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Lopingian

The Lopingian epoch, also referred to as the Lopingian Series, is the final chronostratigraphic division of the Permian Period and the Paleozoic Era, spanning approximately 259.51 ± 0.21 million years ago (Ma) to 251.902 ± 0.024 Ma.^[1] It is subdivided into two stages: the Wuchiapingian (259.51 ± 0.21 to 254.14 ± 0.07 Ma) and the Changhsingian (254.14 ± 0.07 to 251.902 ± 0.024 Ma).^[1] Named after the Loping (Laoping) region in Guangxi Province, southern China, where the marine Permian strata were first extensively studied, with the Global Stratotype Section and Point (GSSP) at the Penglaitan section near Laibin, the Lopingian marks a critical interval in Earth's history characterized by the assembly of the supercontinent Pangaea and culminating in the most devastating mass extinction event recorded.^[2]^[3] During the Lopingian, the global paleogeography was dominated by the vast supercontinent Pangaea, which extended from polar to equatorial latitudes and was surrounded by the superocean Panthalassa, with the Paleo-Tethys Ocean to the east.^[4] Sedimentary records from this epoch include widespread evaporites, red beds, and coal measures in continental settings, reflecting a shift from earlier glacial influences toward more arid conditions in the continental interiors.^[4] Marine environments featured extensive carbonate platforms and deep-water basins, with key fossil-bearing sections preserved in South China, the Salt Range of Pakistan, and the Russian Platform.^[5] The epoch's boundaries are defined by Global Stratotype Sections and Points (GSSPs) based on conodont biostratigraphy: the base at the first appearance of Clarkina postbitteri postbitteri at the Penglaitan section in South China, and the top at the Permian-Triassic boundary marked by the first appearance of Hindeodus parvus at the Meishan section in South China.^[3]^[6] Paleoclimatic conditions in the Lopingian transitioned from relatively cooler, humid phases in the early part to increasingly hot and dry regimes, particularly in the equatorial regions of Pangaea, with evidence of monsoonal circulation driving seasonal aridity.^[4] Oxygen isotope data from brachiopod shells indicate warming trends, while the decline in glacial deposits signals the end of the late Paleozoic ice age.^[7] Volcanic activity, including the formation of the Siberian Traps toward the epoch's close, contributed to environmental perturbations such as ocean anoxia and carbon cycle disruptions.^[5] Lopingian biota were diverse but showed signs of stress, with marine ecosystems dominated by fusulinid foraminifers, rugose and tabulate corals, brachiopods, ammonoids, and conodonts, many of which peaked in abundance before declining.^[5] On land, gymnosperms such as conifers and ginkgophytes formed extensive forests, while therapsid synapsids and early archosauromorph diapsids represented the dominant vertebrates; insects and early lissamphibians also proliferated.^[4] A pre-Lopingian (Capitanian) biodiversity bottleneck at the Guadalupian-Lopingian boundary preceded the epoch's end, but the defining event was the end-Permian mass extinction at the end of the Changhsingian, which eradicated about 95% of marine species and 70% of terrestrial vertebrate genera, fundamentally reshaping global ecosystems and paving the way for Mesozoic dominance by dinosaurs and mammals.^[5]^[4]

Geological Context

Temporal Boundaries

The Lopingian epoch spans an absolute age range of 259.51 ± 0.21 Ma to 251.902 ± 0.024 Ma, as established by the International Chronostratigraphic Chart.^[8] This interval represents a duration of approximately 7.6 million years, calibrated through radiometric dating integrated with biostratigraphic markers.^[8] The lower temporal boundary, the Guadalupian-Lopingian boundary (GLB), occurs at 259.51 ± 0.21 Ma and is defined by the Global Stratotype Section and Point (GSSP) at the base of Bed 6k in the Penglaitan section, Laibin, Guangxi Province, South China.^[9] This boundary is formally recognized by the first appearance datum (FAD) of the conodont *Clarkina postbitteri postbitteri* within a chronomorphocline of Clarkina species in a continuous carbonate succession.^[9] The precise age derives from U-Pb dating of zircon grains extracted from volcanic ash beds near the boundary, including tuffs linked to the waning phase of the Emeishan Large Igneous Province.^[10] The upper boundary aligns with the Permian-Triassic boundary (PTB) at 251.902 ± 0.024 Ma, coinciding with the end-Permian mass extinction, and is defined by the GSSP at the base of Bed 27c in the Meishan D section, Changxing County, Zhejiang Province, South China.^[11] Here, the boundary is marked by the FAD of the conodont Hindeodus parvus in a cherty limestone bed.^[11] High-precision geochronology, utilizing U-Pb isotope analysis on zircons from intercalated ash beds in South China sections, provides the refined age estimate for this critical transition.^[12]

