The great catastrophe: causes of the Permo-Triassic marine mass extinction.

The marine losses during the Permo-Triassic mass extinction were the worst ever experienced. All groups were badly affected, especially amongst the benthos (e.g. brachiopods, corals, bryozoans, foraminifers, ostracods). Planktonic populations underwent a fundamental change with eukaryotic algae being replaced by nitrogen-fixing bacteria, green-sulphur bacteria, sulphate-reducing bacteria and prasinophytes. Detailed studies of boundary sections, especially those in South China, have resolved the crisis to a ∼55 kyr interval straddling the Permo-Triassic boundary. Many of the losses occur at the beginning and end of this interval painting a picture of a two-phase extinction. Improved knowledge of the extinction has been supported by numerous geochemical studies that allow diverse proposed extinction mechanisms to be studied. A transition from oxygenated to anoxic-euxinic conditions is seen in most sections globally, although the intensity and timing shows regional variability. Decreased ocean ventilation coincides with rapidly rising temperatures and many extinction scenarios attribute the losses to both anoxia and high temperatures. Other kill mechanisms include ocean acidification for which there is conflicting support from geochemical proxies and, even less likely, siltation (burial under a massive influx of terrigenous sediment) which lacks substantive sedimentological evidence. The ultimate driver of the catastrophic changes at the end of the Permian was likely Siberian Trap eruptions and their associated carbon dioxide emissions with consequences such as warming, ocean stagnation and acidification. Volcanic winter episodes stemming from Siberian volcanism have also been linked to the crisis, but the short-term nature of these episodes (

Dynamic redox and nutrient cycling response to climate forcing in the Mesoproterozoic ocean.

Controls on Mesoproterozoic ocean redox heterogeneity, and links to nutrient cycling and oxygenation feedbacks, remain poorly resolved. Here, we report ocean redox and phosphorus cycling across two high-resolution sections from the ~1.4 Ga Xiamaling Formation, North China Craton. In the lower section, fluctuations in trade wind intensity regulated the spatial extent of a ferruginous oxygen minimum zone, promoting phosphorus drawdown and persistent oligotrophic conditions. In the upper section, high but variable continental chemical weathering rates led to periodic fluctuations between highly and weakly euxinic conditions, promoting phosphorus recycling and persistent eutrophication. Biogeochemical modeling demonstrates how changes in geographical location relative to global atmospheric circulation cells could have driven these temporal changes in regional ocean biogeochemistry. Our approach suggests that much of the ocean redox heterogeneity apparent in the Mesoproterozoic record can be explained by climate forcing at individual locations, rather than specific events or step-changes in global oceanic redox conditions.

Paleobiology: Anatomy of a mass extinction double whammy.

The Permo-Triassic mass extinction has been resolved into two closely spaced crises that both saw enormous extinction losses. However, food web modelling suggests they were not ecologically equivalent, only the second destabilised communities.

Volcanically driven lacustrine ecosystem changes during the Carnian Pluvial Episode (Late Triassic).

The Late Triassic Carnian Pluvial Episode (CPE) saw a dramatic increase in global humidity and temperature that has been linked to the large-scale volcanism of the Wrangellia large igneous province. The climatic changes coincide with a major biological turnover on land that included the ascent of the dinosaurs and the origin of modern conifers. However, linking the disparate cause and effects of the CPE has yet to be achieved because of the lack of a detailed terrestrial record of these events. Here, we present a multidisciplinary record of volcanism and environmental change from an expanded Carnian lake succession of the Jiyuan Basin, North China. New U-Pb zircon dating, high-resolution chemostratigraphy, and palynological and sedimentological data reveal that terrestrial conditions in the region were in remarkable lockstep with the large-scale volcanism. Using the sedimentary mercury record as a proxy for eruptions reveals four discrete episodes during the CPE interval (ca. 234.0 to 232.4 Ma). Each eruptive phase correlated with large, negative C isotope excursions and major climatic changes to more humid conditions (marked by increased importance of hygrophytic plants), lake expansion, and eutrophication. Our results show that large igneous province eruptions can occur in multiple, discrete pulses, rather than showing a simple acme-and-decline history, and demonstrate their powerful ability to alter the global C cycle, cause climate change, and drive macroevolution, at least in the Triassic.

