Chinook salmon depth distributions on the continental shelf are shaped by interactions between location, season, and individual condition.

BACKGROUND: Ecological and physical conditions vary with depth in aquatic ecosystems, resulting in gradients of habitat suitability. Although variation in vertical distributions among individuals provides evidence of habitat selection, it has been challenging to disentangle how processes at multiple spatio-temporal scales shape behaviour. METHODS: We collected thousands of observations of depth from > 300 acoustically tagged adult Chinook salmon Oncorhynchus tshawytscha, spanning multiple seasons and years. We used these data to parameterize a machine-learning model to disentangle the influence of spatial, temporal, and dynamic oceanographic variables while accounting for differences in individual condition and maturation stage. RESULTS: The top performing machine learning model used bathymetric depth ratio (i.e., individual depth relative to seafloor depth) as a response. We found that bathymetry, season, maturation stage, and spatial location most strongly influenced Chinook salmon depth. Chinook salmon bathymetric depth ratios were deepest in shallow water, during winter, and for immature individuals. We also identified non-linear interactions among covariates, resulting in spatially-varying effects of zooplankton concentration, lunar cycle, temperature and oxygen concentration. CONCLUSIONS: Our results suggest Chinook salmon vertical habitat use is a function of ecological interactions, not physiological constraints. Temporal and spatial variation in depth distributions could be used to guide management decisions intended to reduce fishery impacts on Chinook salmon. More generally, our findings demonstrate how complex interactions among bathymetry, seasonality, location, and life history stage regulate vertical habitat selection.

Marine heatwaves disrupt ecosystem structure and function via altered food webs and energy flux.

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.

An updated end-to-end ecosystem model of the Northern California Current reflecting ecosystem changes due to recent marine heatwaves.

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.

Climate vulnerability assessment for Pacific salmon and steelhead in the California Current Large Marine Ecosystem.

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.

Past and estimated future impact of invasive alien mammals on insular threatened vertebrate populations.

Invasive mammals on islands pose severe, ongoing threats to global biodiversity. However, the severity of threats from different mammals, and the role of interacting biotic and abiotic factors in driving extinctions, remain poorly understood at a global scale. Here we model global extirpation patterns for island populations of threatened and extinct vertebrates. Extirpations are driven by interacting factors including invasive rats, cats, pigs, mustelids and mongooses, native species taxonomic class and volancy, island size, precipitation and human presence. We show that controlling or eradicating the relevant invasive mammals could prevent 41-75% of predicted future extirpations. The magnitude of benefits varies across species and environments; for example, managing invasive mammals on small, dry islands could halve the extirpation risk for highly threatened birds and mammals, while doing so on large, wet islands may have little benefit. Our results provide quantitative estimates of conservation benefits and, when combined with costs in a return-on-investment framework, can guide efficient conservation strategies.

A spatially distinct history of the development of california groundfish fisheries.

During the past century, commercial fisheries have expanded from small vessels fishing in shallow, coastal habitats to a broad suite of vessels and gears that fish virtually every marine habitat on the globe. Understanding how fisheries have developed in space and time is critical for interpreting and managing the response of ecosystems to the effects of fishing, however time series of spatially explicit data are typically rare. Recently, the 1933-1968 portion of the commercial catch dataset from the California Department of Fish and Wildlife was recovered and digitized, completing the full historical series for both commercial and recreational datasets from 1933-2010. These unique datasets include landing estimates at a coarse 10 by 10 minute "grid-block" spatial resolution and extends the entire length of coastal California up to 180 kilometers from shore. In this study, we focus on the catch history of groundfish which were mapped for each grid-block using the year at 50% cumulative catch and total historical catch per habitat area. We then constructed generalized linear models to quantify the relationship between spatiotemporal trends in groundfish catches, distance from ports, depth, percentage of days with wind speed over 15 knots, SST and ocean productivity. Our results indicate that over the history of these fisheries, catches have taken place in increasingly deeper habitat, at a greater distance from ports, and in increasingly inclement weather conditions. Understanding spatial development of groundfish fisheries and catches in California are critical for improving population models and for evaluating whether implicit stock assessment model assumptions of relative homogeneity of fisheries removals over time and space are reasonable. This newly reconstructed catch dataset and analysis provides a comprehensive appreciation for the development of groundfish fisheries with respect to commonly assumed trends of global fisheries patterns that are typically constrained by a lack of long-term spatial datasets.

Green sturgeon distribution in the Pacific Ocean estimated from modeled oceanographic features and migration behavior.

The green sturgeon (Acipenser medirostris), which is found in the eastern Pacific Ocean from Baja California to the Bering Sea, tends to be highly migratory, moving long distances among estuaries, spawning rivers, and distant coastal regions. Factors that determine the oceanic distribution of green sturgeon are unclear, but broad-scale physical conditions interacting with migration behavior may play an important role. We estimated the distribution of green sturgeon by modeling species-environment relationships using oceanographic and migration behavior covariates with maximum entropy modeling (MaxEnt) of species geographic distributions. The primary concentration of green sturgeon was estimated from approximately 41-51.5° N latitude in the coastal waters of Washington, Oregon, and Vancouver Island and in the vicinity of San Francisco and Monterey Bays from 36-37° N latitude. Unsuitably cold water temperatures in the far north and energetic efficiencies associated with prevailing water currents may provide the best explanation for the range-wide marine distribution of green sturgeon. Independent trawl records, fisheries observer records, and tagging studies corroborated our findings. However, our model also delineated patchily distributed habitat south of Monterey Bay, though there are few records of green sturgeon from this region. Green sturgeon are likely influenced by countervailing pressures governing their dispersal. They are behaviorally directed to revisit natal freshwater spawning rivers and persistent overwintering grounds in coastal marine habitats, yet they are likely physiologically bounded by abiotic and biotic environmental features. Impacts of human activities on green sturgeon or their habitat in coastal waters, such as bottom-disturbing trawl fisheries, may be minimized through marine spatial planning that makes use of high-quality species distribution information.