Relation to Permian Period

The Lopingian represents the uppermost epoch and series of the Permian Period, spanning the final approximately 8 million years of the Paleozoic Era, from about 259.5 Ma to 251.9 Ma.^[8] It directly succeeds the Guadalupian Series and forms the late phase of the Permian, which is formally divided into the early Cisuralian Series and the late Guadalupian-Lopingian super-series in the global chronostratigraphic framework.^[8] This positioning underscores the Lopingian's role as the terminal subdivision of the Permian System, encapsulating the period's concluding stratigraphic interval before the transition to the Mesozoic Era.^[2] The Lopingian's significance lies in its stratigraphic placement at the apex of Permian geological history, marking the culmination of biotic and environmental developments prior to the Permian-Triassic mass extinction event, which eliminated over 90% of marine species and signaled the end of the Paleozoic. During this epoch, the Permian reached a peak in the diversification of key groups such as fusulinid foraminifers and rugose corals in marine settings, alongside notable terrestrial ecosystem complexity, before the onset of the extinction's severe impacts.^[13] As defined in the International Chronostratigraphic Chart, the Lopingian is ratified as a formal epoch equivalent to a series, providing a standardized global correlation tool for late Paleozoic rocks through biostratigraphic markers like conodonts and ammonoids.^[8] Historically, the Lopingian originated from Chinese stratigraphy, with the term proposed by Huang Teck-kwang in 1932 to describe upper Permian marine successions in the Loping (Longtan) area of southern China, predating North American equivalents like the Ochoan Series.^[14] It was integrated into Russian schemes as part of the broader Uralian framework, but global unification occurred in the 1990s through efforts of the Subcommission on Permian Stratigraphy, culminating in the 1994 proposal by Jin et al. for a four-series Permian scale that formalized the Lopingian alongside the Cisuralian, Guadalupian, and earlier divisions.^[2] Subsequent ratifications of its global stratotype sections and points (GSSPs)—for the Wuchiapingian Stage in 2005 and the Changhsingian in 2007—solidified its international status by the International Commission on Stratigraphy.^[15]

Nomenclature and Stratigraphy

Etymology and Type Locality

The Lopingian Series derives its name from the Loping (now Leping) area in Jiangxi Province, southeastern China, where Late Permian strata were initially studied and described in the early 20th century.^[9] The term was first proposed by American paleontologist Amadeus W. Grabau in 1923 as the "Loping Series," referring to a distinctive Late Permian lithostratigraphic unit characterized by marine limestones underlying the Changhsing Limestone in South China.^[9] In 1932, Chinese geologist Huang T.K. elevated it to formal series rank as the Lopingian Series, emphasizing its significance in regional stratigraphy as the uppermost division of the Permian System.^[9] This naming reflected the rich fossiliferous successions in the Leping region, which provided key insights into Upper Permian marine faunas and sedimentation.^[16] The Lopingian was initially a regional chronostratigraphic unit developed by Chinese geologists during the 1930s and 1940s, based on biostratigraphic correlations using fusulinids, ammonoids, and brachiopods from South Chinese sections.^[16] Its integration into the global timescale occurred in the mid-1990s, when the Subcommission on Permian Stratigraphy of the International Commission on Stratigraphy (ICS) voted in 1995 to adopt the Lopingian as the international standard for the uppermost Permian series, replacing competing terms like Dzhulfian or Ochoan due to the superior continuity and biostratigraphic utility of South Chinese reference sections.^[17] This decision was formalized in proposals such as Jin et al. (1997), which outlined the three-fold Permian division into Cisuralian, Guadalupian, and Lopingian series.^[16] The Global Stratotype Section and Point (GSSP) defining the base of the Lopingian Series—and thus the Guadalupian-Lopingian boundary (GLB)—is situated at the Penglaitan section along the Hongshui River, approximately 20 km east of Laibin in Guangxi Zhuang Autonomous Region, South China (coordinates: 23°41′43″N, 109°19′16″E).^[3] Ratified by the ICS in 2004 and published in 2006, this GSSP is marked by the first appearance datum (FAD) of the conodont Clarkina postbitteri postbitteri at the base of Bed 6k, within the upper member of the Maokou Formation (Laibin Limestone).^[9] The section exposes a continuous, fossil-rich succession of marine carbonates and cherts spanning the GLB, including thick-bedded crinoidal grainstones, lenticular packstones, and overlying Heshan Formation limestones (up to 250 m thick), which preserve detailed conodont biostratigraphy for global correlation.^[3] This designation underscores the Penglaitan site's exceptional stratigraphic completeness and minimal hiatuses, making it the primary reference for the Lopingian base.^[9]

