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Olson's Extinction

Olson's Extinction was a mass extinction event that occurred approximately 272 million years ago at the Kungurian–Roadian boundary in the early to middle Permian period, primarily affecting terrestrial tetrapods and marking a major faunal turnover from pelycosaur- and amphibian-dominated ecosystems to those dominated by therapsids and parareptiles.^[1]^[2] Named after paleontologist Everett C. Olson, who first noted the abrupt decline in tetrapod diversity in 1982 based on fossil records from North America and Europe, the event has been debated as either a genuine extinction or an artifact of sampling bias known as "Olson's Gap."^[3] Recent analyses using Bayesian tip-dating methods have rejected the gap hypothesis, confirming it as a real extinction with elevated extinction rates persisting for several million years.^[1] The extinction's impacts varied by latitude: in equatorial regions, amphibian species richness plummeted from 14 species to 5 species by the Roadian stage, resulting in depauperate faunas and an inverse latitudinal biodiversity gradient where temperate zones exhibited over twice the diversity of equatorial areas.^[2] In contrast, temperate latitudes experienced a rapid faunal turnover without a significant drop in overall richness, leading to the establishment of more complex therapsid-dominated ecosystems.^[2] Globally, it contributed to the loss of about two-thirds of terrestrial vertebrate families, setting the stage for evolutionary innovations in the middle Permian.^[4] Although the precise causes remain unresolved, evidence points to climatic shifts, including warmer and drier conditions that eliminated everwet biomes essential for many amphibians and early synapsids.^[2] This event, less severe than the later Permian–Triassic extinction but significant in reshaping terrestrial vertebrate evolution, highlights the role of regional environmental heterogeneity in Permian biodiversity dynamics.^[4]

Discovery and Debate

Initial Recognition

The initial recognition of Olson's Extinction is credited to paleontologist Everett C. Olson, who first proposed the event in his 1982 chapter "Extinctions of Permian and Triassic nonmarine vertebrates," published in Geological Society of America Special Paper 190. Olson analyzed the Early Permian tetrapod fossil record, noting a pronounced hiatus and diversity decline that separated pelycosaur-dominated faunas of the lower Permian from therapsid-dominated assemblages of the middle Permian.^[5] Olson's work focused on North American localities, particularly the Texas Red Beds, including formations such as the Clear Fork and Wichita groups, where fossil-bearing horizons revealed abrupt changes in tetrapod communities.^[6] These assemblages documented the sudden absence of numerous basal synapsid lineages (pelycosaurs) and temnospondyl amphibians, alongside a shift toward more advanced therapsid forms, indicating a major biotic turnover rather than gradual replacement.^[5] The pattern was evident in family-level diversity, with a substantial reduction in the number of synapsid and amphibian families crossing the early-middle Permian transition.^[6] Early stratigraphic correlations established by Olson tied this diversity drop to the Kungurian-Roadian boundary in global Permian chronostratigraphy, corresponding to the Leonardian-Guadalupian transition in North American land vertebrate faunachrons.^[5] This boundary, dated to approximately 272 million years ago, marked the end of the Redtankian land vertebrate faunachron and the onset of a low-diversity interval in the fossil record.^[6] Olson's observations laid the foundation for recognizing the event as a distinct extinction pulse within the broader Permian period, which spanned from about 299 to 252 million years ago and featured multiple biotic crises among terrestrial vertebrates.^[5]

