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Extinction event

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Apparent extinction intensity, i.e. the fraction of genera going extinct at any given time, as reconstructed from the fossil record. (Graph not meant to include recent epoch of Holocene extinction)

An extinction event (also known as: mass extinction; extinction-level event, ELE, or biotic crisis) is a sharp decrease in the diversity and abundance of macroscopic life. They occur when the rate of extinction increases with respect to the rate of speciation. Because the majority of diversity and biomass on earth is microbial, and thus difficult to measure, mass extinctions have little effect on the total diversity and abundance of life, but rather affect the easily observed component of the biosphere.[1]

Over 97% of species that ever lived are now extinct, but extinction occurs at an uneven rate. Based on the fossil record, the background rate of extinctions on Earth is about two to five taxonomic families of marine invertebrates and vertebrates every million years. Marine fossils are mostly used to measure extinction rates because of their superior fossil record and stratigraphic range compared to land organisms.

Since life began on Earth, several major mass extinctions have significantly exceeded the background extinction rate. The most recent, the Cretaceous–Tertiary extinction event, occurred 65 million years ago, and has attracted more attention than all others as it marks the extinction of nearly all dinosaur species, which were the dominant animal class of the period. In the past 540 million years there have been five major events when over 50% of animal species died. There probably were mass extinctions in the Archean and Proterozoic Eons, but before the Phanerozoic there were no animals with hard body parts to leave a significant fossil record.

Estimates of the number of major mass extinctions in the last 540 million years range from as few as five to more than twenty. These differences stem from the threshold chosen for describing an extinction event as "major", and the data chosen to measure past diversity.

Major extinction events

The classical "Big Five" mass extinctions identified by Jack Sepkoski and David M. Raup in their 1982 paper are widely agreed upon as some of the most significant: End Ordovician, Late Devonian, End Permian, End Triassic, and End Cretaceous.[2][3]

These and a selection of other extinction events are outlined below. The articles about individual mass extinctions describe their effects in more detail and discuss theories about their causes.

  1. Cretaceous–Tertiary extinction event (K-T extinction) - 65 Ma ago at the Cretaceous-Paleogene transition. About 17% of all families, 50% of all genera and 75% of species went extinct.[4] It ended the reign of dinosaurs and opened the way for mammals and birds to become the dominant land vertebrates. In the seas it reduced the percentage of sessile animals to about 33%. The K-T extinction was rather uneven — some groups of organisms became extinct, some suffered heavy losses and some appear to have been only minimally affected.
  2. Triassic–Jurassic extinction event - 205 Ma at the Triassic-Jurassic transition. About 23% of all families and 48% of all genera (20% of marine families and 55% of marine genera) went extinct.[4] Most non-dinosaurian archosaurs, most therapsids, and most of the large amphibians were eliminated, leaving dinosaurs with little terrestrial competition. Non-dinosaurian archosaurs continued to dominate aquatic environments, while non-archosaurian diapsids continued to dominate marine environments. The Temnospondyl lineage of large amphibians also survived until the Cretaceous in Australia (e.g., Koolasuchus).
  3. Permian–Triassic extinction event - 251 Ma at the Permian-Triassic transition. Earth's largest extinction killed 57% of all families and 83% of all genera (53% of marine families, 84% of marine genera, about 96% of all marine species and an estimated 70% of land species) including vertebrates, insects and plants.[4] The "Great Dying" had enormous evolutionary significance: on land, it ended the primacy of mammal-like reptiles. The recovery of vertebrates took 30 million years,[5] but the vacant niches created the opportunity for archosaurs 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 for at least marine life, even before the "Great Dying".
  4. Late Devonian extinction 360-375 Ma near the Devonian-Carboniferous transition. At the end of the Frasnian Age in the later part(s) of the Devonian Period, a prolonged series of extinctions eliminated about 19% of all families, 50% of all genera and 70% of all species.[4] This extinction event lasted perhaps as long as 20 MY, and there is evidence for a series of extinction pulses within this period.
  5. Ordovician–Silurian extinction event 440-450 Ma at the Ordovician-Silurian transition. Two events occurred that killed off 27% of all families and 57% of all genera.[4] 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 went extinct.