Age Subdivisions

The Lopingian Epoch is subdivided into two chronostratigraphic stages: the lower Wuchiapingian Stage and the upper Changhsingian Stage.^[8] The Wuchiapingian spans from 259.51 ± 0.21 Ma to 254.14 ± 0.07 Ma, lasting approximately 5.4 million years, while the Changhsingian extends from 254.14 ± 0.07 Ma to 251.902 ± 0.024 Ma, with a duration of about 2.2 million years.^[8] These subdivisions provide a framework for correlating Late Permian rocks globally based on biostratigraphic and geochronologic criteria.^[8] The base of the Wuchiapingian Stage, which also marks the base of the Lopingian, is defined by the Global Stratotype Section and Point (GSSP) at the base of Bed 6k in the Penglaitan Section, Laibin City, Guangxi Zhuang Autonomous Region, South China.^[3] This boundary is delineated by the first appearance datum (FAD) of the conodont subspecies Clarkina postbitteri postbitteri.^[3] The section exposes a continuous sequence of cherty limestones and shales from the Heshan Formation, offering excellent preservation for biostratigraphic analysis.^[3] The base of the Changhsingian Stage is defined by the GSSP at 88 cm above the base of the Changxing Limestone in the Meishan Section D, Changxing County, Zhejiang Province, South China.^[18] This horizon corresponds to the FAD of the conodont species Clarkina wangi within the evolutionary lineage from Clarkina changxingensis.^[18] The Meishan sequence, renowned for its well-exposed Permian-Triassic boundary strata, facilitates precise correlation across the Wuchiapingian-Changhsingian transition.^[18] Global correlation of these stages relies primarily on conodont biozonations, supplemented by fusulinid foraminifers and ammonoids.^[17] Conodonts serve as the principal markers due to their abundance, rapid evolution, and wide geographic distribution in marine sediments, enabling high-resolution zonations such as the Clarkina postbitteri Zone for the lower Wuchiapingian and the Clarkina wangi Zone at the base of the Changhsingian.^[19] Fusulinids, like Neoschwagerina and Colaniella, provide additional constraints in shallow-marine carbonates, while ammonoids such as Paleopfaffia and Otoceras aid in interregional ties, particularly near the stage boundaries.^[17]

Paleogeography

Continental Configurations

During the Lopingian epoch, the supercontinent Pangaea dominated global paleogeography, having achieved full assembly by the late Permian as a result of the ongoing collision between Laurussia and Gondwana. This configuration positioned the bulk of Pangaea's landmasses in equatorial to mid-latitudinal belts, spanning approximately 40°N to 40°S, which contributed to widespread aridification in the continental interior due to the supercontinent's equatorial orientation and limited moisture transport.^[13]^[20] Key components of Pangaea included the fused blocks of Laurentia (modern North America) and Gondwana (encompassing the southern continents such as South America, Africa, India, Antarctica, and Australia), forming the primary northern and southern extents, respectively, while Euramerica occupied the northern tropical to subtropical regions around 15–20°N. Siberia remained a distinct cratonic block, positioned at mid-northern latitudes (approximately 28–40°N) and drifting toward the main Pangaean mass, separated by the narrowing Uralian seaway but not yet fully integrated until the Early Triassic. Gondwana, in particular, extended into higher southern latitudes up to 60–65°S, influencing regional floral and faunal distributions.^[13]^[21] Tectonically, the Lopingian marked a phase of relative stability across Pangaea's interior, with minimal rifting or internal deformation, as the supercontinent had largely completed its assembly. Subduction was concentrated along the peripheral margins, forming a nearly continuous circum-Pangaean subduction girdle that encircled the landmass and isolated the underlying mantle thermally, driving limited volcanic arc development without significant continental breakup. This stability persisted as Pangaea underwent a gradual northward migration, stabilizing its axis of symmetry in the equatorial plane by approximately 250 Ma.^[20]^[13] Paleomagnetic studies of volcanic and sedimentary rocks provide primary evidence for these configurations, yielding paleopoles that confirm the equatorial to mid-latitudinal positions of major cratons, such as a Late Permian pole for stable South America aligning with Laurentia's position in a Pangea A reconstruction. Sedimentary paleocurrents from fluvial and deltaic deposits further support the continental orientations, indicating sediment transport directions consistent with the fused Laurentia-Gondwana core and Siberia's proximal but separate placement.^[22]^[13]

