Jonathan L. Payne

Permian–Triassic Boundary Sections from Shallow-Marine Carbonate Platforms of the Nanpanjiang Basin, South China: Implications for Oceanic Conditions Associated with the End-Permian Extinction and Its Aftermath

Abstract Permian-Triassic boundary (PTB) sections from isolated carbonate platforms in the Nanpanjiang Basin of south China contain Upper Permian skeletal packstones with diverse open-marine fossils overlain by a 7–15 m thick boundary horizon composed of calcimicrobial framestone constructed by globular to tufted, calcified cyanobacteria similar to Renalcis. The framestone contains interbeds of lime-grainstone with abundant thin-shelled bivalves and brachiopods. The overlying Lower Triassic strata contain microgastropod lime-packstones followed by a thick succession of thin-bedded lime-mudstones. The PTB event horizon is interpreted to occur at the top of the packstone containing diverse, open-marine fauna and Palaeofusulina and coincident with the abrupt change to calcimicrobial framestone lacking Permian macrofossils. The conformable biostratigraphic boundary occurs at the first appearance of Hindeodus parvus within the basal meter of the calcimicrobial framestone. Intensively studied PTB sections in south China, such as the GSSP at Meishan, primarily are condensed sections from deep-water, basin environments that contain a thin (< 30 cm) boundary horizon of claystone and lime-mudstone or marl. The sections reported herein are fundamentally different in that they consist of shallow-marine carbonate facies, contain a thick boundary horizon composed of calcimicrobial framestone, and lack evidence of an abrupt shift in depositional environment or water depth. The calcimicrobial framestone boundary horizon occurs in all of the isolated carbonate platforms in the Nanpanjiang Basin. A similar microbial facies has been found in the basal Triassic H. parvus zone in the Sichuan Basin and in Japan. Distribution of the calcimicrobial framestone suggests that it may represent an anomalous oceanic event that affected a vast area of the equatorial eastern Tethys and Panthalassa during and/or immediately following the end-Permian mass extinction. The persistence of similar calcimicrobial framestone horizons into the Upper Scythian suggests that detrimental environmental conditions associated with the extinction persisted until the end of the Scythian. Further study of these sections promises to provide constraints on causes of the extinction and the environments in the aftermath.

Timing of recovery from the end-Permian extinction: Geochronologic and biostratigraphic constraints from south China

Four volcanic-ash beds bracket the Early-Middle Triassic boundary, as defined by conodont biostratigraphy, in a stratigraphic section in south China. High-precision U-Pb dates of single zircons allow us to place the Early to Middle Triassic (Olenekian-Anisian) boundary at 247.2 Ma. Magnetic-reversal stratigraphy allows global correlation. The new dates constrain the Early Triassic interval characterized by delayed biotic recovery and carbon-cycle instability to ∼5 m.y. This time constraint must be considered in any model for the end-Permian extinction and subsequent recovery.

Erosional truncation of uppermost Permian shallow-marine carbonates and implications for Permian-Triassic boundary events

On shallow-marine carbonate buildups in south China, Turkey, and Japan, uppermost Permian skeletal limestones are truncated by an erosional surface that exhibits as much as 10 cm of topography, including overhanging relief. Sedimentary facies, microfabrics, carbon isotopes, and cements together suggest that erosion occurred in a submarine setting. Moreover, biostratigraphic data from south China demonstrate that the surface postdates the uppermost Permian sequence boundary at the global stratotype section and truncates strata within the youngest known Permian conodont zone. The occurrences of similar truncation surfaces at the mass-extinction horizon on carbonate platforms across the global tropics, each overlain by microbial buildups, and their association with a large negative excursion in δ 13 C further suggest a causal link between erosion of shallow-marine carbonates and mass extinction. Previously proposed to account for marine extinctions, the hypothesis of rapid carbon release from sedimentary reservoirs or the deep ocean can also explain the petrographic observations. Rapid, unbuffered carbon release would cause submarine carbonate dissolution, accounting for erosion of uppermost Permian skeletal carbonates, and would be followed by a pulse of high carbonate saturation, explaining the precipitation of microbial limestones containing upwardgrowing carbonate crystal fans. Models for other carbon-release events suggest that at least 5 × 10 18 g of carbon, released in <100 k.y., would be required. Of previously hypothesized Permian-Triassic boundary scenarios, thermogenic methane production from heating of coals during Siberian Traps emplacement best accounts for petrographic characteristics and depositional environment of the truncation surface and overlying microbial limestone, as well as an associated carbon isotope excursion and physiologically selective extinction in the marine realm.