The older the fossil record gets, the more difficult it is to read it. This is because:

  • Older fossils are harder to find because they are usually buried at a considerable depth in the rock.
  • Dating older fossils is more difficult.
  • Productive fossil beds are researched more than unproductive ones, therefore leaving certain periods unresearched.
  • Prehistoric environmental disturbances can disturb the deposition process.
  • The preservation of fossils varies on land, but marine fossils tend to be better preserved than their sought after land-based counterparts.[6]

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.[7] However, statistical analysis shows that this can only account for 50% of the observed pattern,[citation needed] and other evidence (such as fungal spikes)[clarification needed] provides reassurance that most widely accepted extinction events are indeed real. A quantification of the rock exposure of Western Europe does indicate that many of the minor events for which a biological explanation has been sought are most readily explained by sampling bias.[8]

Lesser extinctions

Lesser extinction events include:[9]

Precambrian

Cambrian Period

Silurian Period

Carboniferous Period

Jurassic Period

Cretaceous Period

Paleogene Period

Neogene Period

Evolutionary importance

Mass extinctions have sometimes accelerated the evolution of life on Earth. When dominance of particular ecological niches passes from one group of organisms to another, it is rarely because the new dominant group is "superior" to the old and usually because an extinction event eliminates the old dominant group and makes way for the new one.[10][11]

For example mammaliformes ("almost mammals") and then mammals existed throughout the reign of the dinosaurs, but could not compete for the large terrestrial vertebrate niches which dinosaurs monopolized. The end-Cretaceous mass extinction removed the non-avian dinosaurs and made it possible for mammals to expand into the large terrestrial vertebrate niches. Ironically, the dinosaurs themselves had been beneficiaries of a previous mass extinction, the end-Triassic, which eliminated most of their chief rivals, the crurotarsans.

Another point of view put forward in the Escalation hypothesis predicts that species in ecological niches with more organism-to-organism conflict will be less likely to survive extinctions. This is because the very traits that keep a species numerous and viable under fairly static conditions become a burden once population levels fall among competing organisms during the dynamics of an extinction event.

Furthermore, many groups which survive mass extinctions do not recover in numbers or diversity, and many of these go into long-term decline, and these are often referred to as "Dead Clades Walking".[12] So analysing extinctions in terms of "what died and what survived" often fails to tell the full story.

Apparent decreasing frequency

All genera
"Well-defined" genera
Trend line
"Big Five" mass extinctions
Other mass extinctions
Million years ago
Thousands of genera
Phanerozoic biodiversity as shown by the fossil record

The gaps between mass extinctions appear to be becoming longer, while the average and background rates of extinction are decreasing. Mass extinctions are thought to result when a long-term stress is compounded by a short term shock.[13] Over the course of the Phanerozoic, individual taxa appear to be less likely to become extinct at any time,[14] which may reflect more robust food webs as well as less extinction-prone species and other factors such as continental distribution.[14] However the taxonomic susceptibility to extinction does not appear to make mass extinctions more or less probable.[14]

The idea that mass extinctions are becoming less frequent is rather speculative – from a statistical point of view a sample of about 10 extinction events is too small to be a reliable sign of any actual trend.

Both of these phenomena could be explained in one or more ways:[15]

  • Reasonably complete fossils are very rare, most extinct organisms are represented only by partial fossils, and complete fossils are rarest in the oldest rocks. So paleontologists have mistakenly assigned parts of the same organism to different genera which were often defined solely to accommodate these finds (an example is the story of Anomalocaris). The risk of this mistake is higher for older fossils because these are often unlike parts of any living organism. Many of the "superfluous" genera are represented by fragments which are not found again and the "superfluous" genera appear to become extinct very quickly.
  • Martin (1994, 1996) has argued that the oceans have become more hospitable to life over the last 500M years and less vulnerable to mass extinctions: dissolved oxygen became more widespread and penetrated to greater depths; the development of life on land reduced the run-off of nutrients and hence the risk of eutrophication and anoxic events; and marine ecosystems became more diversified so that food chains were less likely to be disrupted.[16][17]

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.[18]

Looking for the causes of particular mass extinctions

A good theory for a particular mass extinction should: (i) explain all of the losses, not just focus on a few groups (such as dinosaurs); (ii) explain why particular groups of organisms died out and why others survived; (iii) provide mechanisms which are strong enough to cause a mass extinction but not a total extinction; (iv) be based on events or processes that can be shown to have happened, not just inferred from the extinction.

