Non-stationary and interactive effects of climate and competition on pink salmon productivity.

Pacific salmon (Oncorhynchus spp.) are exposed to increased environmental change and multiple human stressors. To anticipate future impacts of global change and to improve sustainable resource management, it is critical to understand how wild salmon populations respond to stressors associated with human-caused changes such as climate warming and ocean acidification, as well as competition in the ocean, which is intensified by the large-scale production and release of hatchery reared salmon. Pink salmon (O. gorbuscha) are a keystone species in the North Pacific Ocean and support highly valuable commercial fisheries. We investigated the joint effects of changes in ocean conditions and salmon abundances on the productivity of wild pink salmon. Our analysis focused on Prince William Sound in Alaska, because the region accounts for ~50% of the global production of hatchery pink salmon with local hatcheries releasing 600-700 million pink salmon fry annually. Using 60 years of data on wild pink salmon abundances, hatchery releases, and ecological conditions in the ocean, we find evidence that hatchery pink salmon releases negatively affect wild pink salmon productivity, likely through competition between wild and hatchery juveniles in nearshore marine habitats. We find no evidence for effects of ocean acidification on pink salmon productivity. However, a change in the leading mode of North Pacific climate in 1988-1989 weakened the temperature-productivity relationship and altered the strength of intraspecific density dependence. Therefore, our results suggest non-stationary (i.e., time varying) and interactive effects of ocean climate and competition on pink salmon productivity. Our findings further highlight the need for salmon management to consider potential adverse effects of large-scale hatchery production within the context of ocean change.

The changing physical and ecological meanings of North Pacific Ocean climate indices.

Climate change is likely to change the relationships between commonly used climate indices and underlying patterns of climate variability, but this complexity is rarely considered in studies using climate indices. Here, we show that the physical and ecological conditions mapping onto the Pacific Decadal Oscillation (PDO) index and North Pacific Gyre Oscillation (NPGO) index have changed over multidecadal timescales. These changes apparently began around a 1988/1989 North Pacific climate shift that was marked by abrupt northeast Pacific warming, declining temporal variance in the Aleutian Low (a leading atmospheric driver of the PDO), and increasing correlation between the PDO and NPGO patterns. Sea level pressure and surface temperature patterns associated with each climate index changed after 1988/1989, indicating that identical index values reflect different states of basin-scale climate over time. The PDO and NPGO also show time-dependent skill as indices of regional northeast Pacific ecosystem variability. Since the late 1980s, both indices have become less relevant to physical-ecological variability in regional ecosystems from the Bering Sea to the southern California Current. Users of these climate indices should be aware of nonstationary relationships with underlying climate variability within the historical record, and the potential for further nonstationarity with ongoing climate change.

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.

Characterizing driver-response relationships in marine pelagic ecosystems for improved ocean management.

Scientists and resource managers often use methods and tools that assume ecosystem components respond linearly to environmental drivers and human stressors. However, a growing body of literature demonstrates that many relationships are-non-linear, where small changes in a driver prompt a disproportionately large ecological response. We aim to provide a comprehensive assessment of the relationships between drivers and ecosystem components to identify where and when non-linearities are likely to occur. We focused our analyses on one of the best-studied marine systems, pelagic ecosystems, which allowed us to apply robust statistical techniques on a large pool of previously published studies. In this synthesis, we (1) conduct a wide literature review on single driver-response relationships in pelagic systems, (2) use statistical models to identify the degree of non-linearity in these relationships, and (3) assess whether general patterns exist in the strengths and shapes of non-linear relationships across drivers. Overall we found that non-linearities are common in pelagic ecosystems, comprising at least 52% of all driver-response relation- ships. This is likely an underestimate, as papers with higher quality data and analytical approaches reported non-linear relationships at a higher frequency (on average 11% more). Consequently, in the absence of evidence for a linear relationship, it is safer to assume a relationship is non-linear. Strong non-linearities can lead to greater ecological and socioeconomic consequences if they are unknown (and/or unanticipated), but if known they may provide clear thresholds to inform management targets. In pelagic systems, strongly non-linear relationships are often driven by climate and trophodynamic variables but are also associated with local stressors, such as overfishing and pollution, that can be more easily controlled by managers. Even when marine resource managers cannot influence ecosystem change, they can use information about threshold responses to guide how other stressors are managed and to adapt to new ocean conditions. As methods to detect and reduce uncertainty around threshold values improve, managers will be able to better understand and account for ubiquitous non-linear relationships.

