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Abstract The prevalence and intensity of marine heatwaves is increasing globally, disrupting local environmental conditions. The individual and population-level impacts of prolonged heatwaves on marine species have recently been demonstrated, yet whole-ecosystem consequences remain unexplored. We leveraged time series abundance data of 361 taxa, grouped into 86 functional groups, from six long-term surveys, diet information from a new diet database, and previous modeling efforts, to build two food web networks using an extension of the popular Ecopath ecosystem modeling framework, Ecotran. We compare ecosystem models parameterized before and after the onset of recent marine heatwaves to evaluate the cascading effects on ecosystem structure and function in the Northeast Pacific Ocean. While the ecosystem-level contribution (prey) and demand (predators) of most functional groups changed following the heatwaves, gelatinous taxa experienced the largest transformations, underscored by the arrival of northward-expanding pyrosomes. We show altered trophic relationships and energy flux have potentially profound consequences for ecosystem structure and function, and raise concerns for populations of threatened and harvested species.

IntroductionUnderstanding how abundance, productivity and distribution of individual species may respond to climate change is a critical first step towards anticipating alterations in marine ecosystem structure and function, as well as developing strategies to adapt to the full range of potential changes.MethodsThis study applies the NOAA (National Oceanic and Atmospheric Administration) Fisheries Climate Vulnerability Assessment method to 64 federally-managed species in the California Current Large Marine Ecosystem to assess their vulnerability to climate change, where vulnerability is a function of a species’ exposure to environmental change and its biological sensitivity to a set of environmental conditions, which includes components of its resiliency and adaptive capacity to respond to these new conditions.ResultsOverall, two-thirds of the species were judged to have Moderate or greater vulnerability to climate change, and only one species was anticipated to have a positive response. Species classified as Highly or Very Highly vulnerable share one or more characteristics including: 1) having complex life histories that utilize a wide range of freshwater and marine habitats; 2) having habitat specialization, particularly for areas that are likely to experience increased hypoxia; 3) having long lifespans and low population growth rates; and/or 4) being of high commercial value combined with impacts from non-climate stressors such as anthropogenic habitat degradation. Species with Low or Moderate vulnerability are either habitat generalists, occupy deep-water habitats or are highly mobile and likely to shift their ranges.DiscussionAs climate-related changes intensify, this work provides key information for both scientists and managers as they address the long-term sustainability of fisheries in the region. This information can inform near-term advice for prioritizing species-level data collection and research on climate impacts, help managers to determine when and where a precautionary approach might be warranted, in harvest or other management decisions, and help identify habitats or life history stages that might be especially effective to protect or restore.

As we confront novel environmental challenges, a full understanding of the physical and biological processes that govern species responses to climate change will help maintain biodiversity and support conservation measures that are more robust to irreducible uncertainty. However, climate impacts are so complex, and the literature on salmon and trout is so vast that researchers and decision makers scramble to make sense of it all. Therefore, we conducted a systematic literature review of climate impacts on salmon and anadromous trout as a resource for stakeholders, managers, and researchers. We reviewed studies published from 2010 to 2021 that address climate impacts on these fish and organized them in a database of 1169 physical and 1853 biological papers. Papers are labeled with keywords across eight categories related to subject matter and study methods. We compared the literature by biological process and life stage and used these comparisons to assess strengths and weaknesses. We then summarized expected phenotypic and genetic responses and management actions by life stage. Overall, we found the largest research gaps related to species interactions, behavioral responses, and effects that carry over across life stages. With this collection of the literature, we can better apply scarce conservation resources, fill knowledge gaps, and make informed decisions that do not ignore uncertainty.

Lisa Crozier et al. assembled a database for eight wild populations of threatened Chinook salmon in the Salmon River Basin in order to evaluate climate impacts at all life stages, and model future population trajectories. They project that populations rapidly decline in response to increasing sea surface temperatures and conclude that dramatic increases in the number or survival of smolts are needed to overcome the negative impacts of climate change for this threatened species.

