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Mesozoic

The Mesozoic Era, spanning approximately 252 to 66 million years ago, represents the middle era of the Eon and is renowned as the "Age of Reptiles" due to the evolutionary dominance and diversification of dinosaurs, pterosaurs, and marine reptiles during this time. This era followed the , marked by recovery from the Permian-Triassic mass extinction, and preceded the , with its boundaries defined by significant global events including the initial breakup of the and a catastrophic asteroid impact that ended it. Key characteristics include rising sea levels, shifting continents, and profound biological innovations, such as the emergence of early mammals, , and flowering , alongside gymnosperm-dominated forests. The Mesozoic is subdivided into three periods, each reflecting progressive geological and biological transformations. The Triassic Period (252–201 million years ago) began with the aftermath of the largest mass extinction in Earth's history, allowing archosaurs—including the first dinosaurs—to evolve and spread across Pangaea, while early conifers and cycads formed the primary vegetation. Rifting initiated the supercontinent's fragmentation, creating rift basins and altering global climates from arid to more humid conditions. The Jurassic Period (201–145 million years ago) witnessed the further diversification of dinosaurs into massive herbivores and carnivores, the appearance of the first true birds from theropod ancestors, and the expansion of shallow seas over continents like North America, fostering diverse marine ecosystems with ichthyosaurs and plesiosaurs. Conifer forests and ferns continued to thrive, supporting the era's reptilian giants. The Cretaceous Period (145–66 million years ago) marked the Mesozoic's peak in biodiversity, with dinosaurs reaching their greatest variety, including iconic forms like Tyrannosaurus rex and hadrosaurs, alongside the radiation of modern bird groups and small mammals. Flowering plants (angiosperms) emerged and rapidly diversified by the mid-period, transforming terrestrial ecosystems and pollinator interactions, while high sea levels flooded vast inland areas, depositing chalk-rich sediments that form landmarks like the . The period's warmer, greenhouse climate supported ammonites, mosasaurs, and turtles in the oceans. The Mesozoic concluded abruptly with the Cretaceous-Paleogene extinction event around 66 million years ago, triggered by a massive impact near present-day , which caused widespread fires, tsunamis, and a "" effect from atmospheric dust, leading to the demise of approximately 75% of Earth's species, including all non-avian dinosaurs. This event paved the way for mammalian diversification in the subsequent Cenozoic Era, fundamentally reshaping life on the planet.

Introduction

Naming and Definition

The term "Mesozoic" was coined by English geologist John Phillips in 1841 to designate the era of "middle life," derived from the Greek words mesos (middle) and zōē (life), in order to distinguish it from the earlier era ("ancient life") and the later Cenozoic era ("new life"). Phillips introduced this tripartite division of time in his work on geological chronology, grouping rock systems based on their fossil content and recognizing distinct phases of life's development on . The conceptual foundations of the Mesozoic emerged in the early amid rapid advances in and by British geologists. William , Oxford's first reader in geology, advanced early understanding through his 1824 description of , the first scientifically named dinosaur from strata, highlighting the prevalence of large extinct reptiles in post- rocks. Concurrently, Adam collaborated with Roderick to delineate the in 1838–1839, establishing its upper boundary near the end-Permian mass extinction and creating the stratigraphic context for identifying the succeeding era of intermediate faunas. Phillips built directly on these efforts, integrating the emerging systems of the , , and into his broader . The Mesozoic Era is precisely defined as the major division of geological time between the and eras, encompassing approximately 252 to 66 million years ago and marked by the dominance of reptiles as the primary terrestrial vertebrates alongside the progressive breakup of the into the continents recognizable today. This era is subdivided into three periods—Triassic, , and —each reflecting evolutionary and tectonic shifts that shaped Earth's biota and landmasses.

