Non-stationary climate-salmon relationships in the Gulf of Alaska.

Studies of climate effects on ecology often account for non-stationarity in individual physical and biological variables, but rarely allow for non-stationary relationships among variables. Here, we show that non-stationary relationships among physical and biological variables are central to understanding climate effects on salmon (Onchorynchus spp.) in the Gulf of Alaska during 1965-2012. The relative importance of two leading patterns in North Pacific climate, the Pacific Decadal Oscillation (PDO) and North Pacific Gyre Oscillation (NPGO), changed around 1988/1989 as reflected by changing correlations with leading axes of sea surface temperature variability. Simultaneously, relationships between the PDO and Gulf of Alaska environmental variables weakened, and long-standing temperature-salmon and PDO-salmon covariance declined to zero. We propose a mechanistic explanation for changing climate-salmon relationships in terms of non-stationary atmosphere-ocean interactions coinciding with changing PDO-NPGO relative importance. We also show that regression models assuming stationary climate-salmon relationships are inappropriate over the multidecadal time scale we consider. Relaxing assumptions of stationary relationships markedly improved modelling of climate effects on salmon catches and productivity. Attempts to understand the implications of changing climate patterns in other ecosystems might also be aided by the application of models that allow associations among environmental and biological variables to change over time.

Pelagic marine refugia and climatically sensitive areas in an eastern boundary current upwelling system.

Refugia are areas relatively buffered from contemporary climate change that enable the persistence of valued physical, ecological, or sociocultural resources. Spatially identifying refugia is important for conservation and applied management. Yet the concept of refugia has not been broadly extended to marine ecosystems. Here, we analyze data from a unique and long-term (1999-2015) standardized survey of pelagic marine and anadromous species off Oregon and Washington in the northern California Current to identify such refugia. We use quantitative approaches to assess locations with high species richness and community persistence relative to local and basin-scale environmental fluctuations. We have identified a potential climate change refugial zone along the continental shelf of Washington State in the Northeastern Pacific Ocean, characterized by a species-rich community with low interannual temporal community change. This region contrasts with adjacent areas to the south and offshore that have lower species richness, and higher temporal species community change. Also, using spatially variant generalized additive mixed models, we identify areas with species compositions that are more influenced by basin-scale climatic fluctuations than others. We propose that upwelling regions with retentive topographic features, such as wide continental shelves, can function as marine refugia for pelagic fauna, whereas offshore locations are potentially more climatically sensitive and experience high temporal change in species composition. Further identification of these marine refugia using in situ data for pelagic biodiversity and climatically sensitive areas can help guide management in the face of inevitable climatically driven change.

Eight decades of sampling reveal a contemporary novel fish assemblage in coastal nursery habitats.

In order to adequately monitor biodiversity trends through time and their responses to natural or anthropogenic impacts, researchers require long time series that are often unavailable. This general lack of datasets that are several decades or longer makes establishing a background or baseline of diversity metrics difficult - especially when attempting to understand species composition changes against a backdrop of climate and ecological variability. Here, we present an analysis of a community of juvenile nearshore fishes based on nearly 8 decades of highly standardized Norwegian survey records. Using multivariate statistical techniques, we: (i) characterize the change in taxonomic community composition through time, (ii) determine whether there has been an increase in warm-water affinity species relative to their cold water affinity counterparts, and (iii) characterize the temporal change in the species' functional trait assemblage. Our results strongly indicate a shift toward a novel fish assemblage between the late 1990s and 2000s. The context of changes within the most recent two decades is in stark contrast to those during the 1960s and 1970s, but similar to those during the previous warm period during the 1930s and 1940s. This novel assemblage is tightly linked to the warming temperatures in the region portrayed by the increased presence of warm-water species and a higher incidence of pelagic, planktivorous species. The results indicate a clear influence of ocean temperature on the region's juvenile fish community that points to climate-mediated effects on the species assemblages of an important fish nursery area.

Non-additive and non-stationary properties in the spatial distribution of a large marine fish population.

Density-independent and density-dependent variables both affect the spatial distributions of species. However, their effects are often separately addressed using different analytical techniques. We apply a spatially explicit regression framework that incorporates localized, interactive and threshold effects of both density-independent (water temperature) and density-dependent (population abundance) variables, to study the spatial distribution of a well-monitored flatfish population in the eastern Bering Sea. Results indicate that when population biomass was beyond a threshold a further increase in biomass-promoted habitat expansion in a non-additive fashion with water temperature. In contrast, during years of low population size, habitat occupancy was affected positively only by water temperature. These results reveal the spatial signature of intraspecific abundance distribution relationships as well as the non-additive and non-stationary responses of species spatial dynamics. Furthermore, these results underscore the importance of implementing analytical techniques that can simultaneously account for density-dependent and density-independent sources of variability when studying geographical distribution patterns.

