Did impacts, volcanic eruptions, or climate change affect mammalian evolution?

In recent years, it has become popular to attribute faunal change and mass extinction to impacts, volcanic eruptions, or climatic change. How well do these supposed causes compare to the excellent record of Cenozoic life, especially that of fossil mammals? Two different Cenozoic mammal diversity curves were compared, and important climatic, volcanic, and impact horizons were examined in detail. In no case is there a strong correlation between impacts, eruptions, or climatic events and any episode of mammalian turnover. On the contrary, most of the known impact, eruption, and climatic events of the Cenozoic occurred during intervals of faunal stability. Conversely, episodes of high turnover and faunal change among Cenozoic mammals correlate with no known extrinsic causes. Apparently, extrinsic environmental factors such as impacts, eruptions, and climate change have a minimal effect, and intrinsic biological factors must be more important.     

Impact of the Late Triassic mass extinction on functional diversity and composition of marine ecosystems

Mass extinctions have profoundly influenced the history of life, not only through the death of species but also through changes in ecosystem function and structure. Importantly, these events allow us the opportunity to study ecological dynamics under levels of environmental stress for which there are no recent analogues. Here, we examine the impact and selectivity of the Late Triassic mass extinction event on the functional diversity and functional composition of the global marine ecosystem, and test whether post‐extinction communities in the Early Jurassic represent a regime shift away from pre‐extinction communities in the Late Triassic. Our analyses show that, despite severe taxonomic losses, there is no unequivocal loss of global functional diversity associated with the extinction. Even though no functional groups were lost, the extinction event was, however, highly selective against some modes of life, in particular sessile suspension feeders. Although taxa with heavily calcified skeletons suffered higher extinction than other taxa, lightly calcified taxa also appear to have been selected against. The extinction appears to have invigorated the already ongoing faunal turnover associated with the Mesozoic Marine Revolution. The ecological effects of the Late Triassic mass extinction were preferentially felt in the tropical latitudes, especially amongst reefs, and it took until the Middle Jurassic for reef ecosystems to fully recover to pre‐extinction levels.     

Biotic and environmental dynamics through the Late Jurassic–Early Cretaceous transition: evidence for protracted faunal and ecological turnover

The Late Jurassic to Early Cretaceous interval represents a time of environmental upheaval and cataclysmic events, combined with disruptions to terrestrial and marine ecosystems. Historically, the Jurassic/Cretaceous (J/K) boundary was classified as one of eight mass extinctions. However, more recent research has largely overturned this view, revealing a much more complex pattern of biotic and abiotic dynamics than has previously been appreciated. Here, we present a synthesis of our current knowledge of Late Jurassic–Early Cretaceous events, focusing particularly on events closest to the J/K boundary. We find evidence for a combination of short‐term catastrophic events, large‐scale tectonic processes and environmental perturbations, and major clade interactions that led to a seemingly dramatic faunal and ecological turnover in both the marine and terrestrial realms. This is coupled with a great reduction in global biodiversity which might in part be explained by poor sampling. Very few groups appear to have been entirely resilient to this J/K boundary ‘event’, which hints at a ‘cascade model’ of ecosystem changes driving faunal dynamics. Within terrestrial ecosystems, larger, more‐specialised organisms, such as saurischian dinosaurs, appear to have suffered the most. Medium‐sized tetanuran theropods declined, and were replaced by larger‐bodied groups, and basal eusauropods were replaced by neosauropod faunas. The ascent of paravian theropods is emphasised by escalated competition with contemporary pterosaur groups, culminating in the explosive radiation of birds, although the timing of this is obfuscated by biases in sampling. Smaller, more ecologically diverse terrestrial non‐archosaurs, such as lissamphibians and mammaliaforms, were comparatively resilient to extinctions, instead documenting the origination of many extant groups around the J/K boundary. In the marine realm, extinctions were focused on low‐latitude, shallow marine shelf‐dwelling faunas, corresponding to a significant eustatic sea‐level fall in the latest Jurassic. More mobile and ecologically plastic marine groups, such as ichthyosaurs, survived the boundary relatively unscathed. High rates of extinction and turnover in other macropredaceous marine groups, including plesiosaurs, are accompanied by the origin of most major lineages of extant sharks. Groups which occupied both marine and terrestrial ecosystems, including crocodylomorphs, document a selective extinction in shallow marine forms, whereas turtles appear to have diversified. These patterns suggest that different extinction selectivity and ecological processes were operating between marine and terrestrial ecosystems, which were ultimately important in determining the fates of many key groups, as well as the origins of many major extant lineages. We identify a series of potential abiotic candidates for driving these patterns, including multiple bolide impacts, several episodes of flood basalt eruptions, dramatic climate change, and major disruptions to oceanic systems. The J/K transition therefore, although not a mass extinction, represents an important transitional period in the co‐evolutionary history of life on Earth.     

