MASS EXTINCTIONS AND CLADE EXTINCTIONS IN THE HISTORY OF BRACHIOPODS: BRIEF REVIEW AND A POST-PALEOZOIC CASE STUDY

Brachiopods are a key group in Phanerozoic marine diversity analyses for their excellent fossil record and distinctive evolutionary history. A genus-level survey of raw diversity trajectories allows the identification of the Brachiopod Big Five, episodes of major genus losses in the phylum which are compared with the established Big Five mass extinctions of Phanerozoic marine invertebrates. The two lists differ in that the end-Cretaceous extinction appears subdued for brachiopods, whereas the mid-Carboniferous is recognized as an event with significant loss of brachiopod genera. At a higher taxonomic level, a review of temporal ranges of rhynchonelliform orders reveals episodes of synchronous termination of multiple orders, here termed clade extinctions. The end-Ordovician, Late Devonian and end-Permian events are registered as both mass extinctions and clade extinctions. The Late Cambrian and the Early Jurassic are identified as the other two clade extinction events. Coincident with the Early Toarcian oceanic anoxic event, the last clade extinction of brachiopods is defined by the disappearance of the last two spire-bearing orders, Athyridida and Spiriferinida. Their diversity trajectory through the recovery after the end-Permian crisis parallels that of the extant terebratulides and rhynchonellides until a Late Triassic peak but diverge afterwards. The end-Triassic diversity decline and Toarcian vanishing of spire-bearers correspond with contraction in their spatial distribution. The observed patterns and extinction selectivity may be explained both ecologically and physiologically. The specialized adaptation of morphologically diverse spire-bearers, as well as their fixed lophophore and passive feeding put them at a disadvantage at times of environmental crises, manifest in their end-Triassic near-extinction and Toarcian demise. Similar analyses of other clade extinctions may further improve our understanding of drivers and processes of extinction

Not all biodiversity rich spots are climate refugia

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BioDeepTime : a database of biodiversity time series for modern and fossil assemblages

We thank the Paleosynthesis Project and the Volkswagen Stiftung for funding that supported this project (Az 96 796). M.C.R. acknowledges the German Research Foundation (DFG) for funding through the Cluster of Excellence ‘The Ocean Floor – Earth's Uncharted Interface’ (EXC 2077, grant no. 390741603). E.E.S. acknowledges funding from Leverhulme Trust grant RPG-201170, the Leverhulme Prize and the National Science Research Council grant NE/V011405/1. Q.J.L. and L.N. acknowledge support from the Youth Innovation Promotion Association (2019310) and the Chinese Academy of Sciences (CAS-WX2021SF-0205). A.M.P. acknowledges funding from the Leverhulme Trust through research grant RPG-2019-402. M.D. acknowledges funding from Leverhulme Trust through the Leverhulme Centre for Anthropocene Biodiversity (RC-2018-021) and a research grant (RPG-2019-402), and the European Union (ERC coralINT, 101044975). L. H. L. acknowledges funding from the European Research Council (macroevolution.abc ERC grant no. 724324). K.H.P acknowledges funding from the National Science Foundation Graduate Research Fellowship Program (DGE-2139841). H.H.M.H. acknowledges support from Peter Buck Postdoc Fellowship, Smithsonian Institution. A.T. acknowledges funding from the Slovak Research and Development Agency (APVV 22-0523) and the Slovak Scientific Grant Agency (VEGA 02/0106/23).Motivation We have little understanding of how communities respond to varying magnitudes and rates of environmental perturbations across temporal scales. BioDeepTime harmonizes assemblage time series of presence and abundance data to help facilitate investigations of community dynamics across timescales and the response of communities to natural and anthropogenic stressors. BioDeepTime includes time series of terrestrial and aquatic assemblages of varying spatial and temporal grain and extent from the present-day to millions of years ago. Main Types of Variables Included BioDeepTime currently contains 7,437,847 taxon records from 10,062 assemblage time series, each with a minimum of 10 time steps. Age constraints, sampling method, environment and taxonomic scope are provided for each time series. Spatial Location and Grain The database includes 8752 unique sampling locations from freshwater, marine and terrestrial ecosystems. Spatial grain represented by individual samples varies from quadrats on the order of several cm2 to grid cells of ~100 km2. Time Period and Grain BioDeepTime in aggregate currently spans the last 451?million years, with the 10,062 modern and fossil assemblage time series ranging in extent from years to millions of years. The median extent of modern time series is 18.7?years and for fossil series is 54,872?years. Temporal grain, the time encompassed by individual samples, ranges from days to tens of thousands of years. Major Taxa and Level of Measurement The database contains information on 28,777 unique taxa with 4,769,789 records at the species level and another 271,218 records known to the genus level, including time series of benthic and planktonic foraminifera, coccolithophores, diatoms, ostracods, plants (pollen), radiolarians and other invertebrates and vertebrates. There are to date 7012 modern and 3050 fossil time series in BioDeepTime. Software Format SQLite, Comma-separated values.Publisher PDFPeer reviewe

Data from: Biodiversity dynamics and environmental occupancy of fossil azooxanthellate and zooxanthellate scleractinian corals