Functional responses and scaling in predator-prey interactions of marine fishes: contemporary issues and emerging concepts.

Predator-prey interactions are a primary structuring force vital to the resilience of marine communities and sustainability of the world's oceans. Human influences on marine ecosystems mediate changes in species interactions. This generality is evinced by the cascading effects of overharvesting top predators on the structure and function of marine ecosystems. It follows that ecological forecasting, ecosystem management, and marine spatial planning require a better understanding of food web relationships. Characterising and scaling predator-prey interactions for use in tactical and strategic tools (i.e. multi-species management and ecosystem models) are paramount in this effort. Here, we explore what issues are involved and must be considered to advance the use of predator-prey theory in the context of marine fisheries science. We address pertinent contemporary ecological issues including (1) the approaches and complexities of evaluating predator responses in marine systems; (2) the 'scaling up' of predator-prey interactions to the population, community, and ecosystem level; (3) the role of predator-prey theory in contemporary fisheries and ecosystem modelling approaches; and (4) directions for the future. Our intent is to point out needed research directions that will improve our understanding of predator-prey interactions in the context of the sustainable marine fisheries and ecosystem management.

Ontogenetic and sex-specific differences in density-dependent habitat selection of a marine fish population.

The spatial dynamics of species are the result of complex interactions between density-independent and density-dependent sources of variability. Disentangling these two sources of variability has challenged ecologists working in both terrestrial and aquatic ecosystems. Using a novel spatially explicit statistical model, we tested for the presence of density-independent and density-dependent habitat selection in yellowfin sole (Limanda aspera) in the eastern Bering Sea. We found specificities in the density-dependent processes operating across ontogeny and particularly with gender. Density-dependent habitat expansion occurred primarily in females, and to a lesser degree in males. These patterns were especially evident in adult stages, while juvenile stages of both sexes exhibited a mix of different dynamics. Association of yellowfin sole with substrate type also varied by sex and to a lesser degree with size, with large females distributed over a wider range of substrates than males. Moreover, yellowfin sole expanded northward as cold subsurface waters retracted in summer, suggesting high sensitivity to arctic warming. Our findings illustrate how marginal habitats can play an important role in buffering density-dependent habitat expansion, with direct implications for resource management. Our spatially explicit modeling approach is effective in evaluating density-dependent spatial dynamics, and can easily be used to test similar hypotheses from a variety of aquatic and terrestrial ecosystems.

Small-scale genetic structure in a marine population in relation to water circulation and egg characteristics.

Until the last decade it was assumed that most marine species have pronounced gene flow over vast areas, largely because of their potential for dispersal during early life stages. However, recent genetic, modeling, and field studies have shown that marine populations may be structured at scales that are inconsistent with extensive dispersal of eggs and larvae. Such findings have stimulated the birth of new studies explaining the mechanisms that promote population structure and isolation in the oceans, in the face of high potential for dispersal. Here we study the vertical and horizontal distribution of cod (Gadus morhua) eggs in relation to small-scale circulation and water column hydrography in a coastal location of southern Norway. Previous studies conducted in this region have shown that cod populations inhabiting fjord locations, which are on average 30 km apart, are genetically differentiated, a remarkable outcome considering that Atlantic cod have pelagic egg stages and long pelagic larval duration. We document that cod eggs are found in greater abundance in shallow water layers, which on average are flowing up the fjord (away from the open ocean), and in the inner portion of the fjord, which is subject to lower current speeds compared to the outer or mouth of the fjord. Eggs were found to be neutrally buoyant at shallow depths, a trait that also favors local retention, given the local circulation. The same patterns held during two environmentally contrasting years. These results strongly suggest that population structure of Atlantic cod is favored and maintained by a balance between water circulation and egg characteristics.

Comparative analysis of marine ecosystems: workshop on predator-prey interactions.

