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Ordovician

The Ordovician Period is a geologic period within the Era, spanning from 485.4 to 443.8 million years ago and lasting approximately 41.6 million years, during which marine life underwent significant diversification while continents began to converge toward the formation of supercontinents. This period represents about 0.92% of Earth's total geologic history and follows the Cambrian Period, preceding the . It is named after the , an ancient tribe in where rocks from this time were first studied extensively. Geologically, the Ordovician featured widespread shallow seas covering much of the continents, with the supercontinent positioned over the South Pole and other landmasses like (proto-North America) near the equator. drove major events, including the collision of volcanic island arcs with eastern to form the and mountains, and the onset of that contributed to the later assembly of . was initially warm and stable in the early to mid-period, supporting expansive marine environments, but shifted dramatically in the Late Ordovician to a global as 's position triggered glaciation and sea-level drop. Biologically, the Ordovician is renowned for the "," where marine invertebrate genera expanded fourfold, accounting for 12% of all known marine fauna and marking a transition from Cambrian-style ecosystems to more complex ones. Key groups included trilobites, brachiopods, , , bryozoans, and the first coral reefs, alongside early vertebrates like ostracoderms (jawless armored fish) and . On land, primitive plants resembling mosses and lycophytes emerged around 460 million years ago, while arthropods began colonizing terrestrial habitats. The period ended with the , the first of the "" extinctions, which eliminated about 85% of marine species, including 25% of families such as many trilobites, brachiopods, and , primarily due to cooling climates, glaciation, and habitat loss from falling sea levels. This event reshaped marine ecosystems and set the stage for recovery.

Overview

Etymology and definition

The Ordovician Period derives its name from the Ordovices, an ancient Celtic tribe that inhabited north Wales during the Roman era, as proposed by British geologist Charles Lapworth in 1879 to resolve a longstanding stratigraphic dispute. This controversy arose in the mid-19th century between Adam Sedgwick, who classified certain Welsh rock sequences as Cambrian, and Roderick Murchison, who assigned them to his Silurian System, leading to overlapping definitions of these units. Lapworth's intervention separated the contested strata into a distinct Ordovician System, bridging the Cambrian and Silurian while honoring the regional geological heritage. The Ordovician is defined as the second geological period of the Era, spanning from approximately 485.4 to 443.8 million years ago and succeeding the Period while preceding the . It is characterized by a major episode of marine invertebrate diversification, often termed the , during which shelly faunas such as brachiopods, trilobites, and early corals proliferated across shallow marine environments. This period marks a pivotal phase in life history, with global biodiversity increasing dramatically in response to ecological opportunities in expanding epicontinental seas. The base of the Ordovician System is formally defined by the Global Stratotype Section and Point (GSSP) at Green Point, western Newfoundland, , within the mudstone facies of the Beach Formation. This boundary is marked by the first appearance datum of the conodont Iapetognathus fluctivagus, providing a precise biostratigraphic anchor for international correlation. The section's well-preserved graptolite and conodont assemblages facilitate reliable global synchronization of Ordovician strata.

Timeline and boundaries

The Ordovician Period spans from approximately 485.4 Ma to 443.8 Ma, encompassing a duration of about 41.6 million years. This temporal framework is established through integration of and , providing a precise calibration for the period's boundaries within the Eon. The lower boundary of the Ordovician, marking the Cambrian-Ordovician transition at 485.4 Ma, is defined at the Global Boundary Stratotype Section and Point (GSSP) in the Green Point section of western Newfoundland, . This boundary is characterized by the extinction of late Cambrian taxa, including members of the suborder Eodiscina, alongside the initial radiation of Early Ordovician () graptolites such as Staurograptus and Rhabdinopora. The primary marker for the GSSP is the first appearance datum of the Iapetognathus fluctivagus, which correlates globally with these biotic turnover events. The upper boundary occurs at the Ordovician-Silurian transition, dated to 443.8 Ma, and is delineated by the GSSP for the base of the Stage at the Wangjiawan section near , . This boundary coincides with the onset of the Hirnantian glaciation and the first major pulse of the , defined biostratigraphically by the first appearance of the Normalograptus extraordinarius. Radiometric ages for these boundaries derive primarily from U-Pb dating of crystals in beds (tuffs) interbedded within marine sedimentary sequences. High-precision techniques, such as chemical abrasion-isotope dilution thermal ionization (CA-ID-TIMS), have been applied to ash layers in key sections, including those in and , yielding uncertainties as low as ±0.1 to ±1.0 and anchoring the biostratigraphic framework to absolute time. For instance, ash beds near the Cambrian-Ordovician boundary in Newfoundland and provide U-Pb ages that confirm the 485.4 Ma datum, while similar dating in the Wangjiawan section supports the 443.8 Ma upper limit. The Ordovician is subdivided into three epochs: the Early Ordovician ( and stages), spanning 485.4 to 477.7 Ma; the Middle Ordovician (Dapingian and Darriwilian stages), from 477.7 to 458.4 Ma; and the Late Ordovician (, , and stages), extending to 443.8 Ma. These epoch divisions reflect major evolutionary and environmental transitions, with boundaries ratified through GSSPs that integrate , , and zonations for global correlation.