Evidence for Extinction vs. Fossil Gap

The debate surrounding Olson's Extinction centers on whether the observed faunal turnover between the early and middle Permian represents a genuine mass extinction event or merely "Olson's Gap," an artifact of incomplete fossil preservation in the early Permian record, particularly in North American and European deposits where tetrapod-bearing formations are sparse during the Kungurian-Roadian transition.^[1] This gap hypothesis posits that taphonomic biases, such as limited sedimentation or erosion in equatorial regions, obscure a continuous record, potentially inflating perceived extinction rates without reflecting true biodiversity loss.^[7] A pivotal 2020 Bayesian tip-dating analysis by Angielczyk et al. rejected the gap hypothesis, using phylogenetic clocks calibrated with fossil ages from synapsid lineages (including amniotes, caseids, and captorhinids) to demonstrate continuous diversification across the boundary, with strong Bayes factors favoring an extinction model over gap scenarios (e.g., 244–580 for amniotes).^[1] This approach integrates stratigraphic and molecular-like clock data to resolve temporal uncertainty, revealing elevated extinction rates during the late Kungurian that align with faunal shifts rather than preservational artifacts.^[7] Quantitative analyses further support a real extinction, with elevated per-taxon extinction rates above background levels throughout the Redtankian local vertebrate faunachron (encompassing the extinction interval) in Texas tetrapod assemblages, peaking in the uppermost Choza Formation and resulting in species richness dropping to less than one-quarter of pre-extinction levels.^[3] Phylogenetic methods incorporating ghost lineages—unsampled branches inferred from cladograms—underscore this as authentic loss, as corrected diversity estimates show selective lineage attrition during the event, triggering subsequent radiations like that of therapsids, rather than mere taphonomic gaps.^[8] Recent 2024 research using updated Permo-Carboniferous synapsid datasets reinforces the event's reality, with a new gorgonopsian therapsid fossil from the Mediterranean (dated ~290–265 Ma) documenting rapid post-extinction diversification (~278–268 Ma) following the loss of dominant pelycosaurian groups, highlighting temporal shifts in clade dominance and ecological opportunities arising from the turnover.^[9] These findings, integrated with skyline Fossilized Birth-Death models, depict a protracted decline culminating in the Kungurian-Roadian boundary, affirming elevated extinction as a driver of Permian tetrapod evolution.^[10]

Geological Context

Timing and Stratigraphy

Olson's Extinction occurred approximately 272 million years ago, precisely at the Kungurian-Roadian stage boundary during the Cisuralian Epoch of the Permian Period.^[11] This temporal placement aligns with the transition from late Leonardian time in North American chronostratigraphy to the early Guadalupian, marking a pivotal shift in the Permian timescale.^[12] In North America, the event is primarily documented through stratigraphic units such as the Clear Fork Formation in Texas and equivalent beds like the Hennessey Formation in Oklahoma, where pre-extinction vertebrate assemblages dominate the upper portions.^[13] These units, characterized by red beds and evaporites indicative of semi-arid conditions, preserve the last substantial records of Kungurian tetrapod faunas before the boundary. Global correlations with Russian and European sections rely on biostratigraphic markers, including conodont zones such as Neostreptognathodus prayi for middle Kungurian levels and fusulinid taxa like Schubertella melonica and Parafusulina brooksensis for upper Leonardian assignments, facilitating precise alignment across Pangea.^[13]^[14] The duration of the extinction is estimated at 0.5 to 2 million years, reflecting a relatively brief interval of instability relative to broader Permian timescales.^[2] This timeframe is delineated by abrupt faunal turnover in vertebrate microfossil sites, particularly within the Clear Fork Group and overlying San Angelo Formation, where bone concentrations reveal a sharp decline and replacement of dominant synapsid clades. Such sites, often yielding disarticulated skeletal elements from floodplain deposits, provide high-resolution evidence of the event's rapidity in temperate regions.^[15]