The effect of geographic range on extinction risk during background and mass extinction.

Wide geographic range is generally thought to buffer taxa against extinction, but the strength of this effect has not been investigated for the great majority of the fossil record. Although the majority of genus extinctions have occurred between major mass extinctions, little is known about extinction selectivity regimes during these “background” intervals. Consequently, the question of whether selectivity regimes differ between background and mass extinctions is largely unresolved. Using logistic regression, we evaluated the selectivity of genus survivorship with respect to geographic range by using a global database of fossil benthic marine invertebrates spanning the Cambrian through the Neogene periods, an interval of ≈500 My. Our results show that wide geographic range has been significantly and positively associated with survivorship for the great majority of Phanerozoic time. Moreover, the significant association between geographic range and survivorship remains after controlling for differences in species richness and abundance among genera. However, mass extinctions and several second-order extinction events exhibit less geographic range selectivity than predicted by range alone. Widespread environmental disturbance can explain the reduced association between geographic range and extinction risk by simultaneously affecting genera with similar ecological and physiological characteristics on global scales. Although factors other than geographic range have certainly affected extinction risk during many intervals, geographic range is likely the most consistently significant predictor of extinction risk in the marine fossil record.

Two-phase increase in the maximum size of life over 3.5 billion years reflects biological innovation and environmental opportunity

The maximum size of organisms has increased enormously since the initial appearance of life >3.5 billion years ago (Gya), but the pattern and timing of this size increase is poorly known. Consequently, controls underlying the size spectrum of the global biota have been difficult to evaluate. Our period-level compilation of the largest known fossil organisms demonstrates that maximum size increased by 16 orders of magnitude since life first appeared in the fossil record. The great majority of the increase is accounted for by 2 discrete steps of approximately equal magnitude: the first in the middle of the Paleoproterozoic Era (≈1.9 Gya) and the second during the late Neoproterozoic and early Paleozoic eras (0.6–0.45 Gya). Each size step required a major innovation in organismal complexity—first the eukaryotic cell and later eukaryotic multicellularity. These size steps coincide with, or slightly postdate, increases in the concentration of atmospheric oxygen, suggesting latent evolutionary potential was realized soon after environmental limitations were removed.

Calcium isotope constraints on the end-Permian mass extinction.

The end-Permian mass extinction horizon is marked by an abrupt shift in style of carbonate sedimentation and a negative excursion in the carbon isotope (δ13C) composition of carbonate minerals. Several extinction scenarios consistent with these observations have been put forward. Secular variation in the calcium isotope (δ44/40Ca) composition of marine sediments provides a tool for distinguishing among these possibilities and thereby constraining the causes of mass extinction. Here we report δ44/40Ca across the Permian-Triassic boundary from marine limestone in south China. The δ44/40Ca exhibits a transient negative excursion of ∼0.3‰ over a few hundred thousand years or less, which we interpret to reflect a change in the global δ44/40Ca composition of seawater. CO2-driven ocean acidification best explains the coincidence of the δ44/40Ca excursion with negative excursions in the δ13C of carbonates and organic matter and the preferential extinction of heavily calcified marine animals. Calcium isotope constraints on carbon cycle calculations suggest that the average δ13C of CO2 released was heavier than -28‰ and more likely near -15‰; these values indicate a source containing substantial amounts of mantle- or carbonate-derived carbon. Collectively, the results point toward Siberian Trap volcanism as the trigger of mass extinction.

The Pattern and Timing of Biotic Recovery from the End-Permian Extinction on the Great Bank of Guizhou, Guizhou Province, China

Abstract Microfacies analysis and point counts of thin sections from 608 hand samples were used to track changes in the abundance and diversity of fossil grains through the extended recovery interval following end-Permian mass extinction on the Great Bank of Guizhou (GBG)—an isolated Late Permian to Late Triassic carbonate platform in south China. Exposure of a two-dimensional cross-section of the platform permits the comparison of faunal patterns along an environmental gradient from shallow to deep water. The diverse Late Permian biota was dominated by calcareous sponges, crinoids, articulate brachiopods, foraminifera, and calcareous algae. In contrast, Early Triassic communities were dominated by mollusks, with increasing abundance of crinoids beginning in the Spathian. Increase in the diversity and abundance of fossils on the GBG was confined to a brief interval near the Spathian–Anisian boundary and concentrated along the platform margin. Later Middle Triassic diversification, the return of calcareous algae and calcareous sponges, and the appearance of scleractinian corals did not substantially alter the mollusk-crinoid-Tubiphytes assemblage before the end of the Middle Triassic. The low abundance of skeletal grains in Lower Triassic strata implies: (1) similarities in the relative contributions of micrite, microbialites, and oolites to Neoproterozoic carbonates result, at least in part, from the temporary removal of skeletal sinks for calcium carbonate; and (2) animals with hard skeletons remained at low abundance from the time of the end-Permian extinction through much of the Early Triassic.

