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.

Making spatial-temporal marine ecosystem modelling better - A perspective.

Marine Ecosystem Models (MEMs) provide a deeper understanding of marine ecosystem dynamics. The United Nations Decade of Ocean Science for Sustainable Development has highlighted the need to deploy these complex mechanistic spatial-temporal models to engage policy makers and society into dialogues towards sustainably managed oceans. From our shared perspective, MEMs remain underutilized because they still lack formal validation, calibration, and uncertainty quantifications that undermines their credibility and uptake in policy arenas. We explore why these shortcomings exist and how to enable the global modelling community to increase MEMs' usefulness. We identify a clear gap between proposed solutions to assess model skills, uncertainty, and confidence and their actual systematic deployment. We attribute this gap to an underlying factor that the ecosystem modelling literature largely ignores: technical issues. We conclude by proposing a conceptual solution that is cost-effective, scalable and simple, because complex spatial-temporal marine ecosystem modelling is already complicated enough.

Retrospective analysis of the pelagic ecosystem of the Western Mediterranean Sea: Drivers, changes and effects.

In the Western Mediterranean Sea, forage fishes have changed in abundance, body condition, growth, reproduction, and distribution in the last decades. Different hypotheses have been proposed to explain these changes, including increase in fishing mortality; changes in environmental conditions affecting species fitness, and planktonic productivity and quality; recovery of top predators; and increase in competitors. We investigated the main drivers and changes of the pelagic ecosystem and their effects using an ecosystem-based modelling approach. Specifically, we (1) quantified the potential historical contribution of various drivers of change, (2) investigated changes in temporal trends and spatial distributions of main ecosystem components, and (3) identified ecological consequences of these changes in top predator and competitors, their fisheries and ecosystem traits during 2000-2020. We updated an established Ecopath food-web model representing the Spanish and French Mediterranean sub-areas (GSA06 and GSA07) in 2000 with recent available data. We applied the temporal dynamic Ecosim module, and tested historical time series of fishing effort, fishing mortality and environmental factors as potential drivers. Observed biomass and landings of key species were used to validate model projections. A spatial-temporal Ecospace model was developed to project species distribution changes. Results showed historical biomass and catch changes driven by a combination of high fishing pressure and environmental change (i.e. increase in temperature and salinity, and decline in primary productivity). Small pelagic fish showed significant temporal changes and predicted shifts in their distributions, following a latitudinal gradient. Predators and competitors showed changes as well, displaying heterogeneous spatial patterns, while fisheries landings declined. Overall, results matched observations (e.g., decline of sardine, fluctuations of anchovy and increases in bluefin tuna) and illustrated the need to complement traditional assessments with integrative frameworks to move towards an ecosystem-based approach in the Mediterranean. They also highlighted important knowledge gaps to guide future research in the region.

Evaluating red tide effects on the West Florida Shelf using a spatiotemporal ecosystem modeling framework.

The West Florida Shelf (WFS), located in the eastern Gulf of Mexico, fosters high species richness and supports highly valuable fisheries. However, red tide events occur regularly that can impact fisheries resources as well as ecosystem state, functioning, and derived services. Therefore, it is important to evaluate and quantify the spatiotemporal impacts of red tides to improve population assessments, mitigate potential negative effects through management, and better understand disturbances to support an ecosystem-based management framework. To model red tide effects on the marine community, we used Ecospace, the spatiotemporal module of the ecosystem modeling framework Ecopath with Ecosim. The inclusion of both lethal and sublethal response functions to red tide and a comprehensive calibration procedure allowed to systematically evaluate red tide effects and increased the robustness of the model and management applicability. Our results suggest severe red tide impacts have occurred on the WFS at the ecosystem, community, and population levels in terms of biomass, catch, and productivity. Sublethal and indirect food-web effects of red tide triggered compensatory responses such as avoidance behavior and release from predation and/or competition.. This study represents a step forward to operationalize spatiotemporal ecosystem models for management purposes that may increase the ability of fisheries managers to respond more effectively and be more proactive to episodic mortality events, such as those caused by red tides.

Trophic amplification: A model intercomparison of climate driven changes in marine food webs.

