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Many modern extinction drivers are shared with past mass extinction events, such as rapid climate warming, habitat loss, pollution and invasive species. This commonality presents a key question: can the extinction risk of species during past mass extinction events inform our predictions for a modern biodiversity crisis? To investigate if it is possible to establish which species were more likely to go extinct during mass extinctions, we applied a functional trait-based model of extinction risk using a machine learning algorithm to datasets of marine fossils for the end-Permian, end-Triassic and end-Cretaceous mass extinctions. Extinction selectivity was inferred across each individual mass extinction event, before testing whether the selectivity patterns obtained could be used to 'predict' the extinction selectivity exhibited during the other mass extinctions. Our analyses show that, despite some similarities in extinction selectivity patterns between ancient crises, the selectivity of mass extinction events is inconsistent, which leads to a poor predictive performance. This lack of predictability is attributed to evolution in marine ecosystems, particularly during the Mesozoic Marine Revolution, associated with shifts in community structure alongside coincident Earth system changes. Our results suggest that past extinctions are unlikely to be informative for predicting extinction risk during a projected mass extinction.

Brachiopods dominated the seafloor as a primary member of the Paleozoic fauna. Despite the devastating effects of the end-Permian extinction, the group recovered during the early Mesozoic only to gradually decline from the Jurassic to today. This decline likely had multiple causes, including increased predation and bioturbation-driven substrate disruption, but the role of changing substrate is not well understood. Given the importance of substrate for extant brachiopod habitat, we documented Mesozoic-Cenozoic lithologic preferences and morphological changes to assess how decreasing firm-substrate habitat may have contributed to the brachiopod decline. Compared with bivalves, Mesozoic brachiopods occurred more frequently and were disproportionately abundant in carbonate lithologies. Although patterns in glauconitic or ferruginous sediments are equivocal, brachiopods became more abundant in coarser-grained carbonates and less abundant in fine-grained siliciclastics. During the Jurassic, brachiopod species rarely had abraded beaks but tended to be more convex with a high beak, potentially consistent with a non-analogue lifestyle resting on the seafloor. However, those highly convex morphotypes largely disappeared by the Cenozoic, when more terebratulides had abraded beaks, suggesting closer attachment to hard substrates. Rhynchonellides disproportionately declined to become a minor component of Cenozoic faunas, perhaps because of less pronounced morphological shifts. Trends in lithologic preferences and morphology are consistent with bioturbation-driven substrate disruption, with brachiopods initially using firmer carbonate sediments as refugia before adapting to live primarily attached to hard surfaces. This progressive habitat restriction likely played a role in the final brachiopod decline, as bioturbating ecosystem engineers transformed benthic habitats in the Mesozoic and Cenozoic.

The end-Permian mass extinction (EPME) led to a severe terrestrial ecosystem collapse. However, the ecological response of insects-the most diverse group of organisms on Earth-to the EPME remains poorly understood. Here, we analyse beetle evolutionary history based on taxonomic diversity, morphological disparity, phylogeny, and ecological shifts from the Early Permian to Middle Triassic, using a comprehensive new dataset. Permian beetles were dominated by xylophagous stem groups with high diversity and disparity, which probably played an underappreciated role in the Permian carbon cycle. Our suite of analyses shows that Permian xylophagous beetles suffered a severe extinction during the EPME largely due to the collapse of forest ecosystems, resulting in an Early Triassic gap of xylophagous beetles. New xylophagous beetles appeared widely in the early Middle Triassic, which is consistent with the restoration of forest ecosystems. Our results highlight the ecological significance of insects in deep-time terrestrial ecosystems.

