Sea temperature is the primary driver of recent and predicted fish community structure across Northeast Atlantic shelf seas.

Climate change has strongly influenced the distribution and abundance of marine fish species, leading to concern about effects of future climate on commercially harvested stocks. Understanding the key drivers of large-scale spatial variation across present-day marine assemblages enables predictions of future change. Here we present a unique analysis of standardised abundance data for 198 marine fish species from across the Northeast Atlantic collected by 23 surveys and 31,502 sampling events between 2005 and 2018. Our analyses of the spatially comprehensive standardised data identified temperature as the key driver of fish community structure across the region, followed by salinity and depth. We employed these key environmental variables to model how climate change will affect both the distributions of individual species and local community structure for the years 2050 and 2100 under multiple emissions scenarios. Our results consistently indicate that projected climate change will lead to shifts in species communities across the entire region. Overall, the greatest community-level changes are predicted at locations with greater warming, with the most pronounced effects at higher latitudes. Based on these results, we suggest that future climate-driven warming will lead to widespread changes in opportunities for commercial fisheries across the region.

Growth portfolios buffer climate-linked environmental change in marine systems.

Large-scale, climate-induced synchrony in the productivity of fish populations is becoming more pronounced in the world's oceans. As synchrony increases, a population's "portfolio" of responses can be diminished, in turn reducing its resilience to strong perturbation. Here we argue that the costs and benefits of trait synchronization, such as the expression of growth rate, are context dependent. Contrary to prevailing views, synchrony among individuals could actually be beneficial for populations if growth synchrony increases during favorable conditions, and then declines under poor conditions when a broader portfolio of responses could be useful. Importantly, growth synchrony among individuals within populations has seldom been measured, despite well-documented evidence of synchrony across populations. Here, we used century-scale time series of annual otolith growth to test for changes in growth synchronization among individuals within multiple populations of a marine keystone species (Atlantic cod, Gadus morhua). On the basis of 74,662 annual growth increments recorded in 13,749 otoliths, we detected a rising conformity in long-term growth rates within five northeast Atlantic cod populations in response to both favorable growth conditions and a large-scale, multidecadal mode of climate variability similar to the East Atlantic Pattern. The within-population synchrony was distinct from the across-population synchrony commonly reported for large-scale environmental drivers. Climate-linked, among-individual growth synchrony was also identified in other Northeast Atlantic pelagic, deep-sea and bivalve species. We hypothesize that growth synchrony in good years and growth asynchrony in poorer years reflects adaptive trait optimization and bet hedging, respectively, that could confer an unexpected, but pervasive and stabilizing, impact on marine population productivity in response to large-scale environmental change.

Tradeoffs of managing cod as a sustainable resource in fluctuating environments.

Sustainable human exploitation of living marine resources stems from a delicate balance between yield stability and population persistence to achieve socioeconomic and conservation goals. But our imperfect knowledge of how oceanic oscillations regulate temporal variation in an exploited species can obscure the risk of missing management targets. We illustrate how applying a management policy to suppress fluctuations in fishery yield in variable environments (prey density and regional climate) can present unintended outcomes in harvested predators and the sustainability of harvesting. Using Atlantic cod (Gadus morhua, an apex predatory fish) in the Barents Sea as a case study we simulate age-structured population and harvest dynamics through time-varying, density-dependent and density-independent processes with a stochastic, process-based model informed by 27-year monitoring data. In this model, capelin (Mallotus villosus, a pelagic forage fish), a primary prey of cod, fluctuations modulate the strength of density-dependent regulation primarily through cannibalistic pressure on juvenile cod survival; sea temperature fluctuations modulate thermal regulation of cod feeding, growth, maturation, and reproduction. We first explore how capelin and temperature fluctuations filtered through cod intrinsic dynamics modify catch stability and then evaluate how management to suppress short-term variability in catch targets alters overharvest risk. Analyses revealed that suppressing year-to-year catch variability impedes management responses to adjust fishing pressure, which becomes progressively out of sync with variations in cod abundance. This asynchrony becomes amplified in fluctuating environments, magnifying the amplitudes of both fishing pressure and cod abundance and then intensifying the density-dependent regulation of juvenile survival through cannibalism. Although these transient dynamics theoretically give higher average catches, emergent, quasicyclic behaviors of the population would increase long-term yield variability and elevate overharvest risk. Management strategies that overlook the interplay of extrinsic (fishing and environment) and intrinsic (life history and demography) fluctuations thus can inadvertently destabilize fish stocks, thereby jeopardizing the sustainability of harvesting. These policy implications underscore the value of ecosystem approaches to designing management measures to sustainably harvest ecologically connected resources while achieving socioeconomic security.

Climate based multi-year predictions of the Barents Sea cod stock.

Predicting fish stock variations on interannual to decadal time scales is one of the major issues in fisheries science and management. Although the field of marine ecological predictions is still in its infancy, it is understood that a major source of multi-year predictability resides in the ocean. Here we show the first highly skilful long-term predictions of the commercially valuable Barents Sea cod stock. The 7-year predictions are based on the propagation of ocean temperature anomalies from the subpolar North Atlantic toward the Barents Sea, and the strong co-variability between these temperature anomalies and the cod stock. Retrospective predictions for the period 1957-2017 capture well multi-year to decadal variations in cod stock biomass, with cross-validated explained variance of over 60%. For lead times longer than one year the statistical long-term predictions show more skill than operational short-term predictions used in fisheries management and lagged persistence forecasts. Our results thus demonstrate the potential for ecosystem-based fisheries management, which could enable strategic planning on longer time scales. Future predictions show a gradual decline in the cod stock towards 2024.

Predator-prey interactions cause apparent competition between marine zooplankton groups.

