The Capitanian mass extinction event began around 262 million years ago with its most intense pulse peaking at 259 million years ago, marking the end of the Capitanian stage and Guadalupian (Middle Permian) epoch of the Permian. Historically conflated with the better-known Permian–Triassic extinction, it was only recognised as a distinct event in 1994. Despite this, the mass extinction is believed to be the third-largest of the Phanerozoic in terms of the percentage of genera (33–35%) and species (60–63%) lost after the end-Permian and Late Ordovician mass extinction, while being the fifth worst in terms of ecological severity. The global nature of the Capitanian mass extinction has been called into question by some palaeontologists as a result of some analyses finding it to have affected only low-latitude taxa in the Northern Hemisphere.
Magnitude
In the aftermath of Olson's Extinction, global diversity rose during the Capitanian. This was probably the result of disaster taxa replacing extinct guilds. The Capitanian mass extinction greatly reduced disparity (the range of different guilds); eight guilds were lost. It impacted the diversity within individual communities more severely than the Permian–Triassic extinction event. Although faunas began recovery immediately after the Capitanian extinction event, rebuilding complex trophic structures and refilling guilds, diversity and disparity fell further until the boundary.
Marine ecosystems
The impact of the Capitanian extinction event on marine ecosystems is still heavily debated by palaeontologists. Early estimates indicated a loss of marine invertebrate genera between 35 and 47%, while an estimate published in 2016 suggested a loss of 33–35% of marine genera when corrected for background extinction, the Signor–Lipps effect and clustering of extinctions in certain taxa. The loss of marine invertebrates during the Capitanian mass extinction was comparable in magnitude to the Cretaceous–Paleogene extinction event. Some studies have considered it the third or fourth greatest mass extinction in terms of the proportion of marine invertebrate genera lost; a different study found the Capitanian extinction event to be only the ninth worst in terms of taxonomic severity (number of genera lost) but found it to be the fifth worst with regard to its ecological impact (i.e., the degree of taxonomic restructuring within ecosystems or the loss of ecological niches or even entire ecosystems themselves).
Terrestrial ecosystems
Few published estimates for the impact on terrestrial ecosystems exist for the Capitanian mass extinction. Among vertebrates, Day and colleagues suggested a 74–80% loss of generic richness in tetrapods of the Karoo Basin in South Africa, including the extinction of the dinocephalians. In land plants, Stevens and colleagues found an extinction of 56% of plant species recorded in the mid-Upper Shihhotse Formation in North China, which was approximately mid-Capitanian in age. 24% of plant species in South China went extinct.
Timing Although it is known that the Capitanian mass extinction occurred after Olson's Extinction and before the Permian–Triassic extinction event, the exact age of the Capitanian mass extinction remains controversial. Part of the reason for this is the poorly resolved biostratigraphy of the Guadalupian-Lopingian boundary interval. It is also partly due to the somewhat circumstantial age of the Capitanian–Wuchiapingian boundary itself, which is currently estimated to be approximately 259.1 million years old, but is subject to change by the Subcommission on Permian Stratigraphy of the International Commission on Stratigraphy.
Additionally, there is a dispute regarding the severity of the extinction and whether the extinction in China happened at the same time as the extinction in Spitsbergen. According to one study, the Capitanian mass extinction was not one discrete event but a continuous decline in diversity that began at the end of the Wordian. Another study examining fossiliferous facies in Svalbard found no evidence for a sudden mass extinction, instead attributing local biotic changes during the Capitanian to the southward migration of many taxa through the Zechstein Sea. Carbonate platform deposits in Hungary and Hydra show no sign of an extinction event at the end of the Capitanian; the extinction event there is recorded in the middle Capitanian.
The volcanics of the Emeishan Traps, which are interbedded with tropical carbonate platforms of the Maokou Formation, are unique for preserving a mass extinction and the cause of that mass extinction. Large phreatomagmatic eruptions occurred when the Emeishan Traps first started to erupt, leading to the extinction of fusulinacean foraminifera and calcareous algae.
Historically, the age of the extinction horizon has been estimated to around 260–262 Ma in the absence of direct radiometric ages in marine sections; this fits broadly with radiometric ages from the terrestrial realm, assuming the two events are contemporaneous. Plant losses occurred either at the same time as the marine extinction or after it. A 2026 study published U-Pb constraints on an extinction horizon in South China, using ash beds from marine sections near the Emeishan Traps. These ash beds set the end of the Guadalupian at 259.857 ± 0.084 Ma. This places the boundary at the start of the second major pulse of volcanic activity.
Marine realm The extinction of fusulinacean foraminifera in Southwest China was originally dated to the end of the Guadalupian, but studies published in 2009 and 2010 dated the extinction of these fusulinaceans to the mid-Capitanian. Brachiopod and coral losses occurred in the middle of the Capitanian stage. The extinction suffered by the ammonoids may have occurred in the early Wuchiapingian.
