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Carnian

The Carnian is the lowermost chronostratigraphic stage of the Upper Triassic Series in the geologic time scale, spanning approximately 237 to 227.3 million years ago (Ma).^[1] It is defined at its base by the first appearance of the ammonoid species Daxatina canadensis in the Stuores Wiesen section of the Italian Dolomites, marking a global biostratigraphic boundary.^[2] During the Carnian, Earth's continents were assembled into the supercontinent Pangaea, with paleogeography featuring widespread arid interiors punctuated by rift basins and shallow epicontinental seas.^[3] A defining event was the Carnian Pluvial Episode (CPE), a major climate perturbation around 234–232 Ma driven by massive volcanism from the Wrangellia Large Igneous Province, which triggered global humidification, increased rainfall, and temperature rises.^[4] This episode led to significant biotic turnovers, including the extinction of dominant herbivores like rhynchosaurs and the explosive diversification of early dinosaurs, which began radiating in terrestrial ecosystems amid floral shifts from gymnosperm-dominated arid communities to more humid-adapted vegetation.^[5] Marine environments saw enhanced sedimentation in deltaic and lagoonal settings, with fossil records revealing ammonoids, conodonts, and early reef-building organisms as key index fossils.^[6] The stage's end transitioned into the Norian, setting the stage for further Triassic developments leading toward the end-Triassic extinction.^[2]

Geological Context

Position within the Triassic

The Carnian stage represents the lowermost division of the Upper Triassic series within the Triassic period, immediately succeeding the Ladinian stage of the Middle Triassic and preceding the Norian stage of the Upper Triassic. This positioning establishes the Carnian as the inaugural stage of the Late Triassic epoch, marking the onset of significant evolutionary and environmental shifts that characterize the latter half of the Triassic. The stage's chronostratigraphic framework is ratified by the International Commission on Stratigraphy (ICS), providing a global standard for correlating rock successions worldwide.^[1] The temporal extent of the Carnian spans approximately 10 million years, from a base dated to about 237 Ma to a top at roughly 227.3 Ma, as delineated in the latest ICS International Chronostratigraphic Chart (version 2024/12). This duration underscores its role as a transitional interval, bridging the more stable Middle Triassic faunas—dominated by Ladinian assemblages—with the diverse and expansive Late Triassic ecosystems that emerge in the Norian and Rhaetian. The stage's boundaries are anchored in biostratigraphic events, with the lower boundary formally defined at the Global Boundary Stratotype Section and Point (GSSP) located at Prati di Stuores in the Dolomites, northern Italy, corresponding to the first appearance datum (FAD) of the ammonoid Daxatina canadensis in the San Cassiano Formation.^[7]^[8] The upper boundary, while not yet formalized by a GSSP for the overlying Norian, is conventionally placed at the FAD of Norian index ammonoids such as Klamathites macrolobatus or Stikinoceras kerri, aligning with the end of late Carnian ammonoid zones like that of Tropites welleri.^[9]^[10] This chronological placement highlights the Carnian's foundational importance in the Triassic timescale, facilitating precise dating of geological events and biotic turnovers during a period of increasing continental fragmentation and marine diversification.

Naming and historical development

The term "Carnian" derives from the Carnic Alps (Karnische Alpen), a mountain range along the Austria-Italy border where key Triassic strata were initially studied in the 19th century.^[11] This naming reflects the region's classical geological significance, particularly for exposures of ammonoid-bearing limestones that facilitated early stratigraphic work. The alternative spelling "Karnian" occasionally appears but is less common, with "Carnian" preferred based on Latin roots.^[11] The Carnian Stage was formally proposed in 1869 by Austrian paleontologist Edmund von Mojsisovics, who defined it based on distinctive ammonoid faunas, such as those containing Trachyceras aonoides, in the Hallstatt Limestones of the Eastern Alps. This initial description established the Carnian as the lowermost division of the Upper Triassic, distinguishing it from underlying Ladinian strata through biozonation of cephalopod fossils. Early correlations relied heavily on ammonoid assemblages, which provided a framework for regional mapping across Alpine sequences.^[9] Throughout the 20th century, the Carnian was integrated into a global Triassic timescale through refinements in ammonoid biostratigraphy, notably via the seminal 1895 work by Mojsisovics, Wilhelm Waagen, and Carl Diener, which outlined a comprehensive zonal scheme still influential today.^[9] Alpheus Hyatt contributed to early transatlantic correlations by studying North American ammonoids, linking them to European Carnian markers and aiding in the recognition of synchronous faunal events.^[12] These efforts solidified the stage's boundaries amid ongoing debates over substage divisions. A major milestone occurred in 2012 with the ratification of the Global Stratotype Section and Point (GSSP) for the Carnian base at Prati di Stuores (Stuores Wiesen) near Cortina d'Ampezzo, Italy, defined by the first appearance of the ammonoid Daxatina canadensis in hemipelagic beds of the Cassian Formation.^[13] This designation anchored the stage in a continuous, fossil-rich section, resolving prior ambiguities in boundary placement. Modern radiometric dating, including U-Pb zircon analyses from ash beds at the GSSP, confirms the base at approximately 237 Ma, providing precise chronological constraints that align biostratigraphic correlations with absolute time.^[13]