A model of onshore-offshore change in faunal diversity

Onshore-offshore patterns of faunal change occurred at many taxonomic scales during the Paleozoic Era, ranging from replacement of the Cambrian evolutionary fauna by the Paleozoic fauna to the environmental expansion of many orders and classes. A simple mathematical model is constructed to investigate such change. The environmental gradient across the marine shelf-slope is treated as a linear array of discrete habitats, each of which holds a set number of species, as observed in the fossil record. During any interval of time, some portion of the species in each habitat becomes extinct by background processes, with rates of extinction varying among both clades and habitats, as also observed in the record. After extinction, species are replaced from within the habitat and from immediately adjacent habitats, with proportions dependent on surviving species. This model leads to the prediction that extinction-resistant clades will always diversify at the expense of extinction-prone clades. But if extinction intensity is highest in nearshore habitats, extinction-resistant clades will expand preferentially in the onshore direction, build up diversity there, and then diversify outward toward the offshore. Thus, onshore-offshore patterns of diversification may be the expectation for faunal change quite independently of whether or not clades originate onshore. When the model is parameterized for Paleozoic trilobites and brachiopods, numerical solutions exhibit both a pattern of faunal change and a time span for diversification similar to that seen in the fossil record. They also generate structure similar to that seen in global diversification, including logistic patterns of growth, declining origination but constant extinction within clades through time, and declining overall extinction across clades through time.     

New insights into the reading of Paleozoic plant fossil record discontinuities

Studying the discontinuity patterns of Paleozoic vascular plants provides a global vision of these key events from the multivariate methods viewpoint. Non-metric multidimensional scaling, detrended correspondence analysis and cluster analysis have been employed together with a set of diversity and abundance measures and an evaluation of the geologic constraints from the plant fossil record data. The results reveal four clear significant discontinuities in terms of taxonomic composition and record representativeness during the early-middle Devonian, Devonian–Carboniferous, Mississippian–Pennsylvanian and early-late Permian. Due to the controversial character of the plant fossil record data and the effect of mass extinction events, the results can be explained in taxonomic turnover and ecological reorganisation terms which emphasise the crucial role of the geologic constrains in paleobiological inference.     

Island life in the Cretaceous - faunal composition,biogeography, evolution,and extinction of land-living vertebrates on the Late Cretaceous European archipelago

