The impact of aragonite saturation variability on shelled pteropods: An attribution study in the California Current System.

Observations from the California Current System (CalCS) indicate that the long-term trend in ocean acidification (OA) and the naturally occurring corrosive conditions for the CaCO3 mineral aragonite (saturation state Ω < 1) have a damaging effect on shelled pteropods, a keystone group of calcifying organisms in the CalCS. Concern is heightened by recent findings suggesting that shell formation and developmental progress are already impacted when Ω falls below 1.5. Here, we quantify the impact of low Ω conditions on pteropods using an individual-based model (IBM) with life-stage-specific mortality, growth, and behavior in a high-resolution regional hindcast simulation of the CalCS between 1984 and 2019. Special attention is paid to attributing this impact to different processes that lead to such low Ω conditions, namely natural variability, long-term trend, and extreme events. We find that much of the observed damage in the CalCS, and specifically >70% of the shell CaCO3 loss, is due to the pteropods' exposure to naturally occurring low Ω conditions as a result of their diel vertical migration (DVM). Over the hindcast period, their exposure to damaging waters (Ω < 1.5) increases from 9% to 49%, doubling their shell CaCO3 loss, and increasing their mortality by ~40%. Most of this increased exposure is due to the shoaling of low Ω waters driven by the long-term trend in OA. Extreme OA events amplify this increase by ~40%. Our approach can quantify the health of pteropod populations under shifting environmental conditions, and attribute changes in fitness or population structure to changes in the stressor landscape across hierarchical time scales.

Global gradients in species richness of marine plankton functional groups.

The patterns of species diversity of plankton functional groups (PFGs) remain poorly understood although they matter greatly for marine ecosystem functioning. Here, we use an ensemble of empirical species distribution models for 845 plankton species to estimate the global species richness of three phytoplankton and 11 zooplankton functional groups as a function of objectively selected environmental predictors. The annual mean species richness of all PFGs decreases from the low to the high latitudes, but the steepness and the shape of this decrease vary significantly across PFGs. Pteropods, small copepods (Oithonids and Poecilostomatoids) and Salps have the steepest latitudinal gradients, whereas Amphipods and the three phytoplankton groups have the weakest ones. Temperature, irradiance and nutrient concentration are the first-order control on the latitudinal richness patterns, whilst the environmental conditions associated to upwelling systems, boundary currents and oxygen minimum zones modulate the position of the peaks and troughs in richness. The species richness of all PFGs increases with net primary production but decreases with particles size and the efficiency of the biological carbon pump. Our study puts forward emergent biodiversity-ecosystem functioning relationships and hypotheses about their underlying drivers for future field-based and modelling research.

Interaction matters: Bottom-up driver interdependencies alter the projected response of phytoplankton communities to climate change.

Phytoplankton growth is controlled by multiple environmental drivers, which are all modified by climate change. While numerous experimental studies identify interactive effects between drivers, large-scale ocean biogeochemistry models mostly account for growth responses to each driver separately and leave the results of these experimental multiple-driver studies largely unused. Here, we amend phytoplankton growth functions in a biogeochemical model by dual-driver interactions (CO2 and temperature, CO2 and light), based on data of a published meta-analysis on multiple-driver laboratory experiments. The effect of this parametrization on phytoplankton biomass and community composition is tested using present-day and future high-emission (SSP5-8.5) climate forcing. While the projected decrease in future total global phytoplankton biomass in simulations with driver interactions is similar to that in control simulations without driver interactions (5%-6%), interactive driver effects are group-specific. Globally, diatom biomass decreases more with interactive effects compared with the control simulation (-8.1% with interactions vs. no change without interactions). Small-phytoplankton biomass, by contrast, decreases less with on-going climate change when the model accounts for driver interactions (-5.0% vs. -9.0%). The response of global coccolithophore biomass to future climate conditions is even reversed when interactions are considered (+33.2% instead of -10.8%). Regionally, the largest difference in the future phytoplankton community composition between the simulations with and without driver interactions is detected in the Southern Ocean, where diatom biomass decreases (-7.5%) instead of increases (+14.5%), raising the share of small phytoplankton and coccolithophores of total phytoplankton biomass. Hence, interactive effects impact the phytoplankton community structure and related biogeochemical fluxes in a future ocean. Our approach is a first step to integrate the mechanistic understanding of interacting driver effects on phytoplankton growth gained by numerous laboratory experiments into a global ocean biogeochemistry model, aiming toward more realistic future projections of phytoplankton biomass and community composition.

Biogeochemical extremes and compound events in the ocean.

The ocean is warming, losing oxygen and being acidified, primarily as a result of anthropogenic carbon emissions. With ocean warming, acidification and deoxygenation projected to increase for decades, extreme events, such as marine heatwaves, will intensify, occur more often, persist for longer periods of time and extend over larger regions. Nevertheless, our understanding of oceanic extreme events that are associated with warming, low oxygen concentrations or high acidity, as well as their impacts on marine ecosystems, remains limited. Compound events-that is, multiple extreme events that occur simultaneously or in close sequence-are of particular concern, as their individual effects may interact synergistically. Here we assess patterns and trends in open ocean extremes based on the existing literature as well as global and regional model simulations. Furthermore, we discuss the potential impacts of individual and compound extremes on marine organisms and ecosystems. We propose a pathway to improve the understanding of extreme events and the capacity of marine life to respond to them. The conditions exhibited by present extreme events may be a harbinger of what may become normal in the future. As a consequence, pursuing this research effort may also help us to better understand the responses of marine organisms and ecosystems to future climate change.