Geological Timescale and Boundaries

The Mesozoic Era constitutes the second of three eras within the Eon, following the and preceding the , and it encompasses a temporal span of approximately 186 million years. This era is positioned between the end of the at around 252 million years ago (Ma) and the onset of the at 66 Ma, marking a pivotal interval of recovery and diversification in Earth's after the Paleozoic's major events and before the mammalian dominance of the . The Mesozoic spans from 251.902 ± 0.024 Ma to 66.043 ± 0.043 Ma, according to the latest calibrations integrated into the International Chronostratigraphic Chart and supporting high-precision geochronology. This duration is established through high-resolution geochronology, providing a robust framework for correlating global stratigraphic records. The era's lower boundary is defined at the base of the Induan Stage of the Triassic Period, coinciding with the Permian-Triassic extinction event at approximately 252 Ma, which represents the most severe mass extinction in Earth's history and is ratified by the Global Stratotype Section and Point (GSSP) at Meishan, China. The upper boundary of the Mesozoic occurs at the Cretaceous-Paleogene (K-Pg) boundary, dated to 66.043 ± 0.043 Ma, and is delineated by a globally distributed iridium-rich clay layer that signals the of a large . This boundary is formally defined by the GSSP at , , where the marks the base of the Stage of the Period, with supporting evidence from the Chicxulub impact crater in , confirmed through U-Pb dating of impact melt rock to align precisely with the horizon. These boundaries are correlated with absolute timescales primarily through techniques, such as uranium-lead (U-Pb) dating of crystals from layers interbedded in boundary sections, which yields high-precision ages essential for anchoring the chronostratigraphic framework. The Mesozoic is subdivided into three periods—, , and —each with their own stage-level boundaries refined by similar geochronologic methods.

Geological Framework

Stratigraphy

The of the Mesozoic Era encompasses the description, classification, and correlation of layers formed between approximately 252 and 66 million years ago, utilizing both lithostratigraphy and to delineate sequences across continents. Lithostratigraphy focuses on rock types and their vertical and lateral relationships, such as the —predominantly arkosic sandstones, siltstones, and mudstones—characteristic of the Period, which reflect continental depositional environments in rift basins and floodplains. In contrast, Jurassic sequences often feature widespread marine limestones and shales, indicative of epicontinental seas and platforms, as seen in formations across the North American and European shelves. complements this by employing index s, particularly ammonites, whose evolutionary successions provide precise zonal divisions; for instance, the rapid morphological changes in ammonite genera like those of the Stage mark horizons globally. These approaches together enable the subdivision of Mesozoic strata into formations, groups, and supergroups based on observable lithologic boundaries and assemblages. Global stratotype sections and points (GSSPs) serve as internationally ratified reference sections for defining the lower boundaries of Mesozoic chronostratigraphic units, ensuring standardized correlations. The base of the is defined at the GSSP in the section, Province, , where the first appearance datum (FAD) of the conodont Hindeodus parvus in Bed 27c marks the Permian- boundary, accompanied by a distinctive negative carbon isotope excursion. The Jurassic base is established at the Kuhjoch section in the Austrian , with the FAD of the ammonite Psiloceras spelae tirolicum at 5.80 meters above the top of the Kössen Formation defining the -Jurassic boundary, correlated via and chemostratigraphy. For the , no GSSP has been ratified for the Berriasian Stage base as of 2025, though proposals center on sections in and using markers like the FAD of the ammonite Berriasella jacobi or the base of calpionellid Zone B, highlighting ongoing debates in boundary placement. These GSSPs integrate multiple stratigraphic tools to anchor the era's divisions. Prominent Mesozoic formations exemplify these principles through their distinctive lithologies and fossil content, facilitating regional to global correlations. The Newark Supergroup in eastern represents basin deposits, comprising interbedded red sandstones, shales, and basalt flows up to 12 kilometers thick, recording syn- sedimentation during the breakup of . The Morrison Formation, spanning the western United States, consists of variegated mudstones, sandstones, and limestones in fluvial-lacustrine settings, renowned for its -bearing horizons like those at . In the , the Hell Creek Formation in the northern features fluvial sandstones and mudstones with abundant non-avian fossils, overlying marine shales and marking the final terrestrial deposits before the Cretaceous-Paleogene boundary. These units are delimited by erosional surfaces or lithologic shifts, with their fossils aiding biostratigraphic ties. Correlating Mesozoic strata across continents presents challenges due to —gaps in the rock record from or non-deposition—and lateral changes, where carbonates transition to terrestrial clastics over short distances. For example, the J-3 in the western interior of separates Middle and Upper units, representing a of millions of years and varying regionally in expression from sharp planar surfaces to gradational contacts. variations, such as the shift from Tethyan platform limestones to mudrocks in the , further complicate direct matching, necessitating integrated approaches like ammonite zonations to bridge these disparities. Such issues are particularly acute in tectonically active margins, where uplift and distort sequences. Mesozoic strata play a critical role in resource exploration, particularly as hydrocarbon reservoirs and source rocks, with Jurassic marine shales like those in the Formation serving as prolific sources for oil and gas in the Basin due to their high content and type II. Triassic red beds often form traps in rift-related structures, while Cretaceous sandstones provide reservoirs, as in the North American Interior. These stratigraphic frameworks guide seismic interpretation and drilling, leveraging biostratigraphic markers for precise depth predictions in petroleum systems.