Marine Basins and Sedimentation

During the Lopingian epoch, the global marine realm was dominated by three principal basins that shaped oceanic circulation and sedimentation. The Paleo-Tethys Ocean, situated along the eastern margins of Pangaea and Eurasia, underwent progressive deepening accompanied by recurrent anoxic events, fostering the accumulation of organic-rich mudstones and black shales in its deeper realms. These conditions were exacerbated by restricted water exchange and high productivity, leading to oxygen-depleted bottom waters across much of the basin. In contrast, the nascent Neo-Tethys Ocean began to widen as Cimmerian continental fragments rifted from northern Gondwana, initiating rift-related volcanism and the formation of nascent spreading centers that influenced early sedimentary infill with basaltic and carbonate deposits. Encompassing Pangaea to the north, west, and south, the expansive Panthalassa superocean served as the primary deep-water repository, where siliceous oozes from radiolarian blooms contributed to widespread chert formation amid generally oxic but episodically anoxic deep-sea environments. Sedimentation patterns varied markedly by basin depth and proximity to landmasses. Shallow sectors of the Paleo-Tethys, including platforms around the South China cratonic blocks, hosted expansive carbonate platforms with skeletal limestones, reefs, and lagoons that supported diverse benthic communities before declining toward the epoch's end. Deeper, open-ocean domains of Panthalassa and the Paleo-Tethys abyssal plains accumulated bedded cherts and fine-grained shales, reflecting biogenic silica precipitation under stable, low-energy conditions with minimal clastic input. Marginal and intra-shelf basins along Tethyan borders, subject to episodic isolation due to tectonic uplift and eustatic controls, saw the precipitation of evaporites such as gypsum and halite in hypersaline lagoons and sabkhas, driven by arid climates and restricted circulation that promoted brine concentration. Prominent stratigraphic units exemplify these patterns. The Longtan Formation in southern China records transitional marine to paralic sedimentation, featuring coal seams interbedded with phosphorite nodules and shales in deltaic and tidal-flat settings, indicative of nutrient-enriched coastal waters. Similarly, the Phosphoria Formation across the western United States preserves a vast phosphate province in a semi-restricted epicontinental sea, with nodular phosphorites and cherty limestones deposited via upwelling of nutrient-laden waters from Panthalassa. These formations highlight localized productivity hotspots amid broader basin dynamics. Sea-level dynamics played a pivotal role, with a relative highstand prevailing in the early Lopingian (Wuchiapingian stage) that expanded epeiric seas over continental shelves and facilitated platform aggradation, followed by a pronounced regression in the late Lopingian (Changhsingian stage) that exposed vast areas to subaerial erosion and progradation of terrigenous clastics toward the Permian-Triassic boundary. Facies analysis from global reference sections, including those in South China and the Tethyan margins, documents these shifts through lateral transitions from offshore carbonates to nearshore siliciclastics and evaporites, underscoring the interplay of eustasy, tectonics, and climate in controlling depositional geometries along Pangaea's flanks.

Paleoclimate and Environment

Climatic Conditions

The Lopingian epoch was characterized by a global greenhouse climate, with warm temperatures prevailing across latitudes and minimal polar glaciation by its latter stages. Equatorial sea surface temperatures were approximately 5–10°C higher than modern values, reaching up to 32–35°C in low-latitude regions, as inferred from oxygen isotope analyses of brachiopod shells preserved in Tethyan sediments. This warmth extended poleward, with high-latitude temperatures in Gondwana and northern Pangaea estimated at 10–15°C, supporting the near-complete disappearance of continental ice sheets following the Late Paleozoic Ice Age. Paleotemperature reconstructions from conodont apatite oxygen isotopes in South China sections further confirm a progressive warming trend through the epoch, with seawater temperatures rising by about 4°C from the early to late Lopingian.^[23]^[24]^[25] Atmospheric CO₂ concentrations were elevated during the Lopingian relative to pre-industrial levels, with estimates varying between proxies from approximately 400 to 1,000 ppm; this range is inferred from paleosol carbonate profiles across Pangaean continents as well as carbon isotope ratios in marine carbonates.^[26]^[27] These levels contributed to the greenhouse conditions. Paleosol evidence from interior North America and Europe reveals pedogenic features consistent with elevated CO₂ driving soil respiration and carbonate formation.^[26] Precipitation patterns exhibited strong regional contrasts influenced by Pangaea's configuration, with arid to semi-arid conditions dominating the supercontinent's interior due to its position in subtropical high-pressure belts. Red bed deposits and eolian dunes in the North American midcontinent and European basins indicate low rainfall, often less than 500 mm annually, fostering evaporative sedimentation and soil calcretization as documented in paleosol profiles. In contrast, the eastern margins of Pangaea, particularly in South China, experienced monsoonal regimes with seasonal heavy precipitation, evidenced by extensive coal swamps and fluvial-deltaic sediments in the Yangtze platform, where annual rainfall likely exceeded 1,000 mm supporting lush Cathaysian flora. These patterns reflect a megamonsoonal circulation driven by land-sea thermal contrasts across the vast continent.^[28]^[29] Ocean circulation during the Lopingian was sluggish and regionally restricted owing to the near-closure of equatorial seaways between Laurussia and Gondwana, limiting inter-basin exchange and promoting water column stratification. Model simulations of Pangaean paleogeography show weakened thermohaline circulation, with reduced deep-water formation in the Panthalassic Ocean and stagnant conditions in marginal basins like the Tethys, leading to oxygen-depleted bottom waters as traced by trace element geochemistry in black shales. Oxygen isotope data from brachiopod shells in these basins corroborate stable, warm surface waters with minimal vertical mixing, exacerbating anoxic tendencies without invoking later perturbations.^[30]^[31]^[32]