Climate and human influences on marine ecosystems are largely manifested by changes in predator-prey interactions. It follows that ecosystem-based management of the world's oceans requires a better understanding of food web relationships. An international workshop on predator-prey interactions in marine ecosystems was held at the Oregon State University, Corvallis, OR, USA on 16-18 March 2010. The meeting brought together scientists from diverse fields of expertise including theoretical ecology, animal behaviour, fish and seabird ecology, statistics, fisheries science and ecosystem modelling. The goals of the workshop were to critically examine the methods of scaling-up predator-prey interactions from local observations to systems, the role of shifting ecological processes with scale changes, and the complexity and organizational structure in trophic interactions.

Spatial, interannual, and generational sources of trait variability in a marine population.

Life-history traits of individuals in marine populations exhibit large sources of variability. In marine fish, variation of individual size at a given age has three main components: (1) spatial, correlated with the location in which individuals are caught, (2) temporal, correlated with the time when individuals are caught, and (3) generational, correlated with the year of birth of the examined individuals. These variations, if present, have practical implications for individual fitness as well as for sampling, survey design, and population assessment. Disentangling these variations and understanding their sources is hard, given the potentially correlated nature of their effects on individual traits. This study examines the size-at-age relationship of the Bering Sea Pacific cod, an economically and ecologically important groundfish. We used extensive records spanning 1994 to 2016 (inclusive) of 25,213 observations of both environmental variables and catch, lengths, and ages. We found that the average size of individuals of the same age could differ up to 7 cm. Notably, we found that the cohort composition of the sampled population explained >75% of the year effect and that individuals caught in the northwest and shallower portion of the sampling area were on average 5 cm smaller than individuals caught in the southern and deeper portion. We further found that northwest movement of young cod (age 1-5) as a result of warming places individuals in areas where we predict them to have smaller size at age. Smaller and less conditioned individuals are less fecund and may not be able to perform long migrations to return to their distant spawning grounds. Both the spatial distribution and water temperature experienced by Pacific cod in the Bering Sea are changing, and this study provides a mechanism for how these changes affect Pacific cod life-history traits and individual fitness.

A simulation framework for evaluating multi-stage sampling designs in populations with spatially structured traits.

Selecting an appropriate and efficient sampling strategy in biological surveys is a major concern in ecological research, particularly when the population abundance and individual traits of the sampled population are highly structured over space. Multi-stage sampling designs typically present sampling sites as primary units. However, to collect trait data, such as age or maturity, only a sub-sample of individuals collected in the sampling site is retained. Therefore, not only the sampling design, but also the sub-sampling strategy can have a major impact on important population estimates, commonly used as reference points for management and conservation. We developed a simulation framework to evaluate sub-sampling strategies from multi-stage biological surveys. Specifically, we compare quantitatively precision and bias of the population estimates obtained using two common but contrasting sub-sampling strategies: the random and the stratified designs. The sub-sampling strategy evaluation was applied to age data collection of a virtual fish population that has the same statistical and biological characteristics of the Eastern Bering Sea population of Pacific cod. The simulation scheme allowed us to incorporate contributions of several sources of error and to analyze the sensitivity of the different strategies in the population estimates. We found that, on average across all scenarios tested, the main differences between sub-sampling designs arise from the inability of the stratified design to reproduce spatial patterns of the individual traits. However, differences between the sub-sampling strategies in other population estimates may be small, particularly when large sub-sample sizes are used. On isolated scenarios (representative of specific environmental or demographic conditions), the random sub-sampling provided better precision in all population estimates analyzed. The sensitivity analysis revealed the important contribution of spatial autocorrelation in the error of population trait estimates, regardless of the sub-sampling design. This framework will be a useful tool for monitoring and assessment of natural populations with spatially structured traits in multi-stage sampling designs.

Nonstationary environmental and community relationships in the North Pacific Ocean.

Common approaches for summarizing multivariate environmental or community data assume that relationships among variables are stationary over time, and this assumption is often not tested. Here we test the hypothesis that relationships among environmental and community time series are nonstationary in the Gulf of Alaska ecosystem (North Pacific Ocean) over multidecadal time scales. Dynamic factor analysis (DFA) is applied to environmental and community data from before and after 1988/1989, corresponding to the timing of an abrupt decline in temporal variance of the Aleutian Low atmospheric pattern, a leading driver of Gulf of Alaska climate. Results show that covariance among local atmosphere and ocean environmental variables weakened simultaneous to the decline in Aleutian Low variance. At the same time, community-wide responses of 14 fish and crustacean populations to physical forcing weakened, as indicated by nonstationary environment-biology regression coefficients. In line with theoretical predictions, this loss of a shared response to environmental variability was accompanied by weakening community covariance. Individual populations also showed nonstationary relationships with shared trends of community variability. We conclude that assumptions of fixed environmental and community relationships are likely to produce mistaken inference in this ecosystem. Similar concerns may apply in other ecosystems subject to changing climate patterns.

