Stream hydrology and a pulse subsidy shape patterns of fish foraging.

Pulsed subsidy events create ephemeral fluxes of hyper-abundant resources that can shape annual patterns of consumption and growth for recipient consumers. However, environmental conditions strongly affect local resource availability for much of the year, and can heavily impact consumer foraging and growth patterns prior to pulsed subsidy events. Thus, a consumer's capacity to exploit pulse subsidy resources may be influenced by antecedent environmental conditions, but this has rarely been shown in nature and is unknown in aquatic ecosystems. Here, we sought to understand the importance of hydrologic variation and a salmon pulse subsidy on the foraging and growth patterns of two stream salmonids in a coastal southeast Alaska drainage. To do this, we sampled fish stomach contents at a high temporal frequency (daily-weekly measurements) and analyzed fish consumption rates in relation to streamflow and pulse subsidy resource availability. We then explored the influence of interannual hydrologic variation on access to pulse subsidy resources (i.e. whether fish exceeded an egg consumption gape limit) in a bioenergetic simulation. Prior to Pink Salmon spawning, Dolly Varden and Coho Salmon displayed distinct and nonlinear flow-foraging relationships, where forage for both species consisted primarily of macroinvertebrates. During this time period, consumption maxima coincided with baseflow and the highest observed flow conditions, and consumption minima were observed at severe low-water and intermediate flow values. After salmon spawning began, forage was not significantly related to flow and consisted primarily of salmon eggs. Further, consumption rates increased overall, and foraging patterns did not appear to be affected by flow in either species. Bioenergetic simulations revealed that patterns of interannual hydrologic variation may shift Coho Salmon growth trajectories among years. Together, our results suggest that access to marine pulse subsidy resources may depend on whether antecedent hydrologic conditions are suitable for juvenile salmonids to grow large enough to consume salmon eggs by the onset of spawning.

Species distributions and the recognition of risk in restoration planning: A case study of salmonid fishes.

One of the risks faced by habitat restoration practitioners is whether habitats included in restoration planning will be used by the target species or, conversely, whether habitats excluded from restoration planning would have benefited the target species. With the goal of providing a quantitative decision-making approach that represented varying levels of risk tolerance, we used multiple probability decision thresholds (PDT) to predict the range of occurrence for three anadromous fishes (Oncorhynchus spp.) in a watershed in southwestern Washington, USA. For each species, we compared the predicted range of occurrence to the distribution used for restoration planning and quantified the amount of habitat blocked by anthropogenic barriers. Coho salmon (O. kisutch) had the broadest predicted range of occurrence (3061.6-6357.9 km; 0.75-0.25 PDT), followed by steelhead trout (O. mykiss; 1828.8-2836.8 km) and chum salmon (O. keta; 1373.9-1629.1 km). For each species, the predicted range of occurrence was similar or greater than the distribution used for restoration planning, suggesting that the current plan may exclude habitats that would benefit each species. Coho salmon had the greatest percentage of habitat blocked by anthropogenic barriers, followed by steelhead trout and chum salmon, respectively. Modeling species distributions at multiple risk-tolerance scenarios acknowledges uncertainty in restoration planning and allows practitioners to weigh the ecological benefits and budgetary constraints when considering locations for restoration. To effectively communicate restoration science to support practitioners in decision-making, we developed an R Shiny application online user interface available at: https://shiny.wdfw-fish.us/ChehalisRiverBasinSalmonidRangeOfOccurence/.

Groundwater declines are linked to changes in Great Plains stream fish assemblages.

