Early Toarcian Extinction Research Articles

Extinction cascades, community collapse, and recovery across a Mesozoic hyperthermal event

Mass extinctions are considered to be quintessential examples of Court Jester drivers of macroevolution, whereby abiotic pressures drive a suite of extinctions leading to huge ecosystem changes across geological timescales. Most research on mass extinctions ignores species interactions and community structure, limiting inference about which and why species go extinct, and how Red Queen processes that link speciation to extinction rates affect the subsequent recovery of biodiversity, structure and function. Here, we apply network reconstruction, secondary extinction modelling and community structure analysis to the Early Toarcian (Lower Jurassic; 183 Ma) Extinction Event and recovery. We find that primary extinctions targeted towards infaunal guilds, which caused secondary extinction cascades to higher trophic levels, reproduce the empirical post-extinction community most accurately. We find that the extinction event caused a switch from a diverse community with high levels of functional redundancy to a less diverse, more densely connected community of generalists. Recovery was characterised by a return to pre-extinction levels of some elements of community structure and function prior to the recovery of biodiversity. Full ecosystem recovery took ~7 million years at which point we see evidence of dramatically increased vertical structure linked to the Mesozoic Marine Revolution and modern marine ecosystem structure.

Ocean acidification during the early Toarcian extinction event: Evidence from boron isotopes in brachiopods

Abstract The loss of carbonate production during the Toarcian Oceanic Anoxic Event (T-OAE, ca. 183 Ma) is hypothesized to have been at least partly triggered by ocean acidification linked to magmatism from the Karoo-Ferrar large igneous province (southern Africa and Antarctica). However, the dynamics of acidification have never been directly quantified across the T-OAE. Here, we present the first record of temporal evolution of seawater pH spanning the late Pliensbachian and early Toarcian from the Lusitanian Basin (Portugal) reconstructed on the basis of boron isotopic composition (δ11B) of brachiopod shells. δ11B declines by ∼1‰ across the Pliensbachian-Toarcian boundary (Pl-To) and attains the lowest values (∼12.5‰) just prior to and within the T-OAE, followed by fluctuations and a moderately increasing trend afterwards. The decline in δ11B coincides with decreasing bulk CaCO3 content, in parallel with the two-phase decline in carbonate production observed at global scales and with changes in pCO2 derived from stomatal indices. Seawater pH had declined significantly already prior to the T-OAE, probably due to the repeated emissions of volcanogenic CO2. During the earliest phase of the T-OAE, pH increased for a short period, likely due to intensified continental weathering and organic carbon burial, resulting in atmospheric CO2 drawdown. Subsequently, pH dropped again, reaching the minimum in the middle of the T-OAE. The early Toarcian marine extinction and carbonate collapse were thus driven, in part, by ocean acidification, similar to other Phanerozoic events caused by major CO2 emissions and warming.

Upper Toarcian (Lower Jurassic) marine gastropods from the Cleveland Basin, England: systematics, palaeobiogeography and contribution to biotic recovery from the early Toarcian extinction event

AbstractHere we describe a new upper Toarcian (Lower Jurassic) marine gastropod fauna from rocks of the Cleveland Basin exposed on the North Yorkshire coast of England. The fossil assemblage consists of 16 species, of which three are new:Katosira?bicarinatasp. nov.,Turritelloidea stephenisp. nov. andStriactaenonina eleganssp. nov. Four species are described in open nomenclature asTricarilda? sp.,Jurildasp.,Cylindrobullinasp. andCossmanninasp. The other species have previously been described:Coelodiscus minutus(SchüblerinZieten),Procerithium quadrilineatum(Römer),Pseudokatosira undulata(Benzinvon Zieten),Palaeorissoinaaff.acuminata(Gründel),Pietteia unicarinata(Hudleston),Globulariacf.canina(Hudleston),Striactaeoninacf.richterorumSchulbert & Nützel,Striactaenoninaaff.tenuistriata(Hudleston) andSulcoactaeon sedgvici(Phillips). Most of these species are the earliest records of their respective genera and show palaeobiogeographical connections with contemporary gastropod associations from other regions of Europe and South America. The taxonomic composition of the upper Toarcian Cleveland Basin gastropod assemblage differs substantially from the faunas of the upper Pliensbachian and lower ToarcianTenuicostatumZone, showing the strong effect of the early Toarcian mass extinction event on the marine gastropod communities in the basin. Only a few gastropod species are shared between the late Toarcian faunas and the much more diverse Aalenian gastropod faunas in the Cleveland Basin, suggesting that there was a facies control on gastropod occurrences at that time. This is also a potential explanation for the taxonomic differences between the late Toarcian gastropod faunas in the Cleveland Basin and those in France, and northern and southern Germany.