Stratigraphy and subdivisions

Global stages

The Ordovician Period is formally subdivided into seven global stages by the (ICS), forming the basis for international chronostratigraphic correlation. These stages—Early Ordovician (Tremadocian, Floian, Dapingian), Middle Ordovician (Darriwilian, Sandbian), and Late Ordovician (Katian, )—are defined by Global Boundary Stratotype Sections and Points (GSSPs), with boundaries ratified through biostratigraphic criteria emphasizing , , and trilobites as primary index s. Global correlations rely on integrated biozonations from these fossil groups, which provide high-resolution markers across diverse lithofacies and paleogeographic realms. Recent updates to the International Chronostratigraphic Chart (versions 2023–2024) have refined stage durations using astrochronology, particularly for the Middle Ordovician, alongside U-Pb radioisotopic dating to achieve uncertainties as low as ±0.7 Ma for some boundaries. A 2025 study proposes further refinements for the Late Ordovician, dating the Katian- boundary at 442.65 +0.17/−0.23 Ma and the Ordovician-Silurian boundary at 442.33 +0.34/−0.33 Ma using high-precision U-Pb , potentially shortening the Hirnantian stage, though these await official ICS ratification. This framework ensures precise temporal resolution for studying Ordovician events, such as biodiversification and mass extinctions. The following table summarizes the global stages, their durations, and representative key biostratigraphic markers (using official ICS ages as of 2024):
StageDuration (Ma)Key Index Fossils and Biozones
Tremadocian485.4–477.7Graptolites: Rhabdinopora praeparabola (base-defining FAD), Adelograptus and Expansograptus zones; Conodonts: Iapetognathus fluctivagus (base at Green Point GSSP); Trilobites: Symphysurina spp. and Jujuyaspis borealis.
Floian477.7–470.0Graptolites: Paratetragraptus approximatus (base at Diabasbrottet GSSP); Conodonts: Oelandodus elongatus–Acodus deltatus Subzone (Paroistodus proteus Zone); Trilobites: Megistaspis (Paramegistaspis) planilimbata Zone.
Dapingian470.0–467.3Graptolites: Azygograptus ellesi (upper A. suecicus Zone), Isograptus victoriae victoriae Zone; Conodonts: Baltoniodus triangularis (base at Huanghuachang GSSP).
Darriwilian467.3–458.4Graptolites: Levisograptus austrodentatus (base at Huangnitang GSSP); Conodonts: Lenodus antivariabilis and Histiodella sinuosa zones.
Sandbian458.4–453.0Graptolites: Nemagraptus gracilis (base at Sularp Brook GSSP); Conodonts: Pygodus anserinus Zone, Amorphognathus inaequalis Subzone.
Katian453.0–445.2Graptolites: Diplacanthograptus caudatus (base at Black Knob Ridge GSSP); Conodonts: Upper Amorphognathus tvaerensis Zone.
Hirnantian445.2–443.8Graptolites: Normalograptus extraordinarius (base at Wangjiawan GSSP); Trilobites: Hirnantia–Dalmanitina fauna.