Six-fold increase of atmospheric pCO2 during the Permian-Triassic mass extinction.

The Permian-Triassic mass extinction was marked by a massive release of carbon into the ocean-atmosphere system, evidenced by a sharp negative carbon isotope excursion. Large carbon emissions would have increased atmospheric pCO2 and caused global warming. However, the magnitude of pCO2 changes during the PTME has not yet been estimated. Here, we present a continuous pCO2 record across the PTME reconstructed from high-resolution δ13C of C3 plants from southwestern China. We show that pCO2 increased from 426 +133/-96 ppmv in the latest Permian to 2507 +4764/-1193 ppmv at the PTME within about 75 kyr, and that the reconstructed pCO2 significantly correlates with sea surface temperatures. Mass balance modelling suggests that volcanic CO2 is probably not the only trigger of the carbon cycle perturbation, and that large quantities of 13C-depleted carbon emission from organic matter and methane were likely required during complex interactions with the Siberian Traps volcanism.

Hydrodynamic efficiency in sharks: the combined role of riblets and denticles.

We investigate the influence of smooth and ribletted shark skin on a turbulent boundary layer flow. Through laser Doppler anemometry (LDA) the role of riblets in combination with the shark skin denticle is established for the first time. Our results show that smooth denticles behave like a typical rough surface when exposed to an attached boundary layer. Drag is increased for the full range of tested dimensionless denticle widths,w+≈ 25-80, wherew+is the denticle width,w, scaled by the friction velocity,uτ, and the kinematic viscosity,ν. However, when riblets are added to the denticle crown we demonstrate there is a significant reduction in drag, relative to the smooth denticles. We obtain a modest maximum drag reduction of 2% for the ribletted denticles when compared to the flat plate, but when compared to the smooth denticles the difference in drag is in excess of 20% forw+≈ 80. This study enables a new conclusion that riblets have evolved as a mechanism to reduce or eliminate the skin friction increase due to the presence of scales (denticles). The combination of scales and riblets is hydrodynamically efficient in terms of skin-friction drag, while also acting to maintain flow attachment, and providing the other advantages associated with scales, e.g. anti-fouling, abrasion resistance, and defence against parasites.

Extinction and dawn of the modern world in the Carnian (Late Triassic).

The Carnian Pluvial Episode (Late Triassic) was a time of global environmental changes and possibly substantial coeval volcanism. The extent of the biological turnover in marine and terrestrial ecosystems is not well understood. Here, we present a meta-analysis of fossil data that suggests a substantial reduction in generic and species richness and the disappearance of 33% of marine genera. This crisis triggered major radiations. In the sea, the rise of the first scleractinian reefs and rock-forming calcareous nannofossils points to substantial changes in ocean chemistry. On land, there were major diversifications and originations of conifers, insects, dinosaurs, crocodiles, lizards, turtles, and mammals. Although there is uncertainty on the precise age of some of the recorded biological changes, these observations indicate that the Carnian Pluvial Episode was linked to a major extinction event and might have been the trigger of the spectacular radiation of many key groups that dominate modern ecosystems.

An enormous sulfur isotope excursion indicates marine anoxia during the end-Triassic mass extinction.

