Climate change is an important predictor of extinction risk on macroevolutionary timescales.

Anthropogenic climate change is increasing rapidly and already impacting biodiversity. Despite its importance in future projections, understanding of the underlying mechanisms by which climate mediates extinction remains limited. We present an integrated approach examining the role of intrinsic traits versus extrinsic climate change in mediating extinction risk for marine invertebrates over the past 485 million years. We found that a combination of physiological traits and the magnitude of climate change is necessary to explain marine invertebrate extinction patterns. Our results suggest that taxa previously identified as extinction resistant may still succumb to extinction if the magnitude of climate change is great enough.

Human brains preserve in diverse environments for at least 12 000 years.

The brain is thought to be among the first human organs to decompose after death. The discovery of brains preserved in the archaeological record is therefore regarded as unusual. Although mechanisms such as dehydration, freezing, saponification, and tanning are known to allow for the preservation of the brain on short time scales in association with other soft tissues (≲4000 years), discoveries of older brains, especially in the absence of other soft tissues, are rare. Here, we collated an archive of more than 4400 human brains preserved in the archaeological record across approximately 12 000 years, more than 1300 of which constitute the only soft tissue preserved amongst otherwise skeletonized remains. We found that brains of this type persist on time scales exceeding those preserved by other means, which suggests an unknown mechanism may be responsible for preservation particular to the central nervous system. The untapped archive of preserved ancient brains represents an opportunity for bioarchaeological studies of human evolution, health and disease.

Ecological niche conservatism spurs diversification in response to climate change.

Lengthy debate has surrounded the theoretical and empirical science of whether climatic niche evolution is related to increased or decreased rates of biological diversification. Because species can persist for thousands to millions of years, these questions cross broad scales of time and space. Thus, short-term experiments may not provide comprehensive understanding of the system, leading to the emergence of contrasting opinions: niche evolution may increase diversity by allowing species to explore and colonize new geographic areas across which they could speciate; or, niche conservatism might augment biodiversity by supporting isolation of populations that may then undergo allopatric speciation. Here, we use a simulation approach to test how biological diversification responds to different rates and modes of niche evolution. We find that niche conservatism promotes biological diversification, whereas labile niches-whether adapting to the conditions available or changing randomly-generally led to slower diversification rates. These novel results provide a framework for understanding how Earth-life interactions produced such a diverse biota.

Measurement theory and paleobiology.

Measurement theory, a branch of applied mathematics, offers guiding principles for extracting meaning from empirical observations and is applicable to any science involving measurements. Measurement theory is highly relevant in paleobiology because statistical approaches assuming ratio-scaled variables are commonly used on data belonging to nominal and ordinal scale types. We provide an informal introduction to representational measurement theory and argue for its importance in robust scientific inquiry. Although measurement theory is widely applicable in paleobiology research, we use the study of disparity to illustrate measurement theoretical challenges in the quantitative study of the fossil record. Respecting the inherent properties of different measurements enables meaningful inferences about evolutionary and ecological processes from paleontological data.

Explanations for latitudinal diversity gradients must invoke rate variation.

The latitudinal diversity gradient (LDG) describes the pattern of increasing numbers of species from the poles to the equator. Although recognized for over 200 years, the mechanisms responsible for the largest-scale and longest-known pattern in macroecology are still actively debated. I argue here that any explanation for the LDG must invoke differential rates of speciation, extinction, extirpation, or dispersal. These processes themselves may be governed by numerous abiotic or biotic factors. Hypotheses that claim not to invoke differential rates, such as 'age and area' or 'time for diversification', eschew focus from rate variation that is assumed by these explanations. There is still significant uncertainty in how rates of speciation, extinction, extirpation, and dispersal have varied regionally over Earth history. However, to better understand the development of LDGs, we need to better constrain this variation. Only then will the drivers of such rate variation - be they abiotic or biotic in nature - become clearer.

Origination of the modern-style diversity gradient 15 million years ago.