Marine animal biomass is expected to decrease in the 21st century due to climate driven changes in ocean environmental conditions. Previous studies suggest that the magnitude of the decline in primary production on apex predators could be amplified through the trophodynamics of marine food webs, leading to larger decreases in the biomass of predators relative to the decrease in primary production, a mechanism called trophic amplification. We compared relative changes in producer and consumer biomass or production in the global ocean to assess the extent of trophic amplification. We used simulations from nine marine ecosystem models (MEMs) from the Fisheries and Marine Ecosystem Models Intercomparison Project forced by two Earth System Models under the high greenhouse gas emissions Shared Socioeconomic Pathways (SSP5-8.5) and a scenario of no fishing. Globally, total consumer biomass is projected to decrease by 16.7 ± 9.5% more than net primary production (NPP) by 2090-2099 relative to 1995-2014, with substantial variations among MEMs and regions. Total consumer biomass is projected to decrease almost everywhere in the ocean (80% of the world's oceans) in the model ensemble. In 40% of the world's oceans, consumer biomass was projected to decrease more than NPP. Additionally, in another 36% of the world's oceans consumer biomass is expected to decrease even as projected NPP increases. By analysing the biomass response within food webs in available MEMs, we found that model parameters and structures contributed to more complex responses than a consistent amplification of climate impacts of higher trophic levels. Our study provides additional insights into the ecological mechanisms that will impact marine ecosystems, thereby informing model and scenario development.

Overfishing species on the move may burden seafood provision in the low-latitude Atlantic Ocean.

Climate and fisheries interact, often synergistically, and may challenge marine ecosystem functioning and management, along with seafood provision. Here, we spatially combine highly resolved assessments of climate-driven changes in optimal environmental conditions (i.e., optimal habitats) for the pelagic fish community with available industrial fishery data to identify highly impacted inshore areas in the Central and Southern Atlantic Ocean. Overall, optimal habitat availability remained stable or decreased over recent decades for most commercial, small and medium size pelagic species, particularly in low-latitude regions. We also find a worrying overlap of these areas with fishing hotspots. Nations near the Equator (particularly along the African coast) have been doubly impacted by climate and industrial fisheries, with ultimate consequences on fish stocks and ecosystems as a whole. Management and conservation actions are urgently required to prevent species depletions and ensure seafood provisioning in these highly impacted, and often socioeconomically constrained areas. These actions may include redistributing fishing pressure and reducing it in local areas where climate forcing is particularly high, balancing resource exploitation and the conservation of marine life-supporting services in the face of climate change.

Modelling the Mediterranean Sea ecosystem at high spatial resolution to inform the ecosystem-based management in the region.

Cumulative pressures are rapidly expanding in the Mediterranean Sea with consequences for marine biodiversity and marine resources, and the services they provide. Policy makers urge for a marine ecosystem assessment of the region in space and time. This study evaluates how the whole Mediterranean food web may have responded to historical changes in the climate, environment and fisheries, through the use of an ecosystem modelling over a long time span (decades) at high spatial resolution (8 × 8 km), to inform regional and sub-regional management. Results indicate coastal and shelf areas to be the sites with highest marine biodiversity and marine resources biomass, which decrease towards the south-eastern regions. High levels of total catches and discards are predicted to be concentrated in the Western sub-basin and the Adriatic Sea. Mean spatial-temporal changes of total and commercial biomass show increases in offshore waters of the region, while biodiversity indicators show marginal changes. Total catches and discards increase greatly in offshore waters of the Western and Eastern sub-basins. Spatial patterns and temporal mean changes of marine biodiversity, community biomasses and trophic indices, assessed in this study, aim at identifying areas and food web components that show signs of deterioration with the overall goal of assisting policy makers in designing and implementing spatial management actions for the region.

SOS small pelagics: A safe operating space for small pelagic fish in the western Mediterranean Sea.

Sustainable fishing practices must ensure human wellbeing by safeguarding the integrity of marine life-supporting systems. Unfortunately, a significant challenge to fisheries management is that sustainable fishing levels can decline, often synergistically, by co-occurring with climate-driven environmental stressors. Within one of the most impacted marine areas in the world, and encompassing a number of highly targeted commercial species, the small pelagic fish community of the western Mediterranean Sea has recently shown signs of collapse. In this study, we identify a worrying coincidence where fishing hotspots for the commercially valuable European sardine Sardina pilchardus and anchovy Engraulis encrasicolus occur in marine areas mostly affected by climate change. To identify these areas, we overlayed detailed, spatially explicit measurements of fishing pressure with the finest-scale maps of cumulative climate change impacts onto these species. According to our results, doubly impacted marine areas largely occur in the north-western Mediterranean Sea, with climate and fisheries mostly affecting European sardine. Reducing local stressors (i.e., fishing pressure) in highly impacted areas may contribute to maintain these communities within a "safe operating space" (SOS), where they remain resilient to climate change. Accordingly, the redistribution and/or reduction of fishing intensity may alleviate pressure in those areas already affected by climate change. Sustainable fishing strategies may benefit, therefore, from the SOS concept and the spatial assessments provided in this study.

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.

State-of-the-art global models underestimate impacts from climate extremes.