The role of ocean anoxia as a cause of the end-Triassic marine mass extinction is widely debated. Here, we present carbonate-associated sulfate δ34S data from sections spanning the Late Triassic-Early Jurassic transition, which document synchronous large positive excursions on a global scale occurring in ~50 thousand years. Biogeochemical modeling demonstrates that this S isotope perturbation is best explained by a fivefold increase in global pyrite burial, consistent with large-scale development of marine anoxia on the Panthalassa margin and northwest European shelf. This pyrite burial event coincides with the loss of Triassic taxa seen in the studied sections. Modeling results also indicate that the pre-event ocean sulfate concentration was low (<1 millimolar), a common feature of many Phanerozoic deoxygenation events. We propose that sulfate scarcity preconditions oceans for the development of anoxia during rapid warming events by increasing the benthic methane flux and the resulting bottom-water oxygen demand.

Flat latitudinal diversity gradient caused by the Permian-Triassic mass extinction.

The latitudinal diversity gradient (LDG) is recognized as one of the most pervasive, global patterns of present-day biodiversity. However, the controlling mechanisms have proved difficult to identify because many potential drivers covary in space. The geological record presents a unique opportunity for understanding the mechanisms which drive the LDG by providing a direct window to deep-time biogeographic dynamics. Here we used a comprehensive database containing 52,318 occurrences of marine fossils to show that the shape of the LDG changed greatly during the Permian-Triassic mass extinction from showing a significant tropical peak to a flattened LDG. The flat LDG lasted for the entire Early Triassic (∼5 My) before reverting to a modern-like shape in the Middle Triassic. The environmental extremes that prevailed globally, especially the dramatic warming, likely induced selective extinction in low latitudes and accumulation of diversity in high latitudes through origination and poleward migration, which combined together account for the flat LDG of the Early Triassic.

Permo-Triassic boundary carbon and mercury cycling linked to terrestrial ecosystem collapse.

Records suggest that the Permo-Triassic mass extinction (PTME) involved one of the most severe terrestrial ecosystem collapses of the Phanerozoic. However, it has proved difficult to constrain the extent of the primary productivity loss on land, hindering our understanding of the effects on global biogeochemistry. We build a new biogeochemical model that couples the global Hg and C cycles to evaluate the distinct terrestrial contribution to atmosphere-ocean biogeochemistry separated from coeval volcanic fluxes. We show that the large short-lived Hg spike, and nadirs in δ202Hg and δ13C values at the marine PTME are best explained by a sudden, massive pulse of terrestrial biomass oxidation, while volcanism remains an adequate explanation for the longer-term geochemical changes. Our modelling shows that a massive collapse of terrestrial ecosystems linked to volcanism-driven environmental change triggered significant biogeochemical changes, and cascaded organic matter, nutrients, Hg and other organically-bound species into the marine system.

The latitudinal diversity gradient of tetrapods across the Permo-Triassic mass extinction and recovery interval.

The decline in species richness from the equator to the poles is referred to as the latitudinal diversity gradient (LDG). Higher equatorial diversity has been recognized for over 200 years, but the consistency of this pattern in deep time remains uncertain. Examination of spatial biodiversity patterns in the past across different global climate regimes and continental configurations can reveal how LDGs have varied over Earth history and potentially differentiate between suggested causal mechanisms. The Late Permian-Middle Triassic represents an ideal time interval for study, because it is characterized by large-scale volcanic episodes, extreme greenhouse temperatures and the most severe mass extinction event in Earth history. We examined terrestrial and marine tetrapod spatial biodiversity patterns using a database of global tetrapod occurrences. Terrestrial tetrapods exhibit a bimodal richness distribution throughout the Late Permian-Middle Triassic, with peaks in the northern low latitudes and southern mid-latitudes around 20-40° N and 60° S, respectively. Marine reptile fossils are known almost exclusively from the Northern Hemisphere in the Early and Middle Triassic, with highest diversity around 20° N. Reconstructed terrestrial LDGs contrast strongly with the generally unimodal gradients of today, potentially reflecting high global temperatures and prevailing Pangaean super-monsoonal climate system during the Permo-Triassic.

Deep CO2 in the end-Triassic Central Atlantic Magmatic Province.