Scleractinian corals have two fundamentally different life strategies, which can be inferred from morphological criteria in fossil material. In the non-photosymbiotic group nutrition comes exclusively from heterotrophic feeding, whereas the photosymbiotic group achieves a good part of its nutrition from algae hosted in the coral’s tissue. These ecologic differences arose early in the evolutionary history of corals but with repeated evolutionary losses and presumably also gains of symbiosis since then. We assessed the biodiversity dynamics and environmental occupancy of both ecologic groups to identify times when the evolutionary losses of symbiosis as inferred from molecular analyses might have occurred and if these can be linked to environmental change. Two episodes are likely: The first was in the mid-Cretaceous when non-symbiotic corals experienced an origination pulse and started to become more common in deeper, non-reef habitats and on siliciclastic substrates initiating a long-term offshore trend in occupancy. The second was around the Cretaceous/Paleogene boundary with another origination pulse and increased occupancy of deep-water settings in the non-symbiotic group. Environmental factors such as rapid global warming associated with mid-Cretaceous anoxic events and increased nutrient concentrations in Late Cretaceous–Cenozoic deeper waters are plausible mechanisms for the shift. Turnover rates and durations are not significantly different between the two ecologic groups when compared over the entire history of scleractinians. However, the deep-water shift of non-symbiotic corals was accompanied by reduced extinction rates, supporting the view that environmental occupancy is a prominent driver of evolutionary rates

Data from: Adding fossil occupancy trajectories to the assessment of modern extinction risk

Besides helping to identify species traits that are commonly linked to extinction risk, the fossil record may also be directly relevant for assessing the extinction risk of extant species. Standing geographical distribution or occupancy is a strong predictor of both recent and past extinction risk, but the role of changes in occupancy is less widely assessed. Here we demonstrate, based on the Cenozoic fossil record of marine species, that both occupancy and its temporal trajectory are significant determinants of risk. Based on extinct species we develop a model on the additive and interacting effects of occupancy and its temporal changes on extinction risk. We use this model to predict extinction risk of extant species. The predictions suggest a moderate risk for marine species on average. However, some species seem to be on a long-term decline and potentially at a latent extinction risk, which is not considered in current risk assessments

Supplementary Methods and Text from Adding fossil occupancy trajectories to the assessment of modern extinction risk

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Data from: Radiolarian biodiversity dynamics through the Triassic and Jurassic: implications for proximate causes of the end-Triassic mass extinction

Within a ∼60-Myr interval in the Late Triassic to Early Jurassic, a major mass extinction took place at the end of Triassic, and several biotic and environmental events of lesser magnitude have been recognized. Climate warming, ocean acidification, and a biocalcification crisis figure prominently in scenarios for the end-Triassic event and have been also suggested for the early Toarcian. Radiolarians, as the most abundant silica-secreting marine microfossils of the time, provide a control group against marine calcareous taxa in testing selectivity and responses to changing environmental parameters. We analyzed the origination and extinction rates of radiolarians, using data from the Paleobiology Database and employing sampling standardization, the recently developed gap-filler equations and an improved stratigraphic resolution at the substage level. The major end-Triassic event is well-supported by a late Rhaetian peak in extinction rates. Because calcifying and siliceous organisms appear similarly affected, we consider global warming a more likely proximate trigger of the extinctions than ocean acidification. The previously reported smaller events of radiolarian turnover fail to register above background levels in our analyses. The apparent early Norian extinction peak is not significant compared to the long-term trajectory, and is probably a sampling artifact. The Toarcian Oceanic Anoxic Event, previously also thought to have caused a significant radiolarian turnover, did not significantly affect the group. Radiolarian diversity history appears unique and complexly forced, as its trajectory parallels major calcareous fossil groups at some events and deviates at others

Data from: Climate change and the latitudinal selectivity of ancient marine extinctions

Geologically rapid climate change is anticipated to increase extinction risk non-uniformly across the Earth’s surface. Tropical species may be more vulnerable than temperate species to current climate warming because of high tropical climate velocities and reduced seawater oxygen levels. To test if rapid warming indeed preferentially increased the extinction risk of tropical fossil taxa, we combine a robust statistical assessment of latitudinal extinction selectivity (LES) with the dominant views on climate change occurring at ancient extinction crises. Using a global dataset of marine fossil occurrences, we assess extinction rates for tropical and temperate genera, applying log-ratios to assess effect size and Akaike weights for model support. Among the classical ‘Big Five’ mass extinction episodes, the end-Permian mass extinction exhibits temperate preference of extinctions, whereas the Late Devonian and end-Triassic selectively hit tropical genera. Simple links between the inferred direction of climate change and LES are idiosyncratic, both during crisis and background intervals. More complex models, including sampling patterns and changes in the latitudinal distribution of continental shelf area, show tropical LES to be generally associated with raised tropical heat and temperate LES with global cold temperatures. With implications for the future, our paper demonstrates the consistency of high tropical temperatures, habitat loss and the capacity of both to interact in generating geographic patterns in extinctions

The biogeographical imprint of mass extinctions

Mass extinctions are defined by extinction rates significantly above background levels and have had substantial consequences for the evolution of life. Geographically selective extinctions, subsequent originations and species redistributions may have changed global biogeographical structure, but quantification of this change is lacking. In order to assess quantitatively the biogeographical impact of mass extinctions, we outline time-traceable bioregions for benthic marine species across the Phanerozoic using a compositional network. Mass extinction events are visually recognizable in the geographical depiction of bioregions. The end-Permian extinction stands out with a severe reduction of provinciality. Time series of biogeographical turnover represent a novel aspect of the analysis of mass extinctions, confirming concentration of changes in the geographical distribution of benthic marine life