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.

Effects of paleogeographic changes and CO2 variability on northern mid-latitudinal temperature gradients in the Cretaceous.

The Cretaceous 'greenhouse' period (~145 to ~66 million years ago, Ma) in Earth's history is relatively well documented by multiple paleoproxy records, which indicate that the meridional sea surface temperature (SST) gradient increased (non-monotonically) from the Valanginian (~135 Ma) to the Maastrichtian (~68 Ma). Changes in atmospheric CO2 concentration, solar constant, and paleogeography are the primary drivers of variations in the spatiotemporal distribution of SST. However, the particular contribution of each of these drivers (and their underlying mechanisms) to changes in the SST distribution remains poorly understood. Here we use data from a suite of paleoclimate simulations to compare the relative effects of atmospheric CO2 variability and paleogeographic changes on mid-latitudinal SST gradient through the Cretaceous. Further, we use a fundamental model of wind-driven ocean gyres to quantify how changes in the Northern Hemisphere paleogeography weaken the circulation in subtropical ocean gyres, leading to an increase in extratropical SSTs.

Climatic controls on the ecological ascendancy of dinosaurs.

The ascendancy of dinosaurs to become dominant components of terrestrial ecosystems was a pivotal event in the history of life, yet the drivers of their early evolution and biodiversity are poorly understood.1,2,3 During their early diversification in the Late Triassic, dinosaurs were initially rare and geographically restricted, only attaining wider distributions and greater abundance following the end-Triassic mass extinction event.4,5,6 This pattern is consistent with an opportunistic expansion model, initiated by the extinction of co-occurring groups such as aetosaurs, rauisuchians, and therapsids.4,7,8 However, this pattern could instead be a response to changes in global climatic distributions through the Triassic to Jurassic transition, especially given the increasing evidence that climate played a key role in constraining Triassic dinosaur distributions.7,9,10,11,12,13,14,15,16 Here, we test this hypothesis and elucidate how climate influenced early dinosaur distribution by quantitatively examining changes in dinosaur and tetrapod "climatic niche space" across the Triassic-Jurassic boundary. Statistical analyses show that Late Triassic sauropodomorph dinosaurs occupied a more restricted climatic niche space than other tetrapods and dinosaurs, being excluded from the hottest, low-latitude climate zones. A subsequent, earliest Jurassic expansion of sauropodomorph geographic distribution is linked to the expansion of their preferred climatic conditions. Evolutionary model-fitting analyses provide evidence for an important evolutionary shift from cooler to warmer climatic niches during the origin of Sauropoda. These results are consistent with the hypothesis that global abundance of sauropodomorph dinosaurs was facilitated by climatic change and provide support for the key role of climate in the ascendancy of dinosaurs.

The PhanSST global database of Phanerozoic sea surface temperature proxy data.

Paleotemperature proxy data form the cornerstone of paleoclimate research and are integral to understanding the evolution of the Earth system across the Phanerozoic Eon. Here, we present PhanSST, a database containing over 150,000 data points from five proxy systems that can be used to estimate past sea surface temperature. The geochemical data have a near-global spatial distribution and temporally span most of the Phanerozoic. Each proxy value is associated with consistent and queryable metadata fields, including information about the location, age, and taxonomy of the organism from which the data derive. To promote transparency and reproducibility, we include all available published data, regardless of interpreted preservation state or vital effects. However, we also provide expert-assigned diagenetic assessments, ecological and environmental flags, and other proxy-specific fields, which facilitate informed and responsible reuse of the database. The data are quality control checked and the foraminiferal taxonomy has been updated. PhanSST will serve as a valuable resource to the paleoclimate community and has myriad applications, including evolutionary, geochemical, diagenetic, and proxy calibration studies.

Scleromochlus and the early evolution of Pterosauromorpha.

