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Climate and continental configuration combined to make Early Paleozoic animals susceptible to extinction. 

The largest mass extinction event is associated with changes in degassing style of the Siberian Traps volcanism. 

Paleontological reconstructions of plankton community structure during warm periods of the Cenozoic (last 66 million years) reveal that deep-dwelling ‘twilight zone’ (200–1000 m) plankton were less abundant and diverse, and lived much closer to the surface, than in colder, more recent climates. We suggest that this is a consequence of temperature’s role in controlling the rate that sinking organic matter is broken down and metabolized by bacteria, a process that occurs faster at warmer temperatures. In a warmer ocean, a smaller fraction of organic matter reaches the ocean interior, affecting food supply and dissolved oxygen availability at depth. Using an Earth system model that has been evaluated against paleo observations, we illustrate how anthropogenic warming may impact future carbon cycling and twilight zone ecology. Our findings suggest that significant changes are already underway, and without strong emissions mitigation, widespread ecological disruption in the twilight zone is likely by 2100, with effects spanning millennia thereafter.

 

Abstract The fossil record of marine invertebrates has long fuelled the debate as to whether or not there are limits to global diversity in the sea 1–5 . Ecological theory states that, as diversity grows and ecological niches are filled, the strengthening of biological interactions imposes limits on diversity 6,7 . However, the extent to which biological interactions have constrained the growth of diversity over evolutionary time remains an open question 1–5,8–11 . Here we present a regional diversification model that reproduces the main Phanerozoic eon trends in the global diversity of marine invertebrates after imposing mass extinctions. We find that the dynamics of global diversity are best described by a diversification model that operates widely within the exponential growth regime of a logistic function. A spatially resolved analysis of the ratio of diversity to carrying capacity reveals that less than 2% of the global flooded continental area throughout the Phanerozoic exhibits diversity levels approaching ecological saturation. We attribute the overall increase in global diversity during the Late Mesozoic and Cenozoic eras to the development of diversity hotspots under prolonged conditions of Earth system stability and maximum continental fragmentation. We call this the ‘diversity hotspots hypothesis’, which we propose as a non-mutually exclusive alternative to the hypothesis that the Mesozoic marine revolution led this macroevolutionary trend 12,13 . 

The decline in background extinction rates of marine animals through geologic time is an established but unexplained feature of the Phanerozoic fossil record. There is also growing consensus that the ocean and atmosphere did not become oxygenated to near-modern levels until the mid-Paleozoic, coinciding with the onset of generally lower extinction rates. Physiological theory provides us with a possible causal link between these two observations—predicting that the synergistic impacts of oxygen and temperature on aerobic respiration would have made marine animals more vulnerable to ocean warming events during periods of limited surface oxygenation. Here, we evaluate the hypothesis that changes in surface oxygenation exerted a first-order control on extinction rates through the Phanerozoic using a combined Earth system and ecophysiological modeling approach. We find that although continental configuration, the efficiency of the biological carbon pump in the ocean, and initial climate state all impact the magnitude of modeled biodiversity loss across simulated warming events, atmospheric oxygen is the dominant predictor of extinction vulnerability, with metabolic habitat viability and global ecophysiotype extinction exhibiting inflection points around 40% of present atmospheric oxygen. Given this is the broad upper limit for estimates of early Paleozoic oxygen levels, our results are consistent with the relative frequency of high-magnitude extinction events (particularly those not included in the canonical big five mass extinctions) early in the Phanerozoic being a direct consequence of limited early Paleozoic oxygenation and temperature-dependent hypoxia responses.

 

Abstract. Temperature is a master parameter in the marine carbon cycle, exerting a critical control on the rate of biological transformation of a variety of solid and dissolved reactants and substrates. Although in the construction of numerical models of marine carbon cycling, temperature has been long recognised as a key parameter in the production and export of organic matter at the ocean surface, its role in the ocean interior is much less frequently accounted for. There, bacteria (primarily) transform sinking particulate organic matter (POM) into its dissolved constituents and consume dissolved oxygen (and/or other electron acceptors such as sulfate). The nutrients and carbon thereby released then become available for transport back to the surface, influencing biological productivity and atmospheric pCO2, respectively. Given the substantial changes in ocean temperature occurring in the past, as well as in light of current anthropogenic warming, appropriately accounting for the role of temperature in marine carbon cycling may be critical to correctly projecting changes in ocean deoxygenation and the strength of feedbacks on atmosphericpCO2. Here we extend and calibrate a temperature-dependent representation ofmarine carbon cycling in the cGENIE.muffin Earth system model, intended forboth past and future climate applications. In this, we combine atemperature-dependent remineralisation scheme for sinking organic matterwith a biological export production scheme that also includes a dependenceon ambient seawater temperature. Via a parameter ensemble, we jointlycalibrate the two parameterisations by statistically contrasting model-projected fields of nutrients, oxygen, and the stable carbon isotopicsignature (δ13C) of dissolved inorganic carbon in the oceanwith modern observations. We additionally explore the role of temperature inthe creation and recycling of dissolved organic matter (DOM) and hence itsimpact on global carbon cycle dynamics. We find that for the present day, the temperature-dependent version showsa fit to the data that is as good as or better than the existing tuned non-temperature-dependent version of the cGENIE.muffin. The main impact ofaccounting for temperature-dependent remineralisation of POM is in drivinghigher rates of remineralisation in warmer waters, in turn driving a morerapid return of nutrients to the surface and thereby stimulating organicmatter production. As a result, more POM is exported below 80 m but onaverage reaches shallower depths in middle- and low-latitude warmer waterscompared to the standard model. Conversely, at higher latitudes, colderwater temperature reduces the rate of nutrient resupply to the surface andPOM reaches greater depth on average as a result of slower subsurface ratesof remineralisation. Further adding temperature-dependent DOM processeschanges this overall picture only a little, with a slight weakening ofexport production at higher latitudes. As an illustrative application of the new model configuration andcalibration, we take the example of historical warming and briefly assessthe implications for global carbon cycling of accounting for a more completeset of temperature-dependent processes in the ocean. We find that betweenthe pre-industrial era (ca. 1700) and the present (year 2010), in response to asimulated air temperature increase of 0.9 ∘C and an associatedprojected mean ocean warming of 0.12 ∘C (0.6 ∘C insurface waters and 0.02 ∘C in deep waters), a reduction inparticulate organic carbon (POC) export at 80 m of just 0.3 % occurs (or 0.7 % including a temperature-dependent DOM response). However, due to this increased recycling nearer the surface, the efficiency of the transfer of carbon away from the surface (at 80 m) to the deep ocean (at 1040 m) is reduced by 5 %. In contrast, with no assumed temperature-dependent processes impacting production or remineralisation of either POM or DOM, global POC export at 80 m falls by 2.9 % between the pre-industrial era and the present day as a consequence of ocean stratification and reduced nutrient resupply to the surface. Our analysis suggests that increased temperature-dependent nutrient recycling in the upper ocean has offset much of the stratification-induced restriction in its physical transport. 

The geological record encodes the relationship between climate and atmospheric carbon dioxide (CO₂) over long and short timescales, as well as potential drivers of evolutionary transitions. However, reconstructing CO₂beyond direct measurements requires the use of paleoproxies and herein lies the challenge, as proxies differ in their assumptions, degree of understanding, and even reconstructed values. In this study, we critically evaluated, categorized, and integrated available proxies to create a high-fidelity and transparently constructed atmospheric CO₂record spanning the past 66 million years. This newly constructed record provides clearer evidence for higher Earth system sensitivity in the past and for the role of CO₂thresholds in biological and cryosphere evolution.