Associated Environmental Changes

During the early Permian, central Euramerica experienced a marked shift toward more arid continental climates, as indicated by the widespread development of red bed formations and evaporite deposits. In the western Catalan Pyrenees, the Lower Red Unit, dated to the Artinskian stage (approximately 290 Ma), consists primarily of red mudstones and volcaniclastic deposits with calcic paleosols (calcisols), reflecting semi-arid to arid conditions with annual precipitation likely below 500 mm.^[16] These red beds signify oxidized, well-drained soils under seasonal dryness, contrasting with the preceding humid gray-green beds of the late Carboniferous Grey Unit.^[16] Similarly, in the Permian Basin of southwestern North America, early Permian sequences include thick red beds interspersed with evaporites, such as those in the Wolfcampian Series, deposited in subsiding basins under increasingly evaporative conditions.^[17] Sea-level fluctuations characterized the early Permian sedimentary record, with notable regression events contributing to the contraction of coastal and wetland environments. The Phosphoria Formation in the western United States documents two major transgression-regression cycles during the Leonardian stage (approximately 280–290 Ma), where regressive phases involved gradual sea-level falls that exposed marginal shelves, leading to the deposition of carbonate rocks and sandstones in shallower settings. These regressions restricted marine incursions, thereby diminishing expansive wetland habitats that had persisted from the late Carboniferous, as evidenced by the shift from phosphorite-rich transgressive units to regressive carbonate-dominated sequences. Following the collapse of Carboniferous rainforests around the Carboniferous-Permian boundary, a global cooling trend emerged within the ongoing late Paleozoic icehouse, fostering seasonal aridity in tropical regions. Oxygen isotope analyses of pedogenic carbonates from paleosols in western equatorial Pangea reveal increasing δ¹⁸O values (from -7.8‰ to -1.2‰) from the latest Pennsylvanian to the early Permian, indicating enriched meteoric water δ¹⁸O values due to drier conditions and decreased rainfall by approximately 75 mm per month.^[18] This isotopic shift, observed across low-latitude sites, points to enhanced evaporation and seasonal precipitation patterns, with paleosol profiles transitioning from wet histosols to arid calcisols.^[18] Such trends align with broader early Permian paleoclimatic patterns, including deglaciation in Gondwana that indirectly amplified tropical dryness through atmospheric CO₂ rise and circulation changes.^[19]

Hypothesized Causes

Climatic and Aridification Hypotheses

One leading hypothesis for the causes of Olson's Extinction posits that intensified aridification, driven by the assembly of the supercontinent Pangea, significantly reduced atmospheric humidity and led to widespread habitat fragmentation, particularly affecting moisture-dependent tetrapods such as amphibians and early synapsids.^[20]^[2] The coalescence of Pangea around 320–300 million years ago created vast continental interiors with megamonsoonal circulation patterns, resulting in extreme equatorial aridity and the isolation of wetland habitats that had previously supported diverse tetrapod communities. This environmental stress is thought to have exacerbated drought conditions, limiting water availability and promoting the fragmentation of everwet biomes into isolated refugia, thereby contributing to the selective extinction of taxa reliant on humid environments.^[2] Climate models of the late Paleozoic support this aridification scenario, indicating a transition from the humid, glacially influenced conditions of the Carboniferous to warmer, drier climates in the early Permian, with atmospheric CO₂ levels rising from lows of around 300 ppm during the late Carboniferous glaciation to 500–1,000 ppm or higher by the early Permian, driving deglaciation but also intensifying seasonal dryness in tropical regions. This shift is closely linked to the widespread loss of Carboniferous coal swamps, which were extensive peat-forming wetlands that declined sharply due to reduced precipitation and increased evaporation, marking the end of the "coal forest" ecosystems dominated by lycopsids and ferns.^[20] Paleosol isotopic records, including δ¹⁸O and δD analyses, further corroborate these models by revealing surface temperature increases from approximately 22°C in the late Pennsylvanian to 35°C or more across the Permian-Carboniferous boundary, reflecting cooler initial phases giving way to drier, more variable conditions that stressed vegetation and associated faunas.^[21] The specific impacts of this climatic regime included severe drought stress on amphibians, which saw a dramatic decline in diversity and abundance, and early synapsids, particularly pelycosaur-grade forms, as these agile, insectivorous predators failed to adapt to the shrinking humid habitats.^[22] Varanopids, which had diversified in the humid lowlands of the Cisuralian, underwent a significant decline by the Kungurian-Roadian boundary and became extinct later in the Permian, likely due to their dependence on forested, moisture-rich environments that fragmented under arid conditions.^[22] These findings highlight how prolonged environmental drying selectively eliminated vulnerable lineages, paving the way for therapsid dominance in the post-extinction recovery.