Global impact models represent process-level understanding of how natural and human systems may be affected by climate change. Their projections are used in integrated assessments of climate change. Here we test, for the first time, systematically across many important systems, how well such impact models capture the impacts of extreme climate conditions. Using the 2003 European heat wave and drought as a historical analogue for comparable events in the future, we find that a majority of models underestimate the extremeness of impacts in important sectors such as agriculture, terrestrial ecosystems, and heat-related human mortality, while impacts on water resources and hydropower are overestimated in some river basins; and the spread across models is often large. This has important implications for economic assessments of climate change impacts that rely on these models. It also means that societal risks from future extreme events may be greater than previously thought.

A risk-based approach to cumulative effect assessments for marine management.

Marine ecosystems are increasingly threatened by the cumulative effects of multiple human pressures. Cumulative effect assessments (CEAs) are needed to inform environmental policy and guide ecosystem-based management. Yet, CEAs are inherently complex and seldom linked to real-world management processes. Therefore we propose entrenching CEAs in a risk management process, comprising the steps of risk identification, risk analysis and risk evaluation. We provide guidance to operationalize a risk-based approach to CEAs by describing for each step guiding principles and desired outcomes, scientific challenges and practical solutions. We reviewed the treatment of uncertainty in CEAs and the contribution of different tools and data sources to the implementation of a risk based approach to CEAs. We show that a risk-based approach to CEAs decreases complexity, allows for the transparent treatment of uncertainty and streamlines the uptake of scientific outcomes into the science-policy interface. Hence, its adoption can help bridging the gap between science and decision-making in ecosystem-based management.

Historical changes of the Mediterranean Sea ecosystem: modelling the role and impact of primary productivity and fisheries changes over time.

The Mediterranean Sea has been defined "under siege" because of intense pressures from multiple human activities; yet there is still insufficient information on the cumulative impact of these stressors on the ecosystem and its resources. We evaluate how the historical (1950-2011) trends of various ecosystems groups/species have been impacted by changes in primary productivity (PP) combined with fishing pressure. We investigate the whole Mediterranean Sea using a food web modelling approach. Results indicate that both changes in PP and fishing pressure played an important role in driving species dynamics. Yet, PP was the strongest driver upon the Mediterranean Sea ecosystem. This highlights the importance of bottom-up processes in controlling the biological characteristics of the region. We observe a reduction in abundance of important fish species (~34%, including commercial and non-commercial) and top predators (~41%), and increases of the organisms at the bottom of the food web (~23%). Ecological indicators, such as community biomass, trophic levels, catch and diversity indicators, reflect such changes and show overall ecosystem degradation over time. Since climate change and fishing pressure are expected to intensify in the Mediterranean Sea, this study constitutes a baseline reference for stepping forward in assessing the future management of the basin.

The biodiversity of the Mediterranean Sea: estimates, patterns, and threats.

The Mediterranean Sea is a marine biodiversity hot spot. Here we combined an extensive literature analysis with expert opinions to update publicly available estimates of major taxa in this marine ecosystem and to revise and update several species lists. We also assessed overall spatial and temporal patterns of species diversity and identified major changes and threats. Our results listed approximately 17,000 marine species occurring in the Mediterranean Sea. However, our estimates of marine diversity are still incomplete as yet-undescribed species will be added in the future. Diversity for microbes is substantially underestimated, and the deep-sea areas and portions of the southern and eastern region are still poorly known. In addition, the invasion of alien species is a crucial factor that will continue to change the biodiversity of the Mediterranean, mainly in its eastern basin that can spread rapidly northwards and westwards due to the warming of the Mediterranean Sea. Spatial patterns showed a general decrease in biodiversity from northwestern to southeastern regions following a gradient of production, with some exceptions and caution due to gaps in our knowledge of the biota along the southern and eastern rims. Biodiversity was also generally higher in coastal areas and continental shelves, and decreases with depth. Temporal trends indicated that overexploitation and habitat loss have been the main human drivers of historical changes in biodiversity. At present, habitat loss and degradation, followed by fishing impacts, pollution, climate change, eutrophication, and the establishment of alien species are the most important threats and affect the greatest number of taxonomic groups. All these impacts are expected to grow in importance in the future, especially climate change and habitat degradation. The spatial identification of hot spots highlighted the ecological importance of most of the western Mediterranean shelves (and in particular, the Strait of Gibraltar and the adjacent Alboran Sea), western African coast, the Adriatic, and the Aegean Sea, which show high concentrations of endangered, threatened, or vulnerable species. The Levantine Basin, severely impacted by the invasion of species, is endangered as well. This abstract has been translated to other languages (File S1).