinction smeared over the later half of the
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wh ...
, 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 th ...
stage. Through the 1980s, Raup and Sepkoski continued to elaborate and build upon their extinction and origination data, defining a high-resolution
biodiversity Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic ('' genetic variability''), species ('' species diversity''), and ecosystem ('' ecosystem diversity'') ...
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 astronomers linked this to a hypothetical
brown dwarf Brown dwarfs (also called failed stars) are substellar objects that are not massive enough to sustain nuclear fusion of ordinary hydrogen ( 1H) into helium in their cores, unlike a main-sequence star. Instead, they have a mass between the most ...
in the distant reaches of the solar system, inventing the “ Nemesis hypothesis” which has been strongly disputed by other astronomers. Around the same time, Sepkoski began to devise a compendium of marine animal genera, 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 which 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 which 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 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. Wallister (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 which 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

Sepkoski formally published his marine genera compendium in 2002, prompting 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 in the
Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ...
. 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) found that each of the “Big Five” extinction intervals had 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 which 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 which 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 biases. 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 which 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 relat ...
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 esti ...
”, 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 circumvene 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 a corporation is an individual or legal entity (such as another corporation, a body politic, a trust or partnership) that is registered by the corporation as the legal o ...
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''; plural 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 n ...
) 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” 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 which 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 which 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 mya. ...
and Campanian stages were each used to estimate diversity changes in the
Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the inte ...
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 which 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 plant-based material used as a fuel for heat or electricity production. It can be in the form of wood, wood residues, energy crops, agricultural residues, and waste from industry, farms, and households. Some people use the terms bio ...
on Earth is
microbial A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in olde ...
, 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 rather than the total diversity and abundance of life. For this reason, well-documented extinction events are confined to the
Phanerozoic eon The Phanerozoic Eon is the current geologic eon in the geologic time scale, and the one during which abundant animal and plant life has existed. It covers 538.8 million years to the present, and it began with the Cambrian Period, when anima ...
, before which all living organisms were either microbial or at most soft-bodied; the sole exception is the Great Oxidation Event in the Proterozoic. Perhaps due to the absence of a robust microbial fossil record, mass extinctions ''seem'' mainly to be a Phanerozoic phenomenon, with apparent extinction rates being low before large complex organisms arose.
Extinction Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the Endling, last individual of the species, although the Functional ext ...
occurs at an uneven rate. Based on the fossil record, the background rate of extinctions on Earth is about two to five taxonomic families of marine animals every million years. Marine fossils are mostly used to measure extinction rates because of their superior fossil record and stratigraphic range compared to land animals. The Great Oxidation Event, which occurred around 2.45 billion years ago in the Paleoproterozoic, was probably the first major extinction event. Since the Cambrian explosion, five further major mass extinctions have significantly exceeded the background extinction rate. The most recent and best-known, the
Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event (also known as the Cretaceous–Tertiary extinction) was a sudden mass extinction of three-quarters of the plant and animal species on Earth, approximately 66 million years ago. With the ...
, 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 minor ones as well, and the ongoing mass extinction caused by human activity is sometimes called the sixth extinction.


Evolutionary importance

Mass extinctions have sometimes accelerated the
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
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 in ...
. For example,
mammaliaformes Mammaliaformes ("mammalian forms") is a clade that contains the crown group mammals and their closest extinct relatives; the group radiated from earlier probainognathian cynodonts. It is defined as the clade originating from the most recent c ...
("almost mammals") and then
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur ...
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 period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
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. 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 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 which 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'': : "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 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, 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 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 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 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 provinces by flood basalt events could have: * produced dust and particulate aerosols, which inhibited photosynthesis and thus caused food chains to collapse both on land and at sea * emitted sulfur oxides that were precipitated as acid rain and poisoned many organisms, contributing further to the collapse of food chains * emitted
carbon dioxide Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is t ...
and thus possibly causing sustained global warming once the dust and particulate aerosols dissipated. Flood basalt events occur as pulses of activity punctuated by dormant periods. As a result, they are likely to cause the climate to oscillate between cooling and warming, but with an overall trend towards warming as the carbon dioxide they emit can stay in the atmosphere for hundreds of years. It is speculated that massive volcanism caused or contributed to the End-Permian, End-Triassic and End-Cretaceous extinctions. 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 The Phanerozoic Eon is the current geologic eon in the geologic time scale, and the one during which abundant animal and plant life has existed. It covers 538.8 million years to the present, and it began with the Cambrian Period, when anima ...
.