Stratigraphy

Lithostratigraphic subdivisions

The Carnian stage is informally subdivided into two substages based on biostratigraphic markers that are reflected in lithological transitions and facies variations observed in Tethyan sequences: the lower Carnian (Julian) and upper Carnian (Tuvalian). These divisions reflect cyclic changes in depositional environments, from platform carbonates to deeper marine settings, as established in the Southern Alps type sections.^[10] In the Tethys region, Carnian lithologies are predominantly marine, featuring platform and ramp limestones, hemipelagic shales, and evaporitic deposits that indicate fluctuating salinity and restricted basins.^[14] Volcaniclastic inputs from arc systems are common in basinal successions, as seen in the Wengen Formation of the Southern Alps, which consists of tuffaceous sandstones and shales interbedded with carbonates.^[10] In contrast, the Pangaea interior records continental red beds, fluvial sandstones, and localized volcanics, reflecting arid to semi-arid climates with episodic fluvial systems.^[14] A prominent regional example is the European Alpine sequence, where the Raibl Formation (part of the Raibl Group) exemplifies Julian to Tuvalian lithofacies with three third-order cycles of alternating carbonates (dolomites and limestones) and clastic sandstones, up to 100 m thick, indicating repeated transgressions and regressions in a carbonate platform setting.^[15] In North America, the Dockum Group represents late Carnian to early Norian continental deposits, comprising the basal Santa Rosa Sandstone (conglomeritic fluvial sands), overlain by the Tecovas Formation (red and green mudstones with pedogenic features), and topped by the Trujillo Formation (cross-bedded sandstones), spanning fluvial-lacustrine environments across the Pangaean craton.^[16] Interregional correlation of Carnian units faces challenges from pronounced lateral facies variations driven by eustatic sea-level fluctuations, which caused diachronous shifts between marine and continental deposits.^[14] Magnetostratigraphy provides a key tool for resolving these discrepancies, with composite polarity records from the Germanic Basin and Southern Alps enabling ties between Tethyan and Boreal sequences through identification of consistent reversal patterns, such as the S2n zone at the stage base.^[17]

Biostratigraphic markers

The biostratigraphy of the Carnian Stage relies primarily on marine microfossils and macrofossils to establish zonal schemes and enable global correlations, with ammonoids serving as the cornerstone for defining substages and boundaries. These biological markers allow precise subdivision of the stage into the lower Carnian (Julian Substage) and upper Carnian (Tuvalian Substage), facilitating integration with chemostratigraphic and radiometric data for worldwide stratigraphic frameworks.^[18] Ammonoids provide the most detailed biozonation for the Carnian, with the stage's base defined at the first appearance datum (FAD) of Trachyceras remannianum in the Trachyceras Zone, marking the Julian Substage. Within this zone, the GSSP at Prati di Stuores in the Southern Alps is placed at the FAD of Daxatina canadensis, a subzonal index approximately coincident with T. remannianum, occurring just above a volcanic ash dated to 237.77 ± 0.14 Ma via U-Pb zircon geochronology. The Julian Substage progresses through the Sirenites Zone, characterized by species such as Sirenites lilicus, while the Tuvalian Substage is delimited by the Tropites dilleri Zone at its base, followed by the Tropites fusiformis Zone and culminating in the Paratrachyceras longobardicum Zone. These zones exhibit high ammonoid turnover and are traceable across Tethyan and Panthalassic realms, though boreal equivalents like the Hangardites Zone in northern latitudes require careful correlation.^[19]^[18] Conodonts offer complementary high-resolution markers, particularly for the lower Carnian, where the FAD of Paragondolella polygnathiformis (formerly grouped under Neogondolella) occurs 70 cm above the GSSP level, aligning with the onset of magnetic polarity zone S2n and supporting the ammonoid-defined boundary. Throughout the stage, conodont assemblages dominated by genera like Nicoraella and Metapolygnathus enable subdivision into zones such as the N. budaensis Zone in the Julian and M. primus Zone in the Tuvalian, with diversity peaks and turnovers reflecting environmental shifts. Foraminifera contribute to finer dating in shallow-marine settings, with species like Gladigulina alpina appearing in lower Carnian assemblages alongside nodosariids, providing biostratigraphic control in Tethyan platform carbonates.^[20]^[21] Bivalve and ostracod assemblages, particularly halobiids, are key for event stratigraphy during the Carnian Pluvial Episode (CPE), a mid-stage climatic perturbation. Daonella species, such as D. frami, persist from the late Ladinian into the lowermost Carnian Trachyceras Zone, with their decline marking the Ladinian-Carnian transition, while the FAD of Halobia (e.g., H. zitteli and H. daonellaformis) in the early Julian signals the onset of humid conditions and faunal turnover. These thin-shelled, cosmopolitan bivalves form distinctive lumachelles and enable correlation of the CPE across hemispheres, often integrated with ostracod biofacies showing increased freshwater influence.^[22]^[23]^[24] Global correlations of Carnian strata combine these biozones with radiometric anchors, such as the GSSP's 237 Ma date, and extend to continental sequences via tetrapod biochronology. Recent 2020s refinements incorporate molecular clock estimates for archosauromorph divergences, calibrating late Carnian tetrapod assemblages (e.g., Adamanian land vertebrate faunachron) to approximately 230-225 Ma and aligning them with marine events like the CPE-driven dinosaur radiation. This integrated approach resolves discrepancies between Tethyan marine and Gondwanan terrestrial records, enhancing the stage's chronostratigraphic precision.^[19]^[5]^[25]