Environmental Perturbations

The Lopingian Epoch was marked by significant environmental disturbances, most notably the onset of massive volcanism associated with the Siberian Traps Large Igneous Province in the late Changhsingian substage around 251.9 Ma. This event involved the rapid emission of approximately 36,000 Gt of carbon primarily as volcanic CO₂ over a duration of about 168 kyr, with peak rates reaching 4.5 Gt C yr⁻¹ in short pulses. Concurrent releases of SO₂ from interactions between basaltic magmas and evaporite layers contributed to atmospheric aerosol loading. These emissions drove profound climatic shifts, including a rise in atmospheric pCO₂ from ~440 ppm to ~7,390 ppm and global temperatures from ~25°C to ~40°C, alongside a ~1.1-unit drop in ocean pH that intensified acidification and promoted marine anoxia.^[33] Geochemical records reveal prominent negative excursions in carbon isotopes (δ¹³C) during the Lopingian, signaling disruptions to the global carbon cycle. At the Guadalupian-Lopingian boundary (GLB), δ¹³C values in carbonates and organic matter shifted negatively by up to ~4‰ in some South China sections, likely reflecting enhanced productivity from green sulfur bacteria under expanding photic-zone euxinic conditions rather than widespread methane injection. A more pronounced negative δ¹³C excursion occurred at the Permian-Triassic boundary (PTB) in the latest Changhsingian, with organic δ¹³C declining by ~3.5‰ (e.g., from -25‰ to -28.5‰) and carbonate δ¹³C by ~4.4‰, accompanied by sharp drops in total organic carbon (TOC) from ~6% to ~1%. These shifts indicate a combination of methane release from destabilized clathrates due to volcanic warming and a collapse in primary productivity, exacerbating ecological stress.^[34]^[35] Oceanic anoxia expanded markedly during the Lopingian, particularly in the Changhsingian, as evidenced by widespread deposition of organic-rich shales in intrashelf basins such as the NE Sichuan Basin, where TOC contents reached up to 13.7 wt% (mean ~2.4 wt%). Iron speciation (FeHR/FeT >0.38) and trace metal enrichments (e.g., Mo up to 55 ppm, U up to 21 ppm) document the development of ferruginous to euxinic bottom waters, with oxygen minimum zones (OMZs) intensifying in moderate water depths due to sluggish circulation and high organic flux from upwelling. This anoxic expansion predated the PTB mass extinction and contributed to habitat compression in marine settings.^[36] Sea-level fluctuations characterized the Lopingian, culminating in a notable regression during the Changhsingian that reduced global shallow marine areas from ~43% in the Early Permian to ~13%. In the Upper Yangtze region, this is recorded by paleo-karst surfaces and exposure features in the Changxing Formation, indicating episodic lowstands. The regression stemmed from regional tectonic uplift linked to Emei Taphrogenesis rifting, compounded by ice-free global conditions following the Late Paleozoic Ice Age, which limited eustatic compensation and amplified platform exposure.^[37] Additional perturbations included the development of euxinic (sulfidic) waters in restricted basins, as inferred from elevated molybdenum concentrations (Mo/TOC ratios >10) and iron proxies in Lopingian shales of the Dalong Formation. These conditions, prevalent in silled intrashelf settings like the NE Sichuan Basin, arose from water-column stratification and volcanic nutrient inputs, fostering H₂S accumulation that further stressed marine ecosystems.^[36]