The Late Cretaceous was a time of tremendous global change, as the final stages of the Age of Dinosaurs were shaped by climate and sea level fluctuations and witness to marked paleogeographic and faunal changes, before the end-Cretaceous bolide impact. The terrestrial fossil record of Late Cretaceous Europe is becoming increasingly better understood, based largely on intensive fieldwork over the past two decades, promising new insights into latest Cretaceous faunal evolution. We review the terrestrial Late Cretaceous record from Europe and discuss its importance for understanding the paleogeography, ecology, evolution, and extinction of land-dwelling vertebrates. We review the major Late Cretaceous faunas from Austria, Hungary, France, Spain, Portugal, and Romania, as well as more fragmentary records from elsewhere in Europe. We discuss the paleogeographic background and history of assembly of these faunas, and argue that they are comprised of an endemic ‘core’ supplemented with various immigration waves. These faunas lived on an island archipelago, and we describe how this insular setting led to ecological peculiarities such as low diversity, a preponderance of primitive taxa, and marked changes in morphology (particularly body size dwarfing). We conclude by discussing the importance of the European record in understanding the end-Cretaceous extinction and show that there is no clear evidence that dinosaurs or other groups were undergoing long-term declines in Europe prior to the bolide impact.     

Plant fossil record and survival analyses

Cascales‐Miñana, B. & Cleal, C.J. 2011: Plant fossil record and survival analyses. Lethaia, Vol. 45, pp. 71–82. Survival analysis is a classic palaeobiological method widely used on the animal fossil record. This study reports the first application of survivorship analyses on the plant fossil record from a global viewpoint and provides a new comparative approach of this methodology. The results reveal three important plant extinction events in the history of plant life at a global scale. The results also clearly suggest that the origination events are more intensive than extinction processes and that the origination moment of several lineages of vascular plants is an important factor that conditions their longevity. This study supports the general idea that vascular plants tended to be less affected by the environmental changes that caused mass extinction in other groups of organisms. □Extinction events, fossil record, survival patterns, taxonomic survivorship curves, vascular plants.     

Marine extinction risk shaped by trait–environment interactions over 500 million years

Perhaps the most pressing issue in predicting biotic responses to present and future global change is understanding how environmental factors shape the relationship between ecological traits and extinction risk. The fossil record provides millions of years of insight into how extinction selectivity (i.e., differential extinction risk) is shaped by interactions between ecological traits and environmental conditions. Numerous paleontological studies have examined trait‐based extinction selectivity; however, the extent to which these patterns are shaped by environmental conditions is poorly understood due to a lack of quantitative synthesis across studies. We conducted a meta‐analysis of published studies on fossil marine bivalves and gastropods that span 458 million years to uncover how global environmental and geochemical changes covary with trait‐based extinction selectivity. We focused on geographic range size and life habit (i.e., infaunal vs. epifaunal), two of the most important and commonly examined predictors of extinction selectivity. We used geochemical proxies related to global climate, as well as indicators of ocean acidification, to infer average global environmental conditions. Life‐habit selectivity is weakly dependent on environmental conditions, with infaunal species relatively buffered from extinction during warmer climate states. In contrast, the odds of taxa with broad geographic ranges surviving an extinction (>2500 km for genera, >500 km for species) are on average three times greater than narrow‐ranging taxa (estimate of odds ratio: 2.8, 95% confidence interval = 2.3–3.5), regardless of the prevailing global environmental conditions. The environmental independence of geographic range size extinction selectivity emphasizes the critical role of geographic range size in setting conservation priorities.     

Insect-damaged fossil leaves record food web response to ancient climate change and extinction

Plants and herbivorous insects have dominated terrestrial ecosystems for over 300 million years. Uniquely in the fossil record, foliage with well-preserved insect damage offers abundant and diverse information both about producers and about ecological and sometimes taxonomic groups of consumers. These data are ideally suited to investigate food web response to environmental perturbations, and they represent an invaluable deep-time complement to neoecological studies of global change. Correlations between feeding diversity and temperature, between herbivory and leaf traits that are modulated by climate, and between insect diversity and plant diversity can all be investigated in deep time. To illustrate, I emphasize recent work on the time interval from the latest Cretaceous through the middle Eocene (67-47 million years ago (Ma)), including two significant events that affected life: the end-Cretaceous mass extinction (65.5 Ma) and its ensuing recovery; and globally warming temperatures across the Paleocene-Eocene boundary (55.8 Ma). Climatic effects predicted from neoecology generally hold true in these deep-time settings. Rising temperature is associated with increased herbivory in multiple studies, a result with major predictive importance for current global warming. Diverse floras are usually associated with diverse insect damage; however, recovery from the end-Cretaceous extinction reveals uncorrelated plant and insect diversity as food webs rebuilt chaotically from a drastically simplified state. Calibration studies from living forests are needed to improve interpretation of the fossil data.     