Paleoenvironment

Paleogeography

During the Carnian stage of the Late Triassic, the supercontinent Pangaea reached its maximum latitudinal extent, spanning from high northern to high southern latitudes and encompassing nearly all continental landmasses in a single configuration.^[26] Northern Laurasia, including proto-North America and Eurasia, lay to the north, while southern Gondwana, comprising South America, Africa, India, Antarctica, and Australia, occupied the southern portion, with the two separated by the narrow Paleo-Tethys Ocean that extended eastward from the equatorial region.^[27] This arrangement positioned much of Pangaea in tropical to subtropical latitudes, facilitating extensive arid conditions in continental interiors while the surrounding Panthalassa Ocean dominated the global marine realm.^[28] Key tectonic features included the initiation of rifting along the southern margin of Pangaea, particularly within the Tethys domain, which marked the early stages of Neo-Tethys basin formation as the Cimmerian terranes began drifting northward away from Gondwana.^[29] In western Pangaea, precursors to the Central Atlantic Magmatic Province (CAMP) emerged through incipient extension in rift basins, with early sedimentary fills dating to the early Carnian and signaling the onset of continental stretching that would later lead to Atlantic opening.^[30] These rifting events contributed to the development of elongated basins and fault-controlled highs across the supercontinent. Regionally, the Germanic Basin in northern Pangaea hosted epicontinental seas that accumulated evaporites, such as those in the Keuper Group, reflecting shallow-marine to sabkha environments influenced by Tethyan connections.^[14] In contrast, the interior of western North America featured arid landscapes, as preserved in paleosols of the Chinle Formation, which indicate semiarid to subhumid conditions with calcic horizons and limited fluvial activity.^[31] These basins highlight the variability in depositional settings, from marginal marine incursions in Europe to continental floodplains in North America. Overall, the Carnian witnessed a global highstand in sea level, culminating from earlier Middle Triassic transgressions and leading to widespread marine incursions into continental margins, as evidenced by prograding coastal deposits and expanded carbonate platforms around the Tethys.^[32] This was followed by a eustatic fall at the Ladinian-Carnian boundary, with variable conditions persisting through the stage.^[33]

Climate dynamics and major events

The baseline climate of the Carnian stage was characterized by hot and arid conditions in the interiors of the supercontinent Pangaea, particularly at low to mid-latitudes, with strong monsoonal influences leading to seasonal precipitation primarily along coastal margins.^[34] Oxygen isotope analyses of conodont apatite indicate that marine surface temperatures ranged from 25–30°C during this period, reflecting a generally warm global ocean.^[19] These conditions prevailed prior to the mid-Carnian, with evidence from paleosols and evaporite deposits underscoring the aridity in continental settings.^[35] A pivotal event in Carnian climate dynamics was the Carnian Pluvial Episode (CPE), a major perturbation occurring approximately 234–232 Ma in the mid-Carnian (Julian substage), lasting about 1–2 million years.^[19] This episode was triggered by massive eruptions of the Wrangellia Large Igneous Province, which released voluminous greenhouse gases and disrupted the global carbon cycle, leading to enhanced global warming, increased humidity, and widespread rainfall that penetrated even arid interiors.^[19] The CPE induced oceanic anoxia and acidification, marking a shift from the prevailing arid regime to a more humid one across both terrestrial and marine environments.^[36] Evidence for the CPE includes pronounced sedimentary changes, such as the deposition of coal beds and black shales in terrestrial and marine basins, reflecting heightened runoff and organic matter preservation under anoxic conditions.^[14]^[36] Carbon isotope excursions show negative δ¹³C spikes, indicating perturbations in the global carbon cycle tied to volcanic inputs.^[19] Biotic impacts encompassed a significant turnover, with approximately 33% of marine genera, including ammonoids and conodonts, going extinct amid these environmental stresses.^[19] By the late Carnian (Tuvalian substage), climate trends shifted toward more equable conditions, with a return to aridity in many regions and stabilization of marine ecosystems.^[37] Eustatic sea-level fluctuations, including falls on the order of tens of meters during the CPE, influenced sedimentation patterns worldwide.^[38] This transition facilitated the establishment of more stable, humid-influenced climates that set the stage for Norian developments.^[39]

Biodiversity

Marine invertebrates and microorganisms

During the Carnian, marine invertebrate faunas were dominated by ammonoids, which exhibited significant turnover and high diversity in the late stage, particularly within the Tropites subbullatus zone where genera such as Tropites formed diverse assemblages reflecting post-extinction recovery in Tethyan and Panthalassic settings.^[19] Bivalves, including the opportunistic genus Daonella, were prevalent in black shales that record dysaerobic bottom waters, with Daonella species adapting to low-oxygen conditions through thin-shelled, flat morphologies suited to stressed environments.^[40] Brachiopods and crinoids also contributed to benthic communities, though both groups suffered elevated extinction rates during the Carnian Pluvial Episode (CPE), leading to reduced standing diversity amid anoxic events.^[19] Microfossils provided key insights into paleoenvironmental gradients, with conodonts of the genus Neogondolella distributed across shallow platform interiors to deeper basinal facies in South China and Tethyan sections, enabling reconstruction of carbonate system dynamics.^[41] Foraminifera exhibited similar platform-to-basin variations, often concentrated in shallow-water carbonates while diminishing in deeper, oxygen-poor settings influenced by CPE-related humidity and runoff. Radiolarians, meanwhile, displayed elevated abundances signaling spikes in siliceous productivity, driven by enhanced nutrient delivery to pelagic realms during humid intervals of the CPE.^[42] Ecologically, reef-building sponges and calcareous algae constructed mound-like frameworks in Tethyan carbonates, with sponge-dominated reefs (e.g., involving chaetetids and sphinctozoids) thriving below fair-weather wave base on middle ramps, supporting microbial binding and early metazoan recovery. Opportunistic infaunal bivalves, such as Daonella, proliferated during CPE-induced anoxia, exploiting organic-rich, dysoxic substrates as r-strategists with rapid population blooms in the wake of higher-taxon declines.^[40] Post-CPE recovery in marine ecosystems featured a notable increase in molluscan genera diversity, reflecting humid climate facilitation of continental weathering and nutrient influx that boosted primary productivity and opportunistic colonization.^[19] This rebound, particularly among ammonoids and bivalves, underscored the resilience of soft- and hard-bottom communities amid ongoing environmental instability.