Biotic and Geological Factors

Biotic hypotheses for Olson's Extinction emphasize competitive replacement dynamics among terrestrial vertebrates, particularly the shift from pelycosaurian-grade synapsids and temnospondyl amphibians to more advanced, drought-tolerant therapsid clades. In the early Permian (Cisuralian), faunas were dominated by basal synapsids such as sphenacodontids and edaphosaurids, alongside diverse amphibian groups including diplocaulids, which occupied aquatic and semi-aquatic niches.^[5] This assemblage underwent significant turnover around the Kungurian-Roadian boundary, with therapsids—such as dinocephalians, therocephalians, and early dicynodonts—emerging as dominant herbivores and carnivores in the middle Permian (Guadalupian). The replacement is evidenced by the abrupt decline and extinction of edaphosaurids, which last appear in the Arroyo Formation of the Texas Red Beds, and diplocaulids (e.g., Lysorophia), which vanish by the end of the Choza Formation.^[6] These changes suggest that therapsids, with adaptations for arid conditions like more efficient respiratory systems and burrowing behaviors, outcompeted less resilient clades amid environmental pressures.^[23] Evidence from the Texas Red Beds illustrates this biotic restructuring, where species richness and ecological diversity peaked in the Choza Formation before declining significantly in subsequent assemblages, with elevated extinction rates during the Redtankian local vertebrate faunachron.^[6] Amphibian clades like Eryopidae and Trimerorhachidae also disappeared by this time, while varanopid synapsids showed temporary increases but ultimately gave way to therapsid dominance. This faunal turnover was not uniform; equatorial localities experienced sharper diversity losses compared to temperate regions, supporting a biotic mechanism amplified by regional ecological shifts rather than a global sampling artifact.^[5] Geological factors potentially contributing to Olson's Extinction include localized tectonic activity associated with Pangean assembly, though direct causal links remain tentative. Ongoing uplift in the Appalachians and Urals during the early Permian likely disrupted fluvial systems and habitats, altering sediment deposition and freshwater availability in affected basins. Minor volcanic activity, such as early Permian eruptions in the Korean region, may have contributed localized ash fallout, exacerbating habitat stress through soil acidification and reduced photosynthesis, but these events were not on the scale of later Permian flood basalts. Unlike marine mass extinctions, there is no geochemical or stratigraphic evidence for bolide impacts or widespread anoxia influencing Olson's Extinction, which was primarily terrestrial.^[24] These biotic and geological elements interacted to intensify environmental stress, with competitive pressures from emerging therapsids accelerating the decline of vulnerable clades like edaphosaurids and diplocaulids in tectonically unstable landscapes. Against a backdrop of aridification, such dynamics promoted selective survival of drought-adapted forms, restructuring Permian terrestrial ecosystems without invoking catastrophic extraterrestrial or oceanic drivers.^[23]

Biotic Impacts

Terrestrial Vertebrates and Plants

The Olson's Extinction event, occurring around 272 million years ago during the Kungurian-Roadian transition, profoundly impacted terrestrial vertebrates, particularly those reliant on wetland environments. Among synapsids, numerous basal lineages suffered heavy losses, including the extinction of families such as Ophiacodontidae (e.g., Varanosaurus) and Edaphosauridae (e.g., Edaphosaurus pogonias), which were prominent in early Permian swamp ecosystems but vanished by the upper Redtankian substage in Texas formations.^[6] Similarly, amphibian diversity plummeted, exemplified by the disappearance of Eryopidae and other temnospondyl groups such as Trimerorhachidae that dominated Carboniferous-Permian wetlands.^[6] The impacts varied by latitude, with amphibian genera in equatorial regions declining from 14 to 5 by the Roadian stage, while temperate areas saw faunal turnover without significant overall richness drop.^[2] In contrast, certain synapsid clades like Caseidae (e.g., Cotylorhynchus) persisted and even remained abundant into the middle Permian, while early therapsids began to emerge, setting the stage for their later dominance. Overall, the extinction led to the loss of about two-thirds of terrestrial vertebrate families, with substantial declines in genus richness concentrated among wetland-dependent taxa such as amphibians and basal synapsids.^[4] Plant communities also experienced selective pressures, with a moderate decline in the diversity of lycopsids and ferns that characterized Carboniferous coal swamp floras. In the Texas Red Beds and associated measures, this period marks a shift from lycopsid- and fern-dominated wetlands to increasing gymnosperm prevalence, reflecting adaptations to emerging drier conditions. Evidence from these strata indicates reduced peat accumulation, signaling a contraction of extensive swamp environments and a transition toward more arid-tolerant vegetation assemblages. This floral reorganization paralleled the vertebrate losses, as the diminishing wetland habitats disproportionately affected moisture-dependent species. The extinction's terrestrial bias is evident in its patterns, with greatest impacts on humidity-loving tetrapods and swamp flora, while insects and terrestrial invertebrates showed no comparable mass die-off, maintaining relative stability in their diversity.^[6] In brief contrast, aquatic ecosystems experienced minimal disruption during this interval.

