Filter-feeding gelatinous macrozooplankton response to climate change and implications for benthic food supply and global carbon cycle.

It is often suggested that gelatinous zooplankton may benefit from anthropogenic pressures of all kinds and in particular from climate change. Large pelagic tunicates, for example, are likely to be favored over other types of macrozooplankton due to their filter-feeding mode, which gives them access to small preys thought to be less affected by climate change than larger preys. In this study, we provide model-based estimate of potential community changes in macrozooplankton composition and estimate for the first time their effects on benthic food supply and on the ocean carbon cycle under two 21st-century climate-change scenarios. Forced with output from an Earth System Model climate projections, our ocean biogeochemical model simulates a large reduction in macrozooplankton biomass in response to anthropogenic climate change, but shows that gelatinous macrozooplankton are less affected than nongelatinous macrozooplankton, with global biomass declines estimated at -2.8% and -3.5%, respectively, for every 1°C of warming. The inclusion of gelatinous macrozooplankon in our ocean biogeochemical model has a limited effect on anthropogenic carbon uptake in the 21st century, but impacts the projected decline in particulate organic matter fluxes in the deep ocean. In subtropical oligotrophic gyres, where gelatinous zooplankton dominate macrozooplankton, the decline in the amount of organic matter reaching the seafloor is reduced by a factor of 2 when gelatinous macrozooplankton are considered (-17.5% vs. -29.7% when gelatinous macrozooplankton are not considered, all for 2100 under RCP8.5). The shift to gelatinous macrozooplankton in the future ocean therefore buffers the decline in deep carbon fluxes and should be taken into account when assessing potential changes in deep carbon storage and the risks that deep ecosystems may face when confronted with a decline in their food source.

Ocean iron fertilization may amplify climate change pressures on marine animal biomass for limited climate benefit.

Climate change scenarios suggest that large-scale carbon dioxide removal (CDR) will be required to maintain global warming below 2°C, leading to renewed attention on ocean iron fertilization (OIF). Previous OIF modelling has found that while carbon export increases, nutrient transport to lower latitude ecosystems declines, resulting in a modest impact on atmospheric CO2 . However, the interaction of these CDR responses with ongoing climate change is unknown. Here, we combine global ocean biogeochemistry and ecosystem models to show that, while stimulating carbon sequestration, OIF may amplify climate-induced declines in tropical ocean productivity and ecosystem biomass under a high-emission scenario, with very limited potential atmospheric CO2 drawdown. The 'biogeochemical fingerprint' of climate change, that leads to depletion of upper ocean major nutrients due to upper ocean stratification, is reinforced by OIF due to greater major nutrient consumption. Our simulations show that reductions in upper trophic level animal biomass in tropical regions due to climate change would be exacerbated by OIF within ~20 years, especially in coastal exclusive economic zones (EEZs), with potential implications for fisheries that underpin the livelihoods and economies of coastal communities. Any fertilization-based CDR should therefore consider its interaction with ongoing climate-driven changes and the ensuing ecosystem impacts in national EEZs.

Demystifying global climate models for use in the life sciences.

For each assessment cycle of the Intergovernmental Panel on Climate Change (IPCC), researchers in the life sciences are called upon to provide evidence to policymakers planning for a changing future. This research increasingly relies on highly technical and complex outputs from climate models. The strengths and weaknesses of these data may not be fully appreciated beyond the climate modelling community; therefore, uninformed use of raw or preprocessed climate data could lead to overconfident or spurious conclusions. We provide an accessible introduction to climate model outputs that is intended to empower the life science community to robustly address questions about human and natural systems in a changing world.

High trophic level feedbacks on global ocean carbon uptake and marine ecosystem dynamics under climate change.