Groundwater pumping for agriculture is a major driver causing declines of global freshwater ecosystems, yet the ecological consequences for stream fish assemblages are rarely quantified. We combined retrospective (1950-2010) and prospective (2011-2060) modeling approaches within a multiscale framework to predict change in Great Plains stream fish assemblages associated with groundwater pumping from the United States High Plains Aquifer. We modeled the relationship between the length of stream receiving water from the High Plains Aquifer and the occurrence of fishes characteristic of small and large streams in the western Great Plains at a regional scale and for six subwatersheds nested within the region. Water development at the regional scale was associated with construction of 154 barriers that fragment stream habitats, increased depth to groundwater and loss of 558 km of stream, and transformation of fish assemblage structure from dominance by large-stream to small-stream fishes. Scaling down to subwatersheds revealed consistent transformations in fish assemblage structure among western subwatersheds with increasing depths to groundwater. Although transformations occurred in the absence of barriers, barriers along mainstem rivers isolate depauperate western fish assemblages from relatively intact eastern fish assemblages. Projections to 2060 indicate loss of an additional 286 km of stream across the region, as well as continued replacement of large-stream fishes by small-stream fishes where groundwater pumping has increased depth to groundwater. Our work illustrates the shrinking of streams and homogenization of Great Plains stream fish assemblages related to groundwater pumping, and we predict similar transformations worldwide where local and regional aquifer depletions occur.

Thermal adaptation and phenotypic plasticity in a warming world: Insights from common garden experiments on Alaskan sockeye salmon.

An important unresolved question is how populations of coldwater-dependent fishes will respond to rapidly warming water temperatures. For example, the culturally and economically important group, Pacific salmon (Oncorhynchus spp.), experience site-specific thermal regimes during early development that could be disrupted by warming. To test for thermal local adaptation and heritable phenotypic plasticity in Pacific salmon embryos, we measured the developmental rate, survival, and body size at hatching in two populations of sockeye salmon (Oncorhynchus nerka) that overlap in timing of spawning but incubate in contrasting natural thermal regimes. Using a split half-sibling design, we exposed embryos of 10 families from each of two populations to variable and constant thermal regimes. These represented both experienced temperatures by each population, and predicted temperatures under plausible future conditions based on a warming scenario from the downscaled global climate model (MIROC A1B scenario). We did not find evidence of thermal local adaptation during the embryonic stage for developmental rate or survival. Within treatments, populations hatched within 1 day of each other, on average, and among treatments, did not differ in survival in response to temperature. We did detect plasticity to temperature; embryos developed 2.5 times longer (189 days) in the coolest regime compared to the warmest regime (74 days). We also detected variation in developmental rates among families within and among temperature regimes, indicating heritable plasticity. Families exhibited a strong positive relationship between thermal variability and phenotypic variability in developmental rate but body length and mass at hatching were largely insensitive to temperature. Overall, our results indicated a lack of thermal local adaptation, but a presence of plasticity in populations experiencing contrasting conditions, as well as family-specific heritable plasticity that could facilitate adaptive change.

Modelling dendritic ecological networks in space: an integrated network perspective.

Dendritic ecological networks (DENs) are a unique form of ecological networks that exhibit a dendritic network topology (e.g. stream and cave networks or plant architecture). DENs have a dual spatial representation; as points within the network and as points in geographical space. Consequently, some analytical methods used to quantify relationships in other types of ecological networks, or in 2-D space, may be inadequate for studying the influence of structure and connectivity on ecological processes within DENs. We propose a conceptual taxonomy of network analysis methods that account for DEN characteristics to varying degrees and provide a synthesis of the different approaches within the context of stream ecology. Within this context, we summarise the key innovations of a new family of spatial statistical models that describe spatial relationships in DENs. Finally, we discuss how different network analyses may be combined to address more complex and novel research questions. While our main focus is streams, the taxonomy of network analyses is also relevant anywhere spatial patterns in both network and 2-D space can be used to explore the influence of multi-scale processes on biota and their habitat (e.g. plant morphology and pest infestation, or preferential migration along stream or road corridors).

Hypoxia vulnerability in the salmon watersheds of Southeast Alaska.