Major restructuring of marine plankton assemblages under global warming.

Marine phytoplankton and zooplankton form the basis of the ocean's food-web, yet the impacts of climate change on their biodiversity are poorly understood. Here, we use an ensemble of species distribution models for a total of 336 phytoplankton and 524 zooplankton species to determine their present and future habitat suitability patterns. For the end of this century, under a high emission scenario, we find an overall increase in plankton species richness driven by ocean warming, and a poleward shift of the species' distributions at a median speed of 35 km/decade. Phytoplankton species richness is projected to increase by more than 16% over most regions except for the Arctic Ocean. In contrast, zooplankton richness is projected to slightly decline in the tropics, but to increase strongly in temperate to subpolar latitudes. In these latitudes, nearly 40% of the phytoplankton and zooplankton assemblages are replaced by poleward shifting species. This implies that climate change threatens the contribution of plankton communities to plankton-mediated ecosystem services such as biological carbon sequestration.

Global pattern of phytoplankton diversity driven by temperature and environmental variability.

Despite their importance to ocean productivity, global patterns of marine phytoplankton diversity remain poorly characterized. Although temperature is considered a key driver of general marine biodiversity, its specific role in phytoplankton diversity has remained unclear. We determined monthly phytoplankton species richness by using niche modeling and >540,000 global phytoplankton observations to predict biogeographic patterns of 536 phytoplankton species. Consistent with metabolic theory, phytoplankton richness in the tropics is about three times that in higher latitudes, with temperature being the most important driver. However, below 19°C, richness is lower than expected, with ~8°- 14°C waters (~35° to 60° latitude) showing the greatest divergence from theoretical predictions. Regions of reduced richness are characterized by maximal species turnover and environmental variability, suggesting that the latter reduces species richness directly, or through enhancing competitive exclusion. The nonmonotonic relationship between phytoplankton richness and temperature suggests unanticipated complexity in responses of marine biodiversity to ocean warming.

Chapter 1. Impacts of the oceans on climate change.

The oceans play a key role in climate regulation especially in part buffering (neutralising) the effects of increasing levels of greenhouse gases in the atmosphere and rising global temperatures. This chapter examines how the regulatory processes performed by the oceans alter as a response to climate change and assesses the extent to which positive feedbacks from the ocean may exacerbate climate change. There is clear evidence for rapid change in the oceans. As the main heat store for the world there has been an accelerating change in sea temperatures over the last few decades, which has contributed to rising sea-level. The oceans are also the main store of carbon dioxide (CO2), and are estimated to have taken up approximately 40% of anthropogenic-sourced CO2 from the atmosphere since the beginning of the industrial revolution. A proportion of the carbon uptake is exported via the four ocean 'carbon pumps' (Solubility, Biological, Continental Shelf and Carbonate Counter) to the deep ocean reservoir. Increases in sea temperature and changing planktonic systems and ocean currents may lead to a reduction in the uptake of CO2 by the ocean; some evidence suggests a suppression of parts of the marine carbon sink is already underway. While the oceans have buffered climate change through the uptake of CO2 produced by fossil fuel burning this has already had an impact on ocean chemistry through ocean acidification and will continue to do so. Feedbacks to climate change from acidification may result from expected impacts on marine organisms (especially corals and calcareous plankton), ecosystems and biogeochemical cycles. The polar regions of the world are showing the most rapid responses to climate change. As a result of a strong ice-ocean influence, small changes in temperature, salinity and ice cover may trigger large and sudden changes in regional climate with potential downstream feedbacks to the climate of the rest of the world. A warming Arctic Ocean may lead to further releases of the potent greenhouse gas methane from hydrates and permafrost. The Southern Ocean plays a critical role in driving, modifying and regulating global climate change via the carbon cycle and through its impact on adjacent Antarctica. The Antarctic Peninsula has shown some of the most rapid rises in atmospheric and oceanic temperature in the world, with an associated retreat of the majority of glaciers. Parts of the West Antarctic ice sheet are deflating rapidly, very likely due to a change in the flux of oceanic heat to the undersides of the floating ice shelves. The final section on modelling feedbacks from the ocean to climate change identifies limitations and priorities for model development and associated observations. Considering the importance of the oceans to climate change and our limited understanding of climate-related ocean processes, our ability to measure the changes that are taking place are conspicuously inadequate. The chapter highlights the need for a comprehensive, adequately funded and globally extensive ocean observing system to be implemented and sustained as a high priority. Unless feedbacks from the oceans to climate change are adequately included in climate change models, it is possible that the mitigation actions needed to stabilise CO2 and limit temperature rise over the next century will be underestimated.