Vertebrates

During the Carnian stage of the Late Triassic (approximately 237–227 million years ago), vertebrates underwent significant evolutionary developments, particularly in marine and terrestrial environments, amid the backdrop of the Carnian Pluvial Episode (CPE), a period of increased humidity around 234–232 Ma that influenced biotic turnovers.^[5] Marine reptiles diversified in the Tethys Ocean, while terrestrial archosaurs saw shifts in dominance, with pseudosuchians as key predators and the initial radiation of avemetatarsalians foreshadowing the rise of dinosaurs.^[25] These changes reflected adaptations to changing paleoenvironments, including reef systems and floodplains.^[5] Thalattosaurs, a group of small to medium-sized aquatic reptiles, were also prominent in nearshore Tethyan habitats, filling piscivorous niches alongside other marine forms.^[43] Marine reptiles, including ichthyosaurs and nothosaurs, were prominent in Tethyan marine settings. Ichthyosaurs transitioned from smaller, more primitive forms akin to the Ladinian Mixosaurus to larger species during the Carnian, with notable representatives in the early Carnian Guanling Biota of southwestern China, where they occupied pelagic niches.^[25] This period marked an increase in body size and specialization for open-water hunting, setting the stage for Norian giants like Shonisaurus, though a diversity gap followed in some regions until the Norian.^[44] Nothosaurs, semi-aquatic sauropterygians, adapted to nearshore and reef habitats in the Tethys, preying on fish and invertebrates; while most abundant in the Middle Triassic, they persisted into the Carnian, contributing to coastal ecosystems before their decline.^[43] On land, terrestrial archosaurs dominated, with pseudosuchians such as raurusuchians serving as apex predators in continental Pangean environments. These large, bipedal carnivores, including forms like Postosuchus and Fasolasuchus, reached lengths over 6 meters and filled top predator roles in floodplains and semi-arid landscapes, exerting pressure on smaller herbivores and competing with emerging groups.^[45] Early dinosaurs appeared amid this, with Nyasasaurus parringtoni from Tanzania dated to around 243 Ma, though its dinosaurian status remains debated due to fragmentary remains; it represents a potential pre-Carnian origin, but true unequivocal dinosaurs like Eoraptor lunensis emerged in the late Carnian (around 231 Ma) in the Ischigualasto Formation of Argentina, marking the Dinosaur Diversification Event (DDE) around 234–230 Ma.^[46] This DDE coincided with post-CPE environmental shifts, allowing dinosaurs to increase from less than 5% to over 90% of archosaur assemblages in some localities.^[5] Other vertebrate groups included temnospondyl amphibians, early turtles, and pterosaurs. Temnospondyls like Metoposaurus krasiejowensis inhabited freshwater floodplains in Europe and North America, with M. diagnosticus appearing in the early Carnian of the American Southwest, where they formed mass accumulations indicating gregarious behavior in riverine settings.^[47] The oldest known turtle, Odontochelys semitestacea, from the Carnian (approximately 237–227 Ma) Wayao Member of the Falang Formation in Guizhou, China, featured a partial shell and aquatic adaptations, signaling the initial evolution of chelonian body plans. Pterosaurs, the earliest flying vertebrates, made their first appearances in the late Carnian to early Norian, with Preondactylus buffarinii from the approximately 228 Ma Calcari della Guida Formation in northeastern Italy representing one of the oldest records, characterized by a wingspan of about 50 cm and multicusped teeth for piscivory.^[48] Evolutionary highlights of the Carnian include the radiation of crurotarsans following the CPE, with pseudosuchians diversifying into diverse forms like aetosaurs and early crocodylomorphs, temporarily dominating tetrapod faunas before their Norian peak.^[5] Concurrently, the first appearances of avemetatarsalians—encompassing dinosaurs and pterosaur relatives—signaled the ascent of this clade, driven by post-CPE ecological opportunities such as increased humidity and the decline of competitors like rhynchosaurs, ultimately paving the way for dinosaur dominance in the Mesozoic.^[5]