Seasonal dynamics of spatial distributions and overlap between Northeast Arctic cod (Gadus morhua) and capelin (Mallotus villosus) in the Barents Sea.

The trophic link between cod (Gadus sp.) and capelin (Mallotus sp.) is important in many panarctic ecosystems. Since the early 2000s, the Northeast Arctic cod stock (G. morhua) in the Barents Sea has increased greatly, and the sea has been exceptionally warm. Such changes have potentially large effects on species distributions and overlap, which in turn could affect the strength of species interactions. Due to its high latitude location, the Barents Sea has strong seasonal variation in physical conditions and interactions. To study drivers of variation in cod-capelin overlap, we use data from two annual surveys run in winter and in autumn of 2004-2015. We first model winter and autumn spatial distributions of mature and immature cod and capelin. We then calculate overlap from model predictions on a grid with similar spatial resolution as the survey data. Our approach allowed us to interpret changes in overlap as species-specific effects of stock size and temperature, while accounting for sampling variation due to sampling time and depth. We found that during winter both species expanded their distribution in response to increased stock sizes, but how strongly and where the expansion occurred varied. The effect of temperature on distributions varied in space, and differed for cod and capelin and for different components of the two species. The results for autumn were clearer and more consistent. Both species expanded their distribution areas as their stock sizes increased. A positive effect of temperature was found in the north-eastern Barents Sea, where temperatures were lowest at the start of the study. Overlap increased and shifted north-eastwards during the study period and remained high despite a decline in the capelin stock. The increased overlap during autumn could mainly be attributed to the shift in cod distribution with increased cod stock biomass.

Assessing the potential for competition between Pacific Halibut (Hippoglossus stenolepis) and Arrowtooth Flounder (Atheresthes stomias) in the Gulf of Alaska.

Pacific Halibut (Hippoglossus stenolepis) support culturally and economically important fisheries in the Gulf of Alaska, though recent decreases in mean size-at-age have substantially reduced fishery yields, generating concerns among stakeholders and resource managers. Among the prevailing hypotheses for reduced size-at-age is intensified competition with Arrowtooth Flounder (Atheresthes stomias), a groundfish predator that exhibited nearly five-fold increases in biomass between the 1960s and mid-2010s. To assess the potential for competition between Pacific Halibut and Arrowtooth Flounder, we evaluated their degree of spatiotemporal and dietary overlap in the Gulf of Alaska using bottom trawl survey and food habits data provided by the Alaska Fisheries Science Center (NOAA; 1990 to 2017). We restricted analyses to fish measuring 30 to 69 cm fork length and used a delta modeling approach to quantify species-specific presence-absence and catch-per-unit-effort as a function of survey year, tow location, depth, and bottom temperature. We then calculated an index of spatial overlap across a uniform grid by multiplying standardized predictions of species' abundance. Dietary overlap was calculated across the same uniform grid using Schoener's similarity index. Finally, we assessed the relationship between spatial and dietary overlap as a measure of resource partitioning. We found increases in spatial overlap, moving from east to west in the Gulf of Alaska (eastern: 0.13 ± 0.20; central: 0.21 ± 0.11; western: 0.31 ± 0.13 SD). Dietary overlap was low throughout the study area (0.13 ± 0.20 SD). There was no correlation between spatial and dietary overlap, suggesting an absence of resource partitioning along the niche dimensions examined. This finding provides little indication that competition with Arrowtooth Flounder was responsible for changes in Pacific Halibut alHHsize-at-age in the Gulf of Alaska; however, it does not rule out competitive interactions that may have affected resource use prior to standardized data collection or at different spatiotemporal scales.

Oscillating trophic control induces community reorganization in a marine ecosystem.

Understanding how climate regulates trophic control may help to elucidate the causes of transitions between alternate ecosystem states following climate regime shifts. We used a 34-year time series of the abundance of Pacific cod (Gadus macrocephalus) and five prey species to show that the nature of trophic control in a North Pacific ecosystem depends on climate state. Rapid warming in the 1970s caused an oscillation between bottom-up and top-down control. This shift to top-down control apparently contributed to the transition from an initial, prey-rich ecosystem state to the final, prey-poor state. However, top-down control could not be detected in the final state without reference to the initial state and transition period. Complete understanding of trophic control in ecosystems capable of transitions between alternate states may therefore require observations spanning more than one state.