Despite recurrent emphasis on their ecological and economic roles, the importance of high trophic levels (HTLs) on ocean carbon dynamics, through passive (fecal pellet production, carcasses) and active (vertical migration) processes, is still largely unexplored, notably under climate change scenarios. In addition, HTLs impact the ecosystem dynamics through top-down effects on lower trophic levels, which might change under anthropogenic influence. Here we compare two simulations of a global biogeochemical-ecosystem model with and without feedbacks from large marine animals. We show that these large marine animals affect the evolution of low trophic level biomasses, hence net primary production and most certainly ecosystem equilibrium, but seem to have little influence on the 21st-century anthropogenic carbon uptake under the RCP8.5 scenario. These results provide new insights regarding the expectations for trophic amplification of climate change through the marine trophic chain and regarding the necessity to explicitly represent marine animals in Earth System Models.

Evidence that Pacific tuna mercury levels are driven by marine methylmercury production and anthropogenic inputs.

Pacific Ocean tuna is among the most-consumed seafood products but contains relatively high levels of the neurotoxin methylmercury. Limited observations suggest tuna mercury levels vary in space and time, yet the drivers are not well understood. Here, we map mercury concentrations in skipjack tuna across the Pacific Ocean and build generalized additive models to quantify the anthropogenic, ecological, and biogeochemical drivers. Skipjack mercury levels display a fivefold spatial gradient, with maximum concentrations in the northwest near Asia, intermediate values in the east, and the lowest levels in the west, southwest, and central Pacific. Large spatial differences can be explained by the depth of the seawater methylmercury peak near low-oxygen zones, leading to enhanced tuna mercury concentrations in regions where oxygen depletion is shallow. Despite this natural biogeochemical control, the mercury hotspot in tuna caught near Asia is explained by elevated atmospheric mercury concentrations and/or mercury river inputs to the coastal shelf. While we cannot ignore the legacy mercury contribution from other regions to the Pacific Ocean (e.g., North America and Europe), our results suggest that recent anthropogenic mercury release, which is currently largest in Asia, contributes directly to present-day human mercury exposure.

Next-generation ensemble projections reveal higher climate risks for marine ecosystems.

Projections of climate change impacts on marine ecosystems have revealed long-term declines in global marine animal biomass and unevenly distributed impacts on fisheries. Here we apply an enhanced suite of global marine ecosystem models from the Fisheries and Marine Ecosystem Model Intercomparison Project (Fish-MIP), forced by new-generation Earth system model outputs from Phase 6 of the Coupled Model Intercomparison Project (CMIP6), to provide insights into how projected climate change will affect future ocean ecosystems. Compared with the previous generation CMIP5-forced Fish-MIP ensemble, the new ensemble ecosystem simulations show a greater decline in mean global ocean animal biomass under both strong-mitigation and high-emissions scenarios due to elevated warming, despite greater uncertainty in net primary production in the high-emissions scenario. Regional shifts in the direction of biomass changes highlight the continued and urgent need to reduce uncertainty in the projected responses of marine ecosystems to climate change to help support adaptation planning.

Impact of intensifying nitrogen limitation on ocean net primary production is fingerprinted by nitrogen isotopes.

The open ocean nitrogen cycle is being altered by increases in anthropogenic atmospheric nitrogen deposition and climate change. How the nitrogen cycle responds will determine long-term trends in net primary production (NPP) in the nitrogen-limited low latitude ocean, but is poorly constrained by uncertainty in how the source-sink balance will evolve. Here we show that intensifying nitrogen limitation of phytoplankton, associated with near-term reductions in NPP, causes detectable declines in nitrogen isotopes (δ15N) and constitutes the primary perturbation of the 21st century nitrogen cycle. Model experiments show that ~75% of the low latitude twilight zone develops anomalously low δ15N by 2060, predominantly due to the effects of climate change that alter ocean circulation, with implications for the nitrogen source-sink balance. Our results highlight that δ15N changes in the low latitude twilight zone may provide a useful constraint on emerging changes to nitrogen limitation and NPP over the 21st century.

Around one third of current Arctic Ocean primary production sustained by rivers and coastal erosion.