It may be necessary to consider combinations of causes. For example the marine aspect of the end-Cretaceous extinction appears to have been caused by several processes which partially overlapped in time and may have had different levels of significance in different parts of the world.[19]

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.[20] 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)[21] summarized the relationship between mass extinctions and events which are most often cited as causes of mass extinctions, using data from Courtillot et al. (1996),[22] Hallam (1992)[23] and Grieve et al. (1996):[24]

  • Flood basalt events: 11 occurrences, all associated with significant extinctions[25][26] But Wignall (2001) concluded that only 5 of the major extinctions coincided with flood basalt eruptions and that the main phase of extinctions started before the eruptions.[27]
  • Sea-level falls: 12, of which 7 were associated with significant extinctions.[26]
  • Asteroid impacts producing craters over 100 km wide: one, associated with one mass extinction.
  • Asteroid impacts producing craters less than 100 km wide: over 50, the great majority not associated with significant extinctions.

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:

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-Cretaceous, End-Permian, and End Triassic extinctions.[28][29][30]

Sea-level falls

These are often clearly marked by worldwide sequences of contemporaneous sediments which show all or part of a transition from sea-bed to tidal zone to beach to dry land - and where there is no evidence that the rocks in the relevant areas were raised by geological processes such as orogeny. Sea-level falls could reduce the continental shelf area (the most productive part of the oceans) sufficiently to cause a marine mass extinction, and could disrupt weather patterns enough to cause extinctions on land. But sea-level falls are very probably the result of other events, such as sustained global cooling or the sinking of the mid-ocean ridges.

Sea-level falls are associated with most of the mass extinctions, including all of the "Big Five" — End-Ordovician, Late Devonian, End-Permian, End-Triassic, and End-Cretaceous.

A study, published in the journal Nature (online June 15, 2008) established a relationship between the speed of mass extinction events and changes in sea level and sediment.[31] 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.[32]

Impact events

The impact of a sufficiently large asteroid or comet could have caused food chains to collapse both on land and at sea by producing dust and particulate aerosols and thus inhibiting photosynthesis. Impacts on sulfur-rich rocks could have emitted sulfur oxides precipitating as poisonous acid rain, contributing further to the collapse of food chains. Such impacts could also have caused megatsunamis and / or global forest fires.

Most paleontologists now agree that an asteroid did hit the Earth about 65 Ma, but there is an ongoing dispute whether the impact was the sole cause of the extinctions.[33][34] There is evidence that there was an interval of about 300 ka from the impact to the mass extinction.[33] In 1997, paleontologist Sankar Chatterjee drew attention to the proposed and much larger 600 km (370 mi) Shiva crater and the possibility of a multiple-impact scenario.

In 2007, a hypothesis was put forth that argued the impactor that killed the dinosaurs 65 Ma years ago belonged to the Baptistina family of asteroids.[35] Concerns have been raised regarding the reputed link, in part because very few solid observational constraints exist of the asteroid or family.[36] Indeed, it was recently discovered that 298 Baptistina does not share the same chemical signature as the source of the K-T impact.[37] Although this finding may make the link between the Baptistina family and K-T impactor more difficult to substantiate, it does not preclude the possibility.[37]

In 2010, another hypothesis was offered which implicated the newly-discovered asteroid P/2010 A2, a member of the Flora family of asteroids, as a possible remnant cohort of the K/T impactor. [38]

Sustained and significant global cooling

Sustained global cooling could kill many polar and temperate species and force others to migrate towards the equator; reduce the area available for tropical species; often make the Earth's climate more arid on average, mainly by locking up more of the planet's water in ice and snow. The glaciation cycles of the current ice age are believed to have had only a very mild impact on biodiversity, so the mere existence of a significant cooling is not sufficient on its own to explain a mass extinction.

