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Capitanian

The Capitanian is the uppermost stage of the Guadalupian Series within the Middle Permian Epoch, spanning from approximately 264.3 to 259.5 million years ago.^[1] It is formally defined by the Global Stratotype Section and Point (GSSP) located in the Guadalupe Mountains National Park, Texas, USA, at a horizon 4.5 meters above the base of the Pinery Limestone Member of the Bell Canyon Formation, corresponding to the first appearance datum of the conodont species Jinogondolella postserrata.^[2] Named after the Capitan Reef complex in the same region, this stage represents a period of significant marine carbonate deposition, including reef-building activity by sponges and other organisms, amid a backdrop of global tectonic stability in the supercontinent Pangea.^[3] The Capitanian is characterized by diverse marine faunas, including fusulinid foraminifers, ammonoids, and brachiopods, which provide key biostratigraphic markers for correlation across Pangea and adjacent regions.^[4] On land, terrestrial ecosystems featured advanced therapsid synapsids and early archosauromorph reptiles, reflecting a transitional phase in tetrapod evolution before the subsequent Lopingian.^[5] Climatically, the stage saw warm, arid conditions with episodic sea-level fluctuations influencing sedimentation patterns from shallow shelves to deep basins.^[6] A defining feature of the Capitanian is the associated mass extinction event near its upper boundary, often termed the end-Capitanian or Guadalupian extinction, which resulted in the extinction of approximately 35% of marine genera, including about 82% of fusulinid species and many reef-builders, and impacted terrestrial vertebrate assemblages by shifting dominance from dinocephalians to dicynodonts.^[7]^[4] This crisis, occurring around 260 million years ago, is linked to environmental stressors such as volcanism from the Emeishan Large Igneous Province, ocean anoxia, and possibly hypercapnia, setting the stage for biotic recovery in the subsequent Wuchiapingian Stage.^[8] The event underscores the Capitanian's role as a pivotal interval in Permian biodiversity dynamics, bridging pre- and post-extinction ecosystems.^[5]

Overview

Definition and nomenclature

The Capitanian is the uppermost stage of the Guadalupian Series within the Middle Permian epoch of the Permian period. It represents a chronostratigraphic unit spanning from 264.28 ± 0.16 Ma at its base to 259.51 ± 0.21 Ma at its top, according to the 2024 International Chronostratigraphic Chart.^[9] This interval precedes the Wuchiapingian stage of the Lopingian Series above and succeeds the Wordian stage below.^[9] The name "Capitanian" derives from the Capitan Formation, a prominent reef complex exposed in the Capitan Mountains (also known as Capitan Peak) of southeastern New Mexico, USA, where key stratigraphic sections are preserved.^[10] The term was initially introduced in a lithostratigraphic context by George Burr Richardson in 1904 to describe the massive limestone reefs of the Guadalupe Mountains region spanning New Mexico and West Texas.^[10] Over time, this evolved into a formal chronostratigraphic designation, with the stage ratified by the International Commission on Stratigraphy (ICS) in 2001, establishing its global standardization.^[11] The Capitanian is subdivided into early, middle, and late intervals primarily based on biostratigraphic zonations using conodonts and fusulinid foraminifers. These divisions reflect evolutionary successions, such as the conodont zones of Jinogondolella postserrata (early), J. altudaensis (middle), and Clarkina postbitteri (late), correlated with fusulinid assemblages including Polydiexodina (early) and Parafusulina to Jigulites (middle to late).^[12]

Geological context

The Capitanian Stage represents the uppermost division of the Middle Permian, forming the final stage of the Guadalupian Epoch. It succeeds the Wordian Stage and precedes the Wuchiapingian Stage of the overlying Lopingian Series, marking the transition from the Middle to the Late Permian within the broader Permian Period.^[13] This positioning underscores its role as a pivotal interval in Permian chronostratigraphy, bridging earlier stable depositional regimes with the more volatile conditions of the Late Permian.^[13] Spanning approximately 4.8 million years from 264.28 Ma to 259.51 Ma, the Capitanian embodies a transitional phase in Earth history, characterized by relative tectonic stability amid the consolidation of the supercontinent Pangea while setting the stage for Late Permian disruptions, including biotic crises and climatic shifts.^[9] During this time, the ongoing assembly of Pangea, which had begun in the Carboniferous, reached near-completion, profoundly influencing global sedimentation patterns, ocean circulation, and climate through the coalescence of major landmasses into a single equatorial supercontinent.^[14] Key tectonic processes included the continued effects of the Uralian Orogeny, a major collisional event between Laurussia and Kazakhstania that extended from the Late Devonian through the Late Permian, deforming the eastern margins of the emerging Pangea and generating extensive foreland basins and mountain belts.^[15] Additionally, initial extensional tectonics and rifting episodes in western Pangea, particularly along proto-Atlantic margins, began to manifest as subtle basin development, foreshadowing the supercontinent's eventual fragmentation.^[16] Regionally, the Capitanian correlates with the upper Kazanian Stage on the Russian Platform, where biostratigraphic markers such as conodonts and ammonoids facilitate alignment despite differences in depositional environments.^[17] In China, it aligns with portions of the Middle Permian successions in the Baoshan Block of western Yunnan, integrated through fusulinid and conodont biozonations that link local lithostratigraphy to the global standard.^[18] These correlations highlight the stage's global synchroneity amid varying tectonic influences from Pangea's formation.^[13]

Stratigraphy

Global Stratotype Section and Point (GSSP)