Net primary production (NPP) is the foundation of the oceans' ecosystems and the fisheries they support. In the Arctic Ocean, NPP is controlled by a complex interplay of light and nutrients supplied by upwelling as well as lateral inflows from adjacent oceans and land. But so far, the role of the input from land by rivers and coastal erosion has not been given much attention. Here, by upscaling observations from the six largest rivers and using measured coastal erosion rates, we construct a pan-Arctic, spatio-temporally resolved estimate of the land input of carbon and nutrients to the Arctic Ocean. Using an ocean-biogeochemical model, we estimate that this input fuels 28-51% of the current annual Arctic Ocean NPP. This strong enhancement of NPP is a consequence of efficient recycling of the land-derived nutrients on the vast Arctic shelves. Our results thus suggest that nutrient input from the land is a key process that will affect the future evolution of Arctic Ocean NPP.

A comprehensive quantification of global nitrous oxide sources and sinks.

Nitrous oxide (N2O), like carbon dioxide, is a long-lived greenhouse gas that accumulates in the atmosphere. Over the past 150 years, increasing atmospheric N2O concentrations have contributed to stratospheric ozone depletion1 and climate change2, with the current rate of increase estimated at 2 per cent per decade. Existing national inventories do not provide a full picture of N2O emissions, owing to their omission of natural sources and limitations in methodology for attributing anthropogenic sources. Here we present a global N2O inventory that incorporates both natural and anthropogenic sources and accounts for the interaction between nitrogen additions and the biochemical processes that control N2O emissions. We use bottom-up (inventory, statistical extrapolation of flux measurements, process-based land and ocean modelling) and top-down (atmospheric inversion) approaches to provide a comprehensive quantification of global N2O sources and sinks resulting from 21 natural and human sectors between 1980 and 2016. Global N2O emissions were 17.0 (minimum-maximum estimates: 12.2-23.5) teragrams of nitrogen per year (bottom-up) and 16.9 (15.9-17.7) teragrams of nitrogen per year (top-down) between 2007 and 2016. Global human-induced emissions, which are dominated by nitrogen additions to croplands, increased by 30% over the past four decades to 7.3 (4.2-11.4) teragrams of nitrogen per year. This increase was mainly responsible for the growth in the atmospheric burden. Our findings point to growing N2O emissions in emerging economies-particularly Brazil, China and India. Analysis of process-based model estimates reveals an emerging N2O-climate feedback resulting from interactions between nitrogen additions and climate change. The recent growth in N2O emissions exceeds some of the highest projected emission scenarios3,4, underscoring the urgency to mitigate N2O emissions.

An iron cycle cascade governs the response of equatorial Pacific ecosystems to climate change.

Earth System Models project that global climate change will reduce ocean net primary production (NPP), upper trophic level biota biomass and potential fisheries catches in the future, especially in the eastern equatorial Pacific. However, projections from Earth System Models are undermined by poorly constrained assumptions regarding the biological cycling of iron, which is the main limiting resource for NPP over large parts of the ocean. In this study, we show that the climate change trends in NPP and the biomass of upper trophic levels are strongly affected by modifying assumptions associated with phytoplankton iron uptake. Using a suite of model experiments, we find 21st century climate change impacts on regional NPP range from -12.3% to +2.4% under a high emissions climate change scenario. This wide range arises from variations in the efficiency of iron retention in the upper ocean in the eastern equatorial Pacific across different scenarios of biological iron uptake, which affect the strength of regional iron limitation. Those scenarios where nitrogen limitation replaced iron limitation showed the largest projected NPP declines, while those where iron limitation was more resilient displayed little future change. All model scenarios have similar skill in reproducing past inter-annual variations in regional ocean NPP, largely due to limited change in the historical period. Ultimately, projections of end of century upper trophic level biomass change are altered by 50%-80% across all plausible scenarios. Overall, we find that uncertainties in the biological iron cycle cascade through open ocean pelagic ecosystems, from plankton to fish, affecting their evolution under climate change. This highlights additional challenges to developing effective conservation and fisheries management policies under climate change.