Pterosaurs, the first vertebrates to evolve powered flight, were key components of Mesozoic terrestrial ecosystems from their sudden appearance in the Late Triassic until their demise at the end of the Cretaceous1-6. However, the origin and early evolution of pterosaurs are poorly understood owing to a substantial stratigraphic and morphological gap between these reptiles and their closest relatives6, Lagerpetidae7. Scleromochlus taylori, a tiny reptile from the early Late Triassic of Scotland discovered over a century ago, was hypothesized to be a key taxon closely related to pterosaurs8, but its poor preservation has limited previous studies and resulted in controversy over its phylogenetic position, with some even doubting its identification as an archosaur9. Here we use microcomputed tomographic scans to provide the first accurate whole-skeletal reconstruction and a revised diagnosis of Scleromochlus, revealing new anatomical details that conclusively identify it as a close pterosaur relative1 within Pterosauromorpha (the lagerpetid + pterosaur clade). Scleromochlus is anatomically more similar to lagerpetids than to pterosaurs and retains numerous features that were probably present in very early diverging members of Avemetatarsalia (bird-line archosaurs). These results support the hypothesis that the first flying reptiles evolved from tiny, probably facultatively bipedal, cursorial ancestors1.

African Hydroclimate During the Early Eocene From the DeepMIP Simulations.

The early Eocene (∼56-48 Myr ago) is characterized by high CO2 estimates (1,200-2,500 ppmv) and elevated global temperatures (∼10°C-16°C higher than modern). However, the response of the hydrological cycle during the early Eocene is poorly constrained, especially in regions with sparse data coverage (e.g., Africa). Here, we present a study of African hydroclimate during the early Eocene, as simulated by an ensemble of state-of-the-art climate models in the Deep-time Model Intercomparison Project (DeepMIP). A comparison between the DeepMIP pre-industrial simulations and modern observations suggests that model biases are model- and geographically dependent, however, these biases are reduced in the model ensemble mean. A comparison between the Eocene simulations and the pre-industrial suggests that there is no obvious wetting or drying trend as the CO2 increases. The results suggest that changes to the land sea mask (relative to modern) in the models may be responsible for the simulated increases in precipitation to the north of Eocene Africa. There is an increase in precipitation over equatorial and West Africa and associated drying over northern Africa as CO2 rises. There are also important dynamical changes, with evidence that anticyclonic low-level circulation is replaced by increased south-westerly flow at high CO2 levels. Lastly, a model-data comparison using newly compiled quantitative climate estimates from paleobotanical proxy data suggests a marginally better fit with the reconstructions at lower levels of CO2.

Climatic and tectonic drivers shaped the tropical distribution of coral reefs.

Today, warm-water coral reefs are limited to tropical-to-subtropical latitudes. These diverse ecosystems extended further poleward in the geological past, but the mechanisms driving these past distributions remain uncertain. Here, we test the role of climate and palaeogeography in shaping the distribution of coral reefs over geological timescales. To do so, we combine habitat suitability modelling, Earth System modelling and the ~247-million-year geological record of scleractinian coral reefs. A broader latitudinal distribution of climatically suitable habitat persisted throughout much of the Mesozoic-early Paleogene due to an expanded tropical belt and more equable distribution of shallow marine substrate. The earliest Cretaceous might be an exception, with reduced shallow marine substrate during a 'cold-snap' interval. Climatically suitable habitat area became increasingly skewed towards the tropics from the late Paleogene, likely steepening the latitudinal biodiversity gradient of reef-associated taxa. This was driven by global cooling and increases in tropical shallow marine substrate resulting from the tectonic evolution of the Indo-Australian Archipelago. Although our results suggest global warming might permit long-term poleward range expansions, coral reef ecosystems are unlikely to keep pace with the rapid rate of anthropogenic climate change.

Past terrestrial hydroclimate sensitivity controlled by Earth system feedbacks.

