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.

Long-term variability in spawning stock age structure influences climate-recruitment link for Barents Sea cod.

Fish populations may spawn a vast number of offspring, while only a small and highly variable fraction of a new cohort survives long enough to enter into the fisheries as recruits. It is intuitive that the size and state of the spawning stock, the adult part of the fish population, is important for recruitment. Additionally, environmental conditions can greatly influence survival through vulnerable early life stages until recruitment. To understand what regulates recruitment, an essential part of fish population dynamics, it is thus necessary to explain the impact of fluctuations in both spawning stock and environment, including interactions. Here, we examine if the connection between the environment and recruitment is affected by the state of the spawning stock, including biomass, mean age and age diversity. Specifically, we re-evaluate the hypothesis stating that recruitment from a spawning stock dominated by young fish and few age classes is more vulnerable to environmental fluctuations. We expand upon earlier work on the Barents Sea stock of Atlantic cod, now with data series extended in time both backwards and forwards to cover the period 1922-2019. While our findings are correlative and cannot prove a specific cause and effect mechanism, they support earlier work and strengthen the evidence for the hypothesis above. Furthermore, this study supports that advice to fisheries management should include considerations of environmental status.

Spawning stock and recruitment in North Sea cod shaped by food and climate.

In order to provide better fisheries management and conservation decisions, there is a need to discern the underlying relationship between the spawning stock and recruitment of marine fishes, a relationship which is influenced by the environmental conditions. Here, we demonstrate how the environmental conditions (temperature and the food availability for fish larvae) influence the stock-recruitment relationship and indeed what kind of stock-recruitment relationship we might see under different environmental conditions. Using unique zooplankton data from the Continuous Plankton Recorder, we find that food availability (i.e. zooplankton) in essence determines which model applies for the once large North Sea cod (Gadus morhua) stock. Further, we show that recruitment is strengthened during cold years and weakened during warm years. Our combined model explained 45 per cent of the total variance in cod recruitment, while the traditional Ricker and Beverton-Holt models only explained about 10 per cent. Specifically, our approach predicts that a full recovery of the North Sea cod stock might not be expected until the environment becomes more favourable.

Direct and indirect climate forcing in a multi-species marine system.

Interactions within and between species complicate quantification of climate effects, by causing indirect, often delayed, effects of climate fluctuations and compensation of mortality. Here we identify direct and indirect climate effects by analysing unique Russian time-series data from the Norwegian Sea-Barents Sea ecosystem on the first life stages of cod, capelin, herring and haddock, their predators, competitors and zooplanktonic prey. By analysing growth and survival from one life stage to the next (eggs-larvae-juveniles-recruits), we find evidence for both bottom-up, direct and top-down effects of climate. Ambient zooplankton biomass predicts survival of all species, whereas ambient temperature mainly affects survival through effects on growth. In warm years, all species experienced improved growth and feeding conditions. Cohorts born following a warm year will, however, experience increased predation and competition because of increased densities of subadult cod and herring, leading to delayed climate effects. While climate thus affects early growth and survival through several mechanisms, only some of the identified mechanisms were found to be significant predictors of population growth. In particular, our findings exemplify that climate impacts are barely propagated to later life stages when density dependence is strong.

Contrasting effects of rising temperatures on trophic interactions in marine ecosystems.

In high-latitude marine environments, primary producers and their consumers show seasonal peaks of abundance in response to annual light cycle, water column stability and nutrient availability. Predatory species have adapted to this pattern by synchronising life-history events such as reproduction with prey availability. However, changing temperatures may pose unprecedented challenges by decoupling the predator-prey interactions. Here we build a predator-prey model accounting for the full life-cycle of fish and zooplankton including their phenology. The model assumes that fish production is bottom-up controlled by zooplankton prey abundance and match or mismatch between predator and prey phenology, and is parameterised based on empirical findings of how climate influences phenology and prey abundance. With this model, we project possible climate-warming effects on match-mismatch dynamics in Arcto-boreal and temperate biomes. We find a strong dependence on synchrony with zooplankton prey in the Arcto-boreal fish population, pointing towards a possible pronounced population decline with warming because of frequent desynchronization with its zooplankton prey. In contrast, the temperate fish population appears better able to track changes in prey timing and hence avoid strong population decline. These results underline that climate change may enhance the risks of predator-prey seasonal asynchrony and fish population declines at higher latitudes.

Ice-age survival of Atlantic cod: agreement between palaeoecology models and genetics.