The Global Stratotype Section and Point (GSSP) for the base of the Capitanian Stage is situated at Nipple Hill in the southeastern Guadalupe Mountains National Park, Texas, USA, at coordinates 31°54′32″N 104°47′20″W. It is positioned 4.5 m above the base of the outcrop section within the Pinery Member of the Bell Canyon Formation and was ratified in 2001 by the International Commission on Stratigraphy following a proposal in 1999.^[19]^[20] The lower boundary of the Capitanian Stage is defined by the first appearance datum (FAD) of the conodont Jinogondolella postserrata within the evolutionary lineage from J. aserrata to J. postserrata to J. shannoni. The upper boundary is delineated by the GSSP for the base of the succeeding Wuchiapingian Stage at the Penglaitan section, Laibin, Guangxi, China (coordinates 23°41′43″N 109°19′16″E), marked by the FAD of the conodont Clarkina postbitteri postbitteri and ratified in 2005.^[19]^[21] In 2023, the IUGS ratified a proposal to redefine this GSSP at a nearby, better-exposed section due to flooding and erosion at the original site, though the International Commission on Stratigraphy has not yet updated its official chart as of 2025.^[22] Biostratigraphic correlation is further supported by fusulinids indicative of the Midian regional stage.^[23] At the GSSP locality, the lithostratigraphy comprises thin-bedded limestones of the Pinery Member, featuring chert nodules and interbedded shales, characteristic of deep-water basinal turbidite deposits in the Delaware Basin.^[24] Correlation of the Capitanian boundaries relies on integrated biostratigraphy and radiometric dating to address regional variations; U-Pb CA-ID-TIMS zircon geochronology from ash beds near the GSSP yields an age of 264.28 ± 0.16 Ma for the base, providing a precise anchor for global timescales.^[23]

Biostratigraphy

The biostratigraphy of the Capitanian Stage relies primarily on conodont and fusulinid zonations, which provide robust markers for global correlation due to their wide distribution in marine deposits. Conodont biozones define the stage's boundaries and subdivisions, beginning with the Jinogondolella postserrata Zone at the base (defined by its FAD), marking the transition from the underlying Wordian, followed by the J. shannoni Zone, the J. altudaensis Zone, the J. crofti Zone in the middle to late Capitanian, and culminating in the Clarkina postbitteri hongshuiensis Zone near the top. These zones are characterized by evolutionary changes in platform morphology, such as increasing serration and narrowing in the Jinogondolella lineage, allowing precise identification across equatorial sections.^[25] Fusulinid biozones complement conodonts, with the Parafusulina Zone dominating the early Capitanian, featuring elongate tests and keriothecal wall structures, transitioning to the Polydiexodina Zone in the later part, where genera exhibit advanced axial partitioning and larger sizes indicative of stable, warm-water environments.^[13] Additional biostratigraphic markers include ammonoids, brachiopods, and radiolarians, which support regional correlations. The Timorites Zone characterizes much of the Capitanian in basinal settings, particularly in the Tethyan and Panthalassan realms.^[26] Brachiopod assemblages, such as the Leptodus nobilis-Spiriferella Zone, provide supplementary control in shallow-marine carbonates, while radiolarian zones like the Pseudoalbaillella globosa Zone overlap with conodont intervals in siliceous facies, aiding integration in deep-water sequences.^[27]^[28] Globally, Capitanian zonations facilitate correlation across the Tethys (equivalent to the Midian Stage), Panthalassa (Guadalupe Series), and Gondwana margins, with conodonts and fusulinids achieving near-cosmopolitan distribution in low latitudes.^[13] However, high-latitude regions exhibit discrepancies due to faunal provincialism, where endemic ammonoid and brachiopod taxa hinder direct matching, necessitating auxiliary tools like carbon isotopes for alignment.^[29] In South China, the Lengwu Formation aligns with the J. altudaensis Zone, while Japanese sections correlate via J. crofti, underscoring the stage's utility despite latitudinal biases.^[25] Evolutionary trends in fusulinids highlight a peak in diversity during the early to mid-Capitanian, with Polydiexodina and related genera reaching maximum generic richness in tropical shelves, before a sharp decline toward the stage's end.^[30] This biotic turnover, involving the selective extinction of larger, symbiont-bearing forms, reflects environmental stresses and foreshadows broader Permian disruptions, with over 80% of fusulinoidean genera lost by the Guadalupian-Lopingian boundary.^[31] Conodont faunas show similar patterns, with Jinogondolella diversification giving way to Clarkina dominance, signaling shifts in oceanic conditions.^[32]