Despite tectonic conditions and atmospheric CO2 levels (pCO2) similar to those of present-day, geological reconstructions from the mid-Pliocene (3.3-3.0 Ma) document high lake levels in the Sahel and mesic conditions in subtropical Eurasia, suggesting drastic reorganizations of subtropical terrestrial hydroclimate during this interval. Here, using a compilation of proxy data and multi-model paleoclimate simulations, we show that the mid-Pliocene hydroclimate state is not driven by direct CO2 radiative forcing but by a loss of northern high-latitude ice sheets and continental greening. These ice sheet and vegetation changes are long-term Earth system feedbacks to elevated pCO2. Further, the moist conditions in the Sahel and subtropical Eurasia during the mid-Pliocene are a product of enhanced tropospheric humidity and a stationary wave response to the surface warming pattern, which varies strongly with land cover changes. These findings highlight the potential for amplified terrestrial hydroclimate responses over long timescales to a sustained CO2 forcing.

Impact of global cooling on Early Cretaceous high pCO2 world during the Weissert Event.

The Weissert Event ~133 million years ago marked a profound global cooling that punctuated the Early Cretaceous greenhouse. We present modelling, high-resolution bulk organic carbon isotopes and chronostratigraphically calibrated sea surface temperature (SSTs) based on an organic paleothermometer (the TEX86 proxy), which capture the Weissert Event in the semi-enclosed Weddell Sea basin, offshore Antarctica (paleolatitude ~54 °S; paleowater depth ~500 meters). We document a ~3-4 °C drop in SST coinciding with the Weissert cold end, and converge the Weddell Sea data, climate simulations and available worldwide multi-proxy based temperature data towards one unifying solution providing a best-fit between all lines of evidence. The outcome confirms a 3.0 °C ( ±1.7 °C) global mean surface cooling across the Weissert Event, which translates into a ~40% drop in atmospheric pCO2 over a period of ~700 thousand years. Consistent with geologic evidence, this pCO2 drop favoured the potential build-up of local polar ice.

Orographic evolution of northern Tibet shaped vegetation and plant diversity in eastern Asia.

The growth of the Tibetan Plateau throughout the past 66 million years has profoundly affected the Asian climate, but how this unparalleled orogenesis might have driven vegetation and plant diversity changes in eastern Asia is poorly understood. We approach this question by integrating modeling results and fossil data. We show that growth of north and northeastern Tibet affects vegetation and, crucially, plant diversity in eastern Asia by altering the monsoon system. This northern Tibetan orographic change induces a precipitation increase, especially in the dry (winter) season, resulting in a transition from deciduous broadleaf vegetation to evergreen broadleaf vegetation and plant diversity increases across southeastern Asia. Further quantifying the complexity of Tibetan orographic change is critical for understanding the finer details of Asian vegetation and plant diversity evolution.

Past climates inform our future.

As the world warms, there is a profound need to improve projections of climate change. Although the latest Earth system models offer an unprecedented number of features, fundamental uncertainties continue to cloud our view of the future. Past climates provide the only opportunity to observe how the Earth system responds to high carbon dioxide, underlining a fundamental role for paleoclimatology in constraining future climate change. Here, we review the relevancy of paleoclimate information for climate prediction and discuss the prospects for emerging methodologies to further insights gained from past climates. Advances in proxy methods and interpretations pave the way for the use of past climates for model evaluation-a practice that we argue should be widely adopted.

Asteroid impact, not volcanism, caused the end-Cretaceous dinosaur extinction.

The Cretaceous/Paleogene mass extinction, 66 Ma, included the demise of non-avian dinosaurs. Intense debate has focused on the relative roles of Deccan volcanism and the Chicxulub asteroid impact as kill mechanisms for this event. Here, we combine fossil-occurrence data with paleoclimate and habitat suitability models to evaluate dinosaur habitability in the wake of various asteroid impact and Deccan volcanism scenarios. Asteroid impact models generate a prolonged cold winter that suppresses potential global dinosaur habitats. Conversely, long-term forcing from Deccan volcanism (carbon dioxide [CO2]-induced warming) leads to increased habitat suitability. Short-term (aerosol cooling) volcanism still allows equatorial habitability. These results support the asteroid impact as the main driver of the non-avian dinosaur extinction. By contrast, induced warming from volcanism mitigated the most extreme effects of asteroid impact, potentially reducing the extinction severity.