Terrestrial plants and ecosystems

During the Carnian stage of the Late Triassic, terrestrial floras were characterized by the dominance of seed ferns belonging to the Peltaspermales, which featured pinnate fronds with parallel venation patterns that exhibited early angiosperm-like traits in some taxa.^[49] Conifers of the order Voltziales were also prominent, forming key components of woodland communities with scale-like leaves and woody trunks adapted to varying moisture regimes.^[50] The first appearances of ginkgophytes, including genera like Arberophyllum and Sphenobaiera with fan-shaped or wedge-shaped leaves, marked the initial diversification of this group in high-latitude settings.^[51] Similarly, bennettitales emerged prominently, represented by foliage such as Nilssoniopteris with elongated, pinnate leaves showing cuticular features indicative of their gymnosperm affinities.^[52] In wetland environments, lycopods and ferns thrived, with taxa like marattiacean ferns contributing to understory vegetation in swampy habitats, as evidenced by increased spore abundances during humid intervals.^[6] Terrestrial ecosystems exhibited structured zonation influenced by climatic variability, with riparian forests developing along river systems during humid phases following the Carnian Pluvial Episode (CPE), as indicated by coal deposits in the lower Chinle Formation of western North America.^[53] These forests comprised dense stands of gymnosperms and ferns, supporting peat accumulation in floodplain settings. In contrast, intermontane basins hosted arid-adapted gymnosperm communities, including bennettitales and conifers in upland biotas like that of the Madygen Formation, where drought-tolerant foliage dominated seasonal landscapes.^[54] Herbivory played a role in shaping these ecosystems, with browsing by aetosaurs and early dinosauromorphs exerting pressure on low-lying vegetation such as ferns and seed ferns, influencing community dynamics through selective foraging. Evidence of fungal interactions, including symbiotic associations and decay structures in petrified wood from the Chinle Formation, highlights microbial contributions to nutrient cycling and wood decomposition in these forested settings.^[55] Floral diversity underwent notable turnover during the CPE, with approximately 15% of taxa replaced amid shifts toward more humidity-tolerant forms, reflecting adaptive responses to global wetting. A gnetophyte-like plant from the late Carnian of Texas has been proposed to inform hypotheses on the origin of angiosperm reproductive structures.^[56]

Fossil Localities

Key Lagerstätten

The Monte San Giorgio Lagerstätte, straddling the Swiss-Italian border near Lake Lugano, is a UNESCO World Heritage Site renowned for its exceptional preservation of Middle Triassic (Ladinian) to early Late Triassic marine life, including minor Carnian-aged deposits in the Marne del Monte San Giorgio Formation. The site's primary fossils are embedded in bituminous shales of the Besano Formation (late Anisian to early Ladinian), representing a restricted lagoonal environment with periodic anoxic bottom waters that facilitated the conservation of soft tissues and delicate structures. Over 200 species have been documented, encompassing approximately 50 fish taxa such as the predatory actinopterygian Saurichthys, around 25 reptile species including marine nothosaurs and tanystropheids, diverse invertebrates like ammonites and crustaceans, as well as terrestrial elements such as insects and plants washed into the lagoon.^[57]^[58]^[59] Another prominent Carnian Konservat-Lagerstätte is Polzberg in the Northern Calcareous Alps of Lower Austria, where the Reingraben Shales yield a diverse marine palaeobiota deposited under dysoxic to anoxic conditions during the early Carnian. This site features over 6,000 specimens with remarkable fidelity, including articulated skeletons of fish, thylacocephalan crustaceans, ammonoids, and soft-bodied organisms, alongside plant remains such as conifer twigs and leaves, providing rare insights into Triassic coastal ecosystems. The anoxic bottom waters prevented decay and bioturbation, enabling the preservation of fine details like cephalic cartilage in coleoid cephalopods and gut contents in predators, which reveal trophic interactions within the community.^[60]^[61]^[62] These Lagerstätten offer high-resolution snapshots of Carnian biodiversity hotspots, capturing community structures amid environmental perturbations like the Carnian Pluvial Episode (CPE). At Polzberg, the assemblage documents shifts in marine food webs linked to increased humidity and volcanism during the CPE, highlighting recovery patterns in invertebrates and early ray-finned fishes. Similarly, Monte San Giorgio's lagoonal fossils illustrate pre-CPE stability transitioning into the humid phase, aiding reconstructions of how anoxic events influenced soft-bodied organism diversification, such as primitive elasmobranchs. Their exceptional taphonomic conditions underscore the role of oxygen-depleted basins in preserving evidence of ecological hotspots during this critical interval of Triassic evolution.^[63]^[60]^[59]