Paleoenvironments

Paleogeography

During the Capitanian stage of the Middle Permian, approximately 264.3 to 259.5 million years ago, the supercontinent Pangea was nearly fully assembled, with the suturing of Laurussia (comprising North America and Eurasia) and Gondwana (including South America, Africa, India, Australia, and Antarctica) largely complete following the Late Carboniferous collisions. This configuration formed a vast, C-shaped landmass that extended from high northern to high southern latitudes, enclosing a narrow Paleo-Tethys Ocean along its eastern margin while being surrounded by the expansive Panthalassa superocean to the west and north. The assembly of Pangea marked the culmination of the Variscan-Hercynian orogenic events, stabilizing the central continental core but leaving residual tectonic stresses that influenced peripheral margins.^[33] Much of Pangea occupied equatorial to low-latitude positions, promoting extensive arid interiors due to its elongated shape and continental clustering, which disrupted global moisture distribution. Subduction zones were active along the western margins of Pangea, particularly in the Panthalassa realm, where convergence generated island arcs and volcanic chains, including elements of the future Cordilleran system in western North America. These zones contributed to the initial stages of circum-Pangea subduction, encircling the supercontinent and driving ongoing plate reorganization. In Europe, remnants of the Hercynian Orogeny manifested as elevated highlands and faulted basins, such as those in the Variscan belt, which influenced sediment dispersal into adjacent seas.^[33]^[34] Regional paleogeography featured extensive shallow epicontinental shelves in the Tethys domain, particularly along the northern and southern margins of Pangea, where warm, shallow waters supported carbonate platform development in the Paleo-Tethys. In contrast, the Panthalassa Ocean included deep oceanic basins reaching depths of 2600 to 4400 meters, punctuated by fringing reef buildups along convergent margins, exemplified by the Capitan Reef complex in the Delaware Basin of western Texas and southeastern New Mexico. This reef system formed a prominent barrier along the western Pangean shelf, delineating shallow backreef lagoons from deeper fore-reef slopes in a tropical setting. These spatial contrasts shaped sediment deposition and tectonic stability across the global system.^[29]^[33]^[35]

Climate and sea levels

The Capitanian stage was characterized by a warm greenhouse climate, with arid conditions dominating low-latitude regions and seasonal monsoons influencing precipitation patterns across the supercontinent Pangea. Extensive evaporite deposits in back-reef environments, such as those within the Guadalupian succession of the Permian Basin, provide evidence of heightened aridity, elevated salinities, and minimal freshwater input in shallow marine settings.^[36] Paleosols from contemporaneous terrestrial sequences further indicate semi-arid to arid soils with calcic horizons, reflecting a stable greenhouse state interrupted by episodic wetter phases tied to monsoon variability.^[37] Temperature reconstructions reveal a cooling trend commencing in the late Capitanian, particularly in tropical settings, as derived from oxygen isotope ratios (δ¹⁸O) in well-preserved brachiopod shells from low-latitude sites.^[38] Sea levels experienced notable fluctuations, with highstands in the early Capitanian promoting expansive reef development in equatorial belts, as evidenced by the vertical and lateral growth of carbonate platforms like the Capitan Reef in the Delaware Basin.^[36] A pronounced regression marked the late Capitanian, interpreted as a global eustatic sea-level fall, resulting in widespread subaerial exposure of reefs and the onset of evaporitic and terrestrial sedimentation over marine carbonates.^[39] Carbon isotope records show positive excursions in δ¹³C, including the early Capitanian Kamura event with values rising to around +3‰, signaling enhanced burial of isotopically light organic carbon in anoxic microenvironments of restricted basins and heightened marine productivity through much of the stage.^[40] These shifts, observed in platform-to-basin transects of the Guadalupian Reef Complex, reflect carbon cycle perturbations without widespread oceanic anoxia until the Capitanian-Lopingian boundary.^[41]

Biodiversity

Marine life

The Capitanian stage featured high marine biodiversity, particularly among benthic and reef-dwelling organisms, with fusulinid foraminifera exhibiting exceptional diversity, including over 100 species globally, serving as key index fossils for biostratigraphy.^[42] Rugose and tabulate corals, alongside calcareous sponges and algae, contributed to the construction of extensive reef frameworks, while brachiopods and bryozoans formed prominent components of benthic assemblages, often dominating in shallow-water carbonates.^[43] These groups reflected a thriving Paleozoic marine ecosystem prior to later Permian declines. Key marine ecosystems during the Capitanian included massive sponge-algal reefs, exemplified by the Capitan Reef in the Guadalupe Mountains, which reached thicknesses of up to 600 meters and supported diverse frame-building communities of sponges, phylloid algae, and encrusting organisms like Tubiphytes.^[35]^[43] In back-reef lagoons, low-energy environments hosted abundant fusulinids, brachiopods, and ostracods, while fore-reef talus slopes accumulated debris from bryozoans, crinoids, and rare horn corals. Open-ocean pelagic communities were characterized by ammonoids (e.g., genus Timorites) and conodonts, which inhabited deeper waters beyond the reef margins and provided biostratigraphic markers across the stage.^[44]^[10] Trophic structures emphasized herbivores and filter-feeders, with algae and cyanobacteria as primary producers sustaining grazing fusulinids and deposit-feeding brachiopods; suspension-feeders like bryozoans and crinoids filtered plankton in reef and lagoon settings. Early signals of stress appeared in larger fusulinids toward the stage's end, hinting at environmental shifts affecting herbivorous and symbiotic groups.^[30] Distribution patterns showed pronounced provinciality, with tropical Tethyan faunas—rich in ornate brachiopods, large fusulinids, and compound corals—contrasting cooler-water Boreal assemblages featuring more restricted brachiopod and bryozoan diversity in higher latitudes.^[29] This biogeographic divide underscored the influence of paleogeographic barriers and climatic gradients on Capitanian marine communities.^[45]