Reductions in body size of benthic macroinvertebrates as a precursor of the early Toarcian (Early Jurassic) extinction event in the Lusitanian Basin, Portugal

AbstractReduction of body size is a common response of organisms to environmental stress. Studying the early Toarcian succession in the Lusitanian Basin of Portugal, we tested whether the shell size of benthic marine communities of bivalves and brachiopods changed at and before the global, warming–related Toarcian oceanic anoxic event (T-OAE). Statistical analyses of shell size over time show that the mean shell size of communities decreased significantly before the T-OAE. This trend is distinct in brachiopods and is caused by larger-sized species becoming less abundant over time, whereas it is not significant in bivalves, suggesting a decoupled response to environmental stress. Reductions in shell size precede the decline in standardized sample-level species richness associated with the early Toarcian extinction event. Such decreases in the shell size of marine invertebrates, well before the onset of biodiversity change, suggest that reductions in body size more generally may be a precursor of a subsequent loss of species and turnover at the community level caused by climate change. Sedimentological evidence is against hypoxia as a driver of extinction and the preceding size decrease in the brachiopod fauna in the studied succession, although low oxygen levels are widely held responsible for elevated early Toarcian extinction rates globally. Reduction of mean shell size in brachiopods but stasis in bivalves is difficult to explain with ocean acidification, because experimental work shows that brachiopods can be resilient to lowered pH, albeit long-term metabolic costs and potential evolutionary adaptations are unknown. Rising early Toarcian temperatures in the Lusitanian Basin seem to be a plausible factor in both diversity decline associated with the T-OAE and the preceding reductions in mean shell size, because thermal tolerances in modern bivalves are among the highest within marine invertebrates.

Stratigraphic and environmental control on marine benthic community change through the early Toarcian extinction event (Iberian Range, Spain)

In the Early Jurassic (~183 Ma ago) global warming and associated environmental changes coincided with an extinction event in the marine realm (early Toarcian extinction event). Anoxia was previously considered to have been the main cause of extinction, but extinctions also occur at localities that remained oxygenated throughout the event, suggesting that other factors, such as temperature, may have played a major role. To test this hypothesis, we integrated quantitative analyses of benthic macro-invertebrates with high-resolution geochemical proxies on the bulk rock (TOC, δ13C, δ18O) and on belemnites and brachiopod shells (δ13C, δ18O) from two sections from the Iberian Range, Spain, with no black shale deposition. The sections are orientated SE-NW along an onshore-offshore gradient deepening to the north. The dominant benthic groups, bivalves and brachiopods, show a different response to the extinction: brachiopods go through a complete species-level turnover, while many bivalve species range through the event. In the shallower section, changes in richness and evenness correlate with TOC (Total Organic Carbon), suggesting that variations in nutrient input from runoff, and the possible local onset of low-redox conditions (TOC > 4 wt%), controlled faunal diversity. In contrast, at the deeper section, community change correlates with changes in δ18O, indicating that temperature variations might have influenced faunal change. Different stratigraphic patterns of extinction occur between the two localities, with last-occurrences clustering at the maximum flooding surface in the shallower section, and at the transgressive surface in the deeper one. The observed differences between the two localities highlight the important role of local sedimentary and stratigraphic processes in controlling the shape of the geochemical and fossil record, and the need for studying multiple sections along onshore-offshore gradients in order to extrapolate regional and global patterns.

Modelling determinants of extinction across two Mesozoic hyperthermal events.