Tracking Improvement in Simulated Marine Biogeochemistry Between CMIP5 and CMIP6.

Purpose of Review: The changes or updates in ocean biogeochemistry component have been mapped between CMIP5 and CMIP6 model versions, and an assessment made of how far these have led to improvements in the simulated mean state of marine biogeochemical models within the current generation of Earth system models (ESMs). Recent Findings: The representation of marine biogeochemistry has progressed within the current generation of Earth system models. However, it remains difficult to identify which model updates are responsible for a given improvement. In addition, the full potential of marine biogeochemistry in terms of Earth system interactions and climate feedback remains poorly examined in the current generation of Earth system models. Summary: Increasing availability of ocean biogeochemical data, as well as an improved understanding of the underlying processes, allows advances in the marine biogeochemical components of the current generation of ESMs. The present study scrutinizes the extent to which marine biogeochemistry components of ESMs have progressed between the 5th and the 6th phases of the Coupled Model Intercomparison Project (CMIP).

The impacts of ocean acidification on marine trace gases and the implications for atmospheric chemistry and climate.

Surface ocean biogeochemistry and photochemistry regulate ocean-atmosphere fluxes of trace gases critical for Earth's atmospheric chemistry and climate. The oceanic processes governing these fluxes are often sensitive to the changes in ocean pH (or pCO2) accompanying ocean acidification (OA), with potential for future climate feedbacks. Here, we review current understanding (from observational, experimental and model studies) on the impact of OA on marine sources of key climate-active trace gases, including dimethyl sulfide (DMS), nitrous oxide (N2O), ammonia and halocarbons. We focus on DMS, for which available information is considerably greater than for other trace gases. We highlight OA-sensitive regions such as polar oceans and upwelling systems, and discuss the combined effect of multiple climate stressors (ocean warming and deoxygenation) on trace gas fluxes. To unravel the biological mechanisms responsible for trace gas production, and to detect adaptation, we propose combining process rate measurements of trace gases with longer term experiments using both model organisms in the laboratory and natural planktonic communities in the field. Future ocean observations of trace gases should be routinely accompanied by measurements of two components of the carbonate system to improve our understanding of how in situ carbonate chemistry influences trace gas production. Together, this will lead to improvements in current process model capabilities and more reliable predictions of future global marine trace gas fluxes.

Emergent constraint on Arctic Ocean acidification in the twenty-first century.

The ongoing uptake of anthropogenic carbon by the ocean leads to ocean acidification, a process that results in a reduction in pH and in the saturation state of biogenic calcium carbonate minerals aragonite (Ωarag) and calcite (Ωcalc)1,2. Because of its naturally low Ωarag and Ωcalc (refs. 2,3), the Arctic Ocean is considered the region most susceptible to future acidification and associated ecosystem impacts4-7. However, the magnitude of projected twenty-first century acidification differs strongly across Earth system models8. Here we identify an emergent multi-model relationship between the simulated present-day density of Arctic Ocean surface waters, used as a proxy for Arctic deep-water formation, and projections of the anthropogenic carbon inventory and coincident acidification. By applying observations of sea surface density, we constrain the end of twenty-first century Arctic Ocean anthropogenic carbon inventory to 9.0 ± 1.6 petagrams of carbon and the basin-averaged Ωarag and Ωcalc to 0.76 ± 0.06 and 1.19 ± 0.09, respectively, under the high-emissions Representative Concentration Pathway 8.5 climate scenario. Our results indicate greater regional anthropogenic carbon storage and ocean acidification than previously projected3,8 and increase the probability that large parts of the mesopelagic Arctic Ocean will be undersaturated with respect to calcite by the end of the century. This increased rate of Arctic Ocean acidification, combined with rapidly changing physical and biogeochemical Arctic conditions9-11, is likely to exacerbate the impact of climate change on vulnerable Arctic marine ecosystems.

Trends in tuna carbon isotopes suggest global changes in pelagic phytoplankton communities.