During the Permian, major icehouse-greenhouse climate shifts and tectonic reconfiguration had important biogeographic implications, especially for climate-sensitive organisms such as fusulinids. Here we present multivariate methods on a global fusulinid species dataset including 1546 species from 58 localities in the Early (Asselian, Sakmarian, Artinskian and Kungurian) and Middle (Roadian, Wordian and Capitanian) Permian. Our results show that fusulinid global provincialism was high in the Asselian, Sakmarian, and Artinskian, driven by the development of multiple fusulinid bioregions in and near the Tethys Ocean. During the Asselian, Uralian sites and nearby regions of western Tethys were distinct from eastern Tethys, while stations in Arctic Russia and Norway formed a separate Boreal bioregion. Tectonic closure of the oceanic gateway in the southern Urals resulted in progressive isolation of the Uralian and Boreal bioregions during the Sakmarian and Artinskian and their ultimate disappearance by the Kungurian. Climate warming likely was the most important control on the Sakmarian formation of the distinct peri-Gondwana bioregion, because its development coincided with deglaciation following the Late Paleozoic Ice Age but preceded the separation of the Cimmerian terranes from northern margin of Gondwana. On the other hand, northward movement of the Cimmerian blocks following Artinskian-Kungurian rifting ultimately led to the merger of the peri-Gondwanan bioregion with tropical Tethyan faunas, resulting in lower provincialism in the Guadalupian and minimal faunal differentiation across Tethys. In contrast, faunal similarity between Tethys and eastern Panthalassa (the McCloud region and southwestern United States) was higher in the Asselian-Artinskian but decreased in the Kungurian and Middle Permian, perhaps as the result of sluggish ocean circulation following the warming episode of Late Paleozoic deglaciation.

Conservation of marine species requires the ability to predict the effects of climate-related stressors in an uncertain future. Experiments and observations in modern settings provide crucial information, but lack temporal scale and cannot anticipate emergent effects during ongoing global change. By contrast, the deep-time fossil record contains the long-term perspective at multiple global change events that can be used, at a broad scale, to test hypothesized effects of climate-related stressors. For example, geologically rapid carbon cycle disruption has often caused crises in reef ecosystems, and selective extinctions support the hypothesis that greater activity levels promote survival. Geographical patterns of extinction and extirpation were more variable than predicted from modern physiology, with tropical and temperate extinction peaks observed at different ancient events. Like any data source, the deep-time record has limitations but also provides opportunities that complement the limitations of modern and historical data. In particular, the deep-time record is the best source of information on actual outcomes of climate-related stressors in natural settings and over evolutionary timescales. Closer integration of modern and deep-time evidence can expand the types of hypotheses testable with the fossil record, yielding better predictions of extinction risk as climate-related stressors continue to intensify in future oceans. This article is part of a discussion meeting issue 'The past is a foreign country: how much can the fossil record actually inform conservation?'

The history of insects' taxonomic diversity is poorly understood. The two most common methods for estimating taxonomic diversity in deep time yield conflicting results: the 'range through' method suggests a steady, nearly monotonic increase in family-level diversity, whereas 'shareholder quorum subsampling' suggests a highly volatile taxonomic history with family-level mass extinctions occurring repeatedly, even at the midpoints of geological periods. The only feature shared by these two diversity curves is a steep increase in standing diversity during the Early Cretaceous. This apparent diversification event occurs primarily during the Aptian, the pre-Cenozoic interval with the most described insect occurrences, raising the possibility that this feature of the diversity curves reflects preservation and sampling biases rather than insect evolution and extinction. Here, the capture-mark-recapture (CMR) approach is used to estimate insects' family-level diversity. This method accounts for the incompleteness of the insect fossil record as well as uneven sampling among time intervals. The CMR diversity curve shows extinctions at the Permian/Triassic and Cretaceous/Palaeogene boundaries but does not contain any mass extinctions within geological periods. This curve also includes a steep increase in diversity during the Aptian, which appears not to be an artefact of sampling or preservation bias because this increase still appears when time bins are standardized by the number of occurrences they contain rather than by the amount of time that they span. The Early Cretaceous increase in family-level diversity predates the rise of angiosperms by many millions of years and can be better attributed to the diversification of parasitic and especially parasitoid insect lineages.