The Late Triassic and Early Toarcian extinction events are both associated with greenhouse warming events triggered by massive volcanism. These Mesozoic hyperthermals were responsible for the mass extinction of marine organisms and resulted in significant ecological upheaval. It has, however, been suggested that these events merely involved intensification of background extinction rates rather than significant shifts in the macroevolutionary regime and extinction selectivity. Here, we apply a multivariate modelling approach to a vast global database of marine organisms to test whether extinction selectivity varied through the Late Triassic and Early Jurassic. We show that these hyperthermals do represent shifts in the macroevolutionary regime and record different extinction selectivity compared to background intervals of the Late Triassic and Early Jurassic. The Late Triassic mass extinction represents a more profound change in selectivity than the Early Toarcian extinction but both events show a common pattern of selecting against pelagic predators and benthic photosymbiotic and suspension-feeding organisms, suggesting that these groups of organisms may be particularly vulnerable during episodes of global warming. In particular, the Late Triassic extinction represents a macroevolutionary regime change that is characterized by (i) the change in extinction selectivity between Triassic background intervals and the extinction event itself; and (ii) the differences in extinction selectivity between the Late Triassic and Early Jurassic as a whole.

Brachiopod faunal exchange through an epioceanic-epicontinental transitional area from the Early Jurassic South Iberian platform system

The La Mola region (eastern External Betic Zone) can be regarded as one of the easternmost complete Jurassic successions of the Betic Cordillera in the Iberian Peninsula, but the paleogeographical setting of their outcrops remains widely discussed. Analysis of brachiopod assemblages from the Lower Jurassic improves the accuracy of previous paleogeographical data, enabling identification of a mainly epioceanic transitional area in which influences of epicontinental habitats are also detected. Assemblage 1, mainly with a Mediterranean affinity but also sharing several constituents with the African and Northeastern Iberian basins, typifies the Sinemurian–Pliensbachian transition. Assemblage 2, as a whole, shows a transitional character between epioceanic and epicontinental habitats; it is subdivided into two successive and interrelated sub-assemblages: Ass. 2a (Demonense–Tenuicostatum Zones) reveals a free connection with the epioceanic Subbetic area, whereas Ass. 2b (Uppermost Pliensbachian–Lower Toarcian) shows a closer relationship with epicontinental environments. Assemblage 3 (Uppermost Spinatum–basal Serpentinum Zones) is commonly recorded in the peri-Iberian epicontinental platform system integrated within the NW-European bioprovince, but it can also be regarded as a marginal assemblage that is widespread in the westernmost Tethyan margin prior to the Early Toarcian extinction event. Assemblage 2 constitutes a suitable index for assessing the paleobiogeographical affinity of the La Mola region, as Ass. 2a is progressively replaced by Ass. 2b, thus triggering the arrival of epicontinental taxa to the more intra-epioceanic Subbetic environments, inferring a possible connection through the La Mola transitional slope. Consequently, this region enabled a faunal mixing and exchange between both environments, and La Mola likely remained as an area that would facilitate migration and an effective dispersal seaway or, at least, did not constitute an ecological filter-barrier for brachiopods. Biostratigraphical data from brachiopods and ammonites are correlated for the first time in La Mola, refining and calibrating biochronostratigraphical gaps in the pre-Domerian deposits where biochronological markers are usually scarce, and around the Pliensbachian–Toarcian boundary, a crucial timespan in which the Early Toarcian extinction event took place.

Evolution of the last koninckinids (Athyridida, Koninckinidae), a precursor signal of the early Toarcian mass extinction event in the Western Tethys

Koninckinids are a suitable group to shed light on the biotic crisis suffered by brachiopod fauna in the Early Jurassic. Koninckinid fauna recorded in the late Pliensbachian–early Toarcian from the easternmost Subbetic basin is analyzed and identified as a precursor signal for one of the most conspicuous mass extinction events of the Phylum Brachiopoda, a multi-phased interval with episodes of changing environmental conditions, whose onset can be detected from the Elisa–Mirabile subzones up to the early Toarcian extinction boundary in the lowermost Serpentinum Zone (T-OAE). The koninckinid fauna had a previously well-established migration pattern from the intra-Tethyan to the NW-European basins but a first phase with a progressive warming episode in the Pliensbachian–Toarcian transition triggered a koninckinid fauna exodus from the eastern/central Tethys toward the westernmost Mediterranean margins. A second stage shows an adaptive response to more adverse conditions in the westernmost Tethyan margins and finally, an escape and extinction phase is detected in the Atlantic areas from the mid-Polymorphum Zone onwards up to their global extinction in the lowermost Serpentinum Zone. This migration pattern is independent of the paleogeographic bioprovinciality and is unrelated to a facies-controlled pattern. The anoxic/suboxic environmental conditions should only be considered as a minor factor of partial control since well-oxygenated habitats are noted in the intra-Tethyan basins and this factor is noticeable only in the second westward migratory stage (with dwarf taxa and oligotypical assemblages). The analysis of cold-seep proxies in the Subbetic deposits suggests a radiation that is independent of methane releases in the Subbetic basin.