Significant formations and outcrops

In Europe, the Hallstatt Limestone of Austria represents a key marine formation for Carnian stratigraphy, characterized by thin-bedded, micritic limestones rich in ammonoids that provide essential biostratigraphic markers for the stage.^[9] These outcrops, exposed in the Northern Calcareous Alps, preserve diverse cephalopod assemblages indicative of basinal paleoenvironments and have been instrumental in defining ammonoid zonations across the Julian and Tuvalian substages.^[64] In the Italian Dolomites, the transition from the Werfen Formation to the Cassian Formation (part of the San Cassiano Formation) marks a significant Carnian sequence, featuring cyclic carbonates, volcaniclastic deposits, and reefal buildups that reflect platform-to-basin shifts during the stage.^[65] The Prati di Stuores section in the Southern Alps serves as the Global Stratotype Section and Point (GSSP) for the base of the Carnian, where hemipelagic beds of the Wengen and Cassiano Formations expose the first occurrence of the ammonoid Daxatina canadensis, anchoring the stage's lower boundary at approximately 237 Ma.^[13] In North America, the Santa Rosa Formation in New Mexico exposes fluvial and lacustrine deposits of early late Carnian age, notable for preserving aetosaur osteoderms and trackways that document pseudosuchian dominance in continental settings.^[66] These outcrops, part of the Chinle Group precursor, yield armored archosaur remains alongside metoposaurid fragments, highlighting arid to semi-arid paleolandscapes interrupted by pluvial influences.^[67] Further south, the Ischigualasto Formation in northwestern Argentina contains redbed sandstones and mudstones of late Carnian to early Norian age, renowned for early dinosaur skeletons such as Eoraptor and Herrerasaurus, which illustrate the initial radiation of Dinosauria in alluvial floodplains.^[68] U-Pb dating of intercalated tuffs confirms its span from about 231.4 to 225.9 Ma, making it a critical reference for correlating continental tetrapod assemblages globally.^[69] In Asia, the Ma'antang Formation in the Sichuan Basin of South China features bedded and nodular cherts deposited in a foreland setting during the Carnian, recording siliceous sedimentation linked to heightened humidity and volcanism.^[70] These exposures preserve evidence of marine incursions and geochemical signatures of the Carnian Pluvial Episode (CPE), including mercury anomalies from volcanic inputs.^[71] In southern Africa, the Molteno Formation within the Karoo Basin consists of continental red sandstones and conglomerates of Carnian age, formed in braided river systems that archive braidplain ecosystems with early therapsid and archosaur fossils.^[72] These units, up to 460 m thick in the main basin, provide insights into Gondwanan terrestrial dynamics during the stage's humid phases. Additional notable Carnian localities include the Santa Lucia Formation in central Mexico, which preserves marine reptiles and ammonoids in carbonate platforms, and the Lossiemouth Sandstone Formation in Scotland, yielding terrestrial archosaurs and therapsids from floodplain deposits.^[3] These formations hold substantial research value as type sections for Carnian substages, such as the Julian-Tuvalian boundary in Alpine sequences, and have facilitated recent 2020s excavations uncovering CPE-related volcaniclastic layers, including ash-rich shales in Chinese basins that link large igneous province activity to global climate shifts.^[73] Such discoveries, including deep-water tuff layers in the Nanpanjiang Basin, enhance correlations of the CPE's onset around 232 Ma across marine and terrestrial realms.^[74]