Biological hierarchies and the nature of extinction

Hierarchy theory recognises that ecological and evolutionary units occur in a nested and interconnected hierarchical system, with cascading effects occurring between hierarchical levels. Different biological disciplines have routinely come into conflict over the primacy of different forcing mechanisms behind evolutionary and ecological change. These disconnects arise partly from differences in perspective (with some researchers favouring ecological forcing mechanisms while others favour developmental/historical mechanisms), as well as differences in the temporal framework in which workers operate. In particular, long‐term palaeontological data often show that large‐scale (macro) patterns of evolution are predominantly dictated by shifts in the abiotic environment, while short‐term (micro) modern biological studies stress the importance of biotic interactions. We propose that thinking about ecological and evolutionary interactions in a hierarchical framework is a fruitful way to resolve these conflicts. Hierarchy theory suggests that changes occurring at lower hierarchical levels can have unexpected, complex effects at higher scales due to emergent interactions between simple systems. In this way, patterns occurring on short‐ and long‐term time scales are equally valid, as changes that are driven from lower levels will manifest in different forms at higher levels. We propose that the dual hierarchy framework fits well with our current understanding of evolutionary and ecological theory. Furthermore, we describe how this framework can be used to understand major extinction events better. Multi‐generational attritional loss of reproductive fitness (MALF) has recently been proposed as the primary mechanism behind extinction events, whereby extinction is explainable solely through processes that result in extirpation of populations through a shutdown of reproduction. While not necessarily explicit, the push to explain extinction through solely population‐level dynamics could be used to suggest that environmentally mediated patterns of extinction or slowed speciation across geological time are largely artefacts of poor preservation or a coarse temporal scale. We demonstrate how MALF fits into a hierarchical framework, showing that MALF can be a primary forcing mechanism at lower scales that still results in differential survivorship patterns at the species and clade level which vary depending upon the initial environmental forcing mechanism. Thus, even if MALF is the primary mechanism of extinction across all mass extinction events, the primary environmental cause of these events will still affect the system and result in differential responses. Therefore, patterns at both temporal scales are relevant.     

Trends and patterns in the evolution of vascular plants: macroevolutionary implications of a multilevel taxonomic analysis

Cascales‐Miñana, B., Muñoz‐Bertomeu, J., Ros, R., Segura, J. 2010: Trends and patterns in the evolution of vascular plants: macroevolutionary implications of a multilevel taxonomic analysis. Lethaia, 10.1111/j.1502‐3931.2009.00212.x Studying the macroevolutionary patterns of vascular plants from the Silurian to the present‐day provides a global record of plant life history. Evolutionary rates (origination, extinction and diversification) for families, orders, classes and divisions were analysed, as was abundance and richness for 21 time intervals. An accumulative analysis, based on the total plant fossil record, the accumulated extinctions and relative diversity, was also carried out. The diversification rate shows a uniquely constant and progressive reduction from the end of the Carboniferous to the Permian when the lowest values are registered. Very small peaks seem to reflect Cretaceous extinction for families. At family level, only two time intervals present higher extinctions, than originations. Richness and accumulative analyses reveal that only 32% of the families analysed became extinct, and that approximately 90% of them disappeared at the end of the Palaeozoic. Our results indicate that plants did not undergo mass extinction events in the ‘big five’ sense, but rather, mass ecological reorganization the absence of important extinction events or evolutionary innovations producing diversification patterns without abrupt changes. □Diversification, evolutionary, extinction, fossil record, innovations, radiation, vascular plants.     