The latitudinal diversity gradient (LDG) is a prevalent feature of modern ecosystems across diverse clades1-4. Recognized for well over a century, the causal mechanisms for LDGs remain disputed, in part because numerous putative drivers simultaneously covary with latitude1,3,5. The past provides the opportunity to disentangle LDG mechanisms because the relationships among biodiversity, latitude and possible causal factors have varied over time6-9. Here we quantify the emergence of the LDG in planktonic foraminifera at high spatiotemporal resolution over the past 40 million years, finding that a modern-style gradient arose only 15 million years ago. Spatial and temporal models suggest that LDGs for planktonic foraminifera may be controlled by the physical structure of the water column. Steepening of the latitudinal temperature gradient over 15 million years ago, associated with an increased vertical temperature gradient at low latitudes, may have enhanced niche partitioning and provided more opportunities for speciation at low latitudes. Supporting this hypothesis, we find that higher rates of low-latitude speciation steepened the diversity gradient, consistent with spatiotemporal patterns of depth partitioning by planktonic foraminifera. Extirpation of species from high latitudes also strengthened the LDG, but this effect tended to be weaker than speciation. Our results provide a step change in understanding the evolution of marine LDGs over long timescales.

Integrating geographic ranges across temporal scales.

Geographic ranges are a fundamental unit of biogeography and macroecology. Increasingly, paleontologists and ecologists alike are reconstructing geographic ranges of species from fossils, in order to understand the long-term processes governing biogeographic and macroevolutionary patterns. As these reconstructions have become increasingly common, uncertainty has arisen over the equivalency of paleo-ranges and modern ranges. Here, we argue geographic ranges are time-averaged at all temporal scales, and reflect the biotic and abiotic processes operating across the equivalent range of time and space scales. This conceptual framework integrates the study of geographic ranges reconstructed using modern and ancient data, and highlights the potential for ranges to illuminate processes responsible for diversity patterns over intervals spanning days to tens of millions of years of Earth history.

Climate change research and action must look beyond 2100.

Anthropogenic activity is changing Earth's climate and ecosystems in ways that are potentially dangerous and disruptive to humans. Greenhouse gas concentrations in the atmosphere continue to rise, ensuring that these changes will be felt for centuries beyond 2100, the current benchmark for projection. Estimating the effects of past, current, and potential future emissions to only 2100 is therefore short-sighted. Critical problems for food production and climate-forced human migration are projected to arise well before 2100, raising questions regarding the habitability of some regions of the Earth after the turn of the century. To highlight the need for more distant horizon scanning, we model climate change to 2500 under a suite of emission scenarios and quantify associated projections of crop viability and heat stress. Together, our projections show global climate impacts increase significantly after 2100 without rapid mitigation. As a result, we argue that projections of climate and its effects on human well-being and associated governance and policy must be framed beyond 2100.

Bottom-up controls, ecological revolutions and diversification in the oceans through time.

Animals originated in the oceans and evolved there for hundreds of millions of years before adapting to terrestrial environments. Today, oceans cover more than two-thirds of Earth and generate as much primary production as land. The path from the first macrobiota to modern marine biodiversity involved parallel increases in terrestrial nutrient input, marine primary production, species' abundance, metabolic rates, ecotypic diversity and taxonomic diversity. Bottom-up theories of ecosystem cascades arrange these changes in a causal sequence. At the base of marine food webs, nutrient fluxes and atmosphere-ocean chemistry interact with phytoplankton to regulate production. First-order consumers (e.g., zooplankton) might propagate changes in quantity and quality of phytoplankton to changes in abundance and diversity of larger predators (e.g., nekton). However, many uncertainties remain about the mechanisms and effect size of bottom-up control, particularly in oceans across the entire history of animal life. Here, we review modern and fossil evidence for hypothesized bottom-up pathways, and we assess the ramifications of these processes for four key intervals in marine ecosystems: the Ediacaran-Cambrian (635-485 million years ago), the Ordovician (485-444 million years ago), the Devonian (419-359 million years ago) and the Mesozoic (252-66 million years ago). We advocate for a clear articulation of bottom-up hypotheses to better understand causal relationships and proposed effects, combined with additional ecological experiments, paleontological documentation, isotope geochemistry and geophysical reconstructions. How small-scale ecological change transitions into large-scale evolutionary change remains an outstanding question for empirical and theoretical research.

Triton, a new species-level database of Cenozoic planktonic foraminiferal occurrences.