The common practice of incorporating environmental indices into population models has greatly advanced our understanding of ecological systems. Unfortunately, we are increasingly seeing published correlations between population indicators and environmental indices breaking down when tested with new data. Examining how the correlations among indices change over time could help explain underlying causal mechanisms, which ultimately strengthen the basis for prediction of population indicators. For migratory animals such as anadromous salmon (Oncorhynchus spp.), the habitat conditions they experience can affect their lifetime fitness and population viability. We analyzed 43 freshwater, marine, and climate indices associated with 72 river sites and five coastal ecoregions inhabited by Chinook and coho salmon (O. tshawytscha and O. kisutch) in the western USA. Utilizing long time series (ranging from 32 to 124 years), we examined spatial and temporal patterns in correlations through hierarchical clustering across sites and non-stationarity across time. Individual river sites clustered into two Northwest and one Southwest groups. Northwest sites generally showed stronger correlations between freshwater and climate indices, while Southwest sites showed stronger correlations within freshwater or within marine/climate indices. For a closer examination at shorter periods, we parsed the time series into 10-year windows and showed how pairwise correlations changed over time with spring–summer Pacific Decadal Oscillation index in the Northwest and with spring flow in the Southwest. Stronger correlations across multiple indices tended to occur when large-scale climatic events (e.g., Oceanic Niño and Pacific Decadal Oscillation indices) were in-phase, and phase transitions (e.g., from positive to negative) occurred in the same 10-year window. In a third analysis, we assessed how well indices provided unique vs. confounding/complex information and had consistent vs. varying relationships based on the mean and variance of 10-year correlations. Across index types, the variance in correlations tended to be lowest in marine vs. climate indices, higher among freshwater indices, and highest for freshwater vs. marine/climate indices. Yet, the mean strength of correlations for freshwater vs. marine/climate indices was still comparable to those among freshwater ones. Overall, identifying time periods when correlations tend to change will help interpret historical and projected population indicators. Spatial trends in the strength of correlations also indicate that the level of confounding effects among indices can differ regionally. We emphasize the importance of knowing the strength and variability of correlations among indices, and their representativeness of ecological processes in the context of combined phases of multiple climatic indices.

In 2015, the Pacific marine heat wave, low river flows, and record high water temperatures in the Columbia River Basin contributed to a near-complete failure of the adult migration of endangered Snake River sockeye salmon (Oncorhynchus nerka, NOAA Fisheries 2016). These extreme weather events may become the new normal due to anthropogenic climate change, with catastrophic consequences for endangered species. Existing anthropogenic pressures may amplify vulnerability to climate change, but these potential synergies have rarely been quantified. We examined factors affecting survival of endangered sockeye (Oncorhynchus nerka) and threatened Chinook salmon (O. tshawytscha) as they migrated upstream through eight dams and reservoirs to spawning areas in the Snake River Basin. Our extensive database included histories of 17,279 individual fish that migrated since 2004. A comparison between conditions in 2015 and daily temperatures and flows in a regulated basin forced by output from global climate models showed that 2015 did have many characteristics of projected future mean conditions. To evaluate potential salmon responses, we modeled migration timing and apparent survival under historical and future climate scenarios (2040s). For Chinook salmon, adult survival from the first dam encountered to spawning grounds dropped by 4-15%, depending on the climate scenario. For sockeye, survival dropped by ~80% from their already low levels. Through sensitivity analyses, we observed that the adult sockeye migration would need to shift more than 2 weeks earlier than predicted to maintain survival rates typical of those seen during 2008-2017. Overall, the greater impacts of climate change on adult sockeye compared with adult Chinook salmon reflected differences in life history and environmental sensitivities, which were compounded for sockeye by larger effect sizes from other anthropogenic factors. Compared with Chinook, sockeye was more negatively affected by a history of juvenile transportation and by similar temperatures and flows. The largest changes in temperature and flow were projected to be upstream from the hydrosystem, where direct mitigation through hydrosystem management is not an option. Unfortunately, Snake River sockeye have likely lost much of their adaptive capacity with the loss of the wild population. Further work exploring habitat restoration or additional mitigation actions is urgently needed.