It has been suggested that global cooling caused or contributed to the End-Ordovician, Permian-Triassic, Late Devonian extinctions, and possibly others. Sustained global cooling is distinguished from the temporary climatic effects of flood basalt events or impacts.

Sustained and significant global warming

This would have the opposite effects: expand the area available for tropical species; kill temperate species or force them to migrate towards the poles; possibly cause severe extinctions of polar species; often make the Earth's climate wetter on average, mainly by melting ice and snow and thus increasing the volume of the water cycle. It might also cause anoxic events in the oceans (see below).

Global warming as a cause of mass extinction is supported by several recent studies.[39]

The most dramatic example of sustained warming is the Paleocene-Eocene Thermal Maximum, which was associated with one of the smaller mass extinctions. It has also been suggested to have caused the Triassic-Jurassic extinction event, during which 20% of all marine families went extinct. Furthermore, the Permian–Triassic extinction event has been suggested to have been caused by warming.[40][41][42]

Clathrate gun hypothesis

Main article: Clathrate gun hypothesis

Clathrates are composites in which a lattice of one substance forms a cage round another. Methane clathrates (in which water molecules are the cage) form on continental shelves. These clathrates are likely to break up rapidly and release the methane if the temperature rises quickly or the pressure on them drops quickly — for example in response to sudden global warming or a sudden drop in sea level or even earthquakes. Methane is a much more powerful greenhouse gas than carbon dioxide, so a methane eruption ("clathrate gun") could cause rapid global warming or make it much more severe if the eruption was itself caused by global warming.

The most likely signature of such a methane eruption would be a sudden decrease in the ratio of carbon-13 to carbon-12 in sediments, since methane clathrates are low in carbon-13; but the change would have to be very large, as other events can also reduce the percentage of carbon-13.[43]

It has been suggested that "clathrate gun" methane eruptions were involved in the end-Permian extinction ("the Great Dying") and in the Paleocene-Eocene Thermal Maximum, which was associated with one of the smaller mass extinctions.

Anoxic events

Anoxic events are situations in which the middle and even the upper layers of the ocean become deficient or totally lacking in oxygen. Their causes are complex and controversial, but all known instances are associated with severe and sustained global warming, mostly caused by sustained massive volcanism.

It has been suggested that anoxic events caused or contributed to the Ordovician–Silurian, late Devonian, Permian-Triassic and Triassic-Jurassic extinctions, as well as a number of lesser extinctions (such as the Ireviken, Mulde, Lau, Toarcian and Cenomanian-Turonian events). On the other hand, there are widespread black shale beds from the mid-Cretaceous which indicate anoxic events but are not associated with mass extinctions.

Hydrogen sulfide emissions from the seas

Kump, Pavlov and Arthur (2005) have proposed that during the Permian-Triassic extinction event the warming also upset the oceanic balance between photosynthesising plankton and deep-water sulfate-reducing bacteria, causing massive emissions of hydrogen sulfide which poisoned life on both land and sea and severely weakened the ozone layer, exposing much of the life that still remained to fatal levels of UV radiation.[44][45][46]

Oceanic overturn

Oceanic overturn is a disruption of thermo-haline circulation which lets surface water (which is more saline than deep water because of evaporation) sink straight down, bringing anoxic deep water to the surface and therefore killing most of the oxygen-breathing organisms which inhabit the surface and middle depths. It may occur either at the beginning or the end of a glaciation, although an overturn at the start of a glaciation is more dangerous because the preceding warm period will have created a larger volume of anoxic water.[47]

Unlike other oceanic catastrophes such as regressions (sea-level falls) and anoxic events, overturns do not leave easily-identified "signatures" in rocks and are theoretical consequences of researchers' conclusions about other climatic and marine events.

It has been suggested that oceanic overturn caused or contributed to the late Devonian and Permian-Triassic extinctions.