Large Igneous Province eruptions coincide with many major Phanerozoic mass extinctions, suggesting a cause-effect relationship where volcanic degassing triggers global climatic changes. In order to fully understand this relationship, it is necessary to constrain the quantity and type of degassed magmatic volatiles, and to determine the depth of their source and the timing of eruption. Here we present direct evidence of abundant CO2 in basaltic rocks from the end-Triassic Central Atlantic Magmatic Province (CAMP), through investigation of gas exsolution bubbles preserved by melt inclusions. Our results indicate abundance of CO2 and a mantle and/or lower-middle crustal origin for at least part of the degassed carbon. The presence of deep carbon is a key control on the emplacement mode of CAMP magmas, favouring rapid eruption pulses (a few centuries each). Our estimates suggest that the amount of CO2 that each CAMP magmatic pulse injected into the end-Triassic atmosphere is comparable to the amount of anthropogenic emissions projected for the 21st century. Such large volumes of volcanic CO2 likely contributed to end-Triassic global warming and ocean acidification.

Mercury Spikes Indicate a Volcanic Trigger for the Late Ordovician Mass Extinction Event: An Example from a Deep Shelf of the Peri-Baltic Region.

The Late Ordovician mass extinction (LOME) was the second largest Phanerozoic crisis, but its cause remains elusive. Several triggering mechanisms have been proposed over the years, including bioevolutionary events, oceanographic changes, and geotectonic processes. Here, we report the presence of Hg spikes in the Zbrza PIG-1 borehole from the Upper Ordovician deep shelf sections of the peri-Baltic region. A strong positive anomaly in the lower late Katian (Hg/TOC = 2537.3 ppb/wt%) was noted. No correlation between Hg and TOC (R2 = 0.07) was distinguished in the Hirnantian, although several positive anomalies were found. Because the Hg/Mo ratio showed trends very similar to those of Hg/TOC, it seems likely that TOC values reflect the redox conditions. In order to evaluate the role of anoxia in levels of Hg enrichment several redox indicators were measured. These showed that the elevated mercury values in the Hirnantian are not caused by anoxia/euxinia because euxinic biomarkers (maleimides and aryl isoprenoids) are present in very low abundance and pyrite framboids are absent. In total, positive Hg/TOC anomalies occur in the lower late Katian, at the Katian - Hirnantian boundary, and in the late Hirnantian. The lack of a strong Hg/TOC correlation, Ni enrichments, and the absence of 'anoxic indicators' (no biomarkers, no framboids, low Mo concentration) at these levels, supports the interpretation that Hg enrichment is due to enhanced environmental loading. We conclude that our Hg and Hg/TOC values were associated with volcanic pulses which triggered the massive environmental changes resulting in the Late Ordovician mass extinction.

Decoupled taxonomic and ecological recoveries from the Permo-Triassic extinction.

The Permian-Triassic mass extinction was the worst crisis faced by life; it killed >90% of marine species in less than 0.1 million years (Ma). However, knowledge of its macroecological impact over prolonged time scales is limited. We show that marine ecosystems dominated by non-motile animals shifted to ones dominated by nektonic groups after the extinction. In Triassic oceans, animals at high trophic levels recovered faster than those at lower levels. The top-down rebuilding of marine ecosystems was still underway in the latest Triassic, ~50 Ma after the extinction, and contrasts with the ~5-Ma recovery required for taxonomic diversity. The decoupling between taxonomic and ecological recoveries suggests that a process of vacant niche filling before reaching the maximum environmental carrying capacity is independent of ecosystem structure building.

The onset of widespread marine red beds and the evolution of ferruginous oceans.