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 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, 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 chains to collapse both on land and at sea by producing dust and particulate aerosols and thus inhibiting photosynthesis. Impacts on
sulfur Sulfur (or sulphur in British English) is a chemical element with the symbol S and atomic number 16. It is abundant, multivalent and nonmetallic. Under normal conditions, sulfur atoms form cyclic octatomic molecules with a chemical formul ...
-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, 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 Cretaceous–Tertiary extinction) was a sudden mass extinction of three-quarters of the plant and animal species on Earth, approximately 66 million years ago. With the ...
. Nonetheless, in October 2019, researchers reported that the Cretaceous Chicxulub asteroid impact that resulted in the
extinction Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the Endling, last individual of the species, although the Functional ext ...
of non-avian
dinosaurs Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is the ...
66 Ma, also rapidly acidified the oceans, producing
ecological collapse Ecological collapse refers to a situation where an ecosystem suffers a drastic, possibly permanent, reduction in carrying capacity for all organisms, often resulting in mass extinction. Usually, an ecological collapse is precipitated by a disastr ...
and long-lasting effects on the climate, and was a key reason for end-Cretaceous mass extinction. 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 The Milky Way is the galaxy that includes our Solar System, with the name describing the galaxy's appearance from Earth: a hazy band of light seen in the night sky formed from stars that cannot be individually distinguished by the naked eye. ...
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 The ozone layer or ozone shield is a region of Earth's stratosphere that absorbs most of the Sun's ultraviolet radiation. It contains a high concentration of ozone (O3) in relation to other parts of the atmosphere, although still small in rel ...
, 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.


Global cooling

Sustained and significant global cooling could kill many
polar Polar may refer to: Geography Polar may refer to: * Geographical pole, either of two fixed points on the surface of a rotating body or planet, at 90 degrees from the equator, based on the axis around which a body rotates *Polar climate, the cli ...
and temperate species and force others to migrate towards the
equator The equator is a circle of latitude, about in circumference, that divides Earth into the Northern and Southern hemispheres. It is an imaginary line located at 0 degrees latitude, halfway between the North and South poles. The term can also ...
; reduce the area available for
tropical The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...
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 A glacial period (alternatively glacial or glaciation) is an interval of time (thousands of years) within an ice age that is marked by colder temperatures and glacier advances. Interglacials, on the other hand, are periods of warmer climate bet ...
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 The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...
species; kill temperate species or force them to migrate towards the
poles Poles,, ; singular masculine: ''Polak'', singular feminine: ''Polka'' or Polish people, are a West Slavic nation and ethnic group, who share a common history, culture, the Polish language and are identified with the country of Poland in ...
; 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 The water cycle, also known as the hydrologic cycle or the hydrological cycle, is a biogeochemical cycle that describes the continuous movement of water on, above and below the surface of the Earth. The mass of water on Earth remains fairly co ...
. 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 The Paleocene–Eocene thermal maximum (PETM), alternatively (ETM1), and formerly known as the "Initial Eocene" or "", was a time period with a more than 5–8 °C global average temperature rise across the event. This climate event o ...
, 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 The Permian–Triassic (P–T, P–Tr) extinction event, also known as the Latest Permian extinction event, the End-Permian Extinction and colloquially as the Great Dying, formed the boundary between the Permian and Triassic geologic periods, ...
has been suggested to have been caused by warming.


= Clathrate gun hypothesis

=
Clathrates A clathrate is a chemical substance consisting of a lattice that traps or contains molecules. The word ''clathrate'' is derived from the Latin (), meaning ‘with bars, latticed’. Most clathrate compounds are polymeric and completely envel ...
are composites in which a lattice of one substance forms a cage around another.
Methane clathrate Methane clathrate (CH4·5.75H2O) or (8CH4·46H2O), also called methane hydrate, hydromethane, methane ice, fire ice, natural gas hydrate, or gas hydrate, is a solid clathrate compound (more specifically, a clathrate hydrate) in which a large amou ...
s (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 In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to ...
or a sudden drop in sea level or even
earthquake An earthquake (also known as a quake, tremor or temblor) is the shaking of the surface of the Earth resulting from a sudden release of energy in the Earth's lithosphere that creates seismic waves. Earthquakes can range in intensity, fr ...
s. Methane is a much more powerful
greenhouse A greenhouse (also called a glasshouse, or, if with sufficient heating, a hothouse) is a structure with walls and roof made chiefly of transparent material, such as glass, in which plants requiring regulated climatic conditions are grown.These ...
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 The Paleocene–Eocene thermal maximum (PETM), alternatively (ETM1), and formerly known as the "Initial Eocene" or "", was a time period with a more than 5–8 °C global average temperature rise across the event. This climate event o ...
, 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 Lau or LAU may refer to: People * Lau (surname) * Liu (劉/刘), a common Chinese family name transliterated Lau in Cantonese and Hokkien * Lau clan, one of the Saraswat Brahmin clans of Punjab * LAU (musician): Laura Fares Places * Lebane ...
,
Toarcian The Toarcian is, in the ICS' geologic timescale, an age and stage in the Early or Lower Jurassic. It spans the time between 182.7 Ma (million years ago) and 174.1 Ma. It follows the Pliensbachian and is followed by the Aalenian. The Toar ...
and 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 In pharmacology, bioavailability is a subcategory of absorption and is the fraction (%) of an administered drug that reaches the systemic circulation. By definition, when a medication is administered intravenously, its bioavailability is 100%. H ...
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, 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 organisms. British
oceanologist Oceanography (), also known as oceanology and ocean science, is the scientific study of the oceans. It is an Earth science, which covers a wide range of topics, including ecosystem dynamics; ocean currents, waves, and geophysical fluid dynamic ...
and
atmospheric scientist Atmospheric science is the study of the Earth's atmosphere and its various inner-working physical processes. Meteorology includes atmospheric chemistry and atmospheric physics with a major focus on weather forecasting. Climatology is the study o ...
, Andrew Watson, explained that, while the 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 Permian–Triassic (P–T, P–Tr) extinction event, also known as the Latest Permian extinction event, the End-Permian Extinction and colloquially as the Great Dying, formed the boundary between the Permian and Triassic geologic periods, ...
the warming also upset the oceanic balance between
photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in ...
ing plankton and deep-water sulfate-reducing bacteria, causing massive emissions of
hydrogen sulfide Hydrogen sulfide is a chemical compound with the formula . It is a colorless chalcogen-hydride gas, and is poisonous, corrosive, and flammable, with trace amounts in ambient atmosphere having a characteristic foul odor of rotten eggs. The under ...
, which poisoned life on both land and sea and severely weakened the
ozone layer The ozone layer or ozone shield is a region of Earth's stratosphere that absorbs most of the Sun's ultraviolet radiation. It contains a high concentration of ozone (O3) in relation to other parts of the atmosphere, although still small in rel ...
, exposing much of the life that still remained to fatal levels of
UV radiation Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nm (with a corresponding frequency around 30  PHz) to 400 nm (750  THz), shorter than that of visible light, but longer than X-rays. UV radiatio ...
.