Regional correlations and British stages

The Ordovician System in its type area in has historically been subdivided into six chronostratigraphic units known as the Tremadoc, Arenig, Llanvirn, , Caradoc, and Ashgill series or stages, established primarily on lithostratigraphic and biostratigraphic grounds in the 19th and early 20th centuries. These stages served as a foundational framework for early Ordovician correlations but required revision to align with the global standard ratified by the (). Modern mappings integrate , , and chitinozoan to link them to the global series and stages: the Tremadoc corresponds to the Tremadocian Stage; the Arenig to the Floian and lowermost Dapingian stages; the Llanvirn to the upper Dapingian and Darriwilian stages; the to the upper Darriwilian Stage; the Caradoc to the Sandbian and lowermost Katian stages; and the Ashgill to the upper Katian and stages. This alignment facilitates precise inter-regional comparisons while preserving the historical nomenclature for sections. Correlating these stages to other regional chronostratigraphies presents challenges due to pronounced variations across paleocontinents, which influenced faunal distributions and preservation. For instance, North American sequences like the carbonate-rich and Whiterock successions in the exhibit shallow-shelf environments with endemic and assemblages, contrasting with the deeper-water, clastic-dominated shales and mudstones of European sections in and , where are more abundant. Such lithofacies differences led to initial mismatches in Lower Ordovician correlations between () and (including ), as faunas reflect distinct zoogeographic provinces shaped by ocean barriers and climate gradients. To overcome these issues, trans-regional matching relies heavily on index fossils from and biozones, which provide high-resolution markers less affected by local ; for example, the appearance of like Didymograptus bifidus and such as Baltoniodus triangularis anchors the base of the Llanvirn across continents. These biozonations, combined with chemostratigraphic signals like carbon excursions, enable reliable linkages between disparate belts. The ICS Subcommission on Ordovician Stratigraphy continues to advance global integration of regional schemes through initiatives, including the IGCP Project 735 ("Rocks and the Rise of Ordovician Life"; 2021–2025), with ongoing efforts extending into 2028. Key efforts involve compiling updated regional syntheses, such as a third Geological Society of London Special Publication targeted for mid-2027, and hosting international symposia like the 15th International Symposium on the Ordovician System in , , in August 2027, with field excursions to key Chinese sections. Recent advancements include 2025 discoveries in southern , where new bivalve, , and fossils from the Disi Sandstone and Mudawwara formations have prompted re-evaluation of Middle to Upper Ordovician body fossil records, enhancing correlations with North Gondwanan margins. In , 2025 studies on Ordovician bivalves from the Dali area in reveal faunal affinities bridging and Indochina paleoplates, supporting refined biostratigraphic ties to global stages via and zones. These findings, integrated with the 2024 inauguration of the Xiaoyangqiao auxiliary boundary stratotype section in , bolster high-resolution correlations across Asia and beyond.

Paleogeography and tectonics

Continental configurations

During the Ordovician Period, the global continental configuration featured the as the dominant landmass in the , positioned at high southern latitudes, while (present-day ), ( and parts of ), and (a microcontinent including parts of ) occupied more northerly positions near the . and existed as independent tectonic plates, separate from these major blocks, with located in the and in a transitional position between and . Paleogeographic reconstructions of these configurations rely on paleomagnetic data, which provide latitude estimates through apparent paths, and fossil distributions, such as endemic and faunas that delineate provincial boundaries. For instance, high-latitude glacial deposits preserved in , including tillites and striated pavements in regions like , Arabia, and , confirm its polar position during the Late Ordovician ( stage), supporting reconstructions that place the over northern or eastern around 445 Ma. Gondwana underwent a progressive northward drift toward the throughout the period, shifting from southern polar latitudes in the Early Ordovician to lower southern latitudes by the Late Ordovician, which contributed to by enhancing continental and altering circulation patterns. This movement is evidenced by paleomagnetic poles from n rocks, showing a ~10–15° northward shift between 470 Ma and 450 Ma. Meanwhile, the formed a wide seaway (over 4000 km) between to the west and Baltica- to the east, with paleomagnetic data indicating at ~10–20°S and at ~50–60°S in the Early to Middle Ordovician, setting the stage for later convergence.

