Interpreting the Carbon Isotope Record of Mass Extinctions

Mass extinctions are global-scale environmental crises marked by the loss of numerous species from all habitats. They often coincide with rapid changes in the stable carbon isotope ratios (13C/12C) recorded in sedimentary carbonate and organic matter, ratios which can indicate substantial inputs to the surface carbon reservoirs and/or changes in the cycling of carbon. Models to explain these changes have provided much fuel for debate on the causes and consequences of mass extinctions. For example, the escape of methane from gas hydrate deposits or the emission of huge volumes of gaseous carbon from large-scale volcanic systems, known as large igneous provinces, may have been responsible for decreases of 13C/12C in sedimentary deposits. In this article, we discuss the challenges in distinguishing between these, and other, alternatives

Toarcian oceanic anoxic event: An assessment of global causes using belemnite C isotope records

Two hypotheses have been proposed to explain simultaneous large negative excursions (up to 7% PeeDee belemnite) in bulk carbonate (delta(13)C(carb)) and organic carbon isotope records (delta(13)C(org)) from black shales marking the Toarcian oceanic anoxic event (T-OAE). The first explanation envisions recycling of dissolved inorganic carbon (DIC) with a light isotopic signature into the photic zone from the lower levels of a salinity-stratified water mass, essentially requiring a regional paleoceanographic driver of the carbon cycle. The second involves the rapid and massive dissociation of methane from gas hydrates that effectively renders the T-OAE a global perturbation of the carbon cycle. We present C isotope records from belemnites (delta(13)C(bel)) sampled from two localities, calibrated with high-resolution ammonite biostratigraphy and Sr isotope stratigraphy, in Yorkshire (England) and Dotternhausen (Germany), that can be used to assess which model best explains the observed changes in carbon isotopes. Our records of the delta(13)C composition of belemnite calcite do not show the large negative C isotope excursions shown by coeval records of delta(13)C in sedimentary organic matter or bulk sedimentary carbonate. It follows that isotopically light carbon cannot have dominated the ocean-atmosphere carbon reservoir during the Toarcian OAE, as would be required were the methane release hypothesis correct. On the basis of an evaluation of available carbon isotope records we discuss a model in which the recycling of DIC from the deeper levels of a stratified water body, and shallowing of anoxic conditions into the photic zone, can explain all isotopic profiles. In particular, the model accounts for the higher C isotope values of belemnites that are characteristic of open ocean, well-mixed conditions, and the lower C isotope values of neritic phytoplankton communities that recorded the degree of density stratification and shallowing of anoxia in the photic zone

Pellicle ultrastructure demonstrates that Moyeria is a fossil euglenid

An earlier proposal of euglenid affinity for the acritarch Moyeria was based primarily on the pattern of bi-helical striate ornamentation as seen in scanning electron microscopy and light microscopy. Examination of specimens using transmission electron microscopy reveals that the ‘striae’ are actually integral components of the microfossil wall itself, corresponding to the pellicle strips of some euglenid species today. A Silurian specimen from Scotland preserves an articulated wall composed of thickened arches and thinner U-shaped interconnecting segments paralleling that seen in some modern photosynthetic euglenids. A second specimen from the Moyeria holotype section (Silurian of New York State) shows fused articulation, again compatible with some extant euglenids. This evidence is sufficient to transfer Moyeria out of the Incertae sedis group, Acritarcha, and into the Euglenida. This proposal helps establish the morphological basis for the recognition of euglenid microfossils and ultimately provides evidence of a lengthy fossil record of the eukaryotic supergroup Excavata

Middle Phanerozoic mass extinctions and a tribute to the work of Professor Tony Hallam

Tony Hallam's contributions to mass extinction studies span more than 50 years and this thematic issue provides an opportunity to pay tribute to the many pioneering contributions he has made to this field. Early work (1961) on the Jurassic in Europe revealed a link, during the Toarcian Stage, between extinction and the spread of anoxic waters during transgression – the first time such a common leitmotif had been identified. He also identified substantial sea-level changes during other mass extinction intervals with either regression (end-Triassic) or early transgression (end-Permian) coinciding with the extinction phases. Hallam's (1981) study on bivalves was also the first to elevate the status of the end-Triassic crisis and place it amongst true mass extinctions, changing previous perceptions that it was a part of a protracted period of turnover, although debates on the duration of this crisis continue (Hallam, 2002). Conflicting views on the nature of recovery from mass extinctions have also developed, especially for the aftermath of the end-Permian mass extinction. These discussions can be traced to Hallam's seminal 1991 paper that noted the considerable delay in benthic recovery during Early Triassic time and attributed it to the persistence of the harmful, high-stress conditions responsible for the extinction itself. This idea now forms the cornerstone of one of the more favoured explanations for this ultra-low diversity interval

Basinal restriction, black shales, Re-Os dating, and the Early Toarcian (Jurassic) oceanic anoxic event