A nearby nova, supernova or gamma ray burst

A nearby gamma ray burst (less than 6000 light years away) could sufficiently irradiate the surface of Earth to kill organisms living there and destroy the ozone layer in the process. From statistical arguments, approximately 1 gamma ray burst would be expected to occur close to Earth in the last 540 million years. A proposal that a supernova or gamma ray burst had caused a mass extinction would also have to be backed up by astronomical evidence of such an explosion at the right place and time.

It has been suggested that a supernova or gamma ray burst caused the End-Ordovician extinction.

Continental drift

Movement of the continents into some configurations can cause or contribute to extinctions in several ways: by initiating or ending ice ages; by changing ocean and wind currents and thus altering climate; by opening seaways or land bridges which expose previously isolated species to competition for which they are poorly-adapted (for example, the extinction of most of South America's native ungulates and all of its large metatherians after the creation of a land bridge between North and South America). Occasionally continental drift creates a super-continent which includes the vast majority of Earth's land area, which in addition to the effects listed above is likely to reduce the total area of continental shelf (the most species-rich part of the ocean) and produce a vast, arid continental interior which may have extreme seasonal variations.

It is widely thought that the creation of the super-continent Pangaea contributed to the End-Permian mass extinction. Pangaea was almost fully formed at the transition from mid-Permian to late-Permian, and the "Marine genus diversity" diagram at the top of this article shows a level of extinction starting at that time which might have qualified for inclusion in the "Big Five" if it were not overshadowed by the "Great Dying" at the end of the Permian.

Plate tectonics

Plate tectonics is the mechanism which drives many of the possible causes of mass extinctions, especially volcanism and continental drift. So it is implicated in many extinctions, but in each case it is necessary to specify which manifestations of plate tectonics were involved.

Other hypotheses

Many other hypotheses have been proposed, such as the spread of a new disease, the Shiva hypothesis, or simple out-competition following an especially successful biological innovation. But all have been rejected, usually for one of the following reasons: they require events or processes for which there is no evidence; they assume mechanisms which are contrary to the available evidence; they are based on other theories which have been rejected or superseded.

Postulated extinction cycles

It has been suggested by several sources that biodiversity and/or extinction events may be influenced by cyclic processes. The best-known hypothesis of extinction events by a cyclic process is the 26M to 30M year cycle in extinctions proposed by Raup and Sepkoski (1986).[48] More recently, Rohde and Muller (2005) have suggested that biodiversity fluctuates primarily on 62 ± 3 million year cycles.[49]

It is difficult to evaluate the validity of such claims except through reduction to statistical arguments about how plausible or implausible it is for the observed data to exhibit a particular pattern, as the causes of most extinction events are still too uncertain to attribute to them any specific cause let alone a recurring one. Much early work in this area also suffered from the poor accuracy of geological dating, where errors often exceed 10M years. However, improvements in radiometric dating have reduced the scale of uncertainty to at most 4M years — theoretically adequate for studying these processes.[verification needed]

While the statistics alone have been judged as sufficiently compelling to warrant publication, it is important to consider processes that might be responsible for a cyclic pattern of extinctions and future work may focus on trying to find evidence of such processes.

Hypothetical companion star to the sun

The physicist Richard A. Muller has produced a number of speculative hypotheses for the regularity of mass extinctions. One is that the extinction cycle could be caused by the orbit of a hypothetical companion star dubbed Nemesis that periodically disturbs the Oort cloud, sending storms of large asteroids and comets towards the Solar System.[50]

Galactic plane oscillations

Muller has also speculated the periodicity of mass extinctions may be related to the solar system's oscillation through the plane of our Milky Way galaxy as it rotates around the galactic centre, with a number of possible hypothesized effects including gravitationally-induced comet showers or periods of intense radiation as the solar system hits the galactic shock wave.[51][52]

Passage through galactic spiral arms

It has also been suggested that extinction events correlate to the passage of the solar system through the spiral arms of the Milky Way. The Earth passes through all four arms every 700 million years, and there is some evidence to suggest a cyclicity of extraterrestrial activity back to 2 billion years ago.[53]