Major tectonic events

The closure of the Ocean's margins during the Middle Ordovician initiated the along the eastern margin of , involving of , obduction of volcanic arcs, and accretion of terranes that deformed the continental margin. This event, spanning the Darriwilian stage (approximately 467–458 Ma), resulted from the convergence of with outboard island arcs, such as the Shelburne Falls arc, leading to widespread and sediment deposition in foreland basins. The marked a shift from sedimentation to active tectonism, with volcanic arcs accreted onto by the late Darriwilian, influencing regional basin evolution. Along the northern margin of , subduction processes during the Ordovician established early Andean-type margins characterized by calc-alkaline and arc . This involved intra-oceanic transitioning to arcs along the eastern Proto-Tethys, producing tonalitic batholiths and silicic large igneous provinces as 's margin interacted with peri-Gondwanan terranes. Multiple phases of rifting and occurred from the into the Ordovician, culminating in widespread that built crustal thickness and contributed to the Famatinian and related orogenic belts in regions now part of and . The , centered around 468 Ma in the early Darriwilian stage, represents a pulse of impacts linked to the breakup of the L-chondrite parent body in the , evidenced by meteorites, micrometeorites, and craters across multiple continents. Widespread anomalies in Middle Ordovician sediments, alongside grains in structures like the Lockne and Målingen craters in , indicate enhanced flux, with fragments up to several kilograms preserved in beds. While primarily interpreted as an -related phenomenon from collision, some evidence suggests potential ties to contemporaneous or tectonic disruption, though the dominant view attributes it to asteroidal dynamics rather than solely terrestrial processes. Recent paleomagnetic studies from 2023 to 2025 have refined estimates of Ordovician plate velocities, revealing rapid continental drifts such as Baltica's northward movement from 55°S to 33°S at rates exceeding 10 cm/year during the mid-to-late Ordovician. These analyses, based on high-resolution sampling of limestones and magnetic shifts, highlight faster-than-previously modeled plate motions that influenced global configurations, including the relative positions of and during events. Such velocities, ranging from 0.97 to 5.00 cm/year on average but with peaks up to 30 cm/year in some reconstructions, underscore the dynamic tectonic regime of the period.

Climate and paleoenvironment

Climatic variations

The Ordovician Period began under a pronounced climate, characterized by warm global oceans with low-latitude sea surface temperatures estimated at 10–15°C higher than modern values, fostering extensive marine ecosystems across equatorial regions. This warmth persisted through the Early and Middle Ordovician ( to Darriwilian stages, ca. 485–460 Ma), driven by high atmospheric CO₂ levels of 10–20 times preindustrial concentrations (approximately 2,800–5,600 ppm), which enhanced the and suppressed polar ice formation. However, a gradual cooling trend emerged by the Middle Ordovician, with sea surface temperatures declining by about 10–15°C over several million years at rates of around 1°C per million years, marking the onset of cooler conditions that set the stage for later glacial episodes. Atmospheric CO₂ levels continued to decrease toward the Late Ordovician, falling to 5–10 times preindustrial values (roughly 1,400–2,800 ppm) by the Katian stage (ca. 453–445 Ma), primarily due to enhanced silicate weathering associated with tectonic uplift from the along the Laurentian margin. This drawdown contributed to the overall cooling trajectory, culminating in the glaciation (ca. 445–443 Ma), a brief but intense centered on the southern . Evidence for this glaciation includes widespread tillites and glacial deposits in the Sahara region of , such as sandy diamictites with striated pavements in , and in , where the Cancañiri Formation in southern (with clast provenances extending to ) records multiple ice sheet advances with subglacial deformation features. The ice sheets, which reached temperate grounded conditions, were linked to sea level drops of up to 100 meters, as detailed in records of eustatic fluctuations. Recent cyclostratigraphic analyses from 2025 have revealed that , particularly 1.2-million-year obliquity variations and 405-thousand-year eccentricity cycles, modulated these climate oscillations, driving third- and fourth-order transitions between icehouse and greenhouse states in the Upper Ordovician, as evidenced by tuned records from sections showing correlations with enrichment and sea-level changes. These orbital forcings amplified the Late Ordovician cooling, with obliquity cycles influencing and contributing to the punctuated nature of the glacial maximum.