Competition in slow motion: the unusual case of benthic marine communities in the wake of the end‐Permian mass extinction

Changes of community structure in response to competition usually take place on timescales that are much too short to be visible in the geological record. Here we report the notable exception of a benthic marine community in the wake of the end‐Permian mass extinction, which is associated with the microbial limestone facies of the earliest Triassic of South China. The newly reported fauna is well preserved and extraordinarily rich (30 benthic macroinvertebrate species, including the new species Astartella? stefaniae (Bivalvia) and Eucochlis obliquecostata (Gastropoda)) and stems from an environmentally stable setting providing favourable conditions for benthic organisms. Whereas changes in the taxonomic composition are negligible over the observed time interval of 10–100 ka, three ecological stages are identified, in which relative abundances of initially rare species continuously increased at the cost of previously dominant species. Concomitant with the changes of dominant species is an increase in faunal evenness and heterogeneity. In the absence of both environmental and taxonomic changes, we attribute this pattern to the long‐term effects of interspecific competition, which acted at an unusually slow pace because the number of competing species and potential immigrants was dramatically reduced by the end‐Permian mass extinction. We suggest that these non‐actualistic conditions led to decreased rates of niche differentiation and hence to the delayed rediversification of benthos that characterizes the aftermath of the greatest Phanerozoic mass extinction event. A hyperbolic diversification model is proposed, which accounts for the positive relationship between the intensity of interspecific competition and the rate of niche differentiation and resolves the conundrum of delayed rediversification at a time when niche space was largely vacated.     

Tracking Taphonomic Regimes Using Chemical and Mechanical Damage of Pollen and Spores: An Example from the Triassic–Jurassic Mass Extinction

The interpretation of biotic changes in the geological past relies on the assumption that samples from different time intervals represent an equivalent suite of natural sampling conditions. As a result, detailed investigations of taphonomic regimes during intervals of major biotic upheaval, such as mass extinctions, are crucial. In this paper, we have used variations in the frequency of chemical and mechanical sporomorph (pollen and spore) damage as a guide to taphonomic regimes across the Triassic–Jurassic mass extinction (Tr-J; ∼201.3 Ma) at a boundary section at Astartekløft, East Greenland. We find that the frequency of sporomorph damage is extremely variable in samples from this locality. This likely reflects a combination of taxon-specific susceptibility to damage and the mixing of sporomorphs from a mosaic of environments and taphonomic regimes. The stratigraphic interval containing evidence of plant extinction and compositional change in the source vegetation at Astartekløft is not marked by a consistent rise or fall in the frequency of sporomorph damage. This indicates that natural taphonomic regimes did not shift radically during this critical interval. We find no evidence of a consistent relationship between the taxonomic richness of sporomorph assemblages and the frequency of damage among sporomorphs at Astartekløft. This indicates that previously reported patterns of sporomorph richness across the Tr-J at this locality are likely to be robust. Taken together, our results suggest that the patterns of vegetation change at Astartekløft represent a real biological response to environmental change at the Tr-J.     

When the pond turtle followed the reindeer: effect of the last extreme global warming event on the timing of faunal change in Northern Europe

Faunal communities have been shaped in different ways by past climatic change. The impact of the termination of the last Glacial and the onset of the present (Holocene) Interglacial on large‐scale faunal shifts, extinction dynamics and gene pools of species are of special interest in natural sciences. A general pattern of climate‐triggered range expansion and local extinction of vertebrate species is known for Europe, and shows that in the modern temperate zone the main faunal change took place mainly during the Late Glacial (14 700–11 700 years ago) and Early Holocene (11 700–9 100 years ago). Based on large datasets of new radiocarbon data, we present precise temporal dynamics of climate‐driven disappearance and appearance of reindeer and pond turtle in southern Sweden. These two species are significant climate indicators in Late Quaternary biostratigraphy. Our data reveal that the reindeer disappeared from southern Sweden ca. 10 300 years ago, whereas the pond turtle colonized the area ca. 9 860 years ago, with a 450‐year gap between each species. This provides evidence for a sudden environmental turnover, causing the replacement of an arctic faunal element by a thermophilic species. The postglacial range dynamics of pond turtle and reindeer are a unique model case, allowing insights into the faunal turnover of other vertebrates during the last dramatic natural global warming event at the Pleistocene–Holocene transition.     