Terrestrial life

During the Capitanian stage, terrestrial floras exhibited strong latitudinal provincialism, with Glossopteris-dominated forests characterizing the southern continents of Gondwana. These arborescent gymnosperms formed extensive lowland woodlands, often associated with coal-forming wetlands that supported diverse understory vegetation including ferns and horsetails.^[46] In Gondwana, Glossopteris flora persisted as the dominant vegetation, contributing to peat accumulation in swampy environments, though these wetlands began to decline toward the end of the stage due to increasing aridity.^[47] By contrast, in northern Pangea, encompassing Euramerica and Angara, floras were dominated by conifers such as Walchia and seed ferns like Callipteris, alongside lycopods in more humid, upland settings; these assemblages reflected adaptation to seasonal climates with reduced wetland extent compared to earlier Permian periods.^[48] Terrestrial faunas were overwhelmingly dominated by synapsids, marking a transition from earlier pelycosaur-like forms to more advanced therapsids. In Gondwana, early dicynodonts emerged as herbivorous generalists, while gorgonopsians appeared as apex predators with saber-like canines, filling niches left by the extinction of dinocephalian therapsids during the stage.^[5] These therapsids inhabited floodplain environments, with dicynodonts showing adaptations for browsing in forested or scrubby landscapes.^[49] Smaller tetrapods, including parareptiles and early archosauromorphs, coexisted but were less prominent, underscoring the synapsid hegemony in continental ecosystems. Insect communities retained diversity from early Permian holdovers, such as palaeodictyopteroids and early holometabolous orders like Mecoptera, though overall arthropod richness began to wane amid environmental shifts.^[50] Capitanian ecosystems on land were shaped by increasing aridity across Pangea, featuring arid floodplains traversed by seasonal rivers that supported riparian vegetation and ephemeral water bodies. In Gondwana's Karoo Basin, for instance, meandering fluvial systems graded into playa-like settings, fostering habitats for synapsid assemblages amid periodic droughts.^[51] These environments promoted opportunistic feeding strategies among herbivores and predators, with coal swamps persisting in wetter depressions but diminishing as global climates dried.^[52] Faunal provincialism was pronounced, with distinct synapsid assemblages separating Euramerican and Gondwanan realms despite Pangea's supercontinental configuration. Euramerica hosted burnetiamorph therapsids and varanopid pelycosaurs in its faunas, while Gondwana emphasized dicynodonts and pristerognathid gorgonopsians, reflecting barriers to dispersal posed by equatorial deserts and climatic gradients.^[53] This endemism extended to floral distributions, reinforcing biogeographic divides that limited interprovincial exchange.^[54]

Key Formations

North American formations

The Capitan Formation in the Guadalupe Mountains of west Texas and southeast New Mexico represents a classic Capitanian reef complex, composed primarily of massive, fine-grained fossiliferous limestones built by algal, sponge, and bryozoan frameworks. These reefal deposits form steep cliffs up to 1,000 feet high, such as El Capitan, and reach thicknesses of approximately 2,000 feet (609 m) in exposed sections like McKittrick Canyon, serving as the primary type section for the stage due to their exceptional preservation of Middle Permian reef architecture.^[55]^[35]^[56] Underlying the Capitan Formation, the Bell Canyon Formation consists of basinal shales, fine-grained sandstones, siltstones, and thin limestones (such as the Pinery Member), deposited as turbidites and submarine fan sediments in the deep Delaware Basin. This unit attains thicknesses of 500–600 feet (152–183 m) and hosts the Global Stratotype Section and Point (GSSP) for the base of the Capitanian at Nipple Hill, illustrating the transition from basinal to reef-margin facies.^[55]^[35]^[57]^[56] Further into the Permian Basin shelf, the San Andres Formation comprises cyclic carbonates including limestones, dolomites, and minor sandstones, reflecting repeated transgressive-regressive cycles in shallow shelf environments. These deposits vary in thickness from 300 to 500 feet (91–152 m) in outcrop areas and correlate laterally with basinal equivalents like the upper Cherry Canyon Formation.^[56] Overlying the San Andres, the Grayburg Formation features light-colored dolomites, oolitic limestones, and sandstones indicative of sabkha, tidal flat, and shallow agitated marine settings, with thicknesses reaching up to 475 feet (145 m). This unit marks a progradational phase on the Northwest Shelf and equivalents the middle Capitanian basinal strata.^[56] The San Andres and Grayburg Formations are major hydrocarbon reservoirs within the Permian Basin, hosting billions of barrels of oil through porous carbonates and enhanced recovery techniques, while the Capitan Formation contributes modestly as a marginal aquifer and minor producer.^[58]^[59]

Global formations

The Maokou Formation in South China represents a prominent Capitanian carbonate platform deposit, consisting of thick sequences of limestones exceeding 150 meters in thickness, characterized by massive dark gray bioclastic beds rich in fusulinids such as Yabeina and Neoschwagerina. These fusulinid-rich layers reflect shallow marine shelf environments with high productivity, while the formation's upper portions exhibit karstic features developed during a late Capitanian regression, marked by denudation and exposure surfaces that indicate a significant drop in sea level.^[60]^[61] This regression is evidenced by the transition to overlying continental deposits and the formation of paleokarst landforms preserved in the top strata.^[62] In Antarctica, the Weller Coal Measures exemplify Gondwanan terrestrial deposits of Capitanian age, comprising repetitive coal-bearing cyclothems with interbedded sandstones and shales dominated by Glossopteris flora, including leaves and stems that indicate forested wetlands influenced by regional climatic fluctuations.^[63]^[64] These cyclothems, up to several hundred meters thick, feature coal seams formed from peat accumulation in fluvial and deltaic settings, with Glossopteris as the dominant seed fern contributing to the organic-rich layers that reflect periodic flooding and peat preservation under humid conditions.^[65] The presence of Glossopteris underscores biotic continuity across high-latitude Gondwana, paralleling coal measures in other southern continents during this stage.^[66] The Uralian Basin in Russia hosts equivalent Capitanian strata within the Kazanian Series, featuring a mix of evaporites such as gypsum and halite alongside red beds of siltstones and sandstones that accumulated in restricted marine to continental settings.^[67]^[68] These deposits, reaching thicknesses of over 1 km in depocenters, include lagoonal evaporitic cycles interspersed with terrestrial red beds indicative of aridification and basin restriction, with biostratigraphic markers like conodonts confirming correlation to the global Capitanian.^[69] The evaporites formed during phases of hypersalinity in the eastern Russian Platform, contrasting with more open marine carbonates elsewhere but sharing regression-related features with North American sequences like the Capitan Formation.^[70] Tethyan regions in Italy preserve Capitanian pelagic limestones in formations such as those in the Sosio Valley of western Sicily, where thin-bedded, radiolarian-bearing cherty limestones up to 50 meters thick record deep-water basinal deposition.^[71] These pelagic sequences, dominated by fine-grained micrites with sparse fusulinids and conodonts, reflect open-ocean conditions in the western Tethys, with biostratigraphy assigning them to the Capitanian based on fauna like Follicucullus radiolarians.^[72] The limestones exhibit rhythmic bedding and minor turbidites, highlighting subsidence and sediment starvation in a tectonically active margin.^[39]