Scant scientific attention has been given to the abundance and distribution of marine biota in the face of the lower sea level, and steeper latitudinal gradient in climate, during the ice-age conditions that have dominated the past million years. Here we examine the glacial persistence of Atlantic cod (Gadus morhua) populations using two ecological-niche-models (ENM) and the first broad synthesis of multi-locus gene sequence data for this species. One ENM uses a maximum entropy approach (Maxent ); the other is a new ENM for Atlantic cod, using ecophysiological parameters based on observed reproductive events rather than adult distribution. Both the ENMs were tested for present-day conditions, then used to hindcast ranges at the last glacial maximum (LGM) ca 21kyr ago, employing climate model data. Although the LGM range of Atlantic cod was much smaller, and fragmented, both the ENMs agreed that populations should have been able to persist in suitable habitat on both sides of the Atlantic. The genetic results showed a degree of trans-Atlantic divergence consistent with genealogically continuous populations on both sides of the North Atlantic since long before the LGM, confirming the ENM results. In contrast, both the ENMs and the genetic data suggest that the Greenland G. morhua population post-dates the LGM.

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.

Combined effects of fishing and oil spills on marine fish: Role of stock demographic structure for offspring overlap with oil.

It has been proposed that the multiple pressures of fishing and petroleum activities impact fish stocks in synergy, as fishing-induced demographic changes in a stock may lead to increased sensitivity to detrimental effects of acute oil spills. High fishing pressure may erode the demographic structure of fish stocks, lead to less diverse spawning strategies, and more concentrated distributions of offspring in space and time. Hence an oil spill may potentially hit a larger fraction of a year-class of offspring. Such a link between demographic structure and egg distribution was recently demonstrated for the Northeast Arctic stock of Atlantic cod for years 1959-1993. We here estimate that this variation translates into a two-fold variation in the maximal proportion of cod eggs potentially exposed to a large oil spill. With this information it is possible to quantitatively account for demographic structure in prospective studies of population effects of possible oil spills.

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.

Density dependence and density independence during the early life stages of four marine fish stocks.

Recruitment variability caused by density-dependent and density-independent processes is an important area within the study of fish dynamics. These processes can exhibit nonlinearities and nonadditive properties that may have profound dynamic effects. We investigate the importance of population density (i.e., density dependence) and environmental forcing (i.e., density independence) on the age-0 and age-1 abundance of capelin (Mallotus villosus), northeast Arctic cod (Gadus morhua), northeast Arctic haddock (Melanogrammus aeglefinus), and Norwegian spring spawning herring (Clupea harengus) in the Barents Sea. We use statistical methods that explicitly account for nonlinearities and nonadditive interactions between internal and external variables in the abundance of these two pre-recruitment stages. Our results indicate that, during their first five months of life, cod, haddock, and herring experience higher density-dependent survival than capelin. The abundance of age-0 cod depends on the mean age and biomass of the spawning stock, a result which has implications for the management of the entire cod stock. Temperature is another important factor influencing the abundance at age-0 and age-1 of all four species, except herring at age-1. Between age-0 and age-1, there is an attenuation of density-dependent survival for cod and herring, while haddock and capelin experience density dependence at high and low temperatures, respectively. Predation by subadult cod is important for both capelin and cod at age-1. We found strong indications for interactions among the studied species, pointing to the importance of viewing the problem of species recruitment variability as a community, rather than as a population phenomenon.

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.

Seasonal plankton dynamics along a cross-shelf gradient.

Much interest has recently been devoted to reconstructing the dynamic structure of ecological systems on the basis of time-series data. Using 10 years of monthly data on phyto- and zooplankton abundance from the Bay of Biscay (coastal to shelf-break sites), we demonstrate that the interaction between these two plankton components is approximately linear, whereas the effects of environmental factors (nutrients, temperature, upwelling and photoperiod) on these two plankton population growth rates are nonlinear. With the inclusion of the environmental factors, the main observed seasonal and inter-annual dynamic patterns within the studied plankton assemblage also indicate the prevalence of bottom-up regulatory control.

Review article. Studying climate effects on ecology through the use of climate indices: the North Atlantic Oscillation, El Niño Southern Oscillation and beyond.

Whereas the El Niño Southern Oscillation (ENSO) affects weather and climate variability worldwide, the North Atlantic Oscillation (NAO) represents the dominant climate pattern in the North Atlantic region. Both climate systems have been demonstrated to considerably influence ecological processes. Several other large-scale climate patterns also exist. Although less well known outside the field of climatology, these patterns are also likely to be of ecological interest. We provide an overview of these climate patterns within the context of the ecological effects of climate variability. The application of climate indices by definition reduces complex space and time variability into simple measures, 'packages of weather'. The disadvantages of using global climate indices are all related to the fact that another level of problems are added to the ecology-climate interface, namely the link between global climate indices and local climate. We identify issues related to: (i) spatial variation; (ii) seasonality; (iii) non-stationarity; (iv) nonlinearity; and (v) lack of correlation in the relationship between global and local climate. The main advantages of using global climate indices are: (i) biological effects may be related more strongly to global indices than to any single local climate variable; (ii) it helps to avoid problems of model selection; (iii) it opens the possibility for ecologists to make predictions; and (iv) they are typically readily available on Internet.

