Climate windows of opportunity for plant expansion during the Phanerozoic.

Earth's long-term climate may have profoundly influenced plant evolution. Local climatic factors, including water availability, light, and temperature, play a key role in plant physiology and growth, and have fluctuated substantially over geological time. However, the impact of these key climate variables on global plant biomass across the Phanerozoic has not yet been established. Linking climate and dynamic vegetation modelling, we identify two key 'windows of opportunity' during the Ordovician and Jurassic-Paleogene capable of supporting dramatic expansions of potential plant biomass. These conditions are driven by continental dispersion, paleolatitude of continental area and a lack of glaciation, allowing for an intense hydrological cycle and greater water availability. These windows coincide with the initial expansion of land plants and the later angiosperm radiation. Our findings suggest that the timing and expansion of habitable space for plants played an important role in plant evolution and diversification.

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.

Cyclic evolution of phytoplankton forced by changes in tropical seasonality.

Although the role of Earth's orbital variations in driving global climate cycles has long been recognized, their effect on evolution is hitherto unknown. The fossil remains of coccolithophores, a key calcifying phytoplankton group, enable a detailed assessment of the effect of cyclic orbital-scale climate changes on evolution because of their abundance in marine sediments and the preservation of their morphological adaptation to the changing environment1,2. Evolutionary genetic analyses have linked broad changes in Pleistocene fossil coccolith morphology to species radiation events3. Here, using high-resolution coccolith data, we show that during the last 2.8 million years the morphological evolution of coccolithophores was forced by Earth's orbital eccentricity with rhythms of around 100,000 years and 405,000 years-a distinct spectral signature to that of coeval global climate cycles4. Simulations with an Earth System Model5 coupled with an ocean biogeochemical model6 show a strong eccentricity modulation of the seasonal cycle, which we suggest directly affects the diversity of ecological niches that occur over the annual cycle in the tropical ocean. Reduced seasonality in surface ocean conditions favours species with mid-size coccoliths, increasing coccolith carbonate export and burial; whereas enhanced seasonality favours a larger range of coccolith sizes and reduced carbonate export. We posit that eccentricity pacing of phytoplankton evolution contributed to the strong 405,000-year cyclicity that is seen in global carbon cycle records.

Orbital variations as a major driver of climate and biome distribution during the greenhouse to icehouse transition.

Recent studies suggest increasing sensitivity to orbital variations across the Eocene-Oligocene greenhouse to icehouse climate transition. However, climate simulations and paleoenvironmental studies mostly provide snapshots of the past climate, therefore overlooking the role of this short-term variability in driving major environmental changes and possibly biasing model-data comparisons. We address this problem by performing numerical simulations describing the end-members of eccentricity, obliquity, and precession. The orbitally induced biome variability obtained in our simulations allows to reconcile previous apparent mismatch between models and paleobotanical compilations. We show that precession-driven intermittent monsoon-like climate may have taken place during the Eocene, resulting in biomes shifting from shrubland to tropical forest in the intertropical convergence zone migration region. Our Oligocene simulations suggest that, along with decreased pCO2, orbital variations crucially modulated major faunal dispersal events around the EOT such as the Grande Coupure by creating and fragmenting the biome corridors along several key land bridges.

Quantitative comparison of geological data and model simulations constrains early Cambrian geography and climate.

Marine ecosystems with a diverse range of animal groups became established during the early Cambrian (~541 to ~509 Ma). However, Earth's environmental parameters and palaeogeography in this interval of major macro-evolutionary change remain poorly constrained. Here, we test contrasting hypotheses of continental configuration and climate that have profound implications for interpreting Cambrian environmental proxies. We integrate general circulation models and geological observations to test three variants of the 'Antarctocentric' paradigm, with a southern polar continent, and an 'equatorial' configuration that lacks polar continents. This quantitative framework can be applied to other deep-time intervals when environmental proxy data are scarce. Our results show that the Antarctocentric palaeogeographic paradigm can reconcile geological data and simulated Cambrian climate. Our analyses indicate a greenhouse climate during the Cambrian animal radiation, with mean annual sea-surface temperatures between ~9 °C to ~19 °C and ~30 °C to ~38 °C for polar and tropical palaeolatitudes, respectively.

Response to Comment on "Revised paleoaltimetry data show low Tibetan Plateau elevation during the Eocene".