The Northern California Current is a highly productive marine upwelling ecosystem that is economically and ecologically important. It is home to both commercially harvested species and those that are federally listed under the U.S. Endangered Species Act. Recently, there has been a global shift from single-species fisheries management to ecosystem-based fisheries management, which acknowledges that more complex dynamics can reverberate through a food web. Here, we have integrated new research into an end-to-end ecosystem model (i.e., physics to fisheries) using data from long-term ocean surveys, phytoplankton satellite imagery paired with a vertically generalized production model, a recently assembled diet database, fishery catch information, species distribution models, and existing literature. This spatially-explicit model includes 90 living and detrital functional groups ranging from phytoplankton, krill, and forage fish to salmon, seabirds, and marine mammals, and nine fisheries that occur off the coast of Washington, Oregon, and Northern California. This model was updated from previous regional models to account for more recent changes in the Northern California Current (e.g., increases in market squid and some gelatinous zooplankton such as pyrosomes and salps), to expand the previous domain to increase the spatial resolution, to include data from previously unincorporated surveys, and to add improved characterization of endangered species, such as Chinook salmon (Oncorhynchus tshawytscha) and southern resident killer whales (Orcinus orca). Our model is mass-balanced, ecologically plausible, without extinctions, and stable over 150-year simulations. Ammonium and nitrate availability, total primary production rates, and model-derived phytoplankton time series are within realistic ranges. As we move towards holistic ecosystem-based fisheries management, we must continue to openly and collaboratively integrate our disparate datasets and collective knowledge to solve the intricate problems we face. As a tool for future research, we provide the data and code to use our ecosystem model.

The majority of Columbia River summer-run steelhead encounter high river temperatures (near or > 20°C) during their spawning migration. While some steelhead pass through the mid-Columbia River in a matter of days, others use tributary habitats as temperature refuges for periods that can last months. Using PIT tag detection data from adult return years 2004-2016, we fit 3-component mixture models to differentiate between "fast", "slow", and "overwintering" migration behaviors in five aggregated population groups. Fast fish migrated straight through the reach on average in ~7-9 days while slow fish delayed their migration for weeks to months, and overwintering fish generally took ~150-250 days. We then fit covariate models to examine what factors contributed to the probability of migration delay during summer months (slow or overwintering behaviors), and to explore how migration delay related to mortality. Finally, to account for the impact of extended residence times in the reach for fish that delayed, we compared patterns in estimated average daily rates of mortality between migration behaviors and across population groups. Results suggest that migration delay was primarily triggered by high river temperatures but temperature thresholds for delay were lowest just before the seasonal peak in river temperatures. While all populations groups demonstrated these general patterns, we documented substantial variability in temperature thresholds and length of average delays across population groups. Although migration delay was related to higher reach mortality, it was also related to lower average daily mortality rates due to the proportional increase in reach passage duration being larger than the associated increase in mortality. Lower daily mortality rates suggest that migration delay could help mitigate the impacts of harsh migration conditions, presumably through the use of thermal refuges, despite prolonged exposure to local fisheries. Future studies tracking individual populations from their migration through reproduction could help illuminate the full extent of the tradeoffs between different migration behaviors.

Major ecological realignments are already occurring in response to climate change. To be successful, conservation strategies now need to account for geographical patterns in traits sensitive to climate change, as well as climate threats to species-level diversity. As part of an effort to provide such information, we conducted a climate vulnerability assessment that included all anadromous Pacific salmon and steelhead (Oncorhynchus spp.) population units listed under the U.S. Endangered Species Act. Using an expert-based scoring system, we ranked 20 attributes for the 28 listed units and 5 additional units. Attributes captured biological sensitivity, or the strength of linkages between each listing unit and the present climate; climate exposure, or the magnitude of projected change in local environmental conditions; and adaptive capacity, or the ability to modify phenotypes to cope with new climatic conditions. Each listing unit was then assigned one of four vulnerability categories. Units ranked most vulnerable overall were Chinook (O. tshawytscha) in the California Central Valley, coho (O. kisutch) in California and southern Oregon, sockeye (O. nerka) in the Snake River Basin, and spring-run Chinook in the interior Columbia and Willamette River Basins. We identified units with similar vulnerability profiles using a hierarchical cluster analysis. Life history characteristics, especially freshwater and estuary residence times, interplayed with gradations in exposure from south to north and from coastal to interior regions to generate landscape-level patterns within each species. Nearly all listing units faced high exposures to projected increases in stream temperature, sea surface temperature, and ocean acidification, but other aspects of exposure peaked in particular regions. Anthropogenic factors, especially migration barriers, habitat degradation, and hatchery influence, have reduced the adaptive capacity of most steelhead and salmon populations. Enhancing adaptive capacity is essential to mitigate for the increasing threat of climate change. Collectively, these results provide a framework to support recovery planning that considers climate impacts on the majority of West Coast anadromous salmonids.