Evolutionary dynamics of gastropod size across the end-Permian extinction and through the Triassic recovery interval

Abstract A global database of gastropod sizes from the Permian through the Middle Triassic documents trends in gastropod shell size and permits tests of the suggestion that Early Triassic gastropods were everywhere unusually small. Analysis of the database shows that no specimens of unambiguous Early Triassic age larger than 2.6 cm have been reported, in contrast to common 5– 10-cm specimens of both Permian and Middle Triassic age. The loss of large gastropods is abrupt even at a fine scale of stratigraphic resolution, whereas the return of larger individuals in the Middle Triassic appears gradual when finely resolved. Taphonomic and sampling biases do not adequately explain the absence of large Early Triassic gastropods. Examination of size trends by genus demonstrates that the size decrease across the Permian/Triassic boundary is compatible with both size-selective extinction at the species level and anagenetic size change within lineages. Size increase in the Middle Triassic resulted from the origination of large species within genera that have Early Triassic fossil records and the occurrence of new genera containing large species during the Middle Triassic. Genera recorded from the Permian and Middle Triassic but not the Early Triassic (“Lazarus taxa”) do not contribute to observed size increase in the Middle Triassic. Moreover, Lazarus taxa lack large species and exhibit low species richness during both the Permian and the Middle Triassic, suggesting that they survived as small, rare forms rather than existing at large sizes in Early Triassic refugia. The ecological opportunities and selective pressures that produced large gastropods during most intervals of the Phanerozoic evidently did not operate in Early Triassic oceans. Whether this reflects low predation or competitive pressure, r-selection facilitated by high primary production, or physical barriers to large size remains poorly understood.

Acidification, anoxia, and extinction: A multiple logistic regression analysis of extinction selectivity during the Middle and Late Permian

Patterns of taxonomic and ecologic selectivity are the most direct record of processes infl uencing survival during background and mass extinctions. The Guadalupian (Capitanian) and end-Permian (Changhsingian) extinctions have both been linked to environmental degradation from eruption of large fl ood basalts; however, the extent to which taxonomic selectivity conforms to the expected stresses remains incompletely understood because many of the relevant biological traits are mutually correlated. Here we use a large occurrencebased database to quantify extinction selectivity during background and mass extinction intervals from the Kungurian (latest Early Permian) to Changhsingian. Our multiple logistic regression analysis confi rms that the end-Permian extinction was a physiological crisis, selecting against genera with poorly buffered respiratory physiology and calcareous shells. Genera with unbuffered physiology also fared poorly in the Guadalupian extinction, consistent with recognition of a pronounced crisis only among protists and reef-builders and implying similar respiratory physiological stresses. Despite sharing a similar trigger, the end-Permian extinction was considerably more severe than the Guadalupian or other Phanerozoic physiological crises. Its magnitude may have resulted from a larger environmental perturbation, although the combination of warming, hypercapnia, ocean acidifi cation, and hypoxia during the end-Permian extinction likely exacerbated the crisis because of the multiplicative effects of those stresses. Although ocean carbon cycle and evolutionary changes have reduced the sensitivity of modern ecosystems to physiological stresses, extant marine invertebrates face the same synergistic effects of multiple stressors that were so severe during the end-Permian extinction.

Long-term differences in extinction risk among the seven forms of rarity

Rarity is widely used to predict the vulnerability of species to extinction. Species can be rare in markedly different ways, but the relative impacts of these different forms of rarity on extinction risk are poorly known and cannot be determined through observations of species that are not yet extinct. The fossil record provides a valuable archive with which we can directly determine which aspects of rarity lead to the greatest risk. Previous palaeontological analyses confirm that rarity is associated with extinction risk, but the relative contributions of different types of rarity to extinction risk remain unknown because their impacts have never been examined simultaneously. Here, we analyse a global database of fossil marine animals spanning the past 500 million years, examining differential extinction with respect to multiple rarity types within each geological stage. We observe systematic differences in extinction risk over time among marine genera classified according to their rarity. Geographic range played a primary role in determining extinction, and habitat breadth a secondary role, whereas local abundance had little effect. These results suggest that current reductions in geographic range size will lead to pronounced increases in long-term extinction risk even if local populations are relatively large at present.