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.

Climate change causing phase transitions of walleye pollock (Theragra chalcogramma) recruitment dynamics.

In 1976 the North Pacific climate shifted, resulting in an average increase of the water temperature. In the Gulf of Alaska the climate shift was followed (i.e. early 1980s) by a gradual but dramatic increase in the abundance of groundfish species that typically prey on pre-recruitment stages of walleye pollock. In the present study we used a previously parameterized model to investigate the effect of these climate and biological changes on the recruitment dynamics of walleye pollock in the Gulf of Alaska. Simulations covered the 1970-2000 time frame and emphasized the medium-to-long temporal scale (i.e. about 5-10 years) of environmental variability. Results showed that during periods characterized by high sea surface temperature and high predation on juvenile pollock stages, recruitment variability and magnitude were below average, and recruitment control was delayed to stages older than the 0-group. Opposite dynamics (i.e. high abundance and variability, and early recruitment control) occurred during periods characterized by low temperature and predation. These results are in general agreement with empirical observations, and allowed us to formulate causal explanations for their occurrence. We interpreted the delay of recruitment control and the reduction of variability as an effect of increased constraint on the abundance of post age-0 stages, in turn imposed by high density dependence and predation mortality. On the other hand, low density-dependence and predation favoured post age-0 survival, and allowed for an unconstrained link between larval and recruitment abundance. Our findings demonstrate that the dominant mechanisms of pollock survival change over contrasting climate regimes. Such changes may in turn cause a phase transition of recruitment dynamics with profound implications for the management of the entire stock.

Behavioral responses of Atlantic cod to sea temperature changes.

Understanding responses of marine species to temperature variability is essential to predict impacts of future climate change in the oceans. Most ectotherms are expected to adjust their behavior to avoid extreme temperatures and minimize acute changes in body temperature. However, measuring such behavioral plasticity in the wild is challenging. Combining 4 years of telemetry-derived behavioral data on juvenile and adult (30-80 cm) Atlantic cod (Gadus morhua), and in situ ocean temperature measurements, we found a significant effect of sea temperature on cod depth use and activity level in coastal Skagerrak. During summer, cod were found in deeper waters when sea surface temperature increased. Further, this effect of temperature was stronger on larger cod. Diel vertical migration, which consists in a nighttime rise to shallow feeding habitats, was stronger among smaller cod. As surface temperature increased beyond ∼15°C, their vertical migration was limited to deeper waters. In addition to larger diel vertical migrations, smaller cod were more active and travelled larger distances compared to larger specimens. Cold temperatures during winter tended, however, to reduce the magnitude of diel vertical migrations, as well as the activity level and distance moved by those smaller individuals. Our findings suggest that future and ongoing rises in sea surface temperature may increasingly deprive cod in this region from shallow feeding areas during summer, which may be detrimental for local populations of the species.

Reconstructing source-sink dynamics in a population with a pelagic dispersal phase.

For many organisms, the reconstruction of source-sink dynamics is hampered by limited knowledge of the spatial assemblage of either the source or sink components or lack of information on the strength of the linkage for any source-sink pair. In the case of marine species with a pelagic dispersal phase, these problems may be mitigated through the use of particle drift simulations based on an ocean circulation model. However, when simulated particle trajectories do not intersect sampling sites, the corroboration of model drift simulations with field data is hampered. Here, we apply a new statistical approach for reconstructing source-sink dynamics that overcomes the aforementioned problems. Our research is motivated by the need for understanding observed changes in jellyfish distributions in the eastern Bering Sea since 1990. By contrasting the source-sink dynamics reconstructed with data from the pre-1990 period with that from the post-1990 period, it appears that changes in jellyfish distribution resulted from the combined effects of higher jellyfish productivity and longer dispersal of jellyfish resulting from a shift in the ocean circulation starting in 1991. A sensitivity analysis suggests that the source-sink reconstruction is robust to typical systematic and random errors in the ocean circulation model driving the particle drift simulations. The jellyfish analysis illustrates that new insights can be gained by studying structural changes in source-sink dynamics. The proposed approach is applicable for the spatial source-sink reconstruction of other species and even abiotic processes, such as sediment transport.