Sea level fluctuations

During the mid-Ordovician, particularly in the Darriwilian and Sandbian stages, global eustatic s reached some of their highest points in the , leading to extensive flooding of continental cratons and the development of vast epicontinental seas. In , the largest tropical tectonic plate of the time, these elevated s resulted in shallow, warm epicratonic seas that covered much of the continent, fostering diverse shallow-marine environments. This transgressive phase was driven by a combination of tectonic along passive margins and reduced ice volume under a , allowing seawater to inundate interior lowlands and create broad, low-gradient shelves. In contrast, the Late Ordovician (Katian to stages) witnessed a major eustatic , with global sea levels dropping by approximately 100 meters due to the rapid onset of Gondwanan glaciation. This drained many epicontinental seas, exposing shelves and shifting depositional environments from deep-water carbonates to shallower, terrigenous . The sea-level fall was closely tied to increased ice volume on , which locked up water and amplified cooling effects, as briefly referenced in discussions of Ordovician climatic variations. Stratigraphic records of these fluctuations are well-documented through in various Ordovician basins. In , particularly the Cincinnati Arch region of the , third-order depositional sequences reveal cyclic transgressive-regressive patterns tied to eustatic changes, with mid-Ordovician units showing thick packages indicative of maximum flooding surfaces during highstands. In sections, such as those in the and Middle-Upper Yangtze platform, sequence boundaries marked by paleokarsts and evaporites delineate major sea-level cycles, correlating the mid-Ordovician highstand with widespread platform development and the Late Ordovician with abrupt shifts to siliciclastic input. These records highlight the interplay between eustasy and local tectonic , where cratonic basins amplified global signals through differential accommodation space. Recent advances in astrochronology have refined the timing and drivers of these sea-level . A 2023 review of Ordovician cyclostratigraphy identifies Milankovitch-band periodicities, including a stable 405 kyr , as key pacemakers for eustatic variations through modulation of ice volume and dynamics. Calibration using high-precision U-Pb dating in sections from and has established floating astrochronologies that link (16-19 kyr) and obliquity (31 kyr) to high-frequency sea-level oscillations superimposed on longer-term trends. This framework underscores how contributed to the mid-Ordovician highstand stability and the pulsed nature of the Late Ordovician regression.

Geochemistry and ocean conditions

Isotopic and geochemical records

Stable isotope records from Ordovician carbonates and biogenic provide key proxies for global environmental changes, particularly perturbations in the carbon and oxygen cycles. The δ¹³C record reveals several positive that indicate disruptions in the global , often linked to enhanced burial of organic carbon or changes in inputs. One prominent event is the Mid-Darriwilian Isotope Carbon (MDICE), occurring around 465–460 Ma, characterized by a positive shift of approximately +3‰ in δ¹³C values from baseline levels of -1‰ to +2‰ in records across multiple paleocontinents, including and . This suggests a temporary increase in the fraction of organic carbon burial relative to deposition, potentially driven by expanded anoxic conditions in basins. Another significant δ¹³C perturbation is the Hirnantian Isotope Carbon Excursion (HICE), marking the latest Ordovician around 445–443 Ma, with δ¹³C values rising by +4‰ to +5‰ from pre-excursion levels of about 0‰ to peaks exceeding +4‰ in sections from peri-Gondwanan and Laurentian margins. This excursion, often divided into lower and upper peaks, reflects a major reconfiguration of the , possibly involving increased continental weathering and sequestration during the Hirnantian glaciation. The HICE is globally correlatable and coincides with the onset of the , underscoring its role as a stratigraphic marker for late Hirnantian events. Oxygen isotope (δ¹⁸O) records from and shells document a long-term cooling trend through the Ordovician, with temperatures inferred to decrease from equatorial averages of ~35–40°C in the Early Ordovician to ~25–30°C by the Late Ordovician. Specifically, δ¹⁸O values shift from around -5‰ (VSMOW) in –Floian samples to -2‰ or higher in Katian– records, reflecting a progressive increase in global ice volume and ocean cooling, particularly intensified during the Middle to Late Ordovician. This trend is evident in low-latitude sections and supports models of declining atmospheric CO₂ levels contributing to the transition toward cooler climates. Strontium isotope ratios (⁸⁷Sr/⁸⁶Sr) in Ordovician , preserved in well-screened matrices, exhibit a notable decrease from approximately 0.7091 in the Early Ordovician to 0.7081 by the Late Ordovician, signaling enhanced input of non-radiogenic strontium from hydrothermal sources. This trend, documented in bulk s and from North American and European sections, correlates with periods of mountain building and increased fluxes, which likely buffered CO₂ drawdown and influenced ocean alkalinity. Recent geochronological advancements, including high-precision U-Pb of zircons from ash beds in sections, have refined the timing of these isotopic excursions relative to the , placing the onset of the HICE and associated pulses at approximately 442.76 ± 0.05 Ma. These 2025 studies confirm a rapid tempo for the , with δ¹³C shifts occurring over less than 0.5 million years, linking climatic cooling and disruptions more precisely to the dynamics. Such refinements enhance the of global correlation for Ordovician stage boundaries and environmental events.