Planktonic foraminifera are a major constituent of ocean floor sediments, and thus have one of the most complete fossil records of any organism. Expeditions to sample these sediments have produced large amounts of spatiotemporal occurrence records throughout the Cenozoic, but no single source exists to house these data. We have therefore created a comprehensive dataset that integrates numerous sources for spatiotemporal records of planktonic foraminifera. This new dataset, Triton, contains >500,000 records and is four times larger than the previous largest database, Neptune. To ensure comparability among data sources, we have cleaned all records using a unified set of taxonomic concepts and have converted age data to the GTS 2020 timescale. Where ages were not absolute (e.g. based on biostratigraphic or magnetostratigraphic zones), we have used generalised additive models to produce continuous estimates. This dataset is an excellent resource for macroecological and macroevolutionary studies, particularly for investigating how species responded to past climatic changes.

Thermal niches of planktonic foraminifera are static throughout glacial-interglacial climate change.

Abiotic niche lability reduces extinction risk by allowing species to adapt to changing environmental conditions in situ. In contrast, species with static niches must keep pace with the velocity of climate change as they track suitable habitat. The rate and frequency of niche lability have been studied on human timescales (months to decades) and geological timescales (millions of years), but lability on intermediate timescales (millennia) remains largely uninvestigated. Here, we quantified abiotic niche lability at 8-ka resolution across the last 700 ka of glacial-interglacial climate fluctuations, using the exceptionally well-known fossil record of planktonic foraminifera coupled with Atmosphere-Ocean Global Climate Model reconstructions of paleoclimate. We tracked foraminiferal niches through time along the univariate axis of mean annual temperature, measured both at the sea surface and at species' depth habitats. Species' temperature preferences were uncoupled from the global temperature regime, undermining a hypothesis of local adaptation to changing environmental conditions. Furthermore, intraspecific niches were equally similar through time, regardless of climate change magnitude on short timescales (8 ka) and across contrasts of glacial and interglacial extremes. Evolutionary trait models fitted to time series of occupied temperature values supported widespread niche stasis above randomly wandering or directional change. Ecotype explained little variation in species-level differences in niche lability after accounting for evolutionary relatedness. Together, these results suggest that warming and ocean acidification over the next hundreds to thousands of years could redistribute and reduce populations of foraminifera and other calcifying plankton, which are primary components of marine food webs and biogeochemical cycles.

High Preservation Potential of Paleogeographic Range Size Distributions in Deep Time.

AbstractReconstructing geographic range sizes from fossil data is a crucial tool in paleoecology, elucidating macroecological and macroevolutionary processes. Studies examining links between range size and extinction risk may also offer a predictive tool for identifying species most vulnerable in the "sixth mass extinction." However, the extent to which paleogeographic ranges can be recorded reliably in the fossil record is unknown. We perform simulation-based extinction experiments to examine (1) the fidelity of paleogeographic range size preservation in deep time, (2) the relative performance of different methods for reconstructing range size, and (3) the reliability of detecting patterns of extinction "selectivity" on range size. Our results suggest both that relative paleogeographic range size can be consistently reconstructed and that selectivity patterns on range size can be preserved under many extinction intensities, even when sedimentary rocks are scarce. By identifying patterns of selectivity across Earth's history, paleontologists can thus augment neontological work that aims to predict and prevent extinctions of living species. Last, we find that introducing "false extinctions" in the fossil record can produce spurious range-selectivity signals. Errors in the temporal ranges of species may pose a larger barrier to reconstructing range size-extinction risk signals than the spatial distribution of fossiliferous sediments.

Acknowledging uncertainty in evolutionary reconstructions of ecological niches.

Reconstructing ecological niche evolution can provide insight into the biogeography and diversification of evolving lineages. However, comparative phylogenetic methods may infer the history of ecological niche evolution inaccurately because (a) species' niches are often poorly characterized; and (b) phylogenetic comparative methods rely on niche summary statistics rather than full estimates of species' environmental tolerances. Here, we propose a new framework for coding ecological niches and reconstructing their evolution that explicitly acknowledges and incorporates the uncertainty introduced by incomplete niche characterization. Then, we modify existing ancestral state inference methods to leverage full estimates of environmental tolerances. We provide a worked empirical example of our method, investigating ecological niche evolution in the New World orioles (Aves: Passeriformes: Icterus spp.). Temperature and precipitation tolerances were generally broad and conserved among orioles, with niche reduction and specialization limited to a few terminal branches. Tools for performing these reconstructions are available in a new R package called nichevol.