Geochronology

Geochronology of the Mesozoic era relies primarily on techniques to assign absolute ages to rock layers and geological events, with uranium-lead (U-Pb) dating of crystals from layers serving as the cornerstone method due to its high precision and resistance to alteration. This technique involves measuring the decay of uranium isotopes to lead in zircons, which crystallize rapidly in volcanic environments and provide ages accurate to within 0.1-0.5 million years for Mesozoic strata. Complementary to U-Pb, argon-argon (⁴⁰Ar/³⁹Ar) dating is applied to basaltic rocks, particularly those associated with large igneous provinces, by analyzing potassium-bearing minerals like to determine eruption timings with uncertainties typically around 0.5-1 million years. A pivotal geochronological is the refinement of the Triassic-Jurassic boundary age to 201.4 ± 0.2 million years ago (Ma), achieved through U-Pb dating of zircons linked to the onset of (CAMP) volcanism, which provides a direct temporal anchor for this period transition. This dating integrates volcanic ash layers interbedded with sedimentary sequences, correlating the massive basaltic eruptions with the boundary's global stratotype section. Error margins in Mesozoic ages have been reduced through calibration with astronomical tuning, which aligns sedimentary cycles driven by Milankovitch orbital variations—such as (periods of ~100 and 405 thousand years), obliquity (~41 thousand years), and (~19-23 thousand years)—to enhance resolution beyond radiometric methods alone. This integration allows for timescales with precisions down to 10-20 thousand years in well-preserved sections, by tuning cyclic patterns in or rhythms to modeled insolation changes. The International Chronostratigraphic Chart provides the current numerical ages for Mesozoic subdivisions, with the Triassic spanning 251.902 ± 0.024 Ma to 201.4 ± 0.2 Ma, the from 201.4 ± 0.2 Ma to 145.0 ± 0.8 Ma, and the from 145.0 ± 0.8 Ma to 66.0 Ma, reflecting ongoing updates from the based on refined radiometric data. Since 2000, advances in have significantly improved precision through cyclostratigraphy, which identifies and calibrates Milankovitch-driven cycles in sedimentary records to construct floating astronomical timescales that anchor radiometric dates across the Mesozoic. Key developments include the integration of these cycles with magnetic chronologies, where reversals in —dated via U-Pb and tuned astronomically—provide a global correlation framework, as demonstrated in revised Middle to timescales with resolutions under 0.1 million years. These methods have enabled continuous astronomical time scales for much of the Mesozoic, resolving durations of stages to within 0.5% uncertainty in some cases.

Periods

Triassic

The Triassic Period spanned from 251.9 to 201.4 million years ago (Ma), marking the initial phase of the Mesozoic Era following the Permian-Triassic boundary. It is subdivided into three epochs: the , encompassing the (251.902–249.9 Ma) and (249.9–246.7 Ma) stages; the , including the (246.7–241.5 Ma) and (241.5–237 Ma) stages; and the , comprising the (237–227.3 Ma), (227.3–205.7 Ma), and (205.7–201.4 Ma) stages. These divisions are based on stratigraphic boundaries defined by global standard sections and points (GSSPs), reflecting gradual biotic recovery and environmental stabilization after the preceding mass extinction. The Triassic represented a prolonged phase of ecological rebound from the Permian-Triassic mass extinction, which eliminated over 90% of marine species and 70% of terrestrial vertebrates around 252 Ma. Recovery was uneven, with marine ecosystems showing delayed diversification until the , while terrestrial environments saw an initial dominance of holdover groups like therapsids before the rise of new clades. Notably, —ancestors to crocodilians, pterosaurs, dinosaurs, and birds—underwent significant diversification starting in the , with stem-archosaurs achieving unappreciated ecological breadth by the stage, predating dinosaur dominance in the . This archosaur radiation filled vacant niches in a recovering , driven by adaptations to varied habitats amid fluctuating climates. Paleogeographically, the Triassic was dominated by the supercontinent , a vast landmass assembled during the late that encompassed nearly all , resulting in extensive arid interiors due to distance from moisture sources. Equatorial and mid-latitude regions experienced hot, dry conditions with seasonal monsoons, as evidenced by widespread and paleosols indicating aridity. Marginal basins began forming along Pangaea's edges, particularly in the , as early initiated the supercontinent's breakup; these basins accumulated thick sequences of fluvial and lacustrine sediments in regions like eastern . A key geological event was the eruption of the (CAMP) around 201 Ma, involving massive flood basalts across what is now region, which released greenhouse gases and triggered rapid climate warming, , and the end-Triassic mass that wiped out about 76% of species. Economically, Triassic strata host significant deposits in the , a subsiding intracratonic feature spanning , where (Muschelkalk) and (Keuper) sequences include thick , , and layers formed in restricted marine to lagoonal settings under arid conditions. These support major and chemical industries, with resources exploited for industrial minerals and fertilizers; deposits, though less prominent than in other periods, occur in localized basins like those in the Saar-Nahe area, contributing to regional energy resources.