Ocean chemistry and anoxia

During the Ordovician, enhanced continental weathering, particularly associated with the Taconic orogeny, delivered substantial nutrient loads, including phosphorus and trace metals, to marine basins, promoting eutrophication and the development of oxygen minimum zones. This nutrient influx triggered frequent plankton blooms, increasing organic matter export and leading to mid-Ordovician anoxic events, as evidenced by the widespread deposition of basinal black shales in regions like the Yangtze Sea. These organic-rich sediments, with total organic carbon (TOC) contents often exceeding 5 wt%, reflect heightened primary productivity under nutrient-replete conditions, exacerbating seafloor anoxia and restricting benthic habitats. Seawater sulfate concentrations were lower than modern levels throughout much of the Ordovician, estimated at 5–16 mM, while availability was enhanced by weathering-derived inputs, supporting elevated productivity. (Mo) isotope records from black shales and carbonates indicate the expansion of sulfidic (euxinic) conditions in ocean basins, particularly during intervals of intensified nutrient cycling, with δ⁹⁸Mo values suggesting quantitative Mo removal under low-oxygen, sulfide-rich waters. These proxies reveal that sulfidic was more prevalent in deeper waters, contributing to the preservation of in shales and influencing budgets. Ocean ventilation exhibited dynamic shifts across the period, with early Ordovician deep waters characterized by persistent ferruginous , as shown by iron data from basinal sections indicating nonsulfidic oxygen-deficient conditions. Mid-Ordovician improved transiently, allowing for expanded oxygenation in shelf settings and supporting the , though deep basins remained prone to intermittent . This was followed by pronounced deoxygenation in the Late Ordovician, marked by global expansion of euxinic zones during the mass extinction intervals, driven by intensified recycling and organic carbon burial. Recent 2025 analyses of exceptionally preserved faunas in lowest black shales from highlight the post-Late Ordovician dynamics under lingering anoxic conditions, revealing a low-diversity assemblage dominated by sponges and cephalopods in deep-water settings. These deposits indicate intermittent oxygenation events within otherwise anoxic environments, facilitating the initial recolonization of seafloor ecosystems and underscoring a protracted phase following the .