Considerable uncertainty remains over how increasing atmospheric CO2 and anthropogenic climate changes are affecting open-ocean marine ecosystems from phytoplankton to top predators. Biological time series data are thus urgently needed for the world's oceans. Here, we use the carbon stable isotope composition of tuna to provide a first insight into the existence of global trends in complex ecosystem dynamics and changes in the oceanic carbon cycle. From 2000 to 2015, considerable declines in δ13 C values of 0.8‰-2.5‰ were observed across three tuna species sampled globally, with more substantial changes in the Pacific Ocean compared to the Atlantic and Indian Oceans. Tuna recorded not only the Suess effect, that is, fossil fuel-derived and isotopically light carbon being incorporated into marine ecosystems, but also recorded profound changes at the base of marine food webs. We suggest a global shift in phytoplankton community structure, for example, a reduction in 13 C-rich phytoplankton such as diatoms, and/or a change in phytoplankton physiology during this period, although this does not rule out other concomitant changes at higher levels in the food webs. Our study establishes tuna δ13 C values as a candidate essential ocean variable to assess complex ecosystem responses to climate change at regional to global scales and over decadal timescales. Finally, this time series will be invaluable in calibrating and validating global earth system models to project changes in marine biota.

Global Trends in Marine Plankton Diversity across Kingdoms of Life.

The ocean is home to myriad small planktonic organisms that underpin the functioning of marine ecosystems. However, their spatial patterns of diversity and the underlying drivers remain poorly known, precluding projections of their responses to global changes. Here we investigate the latitudinal gradients and global predictors of plankton diversity across archaea, bacteria, eukaryotes, and major virus clades using both molecular and imaging data from Tara Oceans. We show a decline of diversity for most planktonic groups toward the poles, mainly driven by decreasing ocean temperatures. Projections into the future suggest that severe warming of the surface ocean by the end of the 21st century could lead to tropicalization of the diversity of most planktonic groups in temperate and polar regions. These changes may have multiple consequences for marine ecosystem functioning and services and are expected to be particularly significant in key areas for carbon sequestration, fisheries, and marine conservation. VIDEO ABSTRACT.

Global ensemble projections reveal trophic amplification of ocean biomass declines with climate change.

While the physical dimensions of climate change are now routinely assessed through multimodel intercomparisons, projected impacts on the global ocean ecosystem generally rely on individual models with a specific set of assumptions. To address these single-model limitations, we present standardized ensemble projections from six global marine ecosystem models forced with two Earth system models and four emission scenarios with and without fishing. We derive average biomass trends and associated uncertainties across the marine food web. Without fishing, mean global animal biomass decreased by 5% (±4% SD) under low emissions and 17% (±11% SD) under high emissions by 2100, with an average 5% decline for every 1 °C of warming. Projected biomass declines were primarily driven by increasing temperature and decreasing primary production, and were more pronounced at higher trophic levels, a process known as trophic amplification. Fishing did not substantially alter the effects of climate change. Considerable regional variation featured strong biomass increases at high latitudes and decreases at middle to low latitudes, with good model agreement on the direction of change but variable magnitude. Uncertainties due to variations in marine ecosystem and Earth system models were similar. Ensemble projections performed well compared with empirical data, emphasizing the benefits of multimodel inference to project future outcomes. Our results indicate that global ocean animal biomass consistently declines with climate change, and that these impacts are amplified at higher trophic levels. Next steps for model development include dynamic scenarios of fishing, cumulative human impacts, and the effects of management measures on future ocean biomass trends.

Consistent trophic amplification of marine biomass declines under climate change.