Jurassic

The Jurassic Period, spanning from approximately 201.3 to 145.0 million years ago, represents the middle interval of the Mesozoic Era and is subdivided into three epochs: the (Hettangian to stages, 201.3–174.1 Ma), (Aalenian to stages, 174.1–163.5 Ma), and (Oxfordian to stages, 163.5–145.0 Ma). This 56.3-million-year duration witnessed significant tectonic reconfiguration as the supercontinent continued to fragment, initiating the rifting that formed the Central and led to the separation of and . A pivotal event was the (OAE-1a) around 183 Ma, triggered by massive volcanic activity from the Karoo-Ferrar , which released CO₂ and , causing , , and widespread marine that expanded into shallow photic zones due to salinity stratification and elevated primary productivity. Paleoenvironments during the Jurassic were dominated by expansive shallow epicontinental seas, with warm, humid climates fostering lush coastal forests and diverse marine habitats. In western North America, the Sundance Seaway—an elongate, mid-latitude epicontinental sea extending over 2,000 km from the to —created shallow, oxygen-variable conditions less than 100 meters deep, supporting bivalve and ammonite assemblages while influencing regional through sea-level fluctuations. These settings contributed to the deposition of economically significant resources, including Minette-type oolitic ironstones in the of (e.g., and ), formed in nearshore, agitated subtidal environments during the Toarcian-Aalenian transition through reworking of ferruginous coated grains rich in and chamosite. Additionally, the Formation in , a marine sequence, served as a major source of oil shales and hydrocarbons due to its organic-rich laminations deposited in oxygen-poor shelf seas. The Jurassic marked the first major radiation of dinosaurs, with sauropods achieving dominance as gigantic herbivores that shaped terrestrial ecosystems through their evolutionary success in body size and feeding adaptations, originating from ancestors and diversifying into clades like the Euhelopodidae in isolated Asian regions. Theropod dinosaurs also underwent significant early diversification, with basal forms like coelophysoids proliferating in the and before more derived carnivorous lineages emerged, reflecting adaptations to predation in increasingly fragmented continental landscapes. This period's surge underscored the reptiles' ecological preeminence amid evolving global configurations.

Cretaceous

The Cretaceous Period, spanning from approximately 145 to 66 million years ago, marked the final phase of the Era and was characterized by significant geological and biological transformations. It is subdivided into the (Berriasian, , , , , and stages) and the (Cenomanian, , , Santonian, , and stages), with these divisions based on ammonite and other fossil as well as . During this time, global sea levels reached their peak, driven by volcanism and , leading to extensive shallow marine inundations across continents. A prominent example was the in , a north-south elongate that bisected the from the to the , facilitating unique biogeographic connections and sediment deposition. The period was punctuated by multiple Oceanic Anoxic Events (OAEs), intervals of widespread that promoted the preservation of organic-rich black shales and disrupted marine ecosystems. Notable OAEs include OAE1a in the Early and OAE2 at the Cenomanian-Turonian boundary, linked to volcanic outgassing, nutrient influx, and perturbations that caused transient and enhanced burial of organic carbon. Paleogeographically, the witnessed the near-complete fragmentation of the , with the opening of the South Atlantic and continued rifting between and , resulting in the configuration of continents approaching their modern positions by the period's close. This tectonic evolution influenced ocean circulation, climate, and biodiversity dispersal, as landmasses drifted toward higher latitudes and isolated biogeographic provinces emerged. The Cretaceous culminated in the Cretaceous-Paleogene (K-Pg) mass extinction event at 66 million years ago, which eradicated approximately 75% of Earth's species, including non-avian dinosaurs. This boundary is attributed to synergistic effects of massive Deccan Traps flood basalt volcanism in India, which released climate-altering gases over hundreds of thousands of years, and the Chicxulub asteroid impact in the Yucatán Peninsula, Mexico, which triggered immediate wildfires, tsunamis, and a "nuclear winter" from atmospheric dust and sulfate aerosols. The iridium-rich clay layer at the K-Pg boundary worldwide serves as a key marker of the impact. Economically, Cretaceous strata host significant resources, including vast chalk deposits formed from coccolithophore blooms in warm, shallow seas—exemplified by the White Cliffs of Dover, composed of Late Cretaceous (Santonian-Campanian) chalk up to 110 meters thick. Additionally, the period's sediments in the Gulf of Mexico form major hydrocarbon reservoirs, with Jurassic-Cretaceous carbonates and sands trapping oil and gas in subsalt and suprasalt traps, contributing to the region's status as a prolific petroleum province.