Valdes et al contest our results, suggesting failings in our modeling approach as well as in our comparison with data. Although their comment points to interesting ideas of improvement, we find that their critique reflects an incomplete understanding of our methods and is not supported by the material they provide.

Revised paleoaltimetry data show low Tibetan Plateau elevation during the Eocene.

Paleotopographic reconstructions of the Tibetan Plateau based on stable isotope paleoaltimetry methods conclude that most of the Plateau's current elevation was already reached by the Eocene, ~40 million years ago. However, changes in atmospheric and hydrological dynamics affect oxygen stable isotopes in precipitation and may thus bias such reconstructions. We used an isotope-equipped general circulation model to assess the influence of changing Eocene paleogeography and climate on paleoelevation estimates. Our simulations indicate that stable isotope paleoaltimetry methods are not applicable in Eocene Asia because of a combination of increased convective precipitation, mixture of air masses, and widespread aridity. Rather, a model-data comparison suggests that the Tibetan Plateau only reached low to moderate (less than 3000 meters) elevations during the Eocene, reconciling oxygen isotope data with other proxies.

An early Cambrian greenhouse climate.

The oceans of the early Cambrian (~541 to 509 million years ago) were the setting for a marked diversification of animal life. However, sea temperatures-a key component of the early Cambrian marine environment-remain unconstrained, in part because of a substantial time gap in the stable oxygen isotope (δ18O) record before the evolution of euconodonts. We show that previously overlooked sources of fossil biogenic phosphate have the potential to fill this gap. Pristine phosphatic microfossils from the Comley Limestones, UK, yield a robust δ18O signature, suggesting sea surface temperatures of 20° to 25°C at high southern paleolatitudes (~65°S to 70°S) between ~514 and 509 million years ago. These sea temperatures are consistent with the distribution of coeval evaporite and calcrete deposits, peak continental weathering rates, and also our climate model simulations for this interval. Our results support an early Cambrian greenhouse climate comparable to those of the late Mesozoic and early Cenozoic, offering a framework for exploring the interplay between biotic and environmental controls on Cambrian animal diversification.

Snowball Earth climate dynamics and Cryogenian geology-geobiology.

Geological evidence indicates that grounded ice sheets reached sea level at all latitudes during two long-lived Cryogenian (58 and ≥5 My) glaciations. Combined uranium-lead and rhenium-osmium dating suggests that the older (Sturtian) glacial onset and both terminations were globally synchronous. Geochemical data imply that CO2 was 102 PAL (present atmospheric level) at the younger termination, consistent with a global ice cover. Sturtian glaciation followed breakup of a tropical supercontinent, and its onset coincided with the equatorial emplacement of a large igneous province. Modeling shows that the small thermal inertia of a globally frozen surface reverses the annual mean tropical atmospheric circulation, producing an equatorial desert and net snow and frost accumulation elsewhere. Oceanic ice thickens, forming a sea glacier that flows gravitationally toward the equator, sustained by the hydrologic cycle and by basal freezing and melting. Tropical ice sheets flow faster as CO2 rises but lose mass and become sensitive to orbital changes. Equatorial dust accumulation engenders supraglacial oligotrophic meltwater ecosystems, favorable for cyanobacteria and certain eukaryotes. Meltwater flushing through cracks enables organic burial and submarine deposition of airborne volcanic ash. The subglacial ocean is turbulent and well mixed, in response to geothermal heating and heat loss through the ice cover, increasing with latitude. Terminal carbonate deposits, unique to Cryogenian glaciations, are products of intense weathering and ocean stratification. Whole-ocean warming and collapsing peripheral bulges allow marine coastal flooding to continue long after ice-sheet disappearance. The evolutionary legacy of Snowball Earth is perceptible in fossils and living organisms.

Palaeogeographic regulation of glacial events during the Cretaceous supergreenhouse.

The historical view of a uniformly warm Cretaceous is being increasingly challenged by the accumulation of new data hinting at the possibility of glacial events, even during the Cenomanian-Turonian (∼95 Myr ago), the warmest interval of the Cretaceous. Here we show that the palaeogeography typifying the Cenomanian-Turonian renders the Earth System resilient to glaciation with no perennial ice accumulation occurring under prescribed CO2 levels as low as 420 p.p.m. Conversely, late Aptian (∼115 Myr ago) and Maastrichtian (∼70 Myr ago) continental configurations set the stage for cooler climatic conditions, favouring possible inception of Antarctic ice sheets under CO2 concentrations, respectively, about 400 and 300 p.p.m. higher than for the Cenomanian-Turonian. Our simulations notably emphasize that palaeogeography can crucially impact global climate by modulating the CO2 threshold for ice sheet inception and make the possibility of glacial events during the Cenomanian-Turonian unlikely.