Significant Events

End-Capitanian extinction

The End-Capitanian extinction, also known as the end-Guadalupian biotic crisis, occurred approximately 260 million years ago at the boundary between the Capitanian stage of the Middle Permian and the Wuchiapingian stage of the Late Permian. This event marked a significant but selective mass extinction that affected marine ecosystems more severely than terrestrial ones, serving as a precursor to the more devastating Permian-Triassic extinction about 7 million years later.^[73] Overall, it resulted in the loss of around 35% of marine genera, with particularly high impacts on certain groups such as fusulinid foraminifers (96% species extinction) and ammonoids (severe generic turnover).^[73]^[74] Terrestrial impacts included significant declines in vertebrate faunas, with a 74–80% loss of tetrapod generic richness in the Karoo Basin of South Africa, involving the extinction of dinocephalians and other groups, followed by a radiation of dicynodonts and therapsids in the subsequent Wuchiapingian.^[5] Selective patterns of the extinction highlight environmental heterogeneity, with cool-water faunas experiencing more abrupt and intense losses compared to tropical assemblages. In Boreal cool-water settings, such as the Kapp Starostin Formation in Spitsbergen, brachiopod diversity plummeted by 87% (28 of 32 species), affecting siliceous sponge- and bryozoan-dominated communities, while bivalves showed resilience with only 50% generic loss.^[75] In contrast, tropical Tethyan faunas exhibited more gradual declines, with 25–35% generic extinction across brachiopods, bivalves, and gastropods, but minimal impact on ecologically tolerant groups like certain gastropods.^[73] Taxonomically, the crisis disproportionately targeted large, specialized reef-builders and pelagic forms, including fusulinids (confined to warm, shallow waters) and ammonoids (high turnover in open-marine habitats), while infaunal and opportunistic taxa survived better.^[76] On land, the event significantly affected synapsid vertebrates, with dinocephalians declining but dicynodonts and therapsids diversifying post-crisis, indicating pervasive stress in continental ecosystems.^[5] Paleontological and geochemical evidence underscores the abrupt nature of the marine crisis. Fossil records show sharp declines in abundance and diversity within thin stratigraphic intervals (e.g., 30 cm in Spitsbergen sections), with post-extinction assemblages dominated by disaster taxa like opportunistic bivalves.^[75] A prominent negative excursion in carbon isotopes (δ¹³C up to -6‰ in carbonates and -3‰ in organics) coincides with the event horizon, signaling perturbations in the global carbon cycle, possibly from enhanced productivity collapse or methane release.^[75] Supporting traces include increased pyrite framboids and elevated trace metals (U, Mo, V), indicative of expanded anoxia in marginal seas.^[75] The causes of the End-Capitanian extinction remain debated, with evidence pointing primarily to global cooling and sea-level regression, though some studies suggest contributions from early Emeishan volcanism to anoxia and other stressors. The Kamura cooling event, marked by positive oxygen isotope shifts (δ¹⁸O up to +2‰), likely expanded polar ice caps and lowered sea levels by 50–100 m, contracting shallow-marine habitats and stressing warm-adapted tropical biotas indirectly through habitat loss.^[76] This regression exacerbated anoxia in restricted basins, particularly impacting cool-water margins where upwelling intensified.^[75] Although contemporaneous with early Emeishan volcanism, the extinction's selectivity against warm-water groups and lack of widespread warming signatures suggest no primary causal link to flood basalts.^[76]

Associated volcanism

The Emeishan Large Igneous Province (LIP), situated in southwest China and extending into northern Vietnam, represents the primary volcanic event associated with the Capitanian stage.^[77] This continental flood basalt province erupted around 260 Ma, with its main phase occurring during the late Capitanian.^[78] The erupted volume is estimated at approximately 1.45 × 10^6 km³, with total emplaced magmas reaching up to 2.39 × 10^6 km³, while the preserved areal extent covers about 0.3–0.5 × 10^6 km².^[79]^[77] The peak activity of the Emeishan LIP involved extensive flood basalts, sill complexes, and intrusive bodies, with magmatism spanning roughly 4–8 million years but intensifying toward the end-Capitanian.^[80] High-precision U-Pb zircon dating of felsic tuffs and ignimbrites places the termination of major eruptions at 259.1 ± 0.5 Ma, confirming temporal synchrony with the end-Capitanian biotic crisis.^[81] This dating aligns the LIP's climax with global environmental perturbations recorded in marine sections worldwide.^[7] Environmental impacts from the Emeishan LIP stemmed largely from the massive release of volatiles during eruptions and sill intrusions into sedimentary basins.^[82] Elevated emissions of CO₂ drove initial global warming through greenhouse forcing, while SO₂ injections into the atmosphere caused short-term cooling via sulfate aerosol formation. Additionally, the CO₂ flux contributed to ocean acidification, exacerbating stress on calcifying marine organisms.^[83] Beyond the Emeishan LIP, volcanic activity during the Capitanian included minor rifting-related magmatism in the Panthalassic Ocean, linked to the initial breakup phases of Pangea.^[84] The Siberian Traps, often associated with later Permian events, did not form until the Lopingian stage around 252 Ma and thus played no role in Capitanian volcanism.^[78]