Paleogeography and Tectonics

Continental Configurations

At the onset of the Triassic Period around 252 million years ago, Earth's landmasses were predominantly assembled into the , which encompassed nearly all continental crust and was divided into the northern landmass —comprising present-day , , , and Asia—and the southern landmass , including , , the , , and . This configuration persisted from the late assembly, with Pangaea's elongated form spanning from polar to equatorial latitudes, influencing global geography at the era's start. The breakup of initiated in the around 200 million years ago through rifting that separated from , marking the opening of the Central and the onset of continental fragmentation driven by mantle dynamics. This process accelerated during the , as widened the rift, further isolating from and establishing the proto-Atlantic basin. By the , around 130 million years ago, additional rifting detached the and other fragments from eastern , initiating the formation of the through . In the Mid-Mesozoic, particularly during the , the widening proto-Atlantic facilitated the northward drift of , while began to subdivide, with separating from around 100 million years ago to expand the South Atlantic. These shifts progressively reshaped global landmass distributions, reducing the extent of connected continental interiors. By the end of the Period approximately 66 million years ago, continental positions had evolved toward their modern arrangements, with positioned near the as part of a dispersing , and more distinctly separated by the maturing Atlantic, and the migrating northward. Laurasia had fragmented into and , while 's remnants—, , , , and —were isolated by expanding ocean basins. These evolving continental configurations are reconstructed primarily from paleomagnetic data, which record the and of crustal blocks through remanent magnetism in rocks, revealing Pangaea's initial equatorial position and subsequent drift paths. Fossil distributions provide corroborating evidence, such as the shared presence of cycad-like gymnosperms and theropod faunas across early Mesozoic and , indicating initial connectivity before vicariance led to biogeographic divergence.

Tectonic Processes and Events

The Mesozoic Era was characterized by dynamic , including widespread , rifting, and continental collisions that reshaped global configurations. zones were prominent along convergent margins, such as the eastern Pacific, where the subducted beneath , driving arc magmatism and deformation. Rifting initiated the breakup of the supercontinent , opening the Central in the and , while the underwent progressive closure through northward drift of Cimmerian terranes. These processes exemplify the , wherein ocean basins opened via continental rifting and closed through and collision, as applied to Mesozoic events like the disassembly of and the consumptive closure of Paleo-Tethys. In western North America, the during the (ca. 155–140 Ma) resulted from of the , causing arc-continent collision and metamorphism of Franciscan Complex rocks in . This event involved east-dipping that accreted oceanic terranes to the , forming a magmatic and fold-thrust belt. Precursors to the Sevier Orogeny emerged in the Middle to along the same margin, with initial thrusting and development linked to ongoing , though peak deformation occurred in the . By the (ca. 80–55 Ma), the marked a shift to intra-continental deformation, driven by shallow-angle of a young, buoyant Farallon slab, uplifting basement-cored arches in the . In , the Indosinian (ca. 250–230 Ma) arose from the collision between the and Indochina blocks following closure of the eastern , producing widespread folding, thrusting, and granitic intrusions across southeastern Asia. This event deformed Paleozoic to Early Mesozoic sedimentary sequences and marked the final assembly of the Indochinese Peninsula. The Cimmerian , spanning to (ca. 230–180 Ma), involved northward and collision of Cimmerian continental fragments (e.g., Qiangtang and terranes) with , closing the and forming the Longmu Co-Suture Zone in . These collisions generated extensive fold-thrust belts and metamorphic core complexes, contributing to the proto-Alpine Himalayan system. Volcanic activity was integral to these tectonic processes, with large igneous provinces (LIPs) linked to mantle plumes and plate boundary dynamics. The Siberian Traps, erupting at the Permian-Triassic boundary (ca. 252 Ma), exerted a lingering influence into the Early Triassic by destabilizing the lithosphere and facilitating subsequent rifting in Siberia, though their primary impact preceded the Mesozoic. The Central Atlantic Magmatic Province (CAMP) at the end-Triassic (ca. 201 Ma) involved flood basalts covering ~7 million km², triggered by the initial rifting of Pangaea and associated with intrusive activity that released volatiles, coinciding with the Triassic-Jurassic boundary. In the Cretaceous (ca. 95–88 Ma), the Caribbean-Colombian Plateau formed as an oceanic LIP from Galápagos hotspot volcanism, producing ~150,000 km³ of basalts that accreted to South America via subduction-related obduction. Overall, these tectonic events—subduction-driven orogenies, plume-related , and continent-continent collisions—drove the Mesozoic's major paleogeographic shifts, such as the northward migration of Gondwanan fragments and the expansion of , within the framework of the Wilson Cycle's basin evolution.