A better-ventilated ocean triggered by Late Cretaceous changes in continental configuration.

Oceanic anoxic events (OAEs) are large-scale events of oxygen depletion in the deep ocean that happened during pre-Cenozoic periods of extreme warmth. Here, to assess the role of major continental configuration changes occurring during the Late Cretaceous on oceanic circulation modes, which in turn influence the oxygenation level of the deep ocean, we use a coupled ocean atmosphere climate model. We simulate ocean dynamics during two different time slices and compare these with existing neodymium isotope data (ɛNd). Although deep-water production in the North Pacific is continuous, the simulations at 94 and 71 Ma show a shift in southern deep-water production sites from South Pacific to South Atlantic and Indian Ocean locations. Our modelling results support the hypothesis that an intensification of southern Atlantic deep-water production and a reversal of deep-water fluxes through the Caribbean Seaway were the main causes of the decrease in ɛNd values recorded in the Atlantic and Indian deep waters during the Late Cretaceous.

Tectonic-driven climate change and the diversification of angiosperms.

In 1879, Charles Darwin characterized the sudden and unexplained rise of angiosperms during the Cretaceous as an "abominable mystery." The diversification of this clade marked the beginning of a rapid transition among Mesozoic ecosystems and floras formerly dominated by ferns, conifers, and cycads. Although the role of environmental factors has been suggested [Coiffard C, Gómez B (2012) Geol Acta 10(2):181-188], Cretaceous global climate change has barely been considered as a contributor to angiosperm radiation, and focus was put on biotic factors to explain this transition. Here we use a fully coupled climate model driven by Mesozoic paleogeographic maps to quantify and discuss the impact of continental drift on angiosperm expansion and diversification. We show that the decrease of desertic belts between the Triassic and the Cretaceous and the subsequent onset of long-lasting humid conditions during the Late Cretaceous were driven by the breakup of Pangea and were contemporaneous with the first rise of angiosperm diversification. Positioning angiosperm-bearing fossil sites on our paleobioclimatic maps shows a strong match between the location of fossil-rich outcrops and temperate humid zones, indicating that climate change from arid to temperate dominance may have set the stage for the ecological expansion of flowering plants.

Carbon cycling and snowball Earth.

The possibility that Earth witnessed episodes of global glaciation during the latest Precambrian challenges our understanding of the physical processes controlling the Earth's climate. Peltier et al. suggest that a 'hard snowball Earth' state may have been prevented owing to the release of CO(2) from the oxidation of dissolved organic carbon (DOC) in the ocean as the temperature decreased. Here we show that the model of Peltier et al. is not self-consistent as it implies large fluctuations of the ocean alkalinity content without providing any processes to account for it. Our findings suggest that the hard snowball Earth hypothesis is still valid.

A 'snowball Earth' climate triggered by continental break-up through changes in runoff.

Geological and palaeomagnetic studies indicate that ice sheets may have reached the Equator at the end of the Proterozoic eon, 800 to 550 million years ago, leading to the suggestion of a fully ice-covered 'snowball Earth'. Climate model simulations indicate that such a snowball state for the Earth depends on anomalously low atmospheric carbon dioxide concentrations, in addition to the Sun being 6 per cent fainter than it is today. However, the mechanisms producing such low carbon dioxide concentrations remain controversial. Here we assess the effect of the palaeogeographic changes preceding the Sturtian glacial period, 750 million years ago, on the long-term evolution of atmospheric carbon dioxide levels using the coupled climate-geochemical model GEOCLIM. In our simulation, the continental break-up of Rodinia leads to an increase in runoff and hence consumption of carbon dioxide through continental weathering that decreases atmospheric carbon dioxide concentrations by 1,320 p.p.m. This indicates that tectonic changes could have triggered a progressive transition from a 'greenhouse' to an 'icehouse' climate during the Neoproterozoic era. When we combine these results with the concomitant weathering effect of the voluminous basaltic traps erupted throughout the break-up of Rodinia, our simulation results in a snowball glaciation.