Amber is an organic multicompound derivative from the polymerization of resin of diverse higher plants. Compared with other modes of fossil preservation, amber records the anatomy of and ecological interactions between ancient soft-bodied organisms with exceptional fidelity. However, it is currently suggested that ambers do not accurately record the composition of arthropod forest paleocommunities, due to crucial taphonomic biases. We evaluated the effects of taphonomic processes on arthropod entrapment by resin from the plant Hymenaea, one of the most important resin-producing trees and a producer of tropical Cenozoic ambers and Anthropocene (or subfossil) resins. We statistically compared natural entrapment by Hymenaea verrucosa tree resin with the ensemble of arthropods trapped by standardized entomological traps around the same tree species. Our results demonstrate that assemblages in resin are more similar to those from sticky traps than from malaise traps, providing an accurate representation of the arthropod fauna living in or near the resiniferous tree, but not of entire arthropod forest communities. Particularly, arthropod groups such as Lepidoptera, Collembola, and some Diptera are underrepresented in resins. However, resin assemblages differed slightly from sticky traps, perhaps because chemical compounds in the resins attract or repel specific insect groups. Ground-dwelling or flying arthropods that use the tree-trunk habitat for feeding or reproduction are also well represented in the resin assemblages, implying that fossil inclusions in amber can reveal fundamental information about biology of the past. These biases have implications for the paleoecological interpretation of the fossil record, principally of Cenozoic amber with angiosperm origin.

Amber is an organic multicompound derivative from the polymerization of resin of diverse higher plants. Compared with other modes of fossil preservation, amber records the anatomy of and ecological interactions between ancient soft-bodied organisms with exceptional fidelity. However, it is currently suggested that ambers do not accurately record the composition of arthropod forest paleocommunities, due to crucial taphonomic biases. We evaluated the effects of taphonomic processes on arthropod entrapment by resin from the plant Hymenaea, one of the most important resin-producing trees and a producer of tropical Cenozoic ambers and Anthropocene (or subfossil) resins. We statistically compared natural entrapment by Hymenaea verrucosa tree resin with the ensemble of arthropods trapped by standardized entomological traps around the same tree species. Our results demonstrate that assemblages in resin are more similar to those from sticky traps than from malaise traps, providing an accurate representation of the arthropod fauna living in or near the resiniferous tree, but not of entire arthropod forest communities. Particularly, arthropod groups such as Lepidoptera, Collembola, and some Diptera are underrepresented in resins. However, resin assemblages differed slightly from sticky traps, perhaps because chemical compounds in the resins attract or repel specific insect groups. Ground-dwelling or flying arthropods that use the tree-trunk habitat for feeding or reproduction are also well represented in the resin assemblages, implying that fossil inclusions in amber can reveal fundamental information about biology of the past. These biases have implications for the paleoecological interpretation of the fossil record, principally of Cenozoic amber with angiosperm origin.

Sclerochronology uses shell growth lines or bands for the construction of environmental timeseries and the measurement of organism growth, but more study is needed to constrain the triggers of the dark cessation bands observed in many bivalve groups. We constructed a database of direct observations of modern growth seasonality across the class Bivalvia and compared the occurrence of seasonal growth bands to environmental data including latitude, temperature, and chlorophyll-a concentration. Bivalves with cold-season (winter) cessations are more common towards the poles, with logistic regression showing that temperature, followed by latitude of occurrence, displays the strongest relationship with occurrence of winter cessation. Remotely sensed and directly measured chlorophyll-a concentration show no significant relationship. Summer cessations are sparse and only weakly associated with environmental controls but are concentrated at the subtropical latitudes among temperate bivalves at their equatorial extremes. The rarity of summer cessations can be explained by the limited annual ranges of temperature in the tropics, combined with the exponential relationship of metabolic rate to temperature leading to a narrow window between normal functioning and mortality at high temperatures. This data suggests that, unless annual temperatures have low variability like in equatorial or polar regions, the season of growth cessation across bivalves is primarily a function of temperature tolerance through restriction of scope for growth. At most latitudes, growth bands can be interpreted as being primarily triggered by temperature stress, rather than seasonal starvation.