Individual to community-level faunal responses to environmental change from a marine fossil record of Early Miocene global warming

Modern climate change has a strong potential to shift earth systems and biological communities into novel states that have no present-day analog, leaving ecologists with no observational basis to predict the likely biotic effects. Fossil records contain long time-series of past environmental changes outside the range of modern observation, which are vital for predicting future ecological responses, and are capable of (a) providing detailed information on rates of ecological change, (b) illuminating the environmental drivers of those changes, and (c) recording the effects of environmental change on individual physiological rates. Outcrops of Early Miocene Newport Member of the Astoria Formation (Oregon) provide one such time series. This record of benthic foraminiferal and molluscan community change from continental shelf depths spans a past interval environmental change (≈ 20.3-16.7 mya) during which the region warmed 2.1-4.5°C, surface productivity and benthic organic carbon flux increased, and benthic oxygenation decreased, perhaps driven by intensified upwelling as on the modern Oregon coast. The Newport Member record shows that (a) ecological responses to natural environmental change can be abrupt, (b) productivity can be the primary driver of faunal change during global warming, (c) molluscs had a threshold response to productivity change while foraminifera changed gradually, and (d) changes in bivalve body size and growth rates parallel changes in taxonomic composition at the community level, indicating that, either directly or indirectly through some other biological parameter, the physiological tolerances of species do influence community change. Ecological studies in modern and fossil records that consider multiple ecological levels, environmental parameters, and taxonomic groups can provide critical information for predicting future ecological change and evaluating species vulnerability.     

Delayed recovery of non-marine tetrapods after the end-Permian mass extinction tracks global carbon cycle

During the end-Permian mass extinction, marine ecosystems suffered a major drop in diversity, which was maintained throughout the Early Triassic until delayed recovery during the Middle Triassic. This depressed diversity in the Early Triassic correlates with multiple major perturbations to the global carbon cycle, interpreted as either intrinsic ecosystem or external palaeoenvironmental effects. In contrast, the terrestrial record of extinction and recovery is less clear; the effects and magnitude of the end-Permian extinction on non-marine vertebrates are particularly controversial. We use specimen-level data from southern Africa and Russia to investigate the palaeodiversity dynamics of non-marine tetrapods across the Permo-Triassic boundary by analysing sample-standardized generic richness, evenness and relative abundance. In addition, we investigate the potential effects of sampling, geological and taxonomic biases on these data. Our analyses demonstrate that non-marine tetrapods were severely affected by the end-Permian mass extinction, and that these assemblages did not begin to recover until the Middle Triassic. These data are congruent with those from land plants and marine invertebrates. Furthermore, they are consistent with the idea that unstable low-diversity post-extinction ecosystems were subject to boom-bust cycles, reflected in multiple Early Triassic perturbations of the carbon cycle.     

How catastrophic was the end‐Triassic mass extinction?

A review of marine and terrestrial animal and plant fossils fails to reveal convincing evidence of a global catastrophe at the Triassic-Jurassic boundary, although this time marked the final disappearance of ceratite ammonites and conodonts, together with the extinction of most calcareous demosponges; important groups of bivalves and brachiopods went extinct. Because of facies problems, however, there is no stratigraphic section that reveals a clear-cut disappearance over a short distance. Other marine animal groups except perhaps the radiolarians fail to reveal a notable extinction of global extent immediately across the boundary. On the other hand, there was a substantially higher extinction rate among marine animals in the Rhaetian as compared with the previous stage. On the land, the record is equivocal. Dramatic changes across the T-J boundary have been claimed for plants in particular areas, such as eastern North America and East Greenland, but only gradual change has been recognized elsewhere. Similarly, claims of a T-J boundary vertebrate mass extinction have not been supported by others. For the Rhaetian as a whole, however, the turnover rate of reptiles was high. Although much remains to be learned, it seems evident that the fossil record of the latest Triassic is more consistent with a gradual scenario extended over time than a 'geologically instantaneous' impact catastrophe.     