Biodiversity and Events

Marine and Terrestrial Life

The Lopingian epoch featured marine ecosystems dominated by fusulinid foraminifers, which, though reduced from earlier Permian peaks, persisted with small-sized genera such as Codonofusiella, Reichelina, and Palaeofusulina in Tethyan realms.^[38] Rugose and tabulate corals, including the Waagenophyllinae subfamily, were prominent in the Palaeoequatorial Tethys, contributing to reef frameworks, but their diversity declined sharply after the early Lopingian, with only 68 species and 20 genera recorded in the Changhsingian stage of China.^[38]^[39] Brachiopods, particularly productid forms, formed diverse assemblages associated with these corals and foraminifers, while ammonoids and conodonts (Clarkina species) provided key biostratigraphic markers across shallow marine basins.^[38] Reef ecosystems reached a relative peak in the early Lopingian before a progressive decline, driven by environmental stresses, transitioning to microbial-dominated structures in later stages.^[39] Terrestrial life during the Lopingian was characterized by synapsid reptiles, particularly therapsids such as dicynodonts, which comprised up to 80% of faunal assemblages in regions like the Junggar Basin of China and dominated Gondwanan ecosystems.^[13] Gorgonopsians, therocephalians, and early cynodonts added to therapsid diversity, with examples including Endothiodon in Brazil and Jimusaria in Asia, reflecting adaptations to varied woodlands.^[13] Early archosaurs, including archosauriforms like Archosaurus in Russia, appeared rarely in low-latitude assemblages, signaling the onset of sauropsid radiation.^[13] Floral communities varied by phytoprovince: glossopterid gymnosperms dominated Gondwanan flora in southern high latitudes, co-occurring with conifers and lycopods in temperate settings, while conifers prevailed in Euramerican lowlands.^[40]^[13] Biodiversity trends showed high marine diversity in the Tethys, where fusulinid genera maintained presence despite a post-Guadalupian decline, with over 100 species documented in South Chinese assemblages.^[41] Terrestrial tetrapod richness exhibited a poleward decline, with tropical low-latitude sites like the Bletterbach Biota in Italy hosting the most diverse ecosystems, including integrated plant-animal communities.^[13] Red beds in regions such as southern Brazil and the Venetian Prealps preserved tetrapod tracks, including Dicynodontipus ichnofacies indicative of dicynodont and parareptile activity in eolian and alluvial environments.^[42]^[43] Key examples include pareiasaurs, which were abundant in Russian localities like Kotel'nich, comprising 14-19% of assemblages and reaching sizes over 3 meters in species such as Scutosaurus karpinskii.^[13]^[44] In Cathaysia, gigantopterid flora, represented by genera like Gigantopteris, formed lowland rainforests with large, reticulate-veined leaves, co-occurring with conifers in subtropical settings.^[45] Fossil evidence derives from global assemblages, including the Karoo Basin in South Africa for Gondwanan therapsids and glossopterids, the Zechstein Basin in Europe for conifer-dominated woodlands, and Tethyan sections like Penglaitan in China for marine biotas, preserved in museum collections worldwide.^[13]^[38]

Mass Extinctions

The Lopingian Epoch was bracketed by two significant mass extinction events, the first marking its onset and the second its close. The Guadalupian-Lopingian, or Capitanian, extinction occurred around 260 Ma during the late Capitanian Stage of the Middle Permian, resulting in substantial biodiversity losses estimated at 58% of marine skeletal genera and 24–56% of plant species across regions like South and North China.^[46] This event disproportionately affected marine groups such as fusulinacean foraminifers, rugose and tabulate corals, bryozoans, brachiopods, and ammonoids, while terrestrial impacts included the extinction of up to 80% of tetrapod genera in the Karoo Basin of South Africa, particularly dinocephalian therapsids and pareiasaurs.^[46]^[47] Evidence from fossil records shows taxonomic turnover, with mercury concentration spikes in sediments indicating intense volcanism from the Emeishan Large Igneous Province as a primary driver, potentially triggering environmental perturbations like acidification and atmospheric disruption.^[48] No iridium anomalies suggestive of bolide impact were observed, reinforcing a volcanic origin.^[48] Selectivity in the Capitanian extinction exhibited patterns of size bias in marine realms, where larger-bodied or heavily calcified invertebrates with limited respiratory capabilities suffered higher losses due to ocean chemistry changes, and on land, large herbivores exceeding 2.5 meters in length—such as certain dicynodonts and pareiasaurs—were particularly vulnerable, possibly owing to dietary disruptions and aridification.^[49]^[50] Following this event, ecosystems showed brief signs of recovery during the early Lopingian Wuchiapingian Stage, with rebuilding of trophic structures and refilling of ecological guilds to about nine functional groups in some regions, as evidenced by increased brachiopod diversity and Lazarus taxa reappearances in fossil assemblages.^[51] However, this recovery was stalled by the subsequent end-Permian crisis, preventing full ecological stabilization.^[51] The end-Permian mass extinction at the Permian-Triassic boundary (PTB), dated precisely to 251.9 Ma, stands as the most severe biotic crisis in Earth history, eliminating over 90% of marine species—including the total extinction of trilobites, blastoids, and most rugose and tabulate corals—and approximately 70% of terrestrial vertebrate families, alongside major losses in glossopterid floras and insect orders.^[51]^[51] Causative factors centered on massive Siberian Traps flood basalt eruptions, which released greenhouse gases leading to global warming of about 10°C, widespread marine anoxia affecting up to 80% of ocean oxygen levels, and acidification that devastated calcifying organisms.^[52]^[52] Selectivity mirrored earlier patterns, with marine losses favoring larger or high-oxygen-demand species in deeper waters and tropical latitudes, while terrestrial impacts heavily targeted herbivores through collapsed food webs and habitat loss.^[49] Fossil evidence includes abrupt taxonomic turnovers in sections like Meishan, China, accompanied by mercury spikes tracing Siberian volcanism, but again without iridium enrichment to indicate extraterrestrial influence.^[48]