The frequency of dissolved oxygen depletion events (hypoxia) in coastal aquatic ecosystems has risen dramatically since the late 20th century, yet the causes and consequences of hypoxia for some culturally and economically important species remain poorly understood. In rivers, oxygen depletion can be caused by high densities of spawning Pacific salmon (Oncorhynchus spp.) consuming oxygen faster than can be replaced by reaeration. This process may be exacerbated when salmon densities are artificially inflated, such as when hatchery-origin salmon stray into rivers instead of returning to hatcheries. In Southeast Alaska, hatchery salmon production has increased rapidly since the 1970s, with over 553 million chum salmon (O. keta) and 64 million pink salmon (O. gorbuscha) released in 2021 alone. Straying is pervasive in streams with outlets <25 km from nearshore marine hatchery release sites. Using a previously ground-truthed mechanistic model of dissolved oxygen dynamics, we examined how water temperature and low-flow channel hydraulics contribute to hypoxia vulnerability. We then applied the model to predict hypoxia vulnerability for watersheds within 25 km of hatchery salmon release points, where straying salmon spawner densities are expected to be higher and promote dissolved oxygen depletion. Our model predicted that low-gradient stream reaches, regardless of water temperature, are the most prone to hypoxia due to low reaeration rates. Our spatial analysis determined that nearly 17,000 km of anadromous-accessible stream reaches are vulnerable to high densities of hatchery-origin salmon based on 2021 release sites. To our knowledge, this study is the first to map the spatial variation of hypoxia vulnerability in anadromous watersheds, identify habitat conditions most likely to promote hypoxia, and provide a repeatable analytical approach to identify hypoxia-prone stream reaches that can be updated as empirical data sets improve.

Genomics reveal the origins and current structure of a genetically depauperate freshwater species in its introduced Alaskan range.

Invasive species are a major threat to global biodiversity, yet also represent large-scale unplanned ecological and evolutionary experiments to address fundamental questions in nature. Here we analyzed both native and invasive populations of predatory northern pike (Esox lucius) to characterize landscape genetic variation, determine the most likely origins of introduced populations, and investigate a presumably postglacial population from Southeast Alaska of unclear provenance. Using a set of 4329 SNPs from 351 individual Alaskan northern pike representing the most widespread geographic sampling to date, our results confirm low levels of genetic diversity in native populations (average 𝝅 of 3.18 × 10-4) and even less in invasive populations (average 𝝅 of 2.68 × 10-4) consistent with bottleneck effects. Our analyses indicate that invasive northern pike likely came from multiple introductions from different native Alaskan populations and subsequently dispersed from original introduction sites. At the broadest scale, invasive populations appear to have been founded from two distinct regions of Alaska, indicative of two independent introduction events. Genetic admixture resulting from introductions from multiple source populations may have mitigated the negative effects associated with genetic bottlenecks in this species with naturally low levels of genetic diversity. Genomic signatures strongly suggest an excess of rare, population-specific alleles, pointing to a small number of founding individuals in both native and introduced populations consistent with a species' life history of limited dispersal and gene flow. Lastly, the results strongly suggest that a small isolated population of pike, located in Southeast Alaska, is native in origin rather than stemming from a contemporary introduction event. Although theory predicts that lack of genetic variation may limit colonization success of novel environments, we detected no evidence that a lack of standing variation limited the success of this genetically depauperate apex predator.

Colonization and extinction in dynamic habitats: an occupancy approach for a Great Plains stream fish assemblage.