Climate and Paleoenvironment

Climatic Patterns

The Mesozoic Era was characterized by a persistent , with atmospheric CO₂ levels ranging from approximately 700 to 1,500 ppm—recent studies suggesting values around 750–1,200 ppm in the Late Mesozoic—contributing to global mean surface temperatures 5–10°C warmer than modern pre-industrial values and the absence of permanent polar ice caps. This warm regime supported reduced latitudinal temperature gradients and elevated sea levels, driven primarily by elevated gases from volcanic and tectonic activity. Temporal variations in climate occurred across the era's periods. The featured overall aridity, particularly in the Pangaea's interior, punctuated by intense megamonsoons that delivered seasonal moisture to coastal and high-latitude regions but left vast continental interiors extremely dry. The maintained general warmth but included minor cooling events, such as around 174 Ma, linked to volcanic activity that temporarily disrupted ocean circulation and reduced global temperatures by several degrees. The reached a super-greenhouse peak during the mid-period (e.g., Cenomanian-Turonian, ~94–90 Ma), with extreme warmth exceeding 20°C globally, amplified by high CO₂ and widespread marine transgressions. Paleoclimate proxies provide key evidence for these patterns. Oxygen isotope ratios (δ¹⁸O) in foraminiferal shells from marine sediments indicate sea surface temperatures and global warmth, with lighter isotopic values reflecting higher temperatures throughout the . Stomatal density and index in leaves from Mesozoic plants inversely correlate with atmospheric CO₂, yielding estimates consistent with 700–1,500 levels and supporting the interpretation. A 2024 reconstruction of temperatures indicates Mesozoic global means often exceeded 20°C, aligning with high CO₂ forcings confirmed by 2025 studies. Regionally, the Pangaean interior hosted expansive desert belts due to its continental positioning far from moisture sources, while the saw lush equatorial rainforests thriving under consistently humid, warm conditions. Recent climate modeling from the 2020s highlights how Mesozoic amplified warming, with simulations showing that eruptions released sufficient CO₂ to drive multi-million-year hyperthermal episodes, enhancing the era's overall state beyond baseline tectonic forcings.

Environmental Factors

The Mesozoic era was characterized by significant sea-level fluctuations, manifesting as transgressive-regressive cycles that influenced continental flooding and sedimentary deposition. These cycles, driven by tectonic subsidence, eustatic changes, and sediment supply variations, included multiple episodes of marine inundation followed by regressions, particularly evident in the stratigraphic records of passive margins. For instance, during the , several third-order cycles (lasting 1-3 million years) led to widespread shallow marine environments, while the saw longer-term trends with a notable mid- highstand reaching approximately 200 above present levels, facilitating epicontinental seas across much of and . Ocean gateways played a pivotal role in shaping Mesozoic circulation patterns, with the progressive opening of the altering global thermohaline dynamics. The rifting between and in the Early Cretaceous established the South Atlantic-Southern Ocean gateway, initially restricting deep-water exchange but gradually enabling enhanced ventilation and nutrient transport southward. Concurrently, the functioned as a major warm equatorial current, channeling heat from the proto-Indian Ocean to the Pacific via trans-Atlantic seaways, which promoted circumglobal flow and contributed to hothouse conditions. By the Late Cretaceous, further widening of the facilitated better ocean ventilation, reducing anoxic tendencies in deeper waters. Geochemical perturbations, notably oceanic anoxic events (OAEs), profoundly impacted Mesozoic marine environments through widespread deposition of organic-rich black shales indicative of bottom-water anoxia. These events, such as the early Toarcian OAE (T-OAE) in the Early Jurassic, were marked by intensified burial of organic carbon under stratified oceans, leading to positive carbon isotope excursions in marine records. The T-OAE featured a prominent negative δ¹³C shift of up to 7‰ in organic matter, reflecting massive carbon injection from volcanic or methane sources, followed by a recovery phase with black shale layers signaling expanded oxygen minimum zones. Similar patterns occurred in Cretaceous OAEs, like OAE1a and OAE2, where anoxia affected vast shelf areas, altering global biogeochemical cycles. Atmospheric oxygen levels during the Mesozoic fluctuated between approximately 15% and 30% by volume, with peaks in the and supporting larger body sizes in certain taxa through enhanced respiratory efficiency. These variations, modeled from carbon and isotope proxies, arose from imbalances in organic burial and , influencing aerobic in vertebrates and . For example, elevated oxygen in the to may have facilitated the observed in early dinosaurs, echoing patterns where high oxygen enabled oversized arthropods, though Mesozoic remained comparatively smaller due to other ecological constraints. Silicate weathering rates in the Mesozoic were closely linked to tectonic activity, which exposed fresh surfaces and modulated delivery to oceans via enhanced . Uplift associated with Pangaean rifting and increased chemical of basaltic and granitic terrains, accelerating the release of , , and other nutrients that fueled primary . Orbital forcing paced these rates, as evidenced by cyclostratigraphic records of biogenic silica accumulation, tying intensity to and influencing long-term through the drawdown of atmospheric CO₂.