Ecological effects of the North Atlantic Oscillation.

Climatic oscillations as reflected in atmospheric modes such as the North Atlantic Oscillation (NAO) may be seen as a proxy for regulating forces in aquatic and terrestrial ecosystems. Our review highlights the variety of climate processes related to the NAO and the diversity in the type of ecological responses that different biological groups can display. Available evidence suggests that the NAO influences ecological dynamics in both marine and terrestrial systems, and its effects may be seen in variation at the individual, population and community levels. The ecological responses to the NAO encompass changes in timing of reproduction, population dynamics, abundance, spatial distribution and interspecific relationships such as competition and predator-prey relationships. This indicates that local responses to large-scale changes may be more subtle than previously suggested. We propose that the NAO effects may be classified as three types: direct, indirect and integrated. Such a classification will help the design and interpretation of analyses attempting to relate ecological changes to the NAO and, possibly, to climate in general.

Productivity in the barents sea--response to recent climate variability.

The temporal and spatial dynamics of primary and secondary biomass/production in the Barents Sea since the late 1990s are examined using remote sensing data, observations and a coupled physical-biological model. Field observations of mesozooplankton biomass, and chlorophyll a data from transects (different seasons) and large-scale surveys (autumn) were used for validation of the remote sensing products and modeling results. The validation showed that satellite data are well suited to study temporal and spatial dynamics of chlorophyll a in the Barents Sea and that the model is an essential tool for secondary production estimates. Temperature, open water area, chlorophyll a, and zooplankton biomass show large interannual variations in the Barents Sea. The climatic variability is strongest in the northern and eastern parts. The moderate increase in net primary production evident in this study is likely an ecosystem response to changes in climate during the same period. Increased open water area and duration of open water season, which are related to elevated temperatures, appear to be the key drivers of the changes in annual net primary production that has occurred in the northern and eastern areas of this ecosystem. The temporal and spatial variability in zooplankton biomass appears to be controlled largely by predation pressure. In the southeastern Barents Sea, statistically significant linkages were observed between chlorophyll a and zooplankton biomass, as well as between net primary production and fish biomass, indicating bottom-up trophic interactions in this region.

Environmental toxicology: population modeling of cod larvae shows high sensitivity to loss of zooplankton prey.

Two factors determine whether pollution is likely to affect a population indirectly through loss of prey: firstly, the sensitivity of the prey to the pollutants, and secondly, the sensitivity of the predator population to loss of prey at the given life stage. We here apply a statistical recruitment model for Northeast Arctic cod to evaluate the sensitivity of cod cohorts to loss of zooplankton prey, for example following an oil spill. The calculations show that cod cohorts are highly sensitive to possible zooplankton biomass reductions in the distribution area of the cod larvae, and point to a need for more knowledge about oil-effects on zooplankton. Our study illustrates how knowledge about population dynamics may guide which indirect effects to consider in environmental impact studies.

Competition among fishermen and fish causes the collapse of Barents Sea capelin.

The vast majority of the world's fisheries are typically managed within a single-species perspective, ignoring the dynamic feedback mechanisms generated by the ecological web of which they are a part. Here we show that the dynamics of the Barents Sea capelin (Mallotus villosus), the world's largest stock of this species, is strongly influenced by both within-system ecological feedback mechanisms and the impact of harvesting. Both overexploitation and predation by herring (Clupea harengus) can cause the population to collapse, whereas predation by cod (Gadus morhua) is demonstrated a delay in the stock's recovery after a collapse. Such collapses, which have occurred twice in 20 years, affect the entire Barents Sea ecosystem, a region that for ages has provided food for all of Europe.

Ecological effects of climate fluctuations.

Climate influences a variety of ecological processes. These effects operate through local weather parameters such as temperature, wind, rain, snow, and ocean currents, as well as interactions among these. In the temperate zone, local variations in weather are often coupled over large geographic areas through the transient behavior of atmospheric planetary-scale waves. These variations drive temporally and spatially averaged exchanges of heat, momentum, and water vapor that ultimately determine growth, recruitment, and migration patterns. Recently, there have been several studies of the impact of large-scale climatic forcing on ecological systems. We review how two of the best-known climate phenomena-the North Atlantic Oscillation and the El Niño-Southern Oscillation-affect ecological patterns and processes in both marine and terrestrial systems.