Despite the importance of habitat in determining species distribution and persistence, habitat dynamics are rarely modeled in studies of metapopulations. We used an integrated habitat-occupancy model to simultaneously quantify habitat change, site fidelity, and local colonization and extinction rates for larvae of a suite of Great Plains stream fishes in the Arikaree River, eastern Colorado, USA, across three years. Sites were located along a gradient of flow intermittency and groundwater connectivity. Hydrology varied across years: the first and third being relatively wet and the second dry. Despite hydrologic variation, our results indicated that site suitability was random from one year to the next. Occupancy probabilities were also independent of previous habitat and occupancy state for most species, indicating little site fidelity. Climate and groundwater connectivity were important drivers of local extinction and colonization, but the importance of groundwater differed between periods. Across species, site extinction probabilities were highest during the transition from wet to dry conditions (range: 0.52-0.98), and the effect of groundwater was apparent with higher extinction probabilities for sites not fed by groundwater. Colonization probabilities during this period were relatively low for both previously dry sites (range: 0.02-0.38) and previously wet sites (range: 0.02-0.43). In contrast, no sites dried or remained dry during the transition from dry to wet conditions, yielding lower but still substantial extinction probabilities (range: 0.16-0.63) and higher colonization probabilities (range: 0.06-0.86), with little difference among sites with and without groundwater. This approach of jointly modeling both habitat change and species occupancy will likely be useful to incorporate effects of dynamic habitat on metapopulation processes and to better inform appropriate conservation actions.

Vulnerability of Pacific salmon to invasion of northern pike (Esox lucius) in Southcentral Alaska.

The relentless role of invasive species in the extinction of native biota requires predictions of ecosystem vulnerability to inform proactive management strategies. The worldwide invasion and range expansion of predatory northern pike (Esox lucius) has been linked to the decline of native fishes and tools are needed to predict the vulnerability of habitats to invasion over broad geographic scales. To address this need, we coupled an intrinsic potential habitat modelling approach with a Bayesian network to evaluate the vulnerability of five culturally and economically vital species of Pacific salmon (Oncorhynchus spp.) to invasion by northern pike. This study was conducted along 22,875 stream km in the Southcentral region of Alaska, USA. Pink salmon (O. gorbuscha) were the most vulnerable species, with 15.2% (2,458 km) of their calculated extent identified as "highly" vulnerable, followed closely by chum salmon (O. keta, 14.8%; 2,557 km) and coho salmon (O. kisutch, 14.7%; 2,536 km). Moreover, all five Pacific salmon species were highly vulnerable in 1,001 stream km of shared habitat. This simple to implement, adaptable, and cost-effective framework will allow prioritizing habitats for early detection and monitoring of invading northern pike.

FiCli, the Fish and Climate Change Database, informs climate adaptation and management for freshwater fishes.

Inland fishes provide important ecosystem services to communities worldwide and are especially vulnerable to the impacts of climate change. Fish respond to climate change in diverse and nuanced ways, which creates challenges for practitioners of fish conservation, climate change adaptation, and management. Although climate change is known to affect fish globally, a comprehensive online, public database of how climate change has impacted inland fishes worldwide and adaptation or management practices that may address these impacts does not exist. We conducted an extensive, systematic primary literature review to identify peer-reviewed journal publications describing projected and documented examples of climate change impacts on inland fishes. From this standardized Fish and Climate Change database, FiCli (pronounced fick-lee), researchers and managers can query fish families, species, response types, or geographic locations to obtain summary information on inland fish responses to climate change and recommended management actions. The FiCli database is updatable and provides access to comprehensive published information to inform inland fish conservation and adaptation planning in a changing climate.

An Interferometric Synthetic Aperture Radar (InSAR) Habitat Suitability Model to Identify Overwinter Conditions for Coregonine Whitefishes in Arctic Lagoons.