Profiles of Mo/total organic carbon (TOC) through the Lower Toarcian black shales of the Cleveland Basin, Yorkshire, United Kingdom, and the Posidonia shale of Germany and Switzerland reveal water mass restriction during the interval from late tenuicostatum Zone times to early bifrons Zone times, times which include that of the putative Early Toarcian oceanic anoxic event. The degree of restriction is revealed by crossplots of Mo and TOC concentrations for the Cleveland Basin, which define two linear arrays with regression slopes (ppm/%) of 0.5 and 17. The slope of 0.5 applies to sediment from the upper semicelatum and exaratum Subzones. This value, which is one tenth of that for modern sediments from the Black Sea (Mo/TOC regression slope 4.5), reveals that water mass restriction during this interval was around 10 times more severe than in the modern Black Sea; the renewal frequency of the water mass was between 4 and 40 ka. The Mo/TOC regression slope of 17 applies to the overlying falciferum and commune subzones: the value shows that restriction in this interval was less severe and that the renewal frequency of the water mass was between 10 and 130 years. The more restricted of the two intervals has been termed the Early Toarcian oceanic anoxic event but is shown to be an event caused by basin restriction local to NW Europe. Crossplots of Re, Os, and Mo against TOC show similar trends of increasing element concentration with increase in TOC but with differing slopes. Together with modeling of Os-187/Os-188 and delta Mo-98, the element/TOC trends show that drawdown of Re, Os, and Mo was essentially complete during upper semicelatum and exaratum Subzone times (Mo/TOC regression slope of 0.5). Drawdown sensitized the restricted water mass to isotopic change forced by freshwater mixing so that continental inputs of Re, Os, and Mo, via a low-salinity surface layer, created isotopic excursions of up to 1.3 parts per thousand in delta Mo-98 and up to 0.6% for Os-187/Os-188. Restriction thereby compromises attempts to date Toarcian black shales, and possibly all black shales, using Re-Os chronology and introduces a confounding influence in the attempts to use delta Mo-98 and initial Os-187/Os-188 for palaeo-oceanographic interpretation

End-Triassic calcification crisis and blooms of organic-walled 'disaster species'

The Triassic–Jurassic (T–J) mass-extinction event is marked by isotope anomalies in organic (δ13Corg) and carbonate carbon (δ13Ccarb) reservoirs. These have been attributed to a (rapid) 4-fold rise in pCO2 as a result of massive flood basalt volcanism and/or methane hydrate dissociation. Here we examine the response of marine photosynthetic phytoplankton to the proposed perturbation in the carbon cycle. Our high-resolution micropalaeontological analysis of T–J boundary beds at St Audrie's Bay in Somerset, UK, provides evidence for a bio-calcification crisis that is characterized by (1) extinction and malformation in calcareous nannoplankton and (2) contemporaneous blooms of organic-walled, green algal 'disaster' species which comprise in one case > 70% of the total palynomorph fraction. Blooms of prasinophytes and acritarchs occur at the onset and in association with a prominent negative shift in δ13Corg values close to the first appearance of the Early Jurassic ammonite Psiloceras planorbis. Across the same interval we obtained palaeotemperature and palaeosalinity estimates from oyster low-Mg calcite based on Mg/Ca, Sr/Ca and δ18O records. The results of our palynological and geochemical analyses strongly suggest that shallow marine basins in NW Europe during this period became salinity stratified, inducing anoxic conditions. The T–J boundary event shows similarities with the Permian–Triassic (P–T) mass-extinction event, which was also marked by extensive flood basalt volcanism, negative excursions in carbon isotope records, a bio-calcification crisis, the development of shallow-marine anoxia and mass abundances of acritarchs in the Early Triassic. This leads us to suggest that the proliferation of green algal phytoplankton may be symptomatic of elevated carbon dioxide levels in the atmosphere and oceans during mass-extinction events

Carbonate-platform response to the Toarcian Oceanic Anoxic Event in the southern hemisphere : Implications for climatic change and biotic platform demise

We are grateful to Zhifei Liu for TOC and analyses at the Tongji University. We thank also Wei An, Bo Zhou and Shiyi Li for their assistance in the field, and Zhicheng Huang, Yiwei Xu and Weiwei Xue for their help in the laboratory, and Chao Chang, Tianchen He and Bolin Zhang for their helpful discussion. Hugh Jenkyns commented on a draft of the manuscript. We would also like to thank Editor Derek Vance, Christopher Pearce and two anonymous reviewers whose comments greatly improved the manuscript. This study was financially supported by the National Natural Science Funds for Distinguished Young Scholar in China (41525007) and the Chinese MOST 973 Project (2012CB822001). DBK acknowledges support of NERC Fellowship NE/I02089X/1. This is a contribution to the IGCP 655.Peer reviewedPostprin

Body size trends and recovery amongst bivalves following the end-Triassic mass extinction

Fossils in the immediate aftermath of mass extinctions are often of small size, a phenomenon attributed to the Lilliput Effect (temporary, size reduction of surviving species). There has been little attempt to study size trends during subsequent recovery intervals nor has the relationship between size, diversity and environmental controls been evaluated. Here we examine the recovery following the end-Triassic mass extinction amongst bivalves of the British Lower and Middle Lias. Three distinct phases of size change are seen that are independent of other recovery metrics: initially bivalves are small but the Lilliput Effect is a minor factor, the majority of small taxa belong to new species that undergo a later within-species size increase (the Brobdingnag Effect) throughout the subsequent Hettangian Stage. New species that appeared during the Hettangian were also progressively larger and Cope's Rule (size increase between successive species) is seen – notably amongst ammonites. The size increase was reversed during the Sinemurian Stage, when bivalves once again exhibited small body sizes. During the Pliensbachian Stage another phase of size increase occurred with further evidence of the Brobdingnag Effect. These three phases of size change are seen across all suspension feeding ecological guilds of bivalve but are not expressed among deposit feeders. Local environmental conditions explain some aspects of size patterns, but factors such as temperature, marine oxygenation and sea level, do not correlate with the long-term size trends. The Brobdingnag Effect may reflect increased availability/quality of food during the recovery interval: a factor that controlled bivalve size but not evolution