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[PDF] Permian chronostratigraphic ...
The Lopingian Series comprises two stages， Wuchiapingian. and Changhsingian． Zhao et al．（1981） formally proposed the D See－. tion in Meishan of Changxing ...
[16]
The Permian chronostratigraphic scale: history, status and prospectus
Jan 1, 2018 · Huang (1932) used the Loping as a series to refer to all Chinese Permian strata above the Maokou Formation.
[17]
GSSP for Wuchiapingian Stage
The Base of the Lopingian Series and Wuchiapingian Stage is defined at the base of Bed 6k in the Penglaitan Section along the Hongshui River in Guanxi Province, ...Missing: Chaotian | Show results with:Chaotian
[18]
GSSP for Changhsingian Stage
The section is located at the Meishan Section D, Changxing County, Zhejiang Province between Nanjing and Shanghai, China at a latitude of 31°4'55"N and a ...Missing: wuchiapingensis Wuchiaping
[19]
Revised Wuchiapingian conodont taxonomy and succession of ...
Sep 15, 2017 · Based on the Penglaitan, Dukou, and Nanjiang sections, seven conodont zones (Clarkina postbitteri postbitteri, C. dukouensis, C. asymmetrica, C.Missing: postbitterii | Show results with:postbitterii
[20]
Pangea Migration - Le Pichon - 2021 - Tectonics - Wiley Online Library
May 17, 2021 · A subduction girdle is predicted to set up lateral temperature gradients from the relatively warm sub-Pangean mantle to a cooler sub-oceanic ...
[21]
An early Pangaean vicariance model for synapsid evolution - Nature
Aug 4, 2020 · The Pangaea supercontinent was an assemblage of continental plates that formed a single landmass during the Late Palaeozoic. The most important ...
[22]
A new Late Permian paleomagnetic pole for stable South America: The Independencia group, eastern Paraguay - Earth, Planets and Space
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[23]
A high-resolution Middle to Late Permian paleotemperature curve ...
Additionally, climate warms further as evidenced by conodont oxygen isotopes 48 . ... OXYGEN ISOTOPIC EVIDENCE FOR CLIMATE CHANGE ACROSS THE PERMIAN-TRIASSIC ...
[24]
Carbon and conodont apatite oxygen isotope records of ...
Aug 6, 2025 · Chen et al. (2011a) applied oxygen isotopes of conodonts to estimate a 4 °C increase in seawater temperature during the J. postserrata to the J.
[25]
Oxygen isotope values from high-latitudes: Clues for Permian sea ...
The transition from ice- to greenhouse conditions is reflected also in the oxygen isotope data of unaltered brachiopods and bivalves from high high-latitudes.Missing: epoch | Show results with:epoch
[26]
Low atmospheric CO2 levels during the Permo - PNAS
Here, I report 24 quantitative Carboniferous and Permian atmospheric CO 2 estimates derived from the stomatal characteristics of fossil arborscent lycopsids.
[27]
Atmospheric CO2 concentrations during ancient greenhouse ...
The CO2 paleobarometer suggests that [CO2]atm values exceeded 3,000 parts per million by volume (ppmV) during Permian (289–251 Ma) and Mesozoic (251–65 Ma) ...
[28]
A 300-million-year record of atmospheric carbon dioxide from fossil ...
Reported values range from less than 100 ppm in the Late Permian to more than 1700 ppm in the Early Triassic, according to the range of 95% confidence interval ...
[29]
Climate of the Supercontinent Pangea | The Journal of Geology
Numerous climate models predict that the geography of the supercontinent Pangea was conducive to the establishment of a "megamonsoonal" circulation.
[30]
Provenance of Permian eolian and related strata in the North ...
Dec 28, 2018 · Permian red beds of the southern midcontinent archive an especially rich record of the Permian of western equatorial Pangea. Depositional ...
[31]
Simulated warm polar currents during the middle Permian - Winguth
Oct 17, 2002 · The land-sea distribution during the Permian (∼290 to 252 Ma years ago) was characterized by a supercontinent Pangaea with an almost closed ...<|control11|><|separator|>
[32]
Changes of the Late Permian Ocean Circulation and Deep-Sea ...