The early Toarcian (early Jurassic) ostracod extinction events in the Iberian Range: The effect of temperature changes and prolonged exposure to low dissolved oxygen concentrations

The early Toarcian extinction event, once regarded as spanning in the late Pliensbachian–early Toarcian boundary period, actually occurred in two distinct phases in the early Toarcian of the Iberian Range. The first episode occurred in the Mirabile Subzone, Tenuicostatum Zone and is characterised by the disappearance of about 12% of ostracod species and one suborder, the Metacopina (Superfamily Healdioidea). This study shows that this suborder disappears during stratigraphical intervals that are considered to be representative of low oxygen conditions in the Iberian Range. The collapse of marine ecosystems (affecting up to 40% of ostracod species) in the Elegantulum Subzone, Serpentinum Zone was most likely triggered by a sudden and widespread lethal increase in seawater temperature. These combined processes of oxygenation and temperature change are considered to be responsible for the profound changes in marine environments that resulted in the early Toarcian ostracod extinction in the Iberian Range.

The two Early Toarcian (Early Jurassic) extinction events in ammonoids

The Early Toarcian (Early Jurassic) biological crisis was one of the ‘minor’ mass extinctions. It is linked with an oceanic anoxic event. Fossil data from sections located in northwestern European (epicontinental platforms and basins) and Tethyan (distal, epioceanic) areas indicate that Late Pliensbachian–Early Toarcian ammonoids experienced two extinction events during the Early Toarcian. The older one is linked with disruption of the Tethyan–Boreal provinciality, whereas the younger event correlates with the onset of anoxia and corresponds with the Early Toarcian mass-extinction event. These two extinctions cannot be interpreted as episodes of a single, stepwise, event. Values of the net diversification, more than the number of extinctions, allow the two extinction events to be clearly recognized and distinguished. Values of regional net diversification for northwestern European and Tethyan faunas point to greater evolutionary dynamics in the epioceanic areas. The inclusion of Mediterranean faunas in the database proves that the ammonite turnover at the Early Toarcian mass-extinction event was more important than previously thought. Progenitor (evolute Neolioceratoides), survivor (Dactylioceras, Polyplectus pluricostatus) and Lazarus (Procliviceras) taxa have been recognized. Different selectivity patterns are shown for the two events. The first one, linked to the disruption of the Tethyan–Boreal provinciality, has mainly affected ammonites adapted to epicontinental platforms. In the mass-extinction event, no selectivity is recognized, because also Phylloceratina and Lytoceratina were deeply affected at species level, although their wide biogeographical distribution at clade level was a significant buffer against extinction. In contrast to Palaeozoic mass extinctions, ammonoid survivors and Lazarus taxa are characterized by complex sutures: Phylloceratina (long-ranging ammonoids) and Polyplectus (relatively long-ranging compared to other Ammonitina).

Early and mid-jurassic molluscan biogeography and the establishment of the central Atlantic seaway

A new analysis of Lower and Middle Jurassic bivalve faunas of western North America and the southern Andes in comparison to Europe, supported by evidence from other molluscs, throws further light on the establishment of a “central Atlantic” epicontinental seaway prior to the formation of a true oceanic strait between eastern North America and Africa-South America. Some restriction to free communication persisted until the early Callovian but there was an episode of relatively free intermigration in the Toarcian to early Bajocian time interval, which is associated with major rise of sea level. One of the most significant consequences of this event was a spread into northwest Europe of taxa which were previously confined to the Pacific margins, to occupy niches vacated as a result of an early Toarcian extinction event.