Banded iron formations were a prevalent feature of marine sedimentation ~3.8-1.8 billion years ago and they provide key evidence for ferruginous oceans. The disappearance of banded iron formations at ~1.8 billion years ago was traditionally taken as evidence for the demise of ferruginous oceans, but recent geochemical studies show that ferruginous conditions persisted throughout the later Precambrian, and were even a feature of Phanerozoic ocean anoxic events. Here, to reconcile these observations, we track the evolution of oceanic Fe-concentrations by considering the temporal record of banded iron formations and marine red beds. We find that marine red beds are a prominent feature of the sedimentary record since the middle Ediacaran (~580 million years ago). Geochemical analyses and thermodynamic modelling reveal that marine red beds formed when deep-ocean Fe-concentrations were > 4 nM. By contrast, banded iron formations formed when Fe-concentrations were much higher (> 50 µM). Thus, the first widespread development of marine red beds constrains the timing of deep-ocean oxygenation.The evolution of oceanic redox state in the past is poorly known. Here, the authors present a temporal record of banded iron formations and marine red beds, which indicate deep-ocean oxygenation occurred in the middle Ediacaran, coinciding with the onset of widespread marine red beds.

Macrofossil evidence for a rapid and severe Cretaceous-Paleogene mass extinction in Antarctica.

Debate continues about the nature of the Cretaceous-Paleogene (K-Pg) mass extinction event. An abrupt crisis triggered by a bolide impact contrasts with ideas of a more gradual extinction involving flood volcanism or climatic changes. Evidence from high latitudes has also been used to suggest that the severity of the extinction decreased from low latitudes towards the poles. Here we present a record of the K-Pg extinction based on extensive assemblages of marine macrofossils (primarily new data from benthic molluscs) from a highly expanded Cretaceous-Paleogene succession: the López de Bertodano Formation of Seymour Island, Antarctica. We show that the extinction was rapid and severe in Antarctica, with no significant biotic decline during the latest Cretaceous, contrary to previous studies. These data are consistent with a catastrophic driver for the extinction, such as bolide impact, rather than a significant contribution from Deccan Traps volcanism during the late Maastrichtian.

Anoxia/high temperature double whammy during the Permian-Triassic marine crisis and its aftermath.

The Permian-Triassic mass extinction was the most severe biotic crisis in the past 500 million years. Many hypotheses have been proposed to explain the crisis, but few account for the spectrum of extinction selectivity and subsequent recovery. Here we show that selective losses are best accounted for by a combination of lethally warm, shallow waters and anoxic deep waters that acted to severely restrict the habitable area to a narrow mid-water refuge zone. The relative tolerance of groups to this double whammy provides the first clear explanation for the selective extinction losses during this double-pulsed crisis and also the fitful recovery. Thus, high temperature intolerant shallow-water dwellers, such as corals, large foraminifers and radiolarians were eliminated first whilst high temperature tolerant ostracods thrived except in anoxic deeper-waters. In contrast, hypoxia tolerant but temperature intolerant small foraminifers were driven from shallow-waters but thrived on dysoxic slopes margins. Only those mollusc groups, which are tolerant of both hypoxia and high temperatures, were able to thrive in the immediate aftermath of the extinction. Limited Early Triassic benthic recovery was restricted to mid-water depths and coincided with intervals of cooling and deepening of water column anoxia that expanded the habitable mid-water refuge zone.

Response to comment on "lethally hot temperatures during the Early Triassic greenhouse".

Goudemand et al. replot a subset of our well-constrained data using a new Early Triassic biostratigraphic scheme based on a lower-resolution ammonoid zonation scheme and hypothetical ammonoid-conodont correlation to produce a less distinct seawater temperature history. We dispute their unsubstantiated correlation and, consequently, their allegations.

Lethally hot temperatures during the Early Triassic greenhouse.

Global warming is widely regarded to have played a contributing role in numerous past biotic crises. Here, we show that the end-Permian mass extinction coincided with a rapid temperature rise to exceptionally high values in the Early Triassic that were inimical to life in equatorial latitudes and suppressed ecosystem recovery. This was manifested in the loss of calcareous algae, the near-absence of fish in equatorial Tethys, and the dominance of small taxa of invertebrates during the thermal maxima. High temperatures drove most Early Triassic plants and animals out of equatorial terrestrial ecosystems and probably were a major cause of the end-Smithian crisis.