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70-589: The greatest mass extinction in the Phanerozoic eon , the Permian–Triassic extinction event , occurred around the end of this age. The Changhsingian is named after Changxing ( Chinese : 长兴 ; pinyin : Chángxīng ; Wade–Giles : Ch’ang-hsing ) in northern Zhejiang , China. The stage was named for the Changhsing Limestone . The name was first used for a stage in 1970 and

140-539: 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

210-531: A paraphyletic group) by therapsids 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

280-401: 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

350-413: A "major" extinction event, and the data chosen to measure past diversity. 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

420-425: A Phanerozoic phenomenon, with merely the observable extinction rates appearing low before large complex organisms with hard body parts arose. Extinction occurs at an uneven rate. Based on the fossil record , the background rate of extinctions on Earth is about two to five taxonomic families of marine animals every million years. The Oxygen Catastrophe, which occurred around 2.45 billion years ago in

490-664: A backdrop of decreasing extinction rates through time. Four of these peaks were statistically significant: the Ashgillian ( end-Ordovician ), Late Permian , Norian ( end-Triassic ), and Maastrichtian (end-Cretaceous). The remaining peak was a broad interval of high extinction smeared over the later half of the Devonian , with its apex in the Frasnian stage. Through the 1980s, Raup and Sepkoski continued to elaborate and build upon their extinction and origination data, defining

560-409: 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 ". 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

630-424: A high-resolution biodiversity 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

700-425: A hypothetical brown dwarf 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

770-470: A lack of consensus on Late Triassic chronology 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

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840-440: 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

910-399: 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 ) parallel to extinction rates in the context of geological stages or substages. A review and re-analysis of Sepkoski's data by Bambach (2006) identified 18 distinct mass extinction intervals, including 4 large extinctions in

980-813: 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

1050-402: 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. Clarkina ? Neogondolella Clarkina is an extinct genus of conodonts . It is considered to be an offshore, outer shelf or basinal, deep-water taxon. The genus name

1120-628: A separate event from the P–T extinction; if so, it would be larger than some of the "Big Five" extinction events.   The End Cretaceous extinction, or the K–Pg extinction (formerly K–T extinction) occurred at the Cretaceous ( Maastrichtian ) – Paleogene ( Danian ) transition. The event was formerly called the Cretaceous-Tertiary or K–T extinction or K–T boundary; it is now officially named

1190-508: 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

1260-586: 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)

1330-544: Is a tribute to David Leigh Clark . Clarkina Soodan 1975 is a junior synonym for the prehistoric sea cucumber Soodanella Huddleston 1982 in the family Priscopedatidae . Clarkina Jordan & Evermann, 1927 is a junior synonym for the cyprinid fish genus Mylocheilus Agassiz, 1855. The top of the Capitanian (or the base of the Wuchiapingian ) is defined as the place in the stratigraphic record where

1400-401: Is also the largest known extinction event for insects . The highly successful marine arthropod, the trilobite , became extinct. The evidence regarding plants 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

1470-447: 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

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1540-648: Is known from Zhejiang province of China, along with the elongate saurichthyiform Eosaurichthys and the coelacanths Changxingia and Youngichthys . Within the Eugeneodontida , the helicoprionids are represented by the genus Sinohelicoprion ; as well as some edestids such as Helicampodus ; and other eugeneodontids. Several fish genera were described from Changhsingian deposits of Russia and South Africa. The Hambast Formation of Iran yielded chondrichthyan faunas of Wuchiapingian to Changhsingian age. The conodont Vjalovognathus carinatus

1610-610: Is known from the Selong Formation of Tibet; more common conodonts include the genera Clarkina and Hindeodus . Changhsingian aged beds of the Tesero Member of the Werfen Formation produced fossils of a crown group echinoid , Eotiaris teseroensis and other taxa . The Paratirolites Limestone near Julfa ( Azerbaijan , Iran ) contains a diverse pre-extinction ammonoid fauna, including

1680-637: Is otherwise known for abundant Bellerophon fossils. Only a few trilobite genera are present by the Changhsingian. The last of the Trilobita include the genus Kathwaia of Pakistan . Perhaps the most widespread and diverse genus was Pseudophillipsia , other genera include Acropyge , Cheiropyge , and Paraphillipsia . In Australia, fossils of one of the last surviving eurypterids , ? Woodwardopterus freemanorum , were found. Mass extinction An extinction event (also known as

1750-464: 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

1820-451: Is the " Pull of the recent ", the fact that the fossil record (and thus known diversity) generally improves closer to the modern day. This means that biodiversity and abundance for older geological periods may be underestimated from raw data alone. Alroy (2010) attempted to circumvent sample size-related biases in diversity estimates using a method he called " shareholder quorum subsampling" (SQS). In this method, fossils are sampled from

1890-423: 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

1960-584: The Cambrian . These fit Sepkoski's definition of extinction, as short substages with large diversity loss and overall high extinction rates relative to their surroundings. Bambach et al. (2004) considered each of the "Big Five" extinction intervals to have a different pattern in the relationship between origination and extinction trends. Moreover, background extinction rates were broadly variable and could be separated into more severe and less severe time intervals. Background extinctions were least severe relative to

2030-520: 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 66 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

2100-601: The Induan Stage and the Triassic System ) is at the first appearance of the conodont species Hindeodus parvus . In the second part of the 20th century, appearance of the ammonite Otoceras , that existed no more than 100,000 years, in the boreal region was considered a marker of the Lower Triassic boundary. However, a more detailed study of Lower Induan biostratigraphy revealed the diachronicity of