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GSSP Table - Paleozoic Era - Geologic TimeScale Foundation
Capitanian Stage, 264.3 ± 0.4, Nipple Hill, SE Guadalupe Mountains, Texas, U.S.A, 31.9091°N 104.7892°W, 4.5m above the base of the outcrop section of the Pinery ...
[19]
(PDF) Chapter 24, The Permian Period - ResearchGate
May 15, 2017 · ... 4.5 m in the outcrop section at. Nipple Hill, in the upper Pinery ... GSSP of the Capitanian Stage was. ratiﬁed in early 2001. The ...
[20]
[PDF] The Global Stratotype Section and Point (GSSP) for the boundary ...
Dec 6, 2006 · The Global Stratotype Section and Point (GSSP) for the boundary between the Capitanian and Wuchiapingian stages, also the boundary between ...
[21]
The Global Stratotype Section and Point (GSSP) for the base of the ...
Sep 1, 2022 · The Global Stratotype Section and Point (GSSP) for the base of the Capitanian Stage was proposed in 1999 and defined by the first appearance ...
[22]
USGS: Geological Survey Professional Paper 446 - NPS History
Feb 13, 2008 · A small isolated outcrop of thin-bedded limestone, probably the McCombs or Pinery Member of the Bell Canyon, grades into the Capitan Limestone ...
[23]
None
### Biostratigraphy of Capitanian (Guadalupian, Middle Permian)
[24]
Permian ammonoid biostratigraphy - GeoScienceWorld
Jan 1, 2018 · Two 'zones' of this scale have been used by stratigraphers for many decades: the Perrinites and Waagenoceras zones, corresponding to the ...
[25]
Integrated Radiolarian and Conodont Biostratigraphy of the Middle ...
May 31, 2021 · Abstract. The Permian radiolarian zones and their correlations with conodont zones or other chronostratigraphic schemes are still under debate.
[26]
Middle Permian (Guadalupian) biostratigraphy in South Primorye ...
These beds are commonly overlain by the Early Capitanian (Middle Midian) fusulinid Parafusulina stricta Zone, brachiopod Leptodus nobilis-Spiriferella Zone ...
[27]
Capitanian (Late Guadalupian, Permian) global brachiopod ...
In this paper, we aim to analyze the global palaeobiogeographical patterns of Capitanian brachiopod faunas using a range of multivariate statistical methods, ...
[28]
End-Guadalupian extinction of larger fusulinids in central Iran and ...
Jul 15, 2020 · The end-Guadalupian (Capitanian) biotic crisis was less intense than its end-Permian counterpart but highly selective against certain groups ...
[29]
Permian biotic turnovers in the northwestern margin of Pangea
Feb 4, 2022 · The Capitanian (Guadalupian, Middle Permian) mass extinction in NW Pangea (Borup Fiord, Arctic Canada): A global crisis driven by volcanism ...
[30]
Permian conodont biostratigraphy | Geological Society, London ...
Forty Permian conodont biozones are summarized and 35 additional regional zones are correlated. The Lower Permian is largely zoned on the basis of partial ...
[31]
(PDF) Map Folio 51, late Middle Permian (Capitanian, 263.1 Ma)
This map folio should be referenced as: Scotese, C.R., 2013. Map Folio 51, late Middle Permian (Capitanian, 263.1 Ma), PALEOMAP PaleoAtlas for ArcGIS, volume 4, ...
[32]
Hercynian Orogeny - an overview | ScienceDirect Topics
Hercynian orogeny refers to the geological process that formed the Hercynian (or Variscan) orogenic belt, resulting from the subduction, accretion, and ...
[33]
Geologic Formations - Guadalupe Mountains National Park (U.S. ...
Oct 11, 2025 · During the middle part of the Permian Period a reef developed along the margin of the Delaware Sea. This was the Capitan Reef, now recognized as ...
[34]
Permian Reef Complex Virtual Field Trip
Jun 29, 2020 · El Capitan and Guadalupe Peak as viewed from the Guadalupe Pass area by Bartlett in 1850 on his journey by stage and wagon from San Antonio to ...Overview · Stratigraphic Setting and... · Depositional Patterns · Diagenetic Patterns
[35]
Permian and Triassic greenhouse crises - ScienceDirect.com
Paleoclimatic time series from Permian and Triassic paleosols reveal transient episodes of unusually warm and wet conditions, interrupting long periods of ...
[36]
δ 18 O and δ 13 C of Permian brachiopods: A record of seawater ...
Two hundred sixty-four δ18O and δ13C values of Permian articulate brachiopod shells were analyzed and 81 of these values were characterized as well ...
[37]
Evolution of the marine Permian carbonate platform in the Salt ...
The second sequence comprises the lower Wargal Formation (Capitanian). A ... This increase in δ18O corresponds to cooling with temperatures decreasing ...
[38]
(PDF) CAPITANIAN (MIDDLE PERMIAN) MASS EXTINCTION AND ...
Aug 5, 2025 · Base level was stable through much of the middle-late Permian transition with the exception of a major regression within the Capitanian Stage.
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Carbon isotopic record of a platform-to-basin transect through the ...
Dec 1, 2024 · δ13C values increase through the Capitanian likely due to basin restriction and carbon burial. Abstract. The Guadalupian Permian Reef Complex of ...
[40]
https://www.sciencedirect.com/science/article/pii/S1871174X07000157
[41]
https://www.eprints.whiterose.ac.uk/id/eprint/152599/
[42]
Capitan Limestone and Associated Formations of New Mexico and ...
Sep 12, 2019 · ... corals were important factors in building up the framework of the reef. Locally the reef rock contains numerous fragments of unidentifiable ...
[43]
Capitanian Stage | Permian Period, Carboniferous & Marine Life
Oct 10, 2025 · Capitanian Stage, last of the three stages of the Middle Permian (Guadalupian) Series, encompassing all rocks deposited during the Capitanian Age.