Past East Asian monsoon evolution controlled by paleogeography, not CO2.

The East Asian monsoon plays an integral role in human society, yet its geological history and controlling processes are poorly understood. Using a general circulation model and geological data, we explore the drivers controlling the evolution of the monsoon system over the past 150 million years. In contrast to previous work, we find that the monsoon is controlled primarily by changes in paleogeography, with little influence from atmospheric CO2. We associate increased precipitation since the Late Cretaceous with the gradual uplift of the Himalayan-Tibetan region, transitioning from an ITCZ-dominated monsoon to a sea breeze-dominated monsoon. The rising region acted as a mechanical barrier to cold and dry continental air advecting into the region, leading to increasing influence of moist air from the Indian Ocean/South China Sea. We show that, apart from a dry period in the middle Cretaceous, a monsoon system has existed in East Asia since at least the Early Cretaceous.

Climatic shifts drove major contractions in avian latitudinal distributions throughout the Cenozoic.

Many higher level avian clades are restricted to Earth's lower latitudes, leading to historical biogeographic reconstructions favoring a Gondwanan origin of crown birds and numerous deep subclades. However, several such "tropical-restricted" clades (TRCs) are represented by stem-lineage fossils well outside the ranges of their closest living relatives, often on northern continents. To assess the drivers of these geographic disjunctions, we combined ecological niche modeling, paleoclimate models, and the early Cenozoic fossil record to examine the influence of climatic change on avian geographic distributions over the last ∼56 million years. By modeling the distribution of suitable habitable area through time, we illustrate that most Paleogene fossil-bearing localities would have been suitable for occupancy by extant TRC representatives when their stem-lineage fossils were deposited. Potentially suitable habitat for these TRCs is inferred to have become progressively restricted toward the tropics throughout the Cenozoic, culminating in relatively narrow circumtropical distributions in the present day. Our results are consistent with coarse-scale niche conservatism at the clade level and support a scenario whereby climate change over geological timescales has largely dictated the geographic distributions of many major avian clades. The distinctive modern bias toward high avian diversity at tropical latitudes for most hierarchical taxonomic levels may therefore represent a relatively recent phenomenon, overprinting a complex biogeographic history of dramatic geographic range shifts driven by Earth's changing climate, variable persistence, and intercontinental dispersal. Earth's current climatic trajectory portends a return to a megathermal state, which may dramatically influence the geographic distributions of many range-restricted extant clades.

Ecological niche modelling does not support climatically-driven dinosaur diversity decline before the Cretaceous/Paleogene mass extinction.

In the lead-up to the Cretaceous/Paleogene mass extinction, dinosaur diversity is argued to have been either in long-term decline, or thriving until their sudden demise. The latest Cretaceous (Campanian-Maastrichtian [83-66 Ma]) of North America provides the best record to address this debate, but even here diversity reconstructions are biased by uneven sampling. Here we combine fossil occurrences with climatic and environmental modelling to quantify latest Cretaceous North American dinosaur habitat. Ecological niche modelling shows a Campanian-to-Maastrichtian habitability decrease in areas with present-day rock-outcrop. However, a continent-wide projection demonstrates habitat stability, or even a Campanian-to-Maastrichtian increase, that is not preserved. This reduction of the spatial sampling window resulted from formation of the proto-Rocky Mountains and sea-level regression. We suggest that Maastrichtian North American dinosaur diversity is therefore likely to be underestimated, with the apparent decline a product of sampling bias, and not due to a climatically-driven decrease in habitability as previously hypothesised.

Placing our current 'hyperthermal' in the context of rapid climate change in our geological past.

'…there are known knowns. These are things we know that we know. There are known unknowns. That is to say, there are things that we know we don't know. But there are also unknown unknowns. There are things we don't know we don't know.' Donald Rumsfeld 12th February 2002.This article is part of a discussion meeting issue 'Hyperthermals: rapid and extreme global warming in our geological past'.