Spatially Explicit Modeling Reveals Cephalopod Distributions Match Contrasting Trophic Pathways in the Western Mediterranean Sea.

Populations of the same species can experience different responses to the environment throughout their distributional range as a result of spatial and temporal heterogeneity in habitat conditions. This highlights the importance of understanding the processes governing species distribution at local scales. However, research on species distribution often averages environmental covariates across large geographic areas, missing variability in population-environment interactions within geographically distinct regions. We used spatially explicit models to identify interactions between species and environmental, including chlorophyll a (Chla) and sea surface temperature (SST), and trophic (prey density) conditions, along with processes governing the distribution of two cephalopods with contrasting life-histories (octopus and squid) across the western Mediterranean Sea. This approach is relevant for cephalopods, since their population dynamics are especially sensitive to variations in habitat conditions and rarely stable in abundance and location. The regional distributions of the two cephalopod species matched two different trophic pathways present in the western Mediterranean Sea, associated with the Gulf of Lion upwelling and the Ebro river discharges respectively. The effects of the studied environmental and trophic conditions were spatially variant in both species, with usually stronger effects along their distributional boundaries. We identify areas where prey availability limited the abundance of cephalopod populations as well as contrasting effects of temperature in the warmest regions. Despite distributional patterns matching productive areas, a general negative effect of Chla on cephalopod densities suggests that competition pressure is common in the study area. Additionally, results highlight the importance of trophic interactions, beyond other common environmental factors, in shaping the distribution of cephalopod populations. Our study presents a valuable approach for understanding the spatially variant ecology of cephalopod populations, which is important for fisheries and ecosystem management.

Implications of scaled δ15N fractionation for community predator-prey body mass ratio estimates in size-structured food webs.

Nitrogen stable isotope ratios (δ(15) N) may be used to estimate community-level relationships between trophic level (TL) and body size in size-structured food webs and hence the mean predator to prey body mass ratio (PPMR). In turn, PPMR is used to estimate mean food chain length, trophic transfer efficiency and rates of change in abundance with body mass (usually reported as slopes of size spectra) and to calibrate and validate food web models. When estimating TL, researchers had assumed that fractionation of δ(15) N (Δδ(15) N) did not change with TL. However, a recent meta-analysis indicated that this assumption was not as well supported by data as the assumption that Δδ(15) N scales negatively with the δ(15) N of prey. We collated existing fish community δ(15) N-body size data for the Northeast Atlantic and tropical Western Arabian Sea with new data from the Northeast Pacific. These data were used to estimate TL-body mass relationships and PPMR under constant and scaled Δδ(15) N assumptions, and to assess how the scaled Δδ(15) N assumption affects our understanding of the structure of these food webs. Adoption of the scaled Δδ(15) N approach markedly reduces the previously reported differences in TL at body mass among fish communities from different regions. With scaled Δδ(15) N, TL-body mass relationships became more positive and PPMR fell. Results implied that realized prey size in these size-structured fish communities are less variable than previously assumed and food chains potentially longer. The adoption of generic PPMR estimates for calibration and validation of size-based fish community models is better supported than hitherto assumed, but predicted slopes of community size spectra are more sensitive to a given change or error in realized PPMR when PPMR is small.

Climate and Demography Dictate the Strength of Predator-Prey Overlap in a Subarctic Marine Ecosystem.

There is growing evidence that climate and anthropogenic influences on marine ecosystems are largely manifested by changes in species spatial dynamics. However, less is known about how shifts in species distributions might alter predator-prey overlap and the dynamics of prey populations. We developed a general approach to quantify species spatial overlap and identify the biotic and abiotic variables that dictate the strength of overlap. We used this method to test the hypothesis that population abundance and temperature have a synergistic effect on the spatial overlap of arrowtooth flounder (predator) and juvenile Alaska walleye pollock (prey, age-1) in the eastern Bering Sea. Our analyses indicate that (1) flounder abundance and temperature are key variables dictating the strength of flounder and pollock overlap, (2) changes in the magnitude of overlap may be largely driven by density-dependent habitat selection of flounder, and (3) species overlap is negatively correlated to juvenile pollock recruitment when flounder biomass is high. Overall, our findings suggest that continued increases in flounder abundance coupled with the predicted long-term warming of ocean temperatures could have important implications for the predator-prey dynamics of arrowtooth flounder and juvenile pollock. The approach used in this study is valuable for identifying potential consequences of climate variability and exploitation on species spatial dynamics and interactions in many marine ecosystems.

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.

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.