Predator-mediated apparent competition is an indirect negative interaction between two prey species mediated by a shared predator. Quantifying such indirect ecosystem effects is methodologically challenging but important for understanding ecosystem functioning. Still, there are few examples of apparent competition from pelagic marine environments. Using state-space statistical modeling, we here provide evidence for apparent competition between two dominant zooplankton groups in a large marine ecosystem, i.e., krill and copepods in the Barents Sea. This effect is mediated by a positive association between krill biomass and survival of the main planktivorous fish in the Barents Sea, capelin Mallotus villosus, and a negative association between capelin and copepod biomasses. The biomass of Atlantic krill species is expected to increase in the Barents Sea due to ongoing climate change, thereby potentially negatively affecting copepods through apparent competition. By demonstrating and quantifying apparent competition in a large marine ecosystem, our study paves the way for more realistic projections of indirect ecosystem effects of climate change and harvesting.

Effect of a fish stock's demographic structure on offspring survival and sensitivity to climate.

Commercial fishing generally removes large and old individuals from fish stocks, reducing mean age and age diversity among spawners. It is feared that these demographic changes lead to lower and more variable recruitment to the stocks. A key proposed pathway is that juvenation and reduced size distribution causes reduced ranges in spawning period, spawning location, and egg buoyancy; this is proposed to lead to reduced spatial distribution of fish eggs and larvae, more homogeneous ambient environmental conditions within each year-class, and reduced buffering against negative environmental influences. However, few, if any, studies have confirmed a causal link from spawning stock demographic structure through egg and larval distribution to year class strength at recruitment. We here show that high mean age and size in the spawning stock of Barents Sea cod (Gadus morhua) is positively associated with high abundance and wide spatiotemporal distribution of cod eggs. We find, however, no support for the hypothesis that a wide egg distribution leads to higher recruitment or a weaker recruitment-temperature correlation. These results are based on statistical analyses of a spatially resolved data set on cod eggs covering a period (1959-1993) with large changes in biomass and demographic structure of spawners. The analyses also account for significant effects of spawning stock biomass and a liver condition index on egg abundance and distribution. Our results suggest that the buffering effect of a geographically wide distribution of eggs and larvae on fish recruitment may be insignificant compared with other impacts.

Synergies between climate and management for Atlantic cod fisheries at high latitudes.

The widespread depletion of commercially exploited marine living resources is often seen as a general failure of management and results in criticism of contemporary management procedures. When populations show dramatic and positive changes in population size, this invariably leads to questions about whether favorable climatic conditions or good management (or both) were responsible. The Barents Sea cod (Gadus morhua) stock has recently increased markedly and the spawning stock biomass is now at an unprecedented high. We identify the crucial social and environmental factors that made this unique growth possible. The relationship between vital rates of Barents Sea cod stock productivity (recruitment, growth, and mortality) and environment is investigated, followed by simulations of population size under different management scenarios. We show that the recent sustained reduction in fishing mortality, facilitated by the implementation of a "harvest control rule," was essential to the increase in population size. Simulations show that a drastic reduction in fishing mortality has resulted in a doubling of the total population biomass compared with that expected under the former management regime. However, management alone was not solely responsible. We document that prevailing climate, operating through several mechanistic links, positively reinforced management actions. Heightened temperature resulted in an increase in the extent of the suitable feeding area for Barents Sea cod, likely offering a release from density-dependent effects (for example, food competition and cannibalism) through prolonged overlap with prey and improved adult stock productivity. Management and climate may thus interact to give a positive outlook for exploited high-latitude marine resources.

Spatial anatomy of species survival: effects of predation and climate-driven environmental variability.

The majority of survival analyses focus on temporal scales. Consequently, there is a limited understanding of how species survival varies over space and, ultimately, how spatial variability in the environment affects the temporal dynamics of species abundance. Using data from the Barents Sea, we study the spatiotemporal variability of the juvenile Atlantic cod (Gadus morhua) survival. We develop an index of spatial survival based on changes of juvenile cod distribution through their first winter of life (from age-0 to age-1) and study its variability in relation to biotic and abiotic factors. Over the 25 years analyzed (1980-2004), we found that, once the effect of passive drift due to dominant currents is accounted for, the area where age-0 cod survival was lowest coincided with the area of highest abundance of older cod. Within this critical region, the survival of age-0 cod was negatively affected by its own abundance, by that of older cod, and by bottom depth. Furthermore, during cold years, age-0 cod survival increased in the eastern and coldest portion of the examined area, which was typically avoided by older conspecifics. Based on these results we propose that within the examined area top-down mechanisms and predation-driven density dependence can strongly affect the spatial pattern of age-0 cod survival. Climate-related variables can also influence the spatial survival of age-0 cod by affecting their distribution and that of their predators. Results from these and similar studies, focusing on the spatial variability of survival rates, can be used to characterize species habitat quality of marine renewable resources.

Food web dynamics affect Northeast Arctic cod recruitment.

Proper management of ecosystems requires an understanding of both the species interactions as well as the effect of climate variation. However, a common problem is that the available time-series are of different lengths. Here, we present a general approach for studying the dynamic structure of such interactions. Specifically, we analyse the recruitment of the world's largest cod stock, the Northeast Arctic cod. Studies based on data starting in the 1970-1980s indicate that this stock is affected by temperature through a variety of pathways. However, the value of such studies is somewhat limited by the fact that they are based on a quite specific ecological and climatic situation. Recently, this stock has consisted of fairly young fish and the spawning stock has consisted of relatively few age groups. In this study, we develop a model for the effect of capelin (the cod's main prey) and herring on cod recruitment since 1973. Based on this model, we analyse data on cod, herring and temperature going back to 1921 and find that food-web effects explain a significant part of the cod recruitment variation back to around 1950.