Explaining the ~5-million-year delay in marine biotic recovery following the latest Permian mass extinction, the largest biotic crisis of the Phanerozoic, is a fundamental challenge for both geological and biological sciences. Ocean redox perturbations may have played a critical role in this delayed recovery. However, the lack of quantitative constraints on the details of Early Triassic oceanic anoxia (for example, time, duration, and extent) leaves the links between oceanic conditions and the delayed biotic recovery ambiguous. We report high-resolution U-isotope (δ238U) data from carbonates of the uppermost Permian to lowermost Middle Triassic Zal section (Iran) to characterize the timing and global extent of ocean redox variation during the Early Triassic. Our δ238U record reveals multiple negative shifts during the Early Triassic. Isotope mass-balance modeling suggests that the global area of anoxic seafloor expanded substantially in the Early Triassic, peaking during the latest Permian to mid-Griesbachian, the late Griesbachian to mid-Dienerian, the Smithian-Spathian transition, and the Early/Middle Triassic transition. Comparisons of the U-, C-, and Sr-isotope records with a modeled seawater PO43- concentration curve for the Early Triassic suggest that elevated marine productivity and enhanced oceanic stratification were likely the immediate causes of expanded oceanic anoxia. The patterns of redox variation documented by the U-isotope record show a good first-order correspondence to peaks in ammonoid extinctions during the Early Triassic. Our results indicate that multiple oscillations in oceanic anoxia modulated the recovery of marine ecosystems following the latest Permian mass extinction.

Ancient mass extinction events such as the end-Permian and end- Triassic crises provide analogues for multistressor global change of ocean warming, pH reduction, and deoxygenation. Organism physiology is hypothesized to be a key trait influencing vulnerability to these stressors, but it is not certain how physiology predicts survival over evolutionary time scales and when organisms are faced with opposing or synergistic stressors. Fishes (bony fishes and chondrichthyan fishes) are active organisms with high aerobic scope for thermal tolerance and well-developed acid-base regulation, traits that should confer resilience to global change. To test this, we compiled a database of fossil marine fish occurrences to quantify extinction rates during background and mass extinctions from the Permian through Early Jurassic, using maximum likelihood estimation to compare extinction trajectories with marine invertebrates. Our results show that fewer chondrichthyan fishes underwent extinction than marine invertebrates during the end-Permian crisis. End-Triassic chondrichthyan extinction rates also were not elevated above background levels. In contrast, bony fishes underwent an end-Triassic extinction comparable to that of marine invertebrates. The differing responses of these two groups imply that a more active physiology can be advantageous during global change, although not uniformly. Permian-Triassic chondrichthyan fishes may have had broader environmental tolerances, facilitating survival. Alternatively, the larger offspring size of chondrichthyan fishes may provide greater energy reserves to offset the demands of warming and acidification. Although more active organisms have adult adaptations for thermal tolerance and pH regulation, some may nevertheless be susceptible to global change during early life stages.

Multistressor global change, the combined influence of ocean warming, acidification, and deoxygenation, poses a serious threat to marine organisms. Experimental studies imply that organisms with higher levels of activity should be more resilient, but testing this prediction and understanding organism vulnerability at a global scale, over evolutionary timescales, and in natural ecosystems remain challenging. The fossil record, which contains multiple extinctions triggered by multistressor global change, is ideally suited for testing hypotheses at broad geographic, taxonomic, and temporal scales. Here, I assess the importance of activity level for survival of well-skeletonized benthic marine invertebrates over a 100-million-year-long interval (Permian to Jurassic periods) containing four global change extinctions, including the end-Permian and end-Triassic mass extinctions. More active organisms, based on a semiquantitative score incorporating feeding and motility, were significantly more likely to survive during three of the four extinction events (Guadalupian, end-Permian, and end-Triassic). In contrast, activity was not an important control on survival during nonextinction intervals. Both the end-Permian and end-Triassic mass extinctions also triggered abrupt shifts to increased dominance by more active organisms. Although mean activity gradually returned toward pre-extinction values, the net result was a permanent ratcheting of ecosystem-wide activity to higher levels. Selectivity patterns during ancient global change extinctions confirm the hypothesis that higher activity, a proxy for respiratory physiology, is a fundamental control on survival, although the roles of specific physiological traits (such as extracellular pCO2 or aerobic scope) cannot be distinguished. Modern marine ecosystems are dominated by more active organisms, in part because of selectivity ratcheting during these ancient extinctions, so on average may be less vulnerable to global change stressors than ancient counterparts. However, ancient extinctions demonstrate that even active organisms can suffer major extinction when the intensity of environmental disruption is intense.