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[PDF] Capitanian (Late Guadalupian, Permian) global brachiopod ...
A global database containing 3365 occurrences, 821 species and 251 genera of the Capitanian (Late Guadalupian,. Permian) brachiopod faunas from 24 stations ...
[45]
South African Lagerstätte reveals middle Permian Gondwanan ...
Oct 30, 2022 · The iconic plant Glossopteris, an arborescent gymnosperm, dominated these floras in the lowlands of Gondwana for over 50 million years, from ...
[46]
Fossil wood from the Permian-Triassic Beaufort Group of South ...
Aug 28, 2025 · The Glossopteris flora dominated Gondwana during the Permian (Ecca and lower Beaufort times) and was later replaced by the dominant ...
[47]
Permian Phytogeographic Patterns and Climate Data/Model ...
During the Permian, Gondwana rotated about a pole near Australia so that the Paraná Basin of South America moved northward into the warm, semiarid biome ...Multivariate Statistical... · Sakmarian And Wordian... · Climate Data And Model...
[48]
Early–middle Permian Mediterranean gorgonopsian suggests an ...
Dec 17, 2024 · Therapsids were a dominant component of middle–late Permian terrestrial ecosystems worldwide, eventually giving rise to mammals during the ...
[49]
Evolution of Insect Diversity in the Permian and Triassic - Biotaxa
Oct 30, 2023 · Insects, being a major component of terrestrial ecosystems, demonstrate varied diversity responses to climatic changes in Permian and Triassic.
[50]
GEOL 102 The Late Paleozoic: The Carboniferous & Permian
Jan 16, 2025 · Fusulinids highly diverse, become important index fossils. Marine life of the Permian: Permian reef community: sponges and bryozoans as primary ...
[51]
Implications for the evolution of tetrapod ecosystems in equatorial ...
The sedimentary deposits show an evolution from fluvial meandering systems to playa-lake floodplains, denoting increased aridification and seasonality under a ...
[52]
Provincialization of terrestrial faunas following the end-Permian ...
Apr 29, 2013 · Here, we test whether the end-Permian mass extinction (252.3 Ma) affected the distribution of tetrapod faunas within the southern hemisphere.Missing: Euramerica | Show results with:Euramerica
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Local provincialism of late Permian plant–arthropod associations in ...
Sep 15, 2023 · This turnover triggered the further development of the four phytogeographic provinces of Angara, Cathaysia, Euramerica, and Gondwana (Feng, 2017) ...
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NPS Geodiversity Atlas—Guadalupe Mountains National Park ...
Jul 5, 2024 · The base of the highest stage (Capitanian) is located at the crest of Nipple Hill, 1.1 km (0.7 mi) east of Frijole Ranch within the Pinery ...
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[PDF] Permian sedimentary facies, central Guadalupe Mountains, New ...
... fusulinid species Parafusulina rothi and the ammonoid genera Pseudogastrioceras and Perrinites. The fusulinid species Parafusulina rothi, considered by Skinner.
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[PDF] The National Park Service Stratotype Inventory
The base of the Roadian, Wordian, and Capitanian Stages of the Guadalupian Series (middle Permian) are recognized at Guadalupe Mountains National Park, Texas ( ...
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[PDF] Characterization of the Grayburg Reservoir of the Mobil University ...
Jul 31, 1985 · Within the Permian Basin, 83 percent (16.6 billion barrels) of the cumulative production is from carbonates (Fig. 3), and 46 percent (7.7 ...Missing: Capitanian | Show results with:Capitanian
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[PDF] Shorter Contributions to Paleontology and Stratigraphy
more sharply defined at younger growth stages; posterior margin truncate. Anterior-dorsal margin straight to very slightly convex and inclined gently away ...
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Karst reservoirs seismic prediction of Lower Permian Maokou ...
The comprehensive study of the karst reservoirs in the Lower Permian Maokou Formation was carried out using new geophysical technology.Missing: regression | Show results with:regression
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Rapid sea-level change in the Late Guadalupian (Permian) on the ...
The Capitanian (Late Guadalupian) Maokou Formation at Chaotian in northern Sichuan, South China, is composed mainly of shallow marine shelf carbonates ...<|separator|>
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A case from Middle Permian Maokou Formation in Northwestern ...
Based on the analysis of deposition and tectonism, “residual thickness method” is used to restore the paleokarst landform of Middle Permian Maokou Formation.Missing: regression | Show results with:regression<|separator|>
[62]
Depositional framework of Permian coal-bearing strata, southern ...
The Permian Weller Coal Measures of Antarctica are exposed in a narrow belt that extends along the edge of the Polar Plateau from the Mawson Glacier to the ...Missing: Capitanian | Show results with:Capitanian
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The Beacon supergroup in the Allan Hills, central Victoria Land ...
Dec 21, 2011 · The coal-bearing cyclothems of the Weller Coal Measures lose their finer divisions in an eastwards lateral transition, and the Coal Measures ...