Biodiversity and Evolution

Flora

The Mesozoic Era, spanning from approximately 252 to 66 million years ago, was characterized by the dominance of gymnosperms as the primary vascular , with , cycads, and ginkgos forming extensive forests and understories across various continents. These seed-bearing , lacking flowers and fruits, adapted to a range of environments, from arid lowlands to humid highlands, and their naked seeds facilitated dispersal in the era's often windy and variable climates. By the , modern such as ancestors of pines and redwoods began to diversify, marking a shift toward more resilient woody vegetation. Key plant radiations during the Triassic included the Bennettitales, an extinct group of gymnosperm-like with fern-like leaves and flower-like reproductive structures, which thrived in subtropical settings and contributed to early Mesozoic . Ferns and horsetails also underwent significant diversification in humid environments, forming dense prairies and riparian zones that supported the era's emerging herbivorous reptiles. These non-seed , reliant on spores for , dominated open landscapes in the wake of the Permian-Triassic extinction, with fossil assemblages from the revealing over ten fern species in western . Ecological shifts progressed markedly through the periods: Triassic landscapes featured expansive fern prairies interspersed with early gymnosperms, transitioning in the to vast conifer-dominated forests that covered supercontinents like , with trees reaching heights over 50 meters in coastal and upland areas. By the , the rise of angiosperms introduced diverse flowering plants into forest understories, enhancing habitat complexity while gymnosperms retained canopy dominance in many regions. This progression reflected increasing atmospheric CO2 levels and continental fragmentation, fostering specialized vegetation zones. Fossil evidence abounds, including petrified forests from the at in , where silicified logs of and cycads preserve details of structure from riverine floodplains. records from sediments further document the angiosperm radiation, showing their pollen comprising over 50% of assemblages by around 100 million years ago, signaling a rapid diversification in lowland and aquatic habitats. A 2015 molecular clock analysis, incorporating fossil calibrations and genomic data from over 200 angiosperm species, estimated their crown-group origins at approximately 140 million years ago, predating the earliest unequivocal fossils by up to 5 million years and suggesting an initial diversification in the Early Cretaceous. However, more recent studies as of 2025 continue to debate this timeline, with some molecular clock estimates pushing crown origins back to the Jurassic or even Triassic, though the fossil record firmly supports first appearances in the Early Cretaceous around 135 million years ago. This revised understanding underscores the role of genetic innovations in enabling angiosperms to outcompete gymnosperms by the era's close.