Lagoons provide critical habitats for many fishes, including coregonine whitefishes, which are a mainstay in many subsistence fisheries of rural communities in Arctic Alaska. Despite their importance, little is known about the overwintering habits of whitefishes in Arctic Alaska due to the challenges associated with sampling during winter. We developed a habitat suitability (HS) model to understand the potential range of physical conditions that whitefishes experience during the Arctic winter, using three indicator lagoons that represent a range of environmental characteristics. The HS model was built using a three-step approach. First, remote sensing that uses interferometric synthetic aperture radar (InSAR) identified areas of floating and bottomfast ice. Second, through in-field ground-truthing, we confirmed the presence and quality of liquid water (water depth, temperature, and dissolved oxygen) beneath the ice cover. Third, we assessed the suitability of that liquid water as habitat for whitefishes based on published literature and expert interpretation of water quality parameters. InSAR determined that 0, 65.4, and 88.2% of the three lagoons were composed of floating ice corresponding with areas of liquid water beneath a layer of ice. The HS model indicated that all three lagoons had reduced suitability as whitefish habitat in winter than in summer due to the loss of habitat because of the presence of bottomfast ice and a reduction in the quality of liquid water due to cold temperatures, high salinities, and low dissolved oxygen levels. However, only the shallowest lagoon had lethal conditions and zero suitability as whitefish habitat. The methods outlined here provide a simple, cost-effective method to identify habitats that consistently provide critical winter habitat and integrate remote sensing in a HS model framework.

The role of density-dependent and -independent processes in spawning habitat selection by salmon in an Arctic riverscape.

Density-dependent (DD) and density-independent (DI) habitat selection is strongly linked to a species' evolutionary history. Determining the relative importance of each is necessary because declining populations are not always the result of altered DI mechanisms but can often be the result of DD via a reduced carrying capacity. We developed spatially and temporally explicit models throughout the Chena River, Alaska to predict important DI mechanisms that influence Chinook salmon spawning success. We used resource-selection functions to predict suitable spawning habitat based on geomorphic characteristics, a semi-distributed water-and-energy balance hydrologic model to generate stream flow metrics, and modeled stream temperature as a function of climatic variables. Spawner counts were predicted throughout the core and periphery spawning sections of the Chena River from escapement estimates (DD) and DI variables. Additionally, we used isodar analysis to identify whether spawners actively defend spawning habitat or follow an ideal free distribution along the riverscape. Aerial counts were best explained by escapement and reference to the core or periphery, while no models with DI variables were supported in the candidate set. Furthermore, isodar plots indicated habitat selection was best explained by ideal free distributions, although there was strong evidence for active defense of core spawning habitat. Our results are surprising, given salmon commonly defend spawning resources, and are likely due to competition occurring at finer spatial scales than addressed in this study.

Spatial ecological processes and local factors predict the distribution and abundance of spawning by steelhead (Oncorhynchus mykiss) across a complex riverscape.

Processes that influence habitat selection in landscapes involve the interaction of habitat composition and configuration and are particularly important for species with complex life cycles. We assessed the relative influence of landscape spatial processes and local habitat characteristics on patterns in the distribution and abundance of spawning steelhead (Oncorhynchus mykiss), a threatened salmonid fish, across ∼15,000 stream km in the John Day River basin, Oregon, USA. We used hurdle regression and a multi-model information theoretic approach to identify the relative importance of covariates representing key aspects of the steelhead life cycle (e.g., site access, spawning habitat quality, juvenile survival) at two spatial scales: within 2-km long survey reaches (local sites) and ecological neighborhoods (5 km) surrounding the local sites. Based on Akaike's Information Criterion, models that included covariates describing ecological neighborhoods provided the best description of the distribution and abundance of steelhead spawning given the data. Among these covariates, our representation of offspring survival (growing-season-degree-days, °C) had the strongest effect size (7x) relative to other predictors. Predictive performances of model-averaged composite and neighborhood-only models were better than a site-only model based on both occurrence (percentage of sites correctly classified = 0.80±0.03 SD, 0.78±0.02 vs. 0.62±0.05, respectively) and counts (root mean square error = 3.37, 3.93 vs. 5.57, respectively). The importance of both temperature and stream flow for steelhead spawning suggest this species may be highly sensitive to impacts of land and water uses, and to projected climate impacts in the region and that landscape context, complementation, and connectivity will drive how this species responds to future environments.