Disparities in the analysis of morphological disparity.

Analyses of morphological disparity have been used to characterize and investigate the evolution of variation in the anatomy, function and ecology of organisms since the 1980s. While a diversity of methods have been employed, it is unclear whether they provide equivalent insights. Here, we review the most commonly used approaches for characterizing and analysing morphological disparity, all of which have associated limitations that, if ignored, can lead to misinterpretation. We propose best practice guidelines for disparity analyses, while noting that there can be no 'one-size-fits-all' approach. The available tools should always be used in the context of a specific biological question that will determine data and method selection at every stage of the analysis.

The latitudinal diversity gradient of tetrapods across the Permo-Triassic mass extinction and recovery interval.

The decline in species richness from the equator to the poles is referred to as the latitudinal diversity gradient (LDG). Higher equatorial diversity has been recognized for over 200 years, but the consistency of this pattern in deep time remains uncertain. Examination of spatial biodiversity patterns in the past across different global climate regimes and continental configurations can reveal how LDGs have varied over Earth history and potentially differentiate between suggested causal mechanisms. The Late Permian-Middle Triassic represents an ideal time interval for study, because it is characterized by large-scale volcanic episodes, extreme greenhouse temperatures and the most severe mass extinction event in Earth history. We examined terrestrial and marine tetrapod spatial biodiversity patterns using a database of global tetrapod occurrences. Terrestrial tetrapods exhibit a bimodal richness distribution throughout the Late Permian-Middle Triassic, with peaks in the northern low latitudes and southern mid-latitudes around 20-40° N and 60° S, respectively. Marine reptile fossils are known almost exclusively from the Northern Hemisphere in the Early and Middle Triassic, with highest diversity around 20° N. Reconstructed terrestrial LDGs contrast strongly with the generally unimodal gradients of today, potentially reflecting high global temperatures and prevailing Pangaean super-monsoonal climate system during the Permo-Triassic.

Marine Biodiversity and Geographic Distributions Are Independent on Large Scales.

Fundamental ecological and evolutionary theories, such as community saturation and diversity-dependent diversification, assume that biotic competition restricts resource use, and thus limits realized niche breadth and geographic range size [1-3]. This principle is called competitive exclusion. The corollary (ecological release) posits that, after competitors disappear from a region, species that were previously excluded will invade. Hundreds of field experiments have demonstrated ecological release in living populations. However, few of these studies were conducted in marine environments, and almost no work extended beyond 10 years and 1,000 km2 [4, 5]. In limited investigation of marine taxa at larger spatiotemporal scales, macroecologists and paleobiologists have observed little evidence of competitive exclusion [6-9]. Here, we quantified spatial trends in the rich and densely sampled fossil history of brachiopods and bivalves, while accounting for inconsistent sampling coverage through time using a new method of spatial standardization. The number of potential competitors in a region did not explain the geographic distribution of constituent species or genera. Furthermore, although ecological release predicts species to expand after extinction events, survivors of intervals with net species loss expanded as rarely as species in other intervals. Regression model estimates indicated different spatial responses of brachiopods and bivalves, and of habitat specialists and generalists, but no effect from changes in number of potential competitors. Biotic competition may control the distribution of populations, but, on larger spatiotemporal scales, non-competitive factors may have driven biogeographic patterns of brachiopods and bivalves.

Addressing priority questions of conservation science with palaeontological data.

Palaeontologists often ask identical questions to those asked by ecologists. Despite this, ecology is considered a core discipline of conservation biology, while palaeontologists are rarely consulted in the protection of species, habitats and ecosystems. The recent emergence of conservation palaeobiology presents a big step towards better integration of palaeontology in conservation science, although its focus on historical baselines may not fully capture the potential contributions of geohistorical data to conservation science. In this essay we address previously defined priority questions in conservation and consider which of these questions may be answerable using palaeontological data. Using a statistical assessment of surveys, we find that conservation biologists and younger scientists have a more optimistic view of potential palaeontological contributions to the field compared to experienced palaeontologists. Participants considered questions related to climate change and marine ecosystems to be the best addressable with palaeontological data. As these categories are also deemed most relevant by ecologists and receive the greatest research effort in conservation, they are the natural choice for future academic collaboration. This article is part of a discussion meeting issue 'The past is a foreign country: how much can the fossil record actually inform conservation?'