Fauna

The Mesozoic era witnessed the dominance of archosaurs, particularly dinosaurs, which originated in the Late Triassic period around 233 million years ago as small, bipedal carnivores within the avemetatarsalian clade. Dinosaurs rapidly diversified following the end-Triassic extinction, evolving into a wide array of forms that occupied terrestrial niches across all continents, from the massive herbivorous sauropods like Diplodocus in the Jurassic to apex predators such as the tyrannosaurid Tyrannosaurus rex in the Late Cretaceous. Pterosaurs, the first vertebrates to achieve powered flight, emerged in the Late Triassic and reached enormous sizes, with species like Pteranodon spanning wingspans up to 7 meters in the Cretaceous, filling aerial predator and scavenger roles. Marine reptiles, independent of dinosaurs, included ichthyosaurs—streamlined, dolphin-like predators that thrived from the Triassic to Cretaceous—and plesiosaurs, long-necked swimmers that hunted in Mesozoic oceans alongside mosasaurs in the Late Cretaceous. Invertebrate faunas were equally diverse and ecologically pivotal, with ammonites serving as key index fossils due to their rapid and widespread in marine environments throughout the era, featuring coiled shells with complex sutures for buoyancy control. Belemnites, squid-like cephalopods with internal bullet-shaped guards, dominated seas as active predators, contributing to the rich trophic webs of shallow marine ecosystems. In the , rudist bivalves formed reef-like structures in tropical waters, replacing coral-dominated systems and supporting diverse marine communities until their extinction. Evolutionary milestones included the origin of avian dinosaurs, or birds, from theropod dinosaurs in the , exemplified by around 150 million years ago, which bridged reptilian and traits with feathers and flight capabilities. Mammals, appearing in the , remained small and nocturnal, diversifying into multituberculates, monotremes, and early therians but overshadowed by larger reptiles until post-Mesozoic opportunities. Dinosaur distributions showed biogeographic patterns tied to , with cosmopolitan groups like sauropods in the giving way to distinct assemblages: Gondwanan faunas rich in theropods such as abelisaurids, contrasted with Laurasian ornithischians like hadrosaurs. The era culminated in the Cretaceous-Paleogene (K-Pg) approximately 66 million years ago, which eradicated non-avian dinosaurs, pterosaurs, and many reptiles due to a combination of asteroid impact and , fundamentally reshaping vertebrate communities. Survivors included crocodilians, which endured as semi-aquatic archosaurs adapted to brackish environments, and , whose shelled bodies provided protection across terrestrial and habitats. This mass paved the way for mammalian radiation in the , while avian dinosaurs proliferated from the surviving bird lineage.

Microbiota

During the Mesozoic Era, prokaryotic microorganisms, particularly , played a foundational role in shallow environments through the formation of , although their abundance declined sharply after the recovery from the end-Permian extinction. This decline is attributed to increased pressure from metazoan herbivores and competition from eukaryotic , yet cyanobacteria persisted in restricted niches, such as hypersaline lagoons and marginal marine settings, where they continued to contribute to primary productivity and carbonate precipitation. In deeper ocean contexts, , including sulfate-reducing prokaryotes, thrived in expanded anoxic layers during Oceanic Anoxic Events (OAEs), such as the OAE in the and OAE2 in the mid-Cretaceous, where they mediated preservation by consuming oxygen and producing . These microbial blooms in oxygen-depleted waters facilitated the deposition of organic-rich black shales, highlighting their influence on global carbon burial. Eukaryotic microbes diversified markedly in the Mesozoic oceans, serving as key planktonic components that reflected and influenced . Planktonic , emerging in the from benthic ancestors, exhibited shell —such as isotopes and elements—that indicated variations in seawater , , and across the era. Similarly, , with their siliceous skeletons, proliferated as indicators of silica availability and nutrient dynamics, particularly during periods of high productivity in the and oceans. Dinoflagellates, another group of eukaryotic microbes, became prominent in Jurassic black shales, where their organic-walled cysts contributed significantly to the accumulation of , signaling enhanced under stratified, nutrient-enriched conditions. Fungi and among the microbiota formed critical symbiotic and structural roles in terrestrial and ecosystems. Arbuscular mycorrhizal fungi established associations with early Mesozoic gymnosperms, such as cycads and , enhancing uptake—particularly —from nutrient-poor soils and supporting the expansion of terrestrial . In settings, calcareous , including dasycladaleans and , constructed biogenic frameworks on carbonate platforms, binding sediments and promoting reef-like structures in shallow, tropical environments. These algae's processes were sensitive to chemistry, contributing to the buildup of vast deposits that preserved records of platform evolution. Microbial communities exerted profound biogeochemical influences, driving nutrient cycles essential for Mesozoic ecosystems. Nitrogen-fixing prokaryotes, including and other diazotrophs, sustained in nutrient-limited oceans by converting atmospheric N₂ into bioavailable forms, particularly during high-productivity intervals like the . This process supported blooms and the broader , with evidence from isotopic records indicating its persistence from the onward. In marine sediments, microbes facilitated sulfur cycling through dissimilatory sulfate reduction, especially during OAEs, where sulfate-reducing bacteria in anoxic zones generated and preserved organic carbon, influencing global budgets and conditions. Recent advances in analysis have illuminated microbial resilience following Mesozoic mass extinctions, using lipid proxies as analogs for to trace community dynamics. For instance, post-end-Triassic extinction biomarkers reveal rapid recovery of bacterial communities, with pristane and phytane indicating cyanobacterial and algal persistence amid environmental upheaval. Similarly, across the , sterane and hopane profiles demonstrate that microbial mats and sulfate-reducers quickly recolonized sediments, underscoring their ecological robustness and role in stabilizing biogeochemical cycles after biotic crises. These findings highlight how buffered disruptions, facilitating the recovery of higher trophic levels.

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