Habitat-related benthic macrofaunal assemblages of bays and estuaries of the western United States.

Data from 7 coastwide and regional benthic surveys were combined and used to assess the number and distribution of estuarine benthic macrofaunal assemblages of the western United States. Q-mode cluster analysis was applied to 714 samples and site groupings were tested for differences in 4 habitat factors (latitude, salinity, sediment grain size, and depth). Eight macrofaunal assemblages, structured primarily by latitude, salinity, and sediment grain size, were identified: (A) Puget Sound fine sediment, (B) Puget Sound coarse sediment, (C) southern California marine bays, (D) polyhaline central San Francisco Bay, (E) shallow estuaries and wetlands, (F) saline very coarse sediment, (G) mesohaline San Francisco Bay, and (H) limnetic and oligohaline. The Puget Sound, southern California, and San Francisco Bay assemblages were geographically distinct, while Assemblages E, F and H were distributed widely along the entire coast. A second Q-mode cluster analysis was conducted after adding replicate samples that were available from some of the sites and temporal replicates that were available for sites that were sampled in successive years. Variabilities due to small spatial scale habitat heterogeneity and temporal change were both low in Puget Sound, but temporal variability was high in the San Francisco estuary where large fluctuations in freshwater inputs and salinity among years leads to spatial relocation of the assemblages.

Green sturgeon physical habitat use in the coastal Pacific Ocean.

The green sturgeon (Acipenser medirostris) is a highly migratory, oceanic, anadromous species with a complex life history that makes it vulnerable to species-wide threats in both freshwater and at sea. Green sturgeon population declines have preceded legal protection and curtailment of activities in marine environments deemed to increase its extinction risk. Yet, its marine habitat is poorly understood. We built a statistical model to characterize green sturgeon marine habitat using data from a coastal tracking array located along the Siletz Reef near Newport, Oregon, USA that recorded the passage of 37 acoustically tagged green sturgeon. We classified seafloor physical habitat features with high-resolution bathymetric and backscatter data. We then described the distribution of habitat components and their relationship to green sturgeon presence using ordination and subsequently used generalized linear model selection to identify important habitat components. Finally, we summarized depth and temperature recordings from seven green sturgeon present off the Oregon coast that were fitted with pop-off archival geolocation tags. Our analyses indicated that green sturgeon, on average, spent a longer duration in areas with high seafloor complexity, especially where a greater proportion of the substrate consists of boulders. Green sturgeon in marine habitats are primarily found at depths of 20-60 meters and from 9.5-16.0°C. Many sturgeon in this study were likely migrating in a northward direction, moving deeper, and may have been using complex seafloor habitat because it coincides with the distribution of benthic prey taxa or provides refuge from predators. Identifying important green sturgeon marine habitat is an essential step towards accurately defining the conditions that are necessary for its survival and will eventually yield range-wide, spatially explicit predictions of green sturgeon distribution.

Mixed-source reintroductions lead to outbreeding depression in second-generation descendents of a native North American fish.

Reintroductions are commonly employed to preserve intraspecific biodiversity in fragmented landscapes. However, reintroduced populations are frequently smaller and more geographically isolated than native populations. Mixing genetically, divergent sources are often proposed to attenuate potentially low genetic diversity in reintroduced populations that may result from small effective population sizes. However, a possible negative tradeoff for mixing sources is outbreeding depression in hybrid offspring. We examined the consequences of mixed-source reintroductions on several fitness surrogates at nine slimy sculpin (Cottus cognatus) reintroduction sites in south-east Minnesota. We inferred the relative fitness of each crosstype in the reintroduced populations by comparing their growth rate, length, weight, body condition and persistence in reintroduced populations. Pure strain descendents from a single source population persisted in a greater proportion than expected in the reintroduced populations, whereas all other crosstypes occurred in a lesser proportion. Length, weight and growth rate were lower for second-generation intra-population hybrid descendents than for pure strain and first-generation hybrids. In the predominant pure strain, young-of the-year size was significantly greater than any other crosstype. Our results suggested that differences in fitness surrogates among crosstypes were consistent with disrupted co-adapted gene complexes associated with beneficial adaptations in these reintroduced populations. Future reintroductions may be improved by evaluating the potential for local adaptation in source populations or by avoiding the use of mixed sources by default when information on local adaptations or other genetic characteristics is lacking.

Calibration and evaluation of five indicators of benthic community condition in two California bay and estuary habitats.

Many types of indices have been developed to assess benthic invertebrate community condition, but there have been few studies evaluating the relative performance of different index approaches. Here we calibrate and compare the performance of five indices: the Benthic Response Index (BRI), Benthic Quality Index (BQI), Relative Benthic Index (RBI), River Invertebrate Prediction and Classification System (RIVPACS), and the Index of Biotic Integrity (IBI). We also examine whether index performance improves when the different indices, which rely on measurement of different properties, are used in combination. The five indices were calibrated for two geographies using 238 samples from southern California marine bays and 125 samples from polyhaline San Francisco Bay. Index performance was evaluated by comparing index assessments of 35 sites to the best professional judgment of nine benthic experts. None of the individual indices performed as well as the average expert in ranking sample condition or evaluating whether benthic assemblages exhibited evidence of disturbance. However, several index combinations outperformed the average expert. When results from both habitats were combined, two four-index combinations and a three-index combination performed best. However, performance differences among several combinations were small enough that factors such as logistics can also become a consideration in index selection.