Natural extinction on islands

Almost all recent extinction of species or subspecies on islands comes from human activities. On the other hand, in local populations there is much natural extinction and immigration, i.e. turnover, on small islands. Most of this turnover occurs in locally rare species, and attests to the phenomenon of minimum viable population size. The MacArthur-Wilson theory is based on this turnover which, from an ecological point of view, is generally trivial. More useful theories of minimum viable population size are being developed. Rarity is the precursor of extinction, and species can be rare in several ways. Models of these phenomena are still primitive, particularly those that relate habitat availability to population density. Models of interactive communities show phenomena that may be relevant to the understanding of extinction in the geological record. Lotka-Volterra equations indicate considerable sensitivity to invasions, sometimes producing a cascade of extinction. Chemostat equations show that the behaviour of food chains can change dramatically with small changes in parameters, suggesting that small environmental effects can sometimes cause large ecological changes, including extinctions, in interactive biotic communities.     

Historical species losses in bumblebee evolution

Investigating how species coped with past environmental changes informs how modern species might face human-induced global changes, notably via the study of historical extinction, a dominant feature that has shaped current biodiversity patterns. The genus Bombus, which comprises 250 mostly cold-adapted species, is an iconic insect group sensitive to current global changes. Through a combination of habitat loss, pathogens and climate change, bumblebees have experienced major population declines, and several species are threatened with extinction. Using a time-calibrated tree of Bombus, we analyse their diversification dynamics and test hypotheses about the role of extinction during major environmental changes in their evolutionary history. These analyses support a history of fluctuating species dynamics with two periods of historical species loss in bumblebees. Dating estimates gauge that one of these events started after the middle Miocene climatic optimum and one during the early Pliocene. Both periods are coincident with global climate change that may have extirpated Bombus species. Interestingly, bumblebees experienced high diversification rates during the Plio-Pleistocene glaciations. We also found evidence for a major species loss in the past one million years that may be continuing today.     

Microbial response to limited nutrients in shallow water immediately after the end-Permian mass extinction

Previous work indicates that a variety of microbes bloomed in the oceans after the end-Permian faunal mass extinction, but evidence is sporadically documented. Thus, the nature and geographic distribution of such microbes and their associations are unclear, addressed in this study using a series of biomarker groups. On the basis of microbial biomarker records of the 2-methylhopane index, evidence is presented for cyanobacterial blooms in both the western and eastern Tethys Sea and in both shallow and deep waters, after the mass extinction. The enhanced relative abundance of C(28) (expressed by the C(28) /C(29) ratio of) regular steranes suggests a bloom of prasinophyte algae occurred immediately after the end-Permian faunal extinction, comparable with those observed in some other mass extinctions in Phanerozoic. Significantly, cyanobacteria and prasinophyte algae show a synchronized onset of bloom in the shallow water Bulla section, north Italy, inferring for the first time their coupled response to the biotic crisis and the associated environmental conditions. However, in Meishan of Zhejiang Province in south China, the bloom declined earlier than in Bulla. The association of increased 2-methylhopane index with a negative shift in the nitrogen isotope composition infers a scenario of enhanced nitrogen fixation by cyanobacteria immediately after the faunal mass extinction. N(2) fixation by cyanobacteria is here interpreted to have provided prasinophyte algae with ammonium in nutrient-limited shallow waters, and thus caused their associated blooms.