Geological instabilities

Other hypotheses are that geological instabilities allow heat to periodically build up deep in the Earth, which is then released through mantle plumes, periods of major volcanism and active plate tectonics.[verification needed]

Effects

The impact of mass extinction events varied widely. The worst event, the Permian-Triassic Extinction Event, devastated life on earth and is estimated to have killed off over 90% of species. Life on Earth seemed to recover quickly after this extinction, but this was mostly in the form of disaster taxa, such as the hardy Lystrosaurus. The most recent research indicates that the specialized animals that formed complex ecosystems, with high biodiversity, complex food webs and a variety of niches, took much longer to recover. It is thought that this long recovery was due to the successive waves of extinction which inhibited recovery, as well as to prolonged environmental stress to organisms which continued into the Early Triassic. Recent research indicates that recovery did not begin until the start of the mid-Triassic, 4M to 6M years after the extinction;[54] and some writers estimate that the recovery was not complete until 30M years after the P-Tr extinction, i.e. in the late Triassic.[55]

See also

References

Bibliography

  • Cowen, R. (1999). "The History of Life". Blackwell Science. The chapter about extinctions is reproduced at [1]
  • Richard Leakey and Roger Lewin, 1996, The Sixth Extinction : Patterns of Life and the Future of Humankind, Anchor, ISBN 0-385-46809-1. Excerpt from this book: The Sixth Extinction
  • Wilson, E.O., 2002, The Future of Life, Vintage (pb), ISBN 0-679-76811-4
  • Raup, D., and J. Sepkoski (1986). "Periodic extinction of families and genera". Science. 231 (4740): 833–836. doi:10.1126/science.11542060. PMID 11542060.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  • Rohde, R.A. & Muller, R.A. (2005). "Cycles in fossil diversity". Nature. 434 (7030): 209–210. doi:10.1038/nature03339.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  • The Current Mass Extinction Event
  • Nemesis — Raup and Sepkoski
  • Richard A. Muller, 1988, Nemesis, Weidenfeld & Nicolson, ISBN 1-55584-173-2
  • Robert J. Sawyer, 2000, Calculating God, TOR, ISBN 0-812-58035-4
  • Ward, P.D., (2000) Rivers In Time: The Search for Clues to Earth's Mass Extinctions
  • Ward, P.D., (2007) Under a Green Sky: Global Warming, the Mass Extinctions of the Past, and What They Can Tell Us About Our Future (2007) ISBN 9780061137921 0061137928
  • Phil Berardelli, Of Cosmic Rays and Dangerous Days at ScienceNOW, August 1, 2007.
  • Stojanowski, J.S., 2008, The Gravity Theory Of Mass Extinction: A new unified theory of mass extinction explains the rise and fall of the dinosaurs, ISBN 978-0-9819221-0-2

