An extinction event (also known as a mass extinction or biotic crisis) is a widespread and rapid decrease in the biodiversity on Earth. Such an event is identified by a sharp fall in the diversity and abundance of multicellular organisms. It occurs when the rate of extinction 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" of the Phanerozoic were anciently preceded by the presumed far more extensive mass extinction of microbial life during the Great Oxidation Event (also known as the Oxygen Catastrophe) early in the Proterozoic Eon. At the end of the Ediacaran and just before the Cambrian explosion, yet another Proterozoic extinction event (of unknown magnitude) is speculated to have ushered in the Phanerozoic. Several events in the Cambrian and early Ordovician meet or exceed the "Big Five" in proportional severity, though overall diversity was rather low until the Great Ordovician Biodiversification Event (GOBE). Sepkoski and Raup (1982) initially tracked absolute (rather than proportional) extinction, so their biodiversity estimates overlooked events prior to the GOBE.
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 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:
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 Fungus 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.
+ Extinction proportions (diversity loss) of marine genera or ecological impact in estimates of mass extinction severity !rowspan=2 | Extinction name !rowspan=2 | Age (Myr) !rowspan=2 | Sepkoski (1996) Multiple-interval genera !rowspan=2 | Bambach (2006) !colspan=2 | McGhee et al. (2013) !rowspan=2 | Stanley (2016) |
Late Ordovician (Ashgillian / Hirnantian) | 445–444 | ~49% | 57% (40%, 31%) | 52% | 7 | 42–46% |
Lau event (Ludfordian) | 424 | ~23% | – | 9% | 9 | – |
Kačák Event (Eifelian) | 388~ | ~24% | – | 32% | 9 | – |
Taghanic Event (Givetian) | 384~ | ~30% | 28.5% | 36% | 8 | – |
Late Devonian/Kellwasser event (Frasnian) | 372 | ~35% | 34.7% | 40% | 4 | 16–20% |
End-Devonian/Hangenberg event (Famennian) | 359 | ~28% | 31% | 50% | 7 | <13% |
Serpukhovian | 330–325~ | ~23% | 31% | 39% | 6 | 13–15% |
Capitanian | 260 | ~47% | 48% | 25% | 5 | 33–35% |
Permian–Triassic (Changhsingian) | 252 | ~58% | 55.7% | 83% | 1 | 62% |
Triassic–Jurassic (Rhaetian) | 201 | ~37% | 47% | 73% | 3 | N/A |
Pliensbachian-Toarcian | 186–178 | ~14% | 25%, 20% | – | – | – |
End-Jurassic (Tithonian) | 145 | ~18% | 20% | – | – | – |
Cenomanian-Turonian | 94 | ~15% | 25% | – | – | – |
Cretaceous–Paleogene (Maastrichtian) | 66 | ~39% | 40–47% | 40% | 2 | 38–40% |
Eocene–Oligocene | 34 | ~11% | 15.6% | – | – | – |
Graphed but not discussed by , 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
Through the 1980s, Raup and Sepkoski continued to elaborate and build upon their extinction and origination data, defining a high-resolution biodiversity curve (the "Sepkoski curve") and successive evolutionary faunas with their own patterns of diversification and extinction.
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 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 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 Anthony 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.
Computer models run by 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, 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.
John Alroy (2010) attempted to circumvent sample size-related biases in diversity estimates using a method he called "shareholder 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) 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 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 and Campanian stages were each used to estimate diversity changes in the Maastrichtian 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.
Extinction occurs at an uneven rate. Based on the fossil record, the background rate of extinctions on Earth is about two to five taxonomic families of every million years.
The Oxygen Catastrophe, which occurred around 2.45 billion years ago in the Paleoproterozoic, 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 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, 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.
For example, mammaliaformes ("almost mammals") and then existed throughout the reign of the , 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. Similarly, within Synapsida, the replacement of taxa that originated in the earliest, Pennsylvanian and Cisuralian evolutionary radiation (often still called "", though this is a paraphyletic group) by Therapsida occurred around the Kungurian/Roadian transition, which is often called Olson's extinction (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 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".
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:
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 either more or less probable.
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.
The most commonly suggested causes of mass extinctions are listed below.
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 End-Guadalupian Extinction Event, the End-Permian Extinction Event, the Smithian-Spathian Extinction, the Triassic-Jurassic Extinction Event, the Toarcian Oceanic Anoxic Event, the Cenomanian-Turonian Oceanic Anoxic Event, 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 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.
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. Nonetheless, in October 2019, researchers reported that the Cretaceous Chicxulub asteroid impact that resulted in the extinction of non-avian dinosaurs 66 Ma, also rapidly 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.
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 as a cause of mass extinction is supported by several recent studies.
The most dramatic example of sustained warming is the Paleocene–Eocene Thermal Maximum, which was associated with one of the smaller mass extinctions. It has also been suggested to have caused the Triassic–Jurassic extinction event, during which 20% of all marine families became extinct. Furthermore, the Permian–Triassic extinction event has been suggested to have been caused by warming.
The most likely signature of such a methane eruption would be a sudden decrease in the Isotope analysis in sediments, since methane clathrates are low in carbon-13; but the change would have to be very large, as other events can also reduce the percentage of carbon-13.
It has been suggested that "clathrate gun" methane eruptions were involved in the end-Permian extinction ("the Great Dying") and in the Paleocene–Eocene Thermal Maximum, which was associated with one of the smaller mass extinctions.
It has been suggested that anoxic events caused or contributed to the Ordovician–Silurian, late Devonian, Capitanian, Permian–Triassic, and Triassic–Jurassic extinctions, as well as a number of lesser extinctions (such as the Ireviken event, Lundgreni Event, Mulde event, Lau event, Smithian-Spathian, Toarcian, 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 of essential (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 organisms.
British oceanologist and atmospheric scientist Andrew Watson explained that, while the Holocene exhibits many processes reminiscent of those that have contributed to past anoxic events, full-scale ocean anoxia would take "thousands of years to develop".
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.
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.
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 . 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).
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 pioneer organism, 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;
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.
New data on genera: Sepkoski's compendium
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.
Tackling biases in the fossil record
Uncertainty in the Proterozoic and earlier eons
Evolutionary importance
Patterns in frequency
Causes
Identifying causes of specific mass extinctions
Most widely supported explanations
Flood basalt events
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.
Sea-level fall
Extraterrestrial threats
Impact events
A nearby nova, supernova or gamma ray burst
Global cooling
Global warming
Clathrate gun hypothesis
Anoxic events
Hydrogen sulfide emissions from the seas
Oceanic overturn
Geomagnetic reversal
Plate tectonics
Human activities
Other hypotheses
Future biosphere extinction/sterilization
Effects and recovery
In media
See also
Footnotes
Further reading
External links
|
|