Insects are a hyper-diverse group, comprising nearly three-quarters of all named animal species on the Earth, but the environmental drivers of their richness and the roles of ecological interactions and evolutionary innovations remain unclear. Previous studies have argued that family-level insect richness increased continuously over the evolutionary history of the group, but inclusion of extant family records artificially inflated the relative richness of younger time intervals. Here we apply sampling-standardization methods to a species-level database of fossil insect occurrences, removing biases present in previous richness curves. We show that insect family-richness peaked 125 Ma and that Recent values are only 1.5-3 times as high as the Late Palaeozoic. Rarefied species-richness data also tentatively suggest little or no net increase in richness over the past 125 Myr. The Cretaceous peak in family richness was coincident with major radiations within extant groups but occurred prior to extinctions within more basal groups. Those extinctions may in part be linked to mid-Cretaceous floral turnover following the evolution of flowering plants. Negligible net richness change over the past 125 Myr implies that major radiations within extant groups were offset by reduced richness within groups that are now relict or extinct.

Macroevolutionary studies of insects at diverse taxonomic scales often reveal dynamic evolutionary patterns, with multiple inferred diversification rate shifts. Responses to major past environmental changes, such as the Cretaceous Terrestrial Revolution, or the development of major key innovations, such as wings or complete metamorphosis are usually invoked as potential evolutionary triggers. However this view is partially contradicted by studies on the family-level fossil record showing that insect diversification was relatively constant through time. In an attempt to reconcile both views, we investigate large-scale insect diversification dynamics at family level using two distinct types of diversification analyses on a molecular timetree representing ca. 82% of the extant families, and reassess the insect fossil diversity using up-to-date records. Analyses focusing on the fossil record recovered an early burst of diversification, declining to low and steady rates through time, interrupted by extinction events. Phylogenetic analyses showed that major shifts of diversification rates only occurred in the four richest holometabolous orders. Both suggest that neither the development of flight or complete metamorphosis nor the Cretaceous Terrestrial Revolution environmental changes induced immediate changes in diversification regimes; instead clade-specific innovations likely promoted the diversification of major insect orders.

The Guadalupian (middle Permian) extinction may have triggered substantial ecological restructuring in level-bottom communities, such as turnover in dominant brachiopod genera or a shift from abundant brachiopods to mollusks, despite comparatively minor taxonomic losses. However, ecological changes in relative abundance have been inferred from limited data; as a result, constraints on important shifts like the brachiopod-mollusk transition are imprecise. Here, I reevaluate the magnitude of ecological shifts during the Guadalupian-Lopingian (G-L) interval by supplementing previous census counts of silicified assemblages with counts from non-silicified assemblages and global occurrence data, both sourced from the Paleobiology Database. Brachiopod occurrences are consistent with more pronounced faunal composition changes from the Guadalupian to Lopingian than among stages within those intervals, but only in Iran and South China, and not in Pakistan or a Tethys-wide data set. In Iran and South China, Bray-Curtis dissimilarity values comparing occurrence frequencies between adjacent stages were elevated across the G-L transition, although other intervals exhibited similarly large shifts. However, genus occurrence frequencies were less strongly correlated or were anti-correlated across the G-L transition, suggesting moderate faunal turnover among dominant brachiopod genera. In contrast to previous inferences from silicified faunas, abundances of brachiopods, bivalves, and gastropods remained consistent from the Guadalupian to Lopingian in non-silicified local counts and global occurrences, implying that the brachiopod-mollusk shift did not occur until the end-Permian extinction. Ecological and taxonomic consequences were both minor in level-bottom settings, suggesting that severe environmental perturbations may not be necessary to explain biotic changes during the Guadalupian-Lopingian transition.