Future climate forcing potentially without precedent in the last 420 million years.

The evolution of Earth's climate on geological timescales is largely driven by variations in the magnitude of total solar irradiance (TSI) and changes in the greenhouse gas content of the atmosphere. Here we show that the slow ∼50 Wm-2 increase in TSI over the last ∼420 million years (an increase of ∼9 Wm-2 of radiative forcing) was almost completely negated by a long-term decline in atmospheric CO2. This was likely due to the silicate weathering-negative feedback and the expansion of land plants that together ensured Earth's long-term habitability. Humanity's fossil-fuel use, if unabated, risks taking us, by the middle of the twenty-first century, to values of CO2 not seen since the early Eocene (50 million years ago). If CO2 continues to rise further into the twenty-third century, then the associated large increase in radiative forcing, and how the Earth system would respond, would likely be without geological precedent in the last half a billion years.

Changing atmospheric CO2 concentration was the primary driver of early Cenozoic climate.

The Early Eocene Climate Optimum (EECO, which occurred about 51 to 53 million years ago), was the warmest interval of the past 65 million years, with mean annual surface air temperature over ten degrees Celsius warmer than during the pre-industrial period. Subsequent global cooling in the middle and late Eocene epoch, especially at high latitudes, eventually led to continental ice sheet development in Antarctica in the early Oligocene epoch (about 33.6 million years ago). However, existing estimates place atmospheric carbon dioxide (CO2) levels during the Eocene at 500-3,000 parts per million, and in the absence of tighter constraints carbon-climate interactions over this interval remain uncertain. Here we use recent analytical and methodological developments to generate a new high-fidelity record of CO2 concentrations using the boron isotope (δ(11)B) composition of well preserved planktonic foraminifera from the Tanzania Drilling Project, revising previous estimates. Although species-level uncertainties make absolute values difficult to constrain, CO2 concentrations during the EECO were around 1,400 parts per million. The relative decline in CO2 concentration through the Eocene is more robustly constrained at about fifty per cent, with a further decline into the Oligocene. Provided the latitudinal dependency of sea surface temperature change for a given climate forcing in the Eocene was similar to that of the late Quaternary period, this CO2 decline was sufficient to drive the well documented high- and low-latitude cooling that occurred through the Eocene. Once the change in global temperature between the pre-industrial period and the Eocene caused by the action of all known slow feedbacks (apart from those associated with the carbon cycle) is removed, both the EECO and the late Eocene exhibit an equilibrium climate sensitivity relative to the pre-industrial period of 2.1 to 4.6 degrees Celsius per CO2 doubling (66 per cent confidence), which is similar to the canonical range (1.5 to 4.5 degrees Celsius), indicating that a large fraction of the warmth of the early Eocene greenhouse was driven by increased CO2 concentrations, and that climate sensitivity was relatively constant throughout this period.

The impact of Cenozoic cooling on assemblage diversity in planktonic foraminifera.

The Cenozoic planktonic foraminifera (PF) (calcareous zooplankton) have arguably the most detailed fossil record of any group. The quality of this record allows models of environmental controls on macroecology, developed for Recent assemblages, to be tested on intervals with profoundly different climatic conditions. These analyses shed light on the role of long-term global cooling in establishing the modern latitudinal diversity gradient (LDG)--one of the most powerful generalizations in biogeography and macroecology. Here, we test the transferability of environment-diversity models developed for modern PF assemblages to the Eocene epoch (approx. 56-34 Ma), a time of pronounced global warmth. Environmental variables from global climate models are combined with Recent environment-diversity models to predict Eocene richness gradients, which are then compared with observed patterns. The results indicate the modern LDG--lower richness towards the poles--developed through the Eocene. Three possible causes are suggested for the mismatch between statistical model predictions and data in the Early Eocene: the environmental estimates are inaccurate, the statistical model misses a relevant variable, or the intercorrelations among facets of diversity--e.g. richness, evenness, functional diversity--have changed over geological time. By the Late Eocene, environment-diversity relationships were much more similar to those found today.