Aquatic Life

Aquatic ecosystems experienced relatively minor disruptions during Olson's Extinction compared to the severe losses on land, highlighting the event's predominantly terrestrial focus. Fish assemblages showed limited turnover, with increased extinction rates among freshwater groups such as lungfish (Dipnoi) and sharks (Chondrichthyes), but balanced by origination leading to overall stability. Lungfish diversity declined during the Cisuralian epoch, reflecting broader reductions in freshwater osteichthyan taxa, while shark populations faced moderate pressures but no catastrophic wipeout at the Cisuralian-Guadalupian boundary. In contrast, palaeoniscid fishes persisted robustly in Permian lake environments, maintaining their role as dominant components of freshwater communities without significant interruption. Marine invertebrate faunas likewise exhibited muted responses, with no pronounced extinction signals in key groups like ammonoids and brachiopods, unlike the more devastating marine crises of the Devonian. Global diversity trends for marine invertebrates indicate stability through the early to middle Permian transition, lacking the sharp declines seen later in the end-Guadalupian and end-Permian events. Conodonts, however, underwent minor faunal turnover at this time, primarily linked to eustatic sea-level fluctuations rather than biotic crisis, as evidenced by correlations between lowstands in the late Kungurian and subsequent radiations in the Roadian.^[25] The subdued aquatic impacts are attributed to relatively stable oceanic conditions, including the absence of widespread anoxia that characterized later Permian crises. Evidence from equatorial shelf deposits, such as those in the Tethyan realm, supports continuous reef development across the Cisuralian-Guadalupian boundary, with sponge-microbial and bryozoan-algal structures showing no major hiatus or collapse.^[26] This persistence underscores how marine habitats buffered the environmental stresses—primarily aridification and habitat fragmentation—that disproportionately affected terrestrial systems.

Recovery Patterns

Short-term Ecological Recovery

Following Olson's Extinction, surviving synapsid groups, particularly early therapsids such as dinocephalians, underwent rapid diversification over the subsequent 1-5 million years, occupying vacated ecological niches previously held by pelycosaurian-grade synapsids and amphibians.^[15] This rebound included a notable increase in herbivorous forms, with dicynodonts and caseids emerging as key browsers and grazers in recovering terrestrial communities, reflecting a shift toward more efficient exploitation of plant resources in post-extinction landscapes.^[15] Plant communities exhibited a swift recovery, marked by the regrowth of floodplain forests where callipterid seed ferns largely replaced moisture-dependent lycopsids as dominant vegetation.^[27] Fossil evidence from the San Angelo Formation in Texas documents this transition, with callipterid pteridosperms forming dense stands in riparian settings, adapted to the increasingly seasonal water availability and supporting the base of recovering food webs.^[27] These changes drove broader ecological shifts toward arid-adapted communities, characterized by diminished amphibian dominance—from 14 to 5 species in Roadian assemblages—and a corresponding rise in reptiliomorphs, including therapsids and parareptiles, which better tolerated the drying climate and expanded into upland and semi-arid habitats.^[15] This reorganization stabilized ecosystems within a few million years, setting the stage for later evolutionary developments.^[15]