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The carbonate—clastic cycles of the East-Alpine Raibl group
The Carnian Raibl group of the Eastern Alps consists of three 50–100 m thick, alternating carbonate and clastic third-order cycles, each of which can be ...Missing: lithostratigraphy | Show results with:lithostratigraphy
[16]
Lithostratigraphy, chemostratigraphy, and vertebrate biostratigraphy ...
The Dockum Group of southern Garza County, West Texas, consists of a basal siliceous conglomeritic sandstone (the Santa Rosa Sandstone) with pedogenic ...
[17]
Carnian (Late Triassic) magnetostratigraphy from the Germanic ...
Jul 1, 2020 · We obtained a composite magnetostratigraphy spanning the entire Carnian from three boreholes in the Germanic Basin.
[18]
https://www.lyellcollection.org/doi/10.1144/SP334.10
[19]
Extinction and dawn of the modern world in the Carnian (Late Triassic)
Sep 16, 2020 · The Carnian is the basal stage of the Late Triassic. Its lower boundary is dated at ca. 237 Ma ago, based on U-Pb radiometric dating of ...
[20]
(PDF) Taxonomy and phylomorphogenesis of the Carnian/Norian ...
Generic turnovers of Carnian/Norian conodonts: Climatic control or competition? ... Neogondolella became extinct by the Late Ladinian, and Paragondolella ...<|separator|>
[21]
New Species of Spiral Nodosariids (Foraminifera) from the Lower ...
Aug 5, 2025 · New Species of Spiral Nodosariids (Foraminifera) from the Lower Carnian (Triassic) Deposits on the Laptev Sea Coast (Russia).
[22]
Biochronology of Triassic bivalves | GeoScienceWorld Books
Jan 1, 2010 · Although they remain poorly documented, several occurrences of Daonella are known from lower Carnian strata overlapping with true Halobia.
[23]
[PDF] middle and upper triassic bivalve biostratigraphy of the shublik
As reported herein, this interval contains two biostratigraphic data points (Ladinian Daonella frami and Early Carnian Halobia zitteli) and can be directly ...
[24]
A PRIMITIVE HALOBIA (BIVALVIA: HALOBIOIDEA) FROM THE ...
Mar 3, 2017 · Halobia daonellaformis new species is described from the lowermost Carnian of northeast British Columbia. Halobia daonellaformis n. sp. is ...
[25]
Extinction and dawn of the modern world in the Carnian (Late Triassic)
Sep 16, 2020 · ... Carnian Pluvial Episode (CPE; Fig. 1). The CPE occurred around 234 to 232 million years (Ma) ago. Its most marked characteristic was a ...
[26]
Pangea: Geochronological correlation of successive environmental ...
235 Ma (Carnian) singularity: Pangea at its widest extent undergoes incipient breakup (Extension II) · 190 Ma initial breakup and oldest seafloor spreading.
[27]
Pangean (Late Carboniferous–Middle Jurassic) paleoenvironment ...
These maps depict the plate tectonic configuration, palaeoenvironment and lithofacies during the Late Triassic (Carnian–Norian) and Early Jurassic (Hettangian– ...
[28]
Late Triassic and Early Jurassic palaeogeography of the world
During Triassic time Pangaea began to stretch, initiating the rifting and future break-up of the supercontinent.
[29]
Carnian (Upper Triassic) Lavas and Tuffites from the Mersin Mélange
Aug 6, 2025 · ... opening mainly during the Carnian—Norian interval. This opening marked the birth of Neo-Tethys behind the Cimmerian continent which, at that ...
[30]
[PDF] the central atlantic magmatic province: an historical perspective
The oldest rift basin sediments around the central Atlantic are early Carnian, possibly older than 230 Ma (Olsen ... Latest Triassic onset of the Central Atlantic ...Missing: precursors | Show results with:precursors<|control11|><|separator|>
[31]
Multi-proxy paleosol evidence for middle and late Triassic climate ...
The upper Chinle Formation (Carnian–Norian) reveals a return to semiarid to subhumid conditions and the formation of Castle Valley, Moab and Salt Valley ...
[32]
PRE-QUATERNARY SEA-LEVEL CHANGES - Annual Reviews
There was a rapid transgression in the early mid-Triassic. (Anisian), with the seas spreading again after a minor interruption to a maximum in Ladinian-Carnian ...<|control11|><|separator|>
[33]
Triassic sea level change and the Ladinian-Carnian stage boundary
Aug 9, 2025 · Recently, it has been suggested that some Triassic stage boundaries are associated with eustatic changes in sea level1,2. Here evidence is ...
[34]
Climates of the Late Triassic: Perspectives, Proxies and Problems
The interior of Pangea was arid or semi-arid at low to mid latitudes. The continental configuration meant the climate was strongly monsoonal and the ...
[35]
A continental record of the Carnian Pluvial Episode (CPE) from the ...
Oct 8, 2018 · Based on the palynological assemblages and the chemostratigraphy, the stratigraphical range of the Dunscombe Mudstone Formation can be extended ...
[36]
Perturbations in the carbon cycle during the Carnian Humid Episode
Oceanic anoxia was widespread from the Julian 2 substage, as documented in the extensive deposition of black shales and organic-rich marls in several Tethyan ...
[37]
Carnian–Norian (Late Triassic) climate change - ScienceDirect.com
Mar 15, 2020 · High-latitude areas were characterized by warm summers (> 20 °C) but near freezing during the winter. Conodont oxygen isotope data indicate that ...
[38]
The Carnian pluvial episode (Late Triassic): new insights into this ...
It is marked by severe extinction among important marine groups such as ammonoids and conodonts, and rapid radiation of key taxa on land and in the ocean, e.g. ...
[39]
Palaeoecology of the Triassic black shale bivalve Daonella
Aug 6, 2025 · Circulation lower in the water column was inhibited by the carbonate platform, however, a dysaerobic transitional layer occasionally mixed with ...<|control11|><|separator|>
[40]
https://www.researchgate.net/publication/223268611_Palaeoecology_of_the_Triassic_black_shale_bivalve_Daonella_-_New_insights_into_an_old_controversy
[41]
Pelagic responses to oceanic anoxia during the Carnian Pluvial ...
Sep 28, 2023 · Such temperature increases during the late Julian are supported by a decrease in conodont apatite oxygen isotope composition (δ18Ophos) across ...
[42]
Ichthyosaurs from the Upper Triassic (Carnian–Norian) of the New ...
Dec 13, 2021 · Triassic ichthyosaurians from Russia are poorly known, despite the country's large territory and wide distribution of Triassic marine deposits ...
[43]
A gigantic nothosaur (Reptilia: Sauropterygia) from the Middle ...
Nov 27, 2014 · Here we report a gigantic nothosaur from the lower Middle Triassic of Luoping in southwest China (eastern Tethyan ocean), which possesses the ...
[44]
Palaeobiological inferences of “rauisuchians” Fasolasuchus tenax ...
Irrespective of their taxonomic validity, they fulfilled an important ecological role as the apex predators in the Middle‐Upper Triassic continental ecosystems ...