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 A glacial period (alternatively glacial or glaciation) is an interval of time (thousands of years) within an ice age that is marked by colder temperatures and glacier advances. Interglacials, on the other hand, are periods of warmer climate bet ...
, 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.


Geomagnetic reversal

One theory is that periods of increased
geomagnetic reversal A geomagnetic reversal is a change in a planet's magnetic field such that the positions of magnetic north and magnetic south are interchanged (not to be confused with geographic north and geographic south). The Earth's field has alternate ...
s will weaken
Earth's magnetic field Earth's magnetic field, also known as the geomagnetic field, is the magnetic field that extends from Earth's interior out into space, where it interacts with the solar wind, a stream of charged particles emanating from the Sun. The magneti ...
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 age An ice age is a long period of reduction in the temperature of Earth's surface and atmosphere, resulting in the presence or expansion of continental and polar ice sheets and alpine glaciers. Earth's climate alternates between ice ages and gre ...
s; 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 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 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 A continental shelf is a portion of a continent that is submerged under an area of relatively shallow water, known as a shelf sea. Much of these shelves were exposed by drops in sea level during glacial periods. The shelf surrounding an island ...
(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 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 that are contrary to the available evidence; they are based on other theories that 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 ''Proceedings of the National Academy of Sciences of the United States of America'' (often abbreviated ''PNAS'' or ''PNAS USA'') is a peer-reviewed multidisciplinary scientific journal. It is the official journal of the National Academy of S ...
'' 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.


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 () is a single-celled organism that lacks a nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Greek πρό (, 'before') and κάρυον (, 'nut' or 'kernel').Campbell, N. "Biology:Concepts & Con ...
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 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 or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic ('' genetic variability''), species ('' species diversity''), and ecosystem ('' ecosystem diversity'') ...
to recover after extinction events. In the most severe mass extinctions it may take 15 to 30 million years. The worst Phanerozoic 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 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.


See also

* Bioevent * Elvis taxon * Endangered species *
Geologic time scale The geologic time scale, or geological time scale, (GTS) is a representation of time based on the rock record of Earth. It is a system of chronological dating that uses chronostratigraphy (the process of relating strata to time) and geochr ...
* Global catastrophic risk * Holocene extinction *
Human extinction Human extinction, also known as omnicide, is the hypothetical end of the human species due to either natural causes such as population decline from sub-replacement fertility, an asteroid impact, or large-scale volcanism, or to anthropogenic ...
* Kačák Event * Lazarus taxon * List of impact craters on Earth * List of largest volcanic eruptions * List of possible impact structures on Earth *
Medea hypothesis The Medea hypothesis is a term coined by paleontologist Peter Ward for a hypothesis that contests the Gaian hypothesis and proposes that multicellular life, understood as a superorganism, may be self-destructive or suicidal. The metaphor refers ...
* Rare species * Signor–Lipps effect * Snowball Earth *
Speculative evolution Speculative evolution is a genre of speculative fiction and an artistic movement focused on hypothetical scenarios in the evolution of life, and a significant form of fictional biology. It is also known as speculative biology and it is referred ...
* ''The Sixth Extinction: An Unnatural History'' (nonfiction book) * Timeline of extinctions in the Holocene


Footnotes


References


Further reading

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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 * History of climate variability and change Evolutionary biology Meteorological hypotheses Natural disasters Terms in science and technology