2170-538: 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

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2240-649: The biosphere rather than the total diversity and abundance of life. For this reason, well-documented extinction events are confined to the Phanerozoic eon – with the sole exception of the Oxygen Catastrophe in the Proterozoic – since before the Phanerozoic, all living organisms were either microbial, or if multicellular then soft-bodied. Perhaps due to the absence of a robust microbial fossil record, mass extinctions might only seem to be mainly

2310-428: The genera Neoaganides , Pseudogastrioceras , Dzhulfites , Paratirolites , Julfotirolites , Alibashites , Abichites , Stoyanowites and Arasella The Bellerophon Formation in northern Italy documents a pre-extinction bivalve community with 26 species adapted to stressful conditions (high temperatures, high salinity, shallow water depths, low oxygen and high terrigenous input). The formation

2380-417: The Changhsingian fauna comprised gorgonopsid synapsids like Inostrancevia , anomodont synapsids like Daptocephalus and Dicynodon , and parareptiles like Elginia , milleretids and Nanoparia . Among fishes, the bobasatraniiforms Bobasatrania and Ebenaqua are known from Changhsingian deposits of Greenland and Australia, respectively. Another deep-bodied fish, Sinoplatysomus ,

2450-480: 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 , plesiosaurs and mosasaurs disappeared and the percentage of sessile animals was reduced to about 33%. All non-avian dinosaurs became extinct during that time. The boundary event was severe with a significant amount of variability in

2520-773: 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.   The End Permian extinction or the "Great Dying" occurred at the Permian – Triassic transition. It was the Phanerozoic Eon's largest extinction: 53% of marine families died, 84% of marine genera, about 81% of all marine species and an estimated 70% of terrestrial vertebrate species. This

2590-430: 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

2660-569: 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

2730-415: The accumulating data, it has been established that in the current, Phanerozoic Eon, multicellular animal life has experienced at least five major and many minor mass extinctions. The "Big Five" cannot be so clearly defined, but rather appear to represent the largest (or some of the largest) of a relatively smooth continuum of extinction events. All of the five in the Phanerozoic Eon were anciently preceded by

2800-467: The appearance of these mollusks in different regions of the Earth. The Changhsingian contains only one ammonoid biozone : that of the genus Iranites . The Changhsingian ended with the Permian–Triassic extinction event , the largest mass extinction event of the Phanerozoic Era , when both global biodiversity and alpha diversity (community-level diversity) were devastated. On land,

2870-400: 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

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2940-413: 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 A. Hallam and P.B. Wignall (1997) summarized the new extinction research of the previous two decades. One chapter in

3010-478: 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. A good theory for a particular mass extinction should: It may be necessary to consider combinations of causes. For example,

3080-458: 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 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

3150-497: 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. Because most diversity and biomass on Earth is microbial , and thus difficult to measure via fossils, extinction events placed on-record are those that affect the easily observed, biologically complex component of

3220-513: The end of the Cretaceous period. The Alvarez hypothesis 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 . 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

3290-752: 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. 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): The most commonly suggested causes of mass extinctions are listed below. 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

3360-462: The entire Phanerozoic. 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

3430-556: 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

3500-429: 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. Though Sepkoski died in 1999, his marine genera compendium was formally published in 2002. This prompted

3570-521: The geological record.   The largest extinction was the Kellwasser Event ( Frasnian - Famennian , 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 and numerous tropical benthic (seabed-living) animals such as jawless fish, brachiopods , and trilobites . The other major extinction

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3640-456: 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 " pelycosaurs ", though this is

3710-422: 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. 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

3780-430: 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

3850-434: The mass extinction were global warming , related to volcanism , and anoxia , and not, as considered earlier, cooling and glaciation . 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

3920-426: The old, dominant group and makes way for the new one, a process known as adaptive radiation . For example, mammaliaformes ("almost mammals") and then mammals existed throughout the reign of the dinosaurs , 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

3990-400: The ongoing mass extinction caused by human activity is sometimes called the sixth mass extinction . 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 newly dominant group is "superior" to the old but usually because an extinction event eliminates

4060-565: 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

4130-686: 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

4200-480: The physical environment. He expressed this in The Origin of Species : 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. Various ideas, mostly regarding astronomical influences, attempt to explain the supposed pattern, including the presence of a hypothetical companion star to

4270-596: The presumed far more extensive mass extinction of microbial life during the Great Oxidation Event (a.k.a. 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. In May 2020, studies suggested that the causes of

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4340-566: The rate of extinction between and among different clades . Mammals , descended from the synapsids , and birds , a side-branch of the theropod dinosaurs, emerged as the two predominant clades of terrestrial tetrapods. Despite the common presentation focusing only on these five events, no measure of extinction shows any definite line separating them from the many other Phanerozoic 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

4410-588: 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 . Research completed after the seminal 1982 paper (Sepkoski and Raup) has concluded that a sixth mass extinction event due to human activities is currently under way: 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

4480-639: 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

4550-602: 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 fungal abundance) provides reassurance that most widely accepted extinction events are real. A quantification of

4620-453: 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

4690-494: The vacant niches created the opportunity for archosaurs to become ascendant . In the seas, the percentage of animals that were sessile (unable to move about) 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

4760-556: Was anchored in the international timescale in 1981. The base of the Changhsingian Stage is at the first appearance of the conodont species Clarkina wangi . The global reference profile is profile D at Meishan , in the type area in Changxing, just below the Changhsingian foraminifer index fossil Palaeofusulina and the first appearance of the ammonoid Tapashanites . The top of the Changhsingian (the base of

4830-457: 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

4900-514: 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

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