By the end of the Late Permian, most continents had collided to form the supercontinent of Pangea. The associated climatic changes at the Permian–Triassic ...Missing: closure | Show results with:closure
[33]
Permian Period - Climate, Extinction, Carboniferous | Britannica
As low-latitude seaways closed, warm surface ocean currents were deflected into much higher latitudes (areas closer to the poles), and cool-water upwelling ...Missing: stratified | Show results with:stratified
[34]
Massive and rapid predominantly volcanic CO2 emission during the ...
Sep 7, 2021 · We find that a massive and rapid, predominantly volcanic CO 2 emission during the Siberian Traps volcanism is likely the trigger for the carbon isotope ...
[35]
High-resolution paired C-isotope variations during the Guadalupian linked to paleo-redox changes in South China
### Summary of Negative δ13C Shift at GLB and Causes
[36]
Stable carbon isotope stratigraphy across the Permian–Triassic boundary in shallow marine carbonate platforms, Nanpanjiang Basin, south China
### Summary of Negative δ13C at PTB, Methane Release, and Productivity Collapse
[37]
Marine redox evolution and organic accumulation in an intrashelf basin, NE Sichuan Basin during the Late Permian
### Evidence for Lopingian Anoxia and Organic-Rich Shales
[38]
Sedimentary records of sea level fall during the end-Permian in the ...
May 30, 2024 · From the Early Permian to Changhsingian, the global shallow marine area decreased sharply from 43 % to 13 %, but it increased again to 34 % in ...
[39]
The Permian timescale: an introduction - Lyell Collection
The most important taxa in this regard are conodonts, fusulinids, non-fusulinid forams and ammonoids. Brachiopods, rugose corals and radiolarians also provide ...Missing: epoch | Show results with:epoch
[40]
Diversity and extinction patterns of the Permian corals in China
Aug 6, 2025 · Many marine benthic organisms, such as fusulinid foraminifera, brachiopods, bivalves, and corals, experienced substantial extinctions during ...<|separator|>
[41]
Refined Permian–Triassic floristic timeline reveals early collapse ...
Nov 14, 2019 · The collapse of late Permian (Lopingian) Gondwanan floras, characterized by the extinction of glossopterid gymnosperms, heralded the end of one of the most ...
[42]
Global cooling drove diversification and warming caused extinction ...
Jun 20, 2025 · The subsequent Guadalupian crisis severely affected fusuline species diversity, and the species diversity in the Lopingian is so low that no ...
[43]
Tetrapod tracks in Permo–Triassic eolian beds of southern Brazil ...
May 18, 2018 · Here, we describe the first tetrapod tracks from the Lopingian–Induan eolian strata of southern Brazil, which are identified as Dicynodontipus ...
[44]
[PDF] Lopingian tetrapod footprints from the Venetian Prealps, Italy
Nov 7, 2017 · Introduction. The Lopingian (late Permian) was a key epoch for the evo- lution of terrestrial vertebrates, which reached a noteworthy abundance ...
[45]
Volumetric Body Mass Estimate and in vivo Reconstruction of the ...
Currently four Russian pareiasaur species are recognized. The large and derived Scutosaurus karpinskii (2.5–3 m long) was the first named Russian pareiasaur ( ...
[46]
Floral dynamics and ecological adaptations in the Lopingian ...
Gigantopterids represents an enigmatic plant plexus from the Permian period and are of utmost significance in South China during the Lopingian prior to the ...
[47]
https://doi.org/10.1098/rspb.2015.0834
[48]
https://doi.org/10.3389/feart.2021.651224
[49]
https://doi.org/10.1017/ext.2023.10
[50]
https://doi.org/10.3389/feart.2021.631198
[51]
https://doi.org/10.1098/rspb.2007.1370
[52]
https://sustainability.stanford.edu/news/what-caused-earths-biggest-mass-extinction
[53]
What caused Earth's biggest mass extinction?
Dec 6, 2018 · New research shows the "Great Dying" was caused by global warming that left ocean animals unable to breathe.<|control11|><|separator|>