[45]
Dating the origin of dinosaurs - PMC - NIH
The oldest undisputed dinosaurs are known from the early Late Triassic, about 231 Ma (5, 6). Late Carnian strata of the lower portion of the Ischigualasto ...
[46]
Cranial bone histology of Metoposaurus krasiejowensis (Amphibia ...
Nov 8, 2016 · In this study, 21 skull bones of Metoposaurus krasiejowensis from the Late Triassic of Poland were investigated histologically.
[47]
Climate drivers and palaeobiogeography of lagerpetids and early ...
Jun 18, 2025 · Furthermore, the modelled Carnian–early Norian pterosaur climatic niche would probably be found in temperate and wet conditions typical of ...
[48]
Parallel evolution of angiosperm‐like venation in Peltaspermales: a ...
Apr 16, 2024 · We suggest that its nearest relatives are Peltaspermales similar to ... Lunz flora in the Austrian Alps—a standard for Carnian floras.
[49]
Palaeophytogeographical Patterns Across the Permian–Triassic ...
Dec 7, 2020 · The dominant plant group are the conifers (especially Voltziales; 14 genera), followed by various gymnosperms including Cordaitales (5 genera) ...
[50]
First record of Ginkgo-related fertile organs (Hamshawvia ...
This paper describes new ginkgophytes recently discovered in a new exposure of the original type section of the Passo das Tropas Member, Santa Maria Formation.
[51]
FIRST RECORD OF NILSSONIOPTERIS (GYMNOSPERMOPHYTA ...
Sep 14, 2007 · We describe three species of Nilssoniopteris, which represent the first records for this genus from Lunz, and one of the earliest accounts of Nilssoniopteris ...
[52]
Record of the Carnian wet episode in strata of the Chinle Group ...
Jul 12, 2018 · The Late Triassic Carnian wet episode was an interval of humid climate evident in the lowermost strata of the Upper Triassic Chinle Group in the western USA.Chinle Group · Chinle Group evidence of the... · Correlation of the Chinle...
[53]
Cycadalean and bennettitalean foliage from the Triassic Madygen ...
The Madygen biota thrived at 35°–40°N in an upland intermontane basin (Voigt et al., 2006). The Triassic has been recognized as one of the four warm time ...
[54]
degraded wood in the upper triassic petrified forest formation (chinle ...
Both forms of degradation are inter-preted as the result of pre-burial biotic activity. The pitting appears to be caused by pathogenic fungi (white rot and ...
[55]
(PDF) A New Gnetophyte from the Late Carnian (Late Triassic) of ...
Even concepts about the time of angiosperm origin have been cyclical, ranging from Permo-Triassic (Axelrod, 1952, Axelrod, 1961) to Early Cretaceous (Wolfe et ...
[56]
[PDF] Monte San Giorgio - UNESCO World Heritage Centre
Jul 5, 2003 · Because the lagoon was near to land, the fossil remains also include some land-based fossils including reptiles, insects and plants. The result ...
[57]
The fossiliferous levels - Monte San Giorgio
To date, approximately 25 species of reptile, 50 species of fish and more than 100 species of invertebrates have been recorded in addition to various species of ...
[58]
The marine conservation deposits of Monte San Giorgio ...
Mar 4, 2024 · Marine conservation deposits ('Konservat-Lagerstätten') are characterized by their mode of fossil preservation, faunal composition and sedimentary facies.
[59]
The Upper Triassic Polzberg palaeobiota from a marine Konservat ...
Aug 17, 2021 · The Upper Triassic Polzberg palaeobiota from a marine Konservat-Lagerstätte deposited during the Carnian Pluvial Episode in Austria.
[60]
New data on the marine Upper Triassic palaeobiota from the ...
Jun 6, 2023 · Here we report new data on the Upper Triassic Polzberg Konservat-Lagerstätte in the Northern Calcarous Alps of Lower Austria.Missing: amber | Show results with:amber
[61]
Mineralized belemnoid cephalic cartilage from the late Triassic ...
Apr 20, 2022 · The exceptional conditions during deposition resulted in excellent preservation in the Polzberg Konservat-Lagerstätte, enabling a detailed ...
[62]
The Upper Triassic Polzberg palaeobiota from a marine Konservat ...
Aug 17, 2021 · The dry Middle and Upper Triassic climate was interrupted by a middle Carnian global phase of increased humidity in the western Tethys and hence ...
[63]
(PDF) Carnian Ammonoids from the Wetterstein Limestone of ...
The first ammonoids of the uppermost Wetterstein limestone were discovered in early mining history. LIPOLT (1856) mentioned ammonoids from the Fladung mining ...
[64]
https://www.researchgate.net/publication/362711880_Carnian_Ammonoids_from_the_Wetterstein_Limestone_of_Hochobir_North_Karawanken_Carinthia
[65]
Bone Histology of Phytosaur, Aetosaur, and Other Archosauriform ...
Dec 24, 2013 · Santa Rosa Formation (Santa Rosa Sandstone?), Chinle Group, Carnian, Conchas Lake, New Mexico ... A new aetosaur from the Upper Triassic ...
[66]
[PDF] TRIASSIC STRATIGRAPHY, BIOSTRATIGRAPHY AND ...
Santa Rosa Formation strata have not yielded tetrapods in east-central New Mexico other than fragmentary metoposau- rid fossils indicative of a Late Triassic ...
[67]
Age constraints on the dispersal of dinosaurs in the Late Triassic ...
May 19, 2014 · The leading candidate for the oldest known occurrence of dinosaurs is the tetrapod assemblage of the Ischigualasto Formation of Argentina (1–4) ...
[68]
The Late Triassic Ischigualasto Formation at Cerro Las Lajas (La ...
Jul 29, 2020 · A Carnian theropod with unexpectedly derived features during the first dinosaur radiation ... avemetatarsalians, in which the head is ...Missing: appearances | Show results with:appearances<|control11|><|separator|>
[69]
Origin of Carnian Ma'antang cherts, northwestern Sichuan Basin ...
May 1, 2022 · Abundant Carnian bedded and nodular cherts occur in the Sichuan foreland basin, South China. These cherts faithfully record the Carnian Pluvial Episode.
[70]
Mercury enrichments during the Carnian Pluvial Event (Late Triassic ...
Mar 2, 2022 · Our study supports the hypothesis that enhanced volcanism played a major role in the evolution of biota and the environment during the CPE.
[71]
Late Triassic ecosystems of the Molteno/Lower Elliot biome of ...
Jan 1, 1998 · A first attempt is made to integrate the plant, insect and tetrapod elements of the Late Triassic (Carnian) intracontinental, braidplain ecosystems of the ...
[72]
Distinctive volcanic ash–rich lacustrine shale deposition related to ...
Mar 15, 2024 · The Late Triassic Carnian Pluvial Episode (CPE) witnessed enormous climate change closely associated with volcanic activity.
[73]
Discovery of Late Triassic volcanic ash layers in the deep-water ...
Nov 1, 2025 · The Carnian age and unique lithology suggest the Niluo Member may record the CPE in the slope environment, potentially linked to increased ...