The impact of climate change on the marine food web is highly uncertain. Nonetheless, there is growing consensus that global marine primary production will decline in response to future climate change, largely due to increased stratification reducing the supply of nutrients to the upper ocean. Evidence to date suggests a potential amplification of this response throughout the trophic food web, with more dramatic responses at higher trophic levels. Here we show that trophic amplification of marine biomass declines is a consistent feature of the Coupled Model Intercomparison Project Phase 5 (CMIP5) Earth System Models, across different scenarios of future climate change. Under the business-as-usual Representative Concentration Pathway 8.5 (RCP8.5) global mean phytoplankton biomass is projected to decline by 6.1% ± 2.5% over the twenty-first century, while zooplankton biomass declines by 13.6% ± 3.0%. All models project greater relative declines in zooplankton than phytoplankton, with annual zooplankton biomass anomalies 2.24 ± 1.03 times those of phytoplankton. The low latitude oceans drive the projected trophic amplification of biomass declines, with models exhibiting variable trophic interactions in the mid-to-high latitudes and similar relative changes in phytoplankton and zooplankton biomass. Under the assumption that zooplankton biomass is prey limited, an analytical explanation of the trophic amplification that occurs in the low latitudes can be derived from generic plankton differential equations. Using an ocean biogeochemical model, we show that the inclusion of variable C:N:P phytoplankton stoichiometry can substantially increase the trophic amplification of biomass declines in low latitude regions. This additional trophic amplification is driven by enhanced nutrient limitation decreasing phytoplankton N and P content relative to C, hence reducing zooplankton growth efficiency. Given that most current Earth System Models assume that phytoplankton C:N:P stoichiometry is constant, such models are likely to underestimate the extent of negative trophic amplification under projected climate change.

Global forest carbon uptake due to nitrogen and phosphorus deposition from 1850 to 2100.

Spatial patterns and temporal trends of nitrogen (N) and phosphorus (P) deposition are important for quantifying their impact on forest carbon (C) uptake. In a first step, we modeled historical and future change in the global distributions of the atmospheric deposition of N and P from the dry and wet deposition of aerosols and gases containing N and P. Future projections were compared between two scenarios with contrasting aerosol emissions. Modeled fields of N and P deposition and P concentration were evaluated using globally distributed in situ measurements. N deposition peaked around 1990 in European forests and around 2010 in East Asian forests, and both increased sevenfold relative to 1850. P deposition peaked around 2010 in South Asian forests and increased 3.5-fold relative to 1850. In a second step, we estimated the change in C storage in forests due to the fertilization by deposited N and P (∆Cν dep ), based on the retention of deposited nutrients, their allocation within plants, and C:N and C:P stoichiometry. ∆Cν dep for 1997-2013 was estimated to be 0.27 ± 0.13 Pg C year-1 from N and 0.054 ± 0.10 Pg C year-1 from P, contributing 9% and 2% of the terrestrial C sink, respectively. Sensitivity tests show that uncertainty of ∆Cν dep was larger from P than from N, mainly due to uncertainty in the fraction of deposited P that is fixed by soil. ∆CPdep was exceeded by ∆CNdep over 1960-2007 in a large area of East Asian and West European forests due to a faster growth in N deposition than P. Our results suggest a significant contribution of anthropogenic P deposition to C storage, and additional sources of N are needed to support C storage by P in some Asian tropical forests where the deposition rate increased even faster for P than for N.

Influence of anthropogenic aerosol deposition on the relationship between oceanic productivity and warming.

Satellite data and models suggest that oceanic productivity is reduced in response to less nutrient supply under warming. In contrast, anthropogenic aerosols provide nutrients and exert a fertilizing effect, but its contribution to evolution of oceanic productivity is unknown. We simulate the response of oceanic biogeochemistry to anthropogenic aerosols deposition under varying climate from 1850 to 2010. We find a positive response of observed chlorophyll to deposition of anthropogenic aerosols. Our results suggest that anthropogenic aerosols reduce the sensitivity of oceanic productivity to warming from -15.2 ± 1.8 to -13.3 ± 1.6 Pg C yr-1 °C-1 in global stratified oceans during 1948-2007. The reducing percentage over the North Atlantic, North Pacific, and Indian Oceans reaches 40, 24, and 25%, respectively. We hypothesize that inevitable reduction of aerosol emissions in response to higher air quality standards in the future might accelerate the decline of oceanic productivity per unit warming.