Biodiversity and evolution

Marine fauna diversification

The Great Ordovician Biodiversification Event (GOBE), spanning much of the Ordovician Period from approximately 485 to 443 million years ago, marked an extraordinary surge in marine animal diversity, with global genera counts rising from around 500 in the Early Ordovician to over 1,200 by the Late Ordovician, achieving the fastest diversification rate in history. This radiation involved profound evolutionary innovations across multiple phyla, transforming marine ecosystems from relatively simple Cambrian-style assemblages to complex, tiered communities that foreshadowed dominance. The event's peak occurred during the Katian stage of the Late Ordovician, where metrics, including family and order-level expansions, reached unprecedented levels before the subsequent mass extinction. Major marine groups exemplified this diversification, with trilobites achieving peak generic richness through adaptive radiations in benthic habitats, including iconic genera like Asaphus that dominated shallow-shelf environments with specialized appendages for sediment feeding and predation. Brachiopods proliferated dramatically, with orthids and strophomenids evolving diverse shell morphologies to exploit varied substrates, from articulate forms in high-energy settings to strophomenid pedunculate species in deeper waters. Cephalopods, particularly nautiloids, underwent rapid innovation in shell complexity and locomotion, as evidenced by newly described Late Ordovician assemblages from the North in , revealing over a dozen genera with reticulated and annulated orthoconic forms indicative of nektonic lifestyles. Concurrently, and filled pelagic niches, with graptolite biserial colonies evolving into efficient filter-feeders and conodont elements showing increased apparatus complexity for enhanced feeding efficiency. Ecological structuring deepened during the GOBE, with benthic tiers expanding from simple epifaunal mats to multi-layered communities where trilobites and echinoderms (such as and cystoids) occupied infaunal and erect tiers above the seafloor, facilitating resource partitioning and bioturbation. In contrast, pelagic realms saw the rise of mobile predators and , including cephalopods as active swimmers and as drifting colonists, which together occupied vertical zones from surface waters to mid-water columns, enhancing trophic complexity. This between benthic and pelagic faunas decoupled diversification pulses, with offshore and deep-water assemblages lagging slightly behind nearshore ones but ultimately contributing to global ecosystem modernization. Recent paleontological discoveries have further illuminated the GOBE's scope, such as 2025 research in southern Jordan describing new bivalve, , and taxa from Ordovician strata and re-evaluating the regional body fossil record. These finds underscore ongoing revelations in understudied regions, reinforcing the event's global scale and the role of localized radiations in overall marine faunal enrichment.

Early terrestrial life and microbiota

The earliest evidence for terrestrial plant colonization during the Ordovician appears in the form of cryptospores, which are permanent dyads or tetrads of spores produced by non-vascular embryophytes. These microfossils first occur in sediments dating to approximately 480 million years ago in the Floian stage of the Early Ordovician, preserved in deposits from . Such cryptospores are interpreted as originating from liverwort-like gametophytes, representing the dominant life stage of these basal , which lacked and were likely small, thalloid organisms adapted to moist terrestrial environments. Vascular did not emerge until the period, marking a later in the greening of the continents. Terrestrial microbiota during the Ordovician was dominated by microbial communities, including fungi and algae, which formed the primary biological cover on land surfaces prior to widespread plant establishment. Fossil evidence includes hyphae and spores of glomalean fungi from approximately 460 million-year-old deposits in Wisconsin, indicating early mycorrhizal-like associations that may have facilitated nutrient cycling in primitive soils. Cyanobacteria contributed to soil crusts and microbial mats, promoting nitrogen fixation and early weathering processes, as evidenced by their preserved filaments in mid-Ordovician paleosols. Chytrid-like fungi, inferred from similar ancient microbial fossils, likely played roles in decomposition and aquatic-to-terrestrial transitions within these ecosystems. Isotopic signatures in Ordovician paleosols, such as elevated δ¹³C values, reflect increased terrestrial productivity from these microbial communities, driven by photosynthetic cyanobacteria and early fungal activity that fractionated carbon isotopes via rubisco enzymes. In marine environments, such as acritarchs—organic-walled microfossils of algal —reached in the Middle Ordovician, with assemblages exceeding 300 species in , correlating to expanded epicontinental seas and nutrient . Chitinozoans, bottle-shaped microfossils possibly related to marine metazoans, also diversified markedly during this interval, providing key index fossils for global , particularly in peri-Gondwanan and Baltoscandian sequences. These microbial groups underpinned in Ordovician oceans, influencing the base of the and enabling the diversification of higher marine organisms. A notable 2025 discovery, the "inside-out" fossil of Keurbos susanae (affectionately named "Sue") from the 444-million-year-old Soom Shale in , exemplifies exceptional preservation of soft tissues in late Ordovician biota, offering insights into contemporaneous microbial environments through its authigenic mineralization under anoxic conditions.