Insect taphonomy is a topic that has drawn interest because of its potential biases on diversity patterns and the ecological information recorded by ancient insect faunas. Other than the onset of common amber fossilization in the Cretaceous, very little is known about long-term trends in the nature and quality of insect preservation and, as a result, the effects of taphonomic biases are poorly constrained. We assembled a database of nearly 7000 Carboniferous-Pliocene insect adpression (compression and impression) species from the primary literature to assess changes in insect taphonomy over time and test biotic and environmental controls on preservation. We grouped the fossils into 10-Myr bins and scored preservation of holotype specimens as either articulated bodies or isolated wings; articulated specimens with a body implied a generally higher quality of preservation. Paleozoic and Triassic insect holotypes are known overwhelmingly from isolated wings (only 12% articulated bodies), but our database shows a significant increase in the percentage preserved as articulated bodies, to more than 70%, beginning about 160 Myr ago in the Late Jurassic. This transition could reflect variations in the robustness of different insect orders and shifts in the taxonomic composition of insect faunas, but all the major orders in the database exhibit significant increases in articulation. Instead, a shift to increased preservation in lacustrine paleoenvironments, which contain a greater proportion of articulated body fossils, explains most of the trend. The pronounced Late Jurassic increase in articulation has implications for evolutionary and ecological reconstructions, for example, suggesting that preserved insect diversity may be biased downward in the earlier part of their history when articulation was poor.

Video abstractAt Mistaken Point, Newfoundland, Canada, rangeomorph "fronds" dominate the earliest (579-565 million years ago) fossil communities of large (0.1 to 2 m height) multicellular benthic eukaryotes. They lived in low-flow environments, fueled by uptake [1-3] of dissolved reactants (osmotrophy). However, prokaryotes are effective osmotrophs, and the advantage of taller eukaryotic osmotrophs in this deep-water community context has not been addressed. We reconstructed flow-velocity profiles and vertical mixing using canopy flow models appropriate to the densities of the observed communities. Further modeling of processes at organismal surfaces documents increasing uptake with height in the community as a function of thinning of the diffusive boundary layer with increased velocity. The velocity profile, produced by canopy flow in the community, generates this advantage of upward growth. Alternative models of upward growth advantage based on redox/resource gradients fail, given the efficiency of vertical mixing. In benthic communities of osmotrophs of sufficient density, access to flow in low-flow settings provides an advantage to taller architecture, providing a selectional driver for communities of tall eukaryotes in contexts where phototropism cannot contribute to upward growth. These Ediacaran deep-sea fossils were preserved during the increasing oxygenation prior to the Cambrian radiation of animals and likely represent an important phase in the ecological and evolutionary transition to more complex eukaryotic forms.

Studies of modern ecological communities demonstrate that climate change may trigger changes in diversity and taxonomic composition; however, these studies are fundamentally limited to short timescales and therefore cannot demonstrate the full impact of major climate change. Understanding the ecological response of marine invertebrate communities to the Late Paleozoic Ice Age (LPIA), the last complete transition from icehouse to greenhouse, can establish a more complete picture of the climate-faunal relationship. We analyzed brachiopod community structure in Moscovian-Sakmarian (mid-Pennsylvanian to Early Permian) samples spanning the greatest extent of the LPIA, collected from four localities of the Copacabana Formation in Bolivia: Ancoraimes, Yaurichambi, Cuyavi, and Yampupata. Cluster analysis reveals three main groups that appear to coincide with pre-, syn-, and post-glacial times. Genus richness was significantly greater in samples during the Asselian glacial episode; however, the difference may be due to a combination of smaller body size and time averaged mixing of genera from different depths during more rapid glacioeustatic sea level change. Genera present in Bolivia consistently had warm-water affinities, even during the main glaciation, but warm-water taxa increased in abundance over time and the samples became increasingly dominated by characteristically North American genera. Overall mean body size and the size of particular genera were smaller in the Asselian cluster. These size changes likely reflect variations in substrate because marine invertebrates should be larger at cooler temperatures due to oxygen limitation at higher temperatures. The monotonic increase in abundance of warm-water genera and increasingly North American biogeographic affinity imply that community change was most likely the result of the northward drift of Bolivia rather than a response to late Paleozoic glacial-nonglacial cycles. This lack of climate related faunal change was probably a result of Bolivia's mid-latitude location during the late Paleozoic because both the rate of temperature change and its magnitude were likely smaller at lower latitudes, reducing the impact of climate change on marine communities. © 2013 Elsevier B.V.