Notes

  1. ^ Attention: This template ({{cite doi}}) is deprecated. To cite the publication identified by doi:10.1371/journal.pbio.0020272, please use {{cite journal}} (if it was published in a bona fide academic journal, otherwise {{cite report}} with |doi=10.1371/journal.pbio.0020272 instead.
  2. ^ Raup, D. & Sepkoski, J. (1982). "Mass extinctions in the marine fossil record". Science. 215: 1501–1503. doi:10.1126/science.215.4539.1501. PMID 17788674.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  3. ^ Morell, V., and Lanting, F., 1999. "The Sixth Extintion," National Geographic Magazine, February.
  4. ^ a b c d e "extinction". Math.ucr.edu. Retrieved 2008-11-09.
  5. ^ Sahney S & Benton MJ (2008). "Recovery from the most profound mass extinction of all time". Proceedings of the Royal Society: Biological. 275 (759).
  6. ^ Sole, R. V., and Newman, M., 2002. "Extinctions and Biodiversity in the Fossil Record - Volume Two, The Earth system: biological and ecological dimensions of global environment change" pp. 297-391, Encyclopedia of Global Enviromental Change John Wilely & Sons.
  7. ^ Smith, A. (2005). "Cyclicity in the fossil record mirrors rock outcrop area". Biology Letters. 1 (4): 443–445. doi:10.1098/rsbl.2005.0345. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  8. ^ ANDREW B. SMITH, ALISTAIR J. McGOWAN (2007). "The Shape Of The Phanerozoic Marine Palaeodiversity Curve: How Much Can Be Predicted From The Sedimentary Rock Record Of Western Europe?". Palaeontology. 50 (4): 765–774. doi:10.1111/j.1475-4983.2007.00693.x. {{cite journal}}: Unknown parameter |jorunal= ignored (help); Unknown parameter |month= ignored (help)
  9. ^ Partial list from Image:Extinction Intensity.png
  10. ^ Benton, M.J. (2004). "6. Reptiles Of The Triassic". Vertebrate Palaeontology. Blackwell.
  11. ^ Van Valkenburgh, B. (1999). "Major patterns in the history of carnivorous mammals". Annual Review of Earth and Planetary Sciences. 26: 463–493. doi:10.1146/annurev.earth.27.1.463.
  12. ^ Jablonski, D. (2002). "Survival without recovery after mass extinctions". PNAS. 99 (12): 8139–8144. doi:10.1073/pnas.102163299. PMID 12060760.
  13. ^ Arens, Nan Crystal (2008). "Press-pulse: a general theory of mass extinction?". Paleobiology. 34: 456. doi:10.1666/07034.1.
  14. ^ a b c Wang, Steve C. (2008). "Adjusting global extinction rates to account for taxonomic susceptibility". Paleobiology. 34: 434. doi:10.1666/07060.1.
  15. ^ MacLeod, Norman (2001-01-06). "Extinction!".
  16. ^ Martin, R.E. (1995). "Cyclic and secular variation in microfossil biomineralization: clues to the biogeochemical evolution of Phanerozoic oceans". Global and Planetary Change. 11 (1): 1. doi:10.1016/0921-8181(94)00011-2.
  17. ^ Martin, R.E. (1996). "Secular increase in nutrient levels through the Phanerozoic: Implications for productivity, biomass, and diversity of the marine biosphere". Palaios. 11: 209–219. doi:10.2307/3515230.
  18. ^ Arens, Nan Crystal (2008). "Press-pulse: a general theory of mass extinction?". Paleobiology. 34: 456. doi:10.1666/07034.1.
  19. ^ Marshall, C.R.; Ward, P.D. (1996). "Sudden and Gradual Molluscan Extinctions in the Latest Cretaceous of Western European Tethys". Science. 274: 1360–1363. doi:10.1126/science.274.5291.1360.
  20. ^ Arens, N.C. and West, I.D. (2006). "Press/Pulse: A General Theory of Mass Extinction?"" 'GSA Conference paper' Abstract
  21. ^ MacLeod, N (2001-01-06). "Extinction!".
  22. ^ Courtillot, V., Jaeger, J-J., Yang, Z., Féraud, G., Hofmann, C. (1996). "The influence of continental flood basalts on mass extinctions: where do we stand?" in Ryder, G., Fastovsky, D., and Gartner, S, eds. "The Cretaceous-Tertiary event and other catastrophes in earth history". The Geological Society of America, Special Paper 307, 513-525.
  23. ^ Hallam, A. (1992). "Phanerozoic sea-level changes". New York; Columbia University Press.
  24. ^ Grieve, R., Rupert, J., Smith, J., Therriault, A. (1996). "The record of terrestrial impact cratering". GSA Today 5: 193-195
  25. ^ The earliest known flood basalt event is the one which produced the Siberian Traps and is associated with the end-Permian extinction.
  26. ^ a b Some of the extinctions associated with flood basalts and sea-level falls were significantly smaller than the "major" extinctions, but still much greater than the background extinction level.
  27. ^ Wignall, P.B. (2001), "Large igneous provinces and mass extinctions", Earth-Science Reviews vol. 53 issues 1-2 pp 1-33
  28. ^ Speculated Causes of the End-Cretaceous Extinction
  29. ^ What was the Permian–Triassic Extinction Event?
  30. ^ What is the Triassic-Jurassic Extinction Event?
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