Long-term Evolutionary Shifts

Olson's Extinction facilitated the radiation of therapsids, a group of synapsids that would eventually give rise to mammals, by eliminating dominant pelycosaurian-grade taxa such as ophiacodontids and edaphosaurids, thereby creating vacant ecological niches.^[9] This event, occurring around 273 million years ago at the Cisuralian-Guadalupian boundary, allowed therapsids—previously minor components of early Permian faunas—to diversify rapidly into diverse phenotypes and roles, including apex predation and herbivory.^[9] The extinction contributed significantly to Permian tetrapod turnover, marking a shift from pelycosaur-dominated assemblages to therapsid-led communities and serving as an early precursor to subsequent events like the Capitanian (end-Guadalupian) extinction around 260 million years ago and the end-Permian mass extinction.^[4] Recent studies, including analyses from 2024, have linked Olson's Extinction to increased synapsid morphological disparity, with therapsids exhibiting expanded functional feeding groups—such as power shearers and speed specialists—beyond the raptorial modes of earlier synapsids, signaling greater trophic dynamism in late Paleozoic terrestrial ecosystems.^[28] On a broader scale, the evolutionary shifts triggered by Olson's Extinction enhanced the resilience of terrestrial ecosystems to aridification, as therapsid radiations produced more adaptable guilds capable of exploiting drought-stressed environments, a trait that influenced the protracted Triassic recovery following the end-Permian extinction.^[4] By restructuring communities toward taxa like dicynodonts and gorgonopsians that tolerated low-diversity, arid conditions, the event preconditioned Permian biotas for the ecological imbalances seen in the latest Permian, ultimately aiding the refilling of guilds over 10-15 million years into the Triassic.^[4]

References

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[2]
Olson's Extinction and the latitudinal biodiversity gradient of ...
Apr 5, 2017 · The terrestrial vertebrate fauna underwent a substantial change in composition between the lower and middle Permian. The lower Permian fauna ...
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May 15, 2018 · Benson & Upchurch (2013) suggested that the mass extinction was an artefact of the geographically patchy fossil record.
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Recognition of Olson's gap does not preclude the possibility of an extinction at the early-middle Permian boundary (“Olson's extinction”). However, the gap ...
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Nov 23, 2015 · The calculated extinction rates are consistent with the hypothesis that Olson's Extinction impacted severely on synapsids (Fig. 3A) ...
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Dec 17, 2024 · ... Olson's Extinction. Our findings place this unambiguous early therapsid in an ancient summer wet biome of equatorial Pangaea, thus ...
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A review of the global record of Permian tetrapod body fossils and footprints reveals that this is a hiatus of global extent. Olson's gap corresponds to a ...
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May 5, 2019 · Recognition of Olson's gap does not preclude the possibility of an extinction at the early-middle Permian boundary (“Olson's extinction”).
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Apr 5, 2017 · Olson's Extinction has been demonstrated in global diversity estimates using sampling correction [4,5,10,16], but even so there is still ...
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### Summary of Olson's Extinction and Related Topics
[22]
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Apr 28, 2024 · Significant global carbon cycle disturbances triggered major environmental and climatic changes and mass extinction events globally.
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[25]
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[26]
Evolution of Permian conodont provincialism and its significance in ...
Extinction and radiation trends correlate well with sea-level changes and sequence boundaries: faunal turnovers correspond to lowstands in the late ...
[27]
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Jan 1, 2013 · The Permian reef facies studied in the EEP contain fusulinids, corals, Tubiphytes and Archaeolithoporella; some of which are probably of ...Missing: continuous | Show results with:continuous
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From wetlands to wet spots: Environmental tracking and the fate of ...
Jan 1, 2006 · This flora occurs in the San Angelo and Blaine Formations of the Pease River Group. These rocks are upper Lower Permian to lower Middle Permian ...
[29]
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Feb 17, 2024 · We find a major morphofunctional shift in synapsid carnivory between the early and middle Permian, via the addition of new feeding modes.