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Glacier-fed(?) sandstone sheets in the Weller Coal Measures ...
The upper member of the Weller Coal Measures (Permian) at Allan Hills, Victoria Land, Antarctica, consists of repetitive coal layers separated by sheetlike ...
[65]
Glossopteris flora in the Permian Weller Formation of Allan Hills ...
Here we describe the first comprehensive record of the Glossopteris flora from the Permian Weller Formation of Allan Hills, South Victoria Land, Antarctica. ...Missing: cyclothem | Show results with:cyclothem<|separator|>
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An overview of the Upper Palaeozoic–Mesozoic stratigraphy of the ...
Aug 12, 2016 · A Kazanian (Wordian–Capitanian) marine connection was established between the Barents Sea, the Jameson Land Basin and the European Zechstein ...
[67]
Stratigraphic data of the Middle – Late Permian on Russian Platform
Jul 8, 2025 · This paper presents the litho– and biostratigraphic data and correlations of the Middle and Late Permian (Ufimian, Kazanian and Tatarian) on ...
[68]
[PDF] Geology of the Volga-Ural Petroleum Province and detailed ...
Continental and marine redbeds, evaporites, dolomite, and alluvial conglomerate beds predominate in the upper part of each cycle. All of the sedimentary cycles.
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Mineralogical composition of the Lower and Upper Kazanian (Mid ...
Aug 9, 2025 · ... Kazanian, representing an additional biostratigraphic marker. The lower boundaries of the stages which are formed by continental red beds ...
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[PDF] Upper Permian Radiolarians from the Sosio Valley Area, Western ...
The largest part of the Lamar Limestone be- tween these two horizons belongs either to the Capitanian or to the Dzhulfian (Wuchiapingian) depending from the.
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CAPITANIAN (MIDDLE PERMIAN) MASS EXTINCTION AND ...
Feb 1, 2012 · The Capitanian (middle Permian) extinction and recovery event is examined in carbonate platform settings from western Tethys (Hungary and ...
[72]
Prolonged Permian–Triassic ecological crisis recorded by ... - PNAS
Aug 7, 2007 · There was a biotic crisis affecting fusulinids (11, 22) and an ecological crisis in reef communities (23) at the end of the Guadalupian, but ...
[73]
Extinction process and patterns of Middle Permian Fusulinaceans in ...
With only seven species of five genera surviving into the Late Permian, the extinction of fusulinaceans in the Guadalupian mass extinction is 96% at species ...
[74]
An abrupt extinction in the Middle Permian (Capitanian) of the ...
Sep 1, 2015 · The combined δ13Corg and 87Sr/86Sr records both suggest a middle Capitanian age for a level high in the Kapp Starostin Formation near the 3YLB.
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The double mass extinction revisited: reassessing the severity ...
Apr 8, 2016 · Previous estimates suggest that the crisis eliminated 58% of marine invertebrate genera during the Capitanian stage and that its selectivity ...
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The Emeishan large igneous province: A synthesis - ScienceDirect
The late Permian Emeishan large igneous province (ELIP) covers ∼0.3 × 10 6 km 2 of the western margin of the Yangtze Block and Tibetan Plateau
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Large igneous provinces and mass extinctions - GeoScienceWorld
The compilation of Liu and Zhu (2009) shows the oldest and youngest U-Pb dates from. Emeishan samples spanning 267–251 Ma, which was recently more tightly ...<|control11|><|separator|>
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Crustal SiO 2 Content of the Emeishan Large Igneous Province and ...
Jul 28, 2021 · It further allows a minimum estimate of the volume of ∼1.45 × 106 km3 for erupted magmas and ∼2.39 × 106 km3 for total emplaced magmas, which is ...
[79]
(PDF) Magmatic duration of the Emeishan large igneous province
Feb 8, 2020 · The eruption of Emeishan lava in southwestern China and northern Vietnam is considered to be a contributing factor to the Capitanian mass ...
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[PDF] Implications for the termination age of Emeishan la - CORE
Mar 12, 2014 · The study used CA-TIMS zircon U-Pb dating on felsic ignimbrite, yielding a 259.1 ± 0.5 Ma age, interpreted as the termination age of Emeishan ...
[81]
New timing and geochemical constraints on the Capitanian (Middle ...
Mar 15, 2019 · Our dating shows that a crisis amongst radiolarian and a subsequent productivity decline occurred during the middle Capitanian, prior to the ...Missing: Baoshan | Show results with:Baoshan
[82]
Surface-wave tomography of the Emeishan large igneous province ...
May 17, 2021 · It has been proposed that the environmental impacts are primarily related to sill emplacement. Therefore, the structure of LIPs' magma storage ...
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(PDF) Recycled carbon degassed from the Emeishan plume as the ...
Massive CO2 and SO2 degassed from large igneous provinces (LIPs) are thought to cause ocean acidification and calcification crisis with a significant loss of ...
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86 Sr ratio in the Capitanian (Permian) mid-oceanic carbonates: A ...
The initial rifting of Pangea probably started in the Capitanian, and the rifting-induced new connection of many intra-supercontinental drainage systems to the ...