Terrestrial realm The existence of change in tetrapod faunas in the mid-Permian has long been known in South Africa and Russia. In Russia, it corresponded to the boundary between what became known as the Titanophoneus Superzone and the Scutosaurus SuperzoneIvakhnenko, M. F., Golubev, V. K., Gubin, Yu. M., Kalandadze, N. N., Novikov, I. V., Sennikov, A. G. & Rautian, A. S. 1997. "Permskiye I Triasovyye tetrapody vostochnoi Evropy (Permian and Triassic Tetrapods of Eastern Europe)". GEOS, Moscow (original in Russian). and later the Dinocephalian Superassemblage and the Theriodontian Superassemblage, respectively. In South Africa, this corresponded to the boundary between the variously named Pareiasaurus, Dinocephalian or Tapinocephalus Assemblage Zone and the overlying assemblages.Keyser, A. W. & Smith, R. H. M. 1979. "Vertebrate biozonation of the Beaufort Group with special reference to the Western Karoo Basin." Annals of the Geological Survey of South Africa 12, pp. 1-36. In both Russia and South Africa, this transition was associated with the extinction of the previously dominant group of therapsid amniotes, the dinocephalians, which led to its later designation as the dinocephalian extinction. Post-extinction origination rates remained low through the Pristerognathus Assemblage Zone for at least 1 million years, which suggests that there was a delayed recovery of Karoo Basin ecosystems.
After the recognition of a separate marine mass extinction at the end of the Guadalupian, the dinocephalian extinction was seen to represent its terrestrial correlate. Though it was subsequently suggested that because the Russian Ischeevo fauna, which was considered the youngest dinocephalian fauna in that region, was constrained to below the Illawarra magnetic reversal and therefore had to have occurred in the Wordian stage, well before the end of the Guadalupian, this constraint applied to the type locality only. The recognition of a younger dinocephalian fauna in Russia (the Sundyr Tetrapod Assemblage) and the retrieval of biostratigraphically radiometric ages via uranium–lead dating of a tuff from the Tapinocephalus Assemblage Zone of the Karoo Basin demonstrated that the dinocephalian extinction did occur in the late Capitanian, around 260 million years ago.
Effects on life
Marine life In the oceans, the Capitanian extinction event led to high extinction rates among ammonoids, corals and calcareous algal reef-building organisms, foraminifera, bryozoans, and brachiopods. It was more severe in restricted marine basins than in the open oceans. It appears to have been particularly selective against shallow-water taxa that relied on photosynthesis or a photosymbiotic relationship; many species with poorly buffered respiratory physiologies also became extinct. The extinction event led to a collapse of the reef carbonate factory in the shallow seas surrounding South China.
The ammonoids, which had been in a long-term decline for a 30 million year period since the Roadian, suffered a selective extinction pulse at the end of the Capitanian. 75.6% of coral families, 77.8% of coral genera and 82.2% of coral species that were in Permian China were lost during the Capitanian mass extinction.
Among bivalve losses, perhaps most prominent were the giant alatoconchids, which occurred abundantly in the shallow seas around South China before the mass extinction event but failed to survive it.
Among foraminifera, the extinction event exhibited a strong size-dependent selectivity; large foraminifera suffered especially heavily during the crisis. The Verbeekinidae, a family of large fusuline foraminifera, went extinct. Morphologically complex miliolids also suffered major losses.
87% of brachiopod species found at the Kapp Starostin Formation on Spitsbergen disappeared over a period of tens of thousands of years; though new brachiopod and bivalve species emerged after the extinction, the dominant position of the brachiopods was taken over by the bivalves. Approximately 70% of other species found at the Kapp Starostin Formation also vanished. The fossil record of East Greenland is similar to that of Spitsbergen; the faunal losses in Canada's Sverdrup Basin are comparable to the extinctions in Spitsbergen and East Greenland, but the post-extinction recovery that happened in Spitsbergen and East Greenland did not occur in the Sverdrup Basin. Whereas rhynchonelliform brachiopods made up 99.1% of the individuals found in tropical carbonates in the Western United States, South China and Greece prior to the extinction, molluscs made up 61.2% of the individuals found in similar environments after the extinction. 87% of brachiopod species and 82% of fusulinacean foraminifer species in South China were lost. Although severe for brachiopods, the Capitanian extinction's impact on their diversity was nowhere near as strong as that of the later end-Permian extinction.
The effects of the extinction event on marine vertebrates is enigmatic. There is no evidence of a diversity decline among bony fish, although a pronounced increase in body mass is documented among 'palaeopterygians' across the Guadalupian-Lopingian boundary.
Biomarker evidence indicates red algae and photoautotrophic bacteria dominated marine microbial communities. Significant turnovers in microbial ecosystems occurred during the Capitanian mass extinction, though they were smaller in magnitude than those associated with the end-Permian extinction.
Most of the marine victims of the extinction were either endemic species of epicontinental seas around Pangaea that died when the seas closed, or were dominant species of the Paleotethys Ocean. Evidence from marine deposits in Japan and Primorye suggests that mid-latitude marine life became affected earlier by the extinction event than marine organisms of the tropics.