Late Ordovician mass extinction

Extinction pulses and impacts

The (LOME) ranks as the second-largest in Earth's history, eliminating approximately 85% of across two distinct pulses separated by roughly 1 million years. The first pulse occurred in the late Katian stage, coinciding with the onset of the glaciation and resulting in the loss of about two-thirds of marine genera, while the second pulse took place during the stage, accounting for the remaining third of genera losses. Recent 2025 geochronological studies refine the timing of these events, linking them to the carbon excursion (HICE) and emphasizing the role of rapid climatic shifts in driving the biotic turnover. Biotic impacts were profound and selective, with pelagic and nektonic groups suffering the most severe declines. experienced near-total species loss among planktic forms, nearly wiping out this key planktonic group and disrupting oceanic food webs. Trilobites saw about 50% of their genera disappear, particularly affecting shallow-water and biofacies-restricted forms, while brachiopods exhibited differential survival, with lingulids among the few clades that persisted relatively unscathed due to their infaunal, opportunistic lifestyle. These losses extended to other invertebrates, including and corals, but spared some resilient benthic taxa. The displayed distinct global and regional patterns, reflecting the heterogeneous environmental stresses. The first disproportionately struck equatorial faunas adapted to warm, stable conditions, as cooling and sea-level drop disrupted tropical ecosystems. In contrast, the second more intensely affected polar and high-latitude assemblages, where post-glacial warming and associated ocean expanded lethal conditions into cooler realms. This latitudinal selectivity highlights how the LOME's pulses amplified vulnerabilities in diverse marine habitats.

Causes and recovery dynamics

The (LOME) is widely regarded as a multifactorial event, with leading hypotheses emphasizing the interplay of environmental stressors rather than a single trigger, though debate persists on the primacy of cooling versus warming/. Glaciation over during the stage initiated the first extinction pulse (LOME I), inducing of approximately 9°C and a significant drop of up to 100 meters, which contracted shallow marine habitats and disrupted benthic communities. This cooling phase, occurring at a mean rate of 26°C per million years, selectively impacted warm-water-adapted taxa, contributing to about 8.4% loss per 100,000 years. Expanding ocean , evidenced by widespread black shales and geochemical proxies like and enrichments, exacerbated habitat loss during both extinction pulses, particularly in epicontinental seas where oxygen minimum zones intensified. from an unidentified , inferred from mercury spikes reaching 550 parts per billion in sections like Dob's Linn, , released greenhouse gases that drove and subsequent warming of 7.3°C during LOME II, at a rapid mean rate of 122°C per million years. This volcanogenic warming expanded anoxic conditions further, collapsing primary productivity and causing 71.6% species loss per 100,000 years in the second pulse, aligning the LOME mechanistically with other extinctions driven by similar processes. The overall event eliminated approximately 85% of marine species. Hypotheses involving impacts, such as a meteor strike triggering initial glaciation, remain debated due to the absence of anomalies or confirmed craters contemporaneous with the LOME, with negligible evidence from global databases. Recent 2025 geochronological analyses highlight synergies between and rapid shifts, where the abrupt transition from icehouse to conditions amplified rates beyond individual factors alone. Earlier proposals of a depleting stratospheric have not gained support from updated stratigraphic or isotopic records. Biotic recovery following the LOME exhibited pronounced ecological selectivity, with slower repopulation in deep-sea environments compared to nearshore settings. In deep waters, marked by Rhuddanian black shales, sponge-dominated assemblages like the Huangshi Fauna in South China—comprising hexactinellid sponges, cephalopods, and rare arthropods—emerged as pioneers, reflecting intermittent seafloor oxygenation but lower diversity (around 20 sponge species) than pre-extinction levels. This contrasts with faster recovery on shallow shelves, where brachiopod-rich faunas such as the Edgewood-Cathay assemblage proliferated within the late Hirnantian, indicating habitat-specific resilience among disaster taxa. Opportunistic groups, including chitinozoans and microbes, rebounded first, with chitinozoan diversity tracking recovery and surviving the crisis to diversify in the early , serving as key biostratigraphic markers. Sponges similarly acted as ecological restorers in post-extinction niches, peaking in abundance during initial recolonization phases. Long-term recovery reshaped ecosystems, leading to a shift toward biofacies characterized by increased dominance of resilient clades like strophomenoid brachiopods and reduced overall diversity until the , when pre-LOME levels were finally approached after approximately 5 million years. Persistent anoxic events delayed full stabilization, fostering evolutionary innovations in surviving lineages.

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