Abstract
Anadromous fishes have been blocked from the middle Snake River basin since the construction of flood control, irrigation, and hydroelectric projects during the 19th and 20th centuries, culminating with the construction of Hells Canyon Dam (river kilometer [rkm] 398) in 1967. Seven large watersheds in the blocked area of the basin are under consideration for Pacific salmon reintroduction. The primary objective of this study was to identify and characterize potential reintroduction sites with high-quality rearing and spawning habitat for Chinook salmon (Oncorhynchus tshawytscha) and steelhead (O. mykiss) upstream of the Hells Canyon Complex in Idaho, Oregon, and Nevada, USA. We created and tested habitat models to predict rearing presence/absence, rearing abundance, and spawning presence/absence using channel morphology, hydrology, and stream temperature variables obtained from regional peer-reviewed datasets. Habitat models were trained with salmonid presence and abundance records collected in the lower Snake River basin (downstream of Hells Canyon Dam), where anadromous fish can currently access, from 1993 to 2011. Model performance was tested with set-aside data comprised of randomly selected reaches and independent environmental DNA data. An index model was created in the middle Snake River basin for each species by combining results from the habitat models. Modeling covariates differed by species and life stage and included different combinations of August stream temperature, summer flow, channel slope, and quadratic terms for temperature and slope. The habitat models predicted high versus low Chinook salmon and steelhead probability and abundance with 66% to 85% accuracy depending on species and life stage. The index models predicted a total of 2,887 km of Chinook salmon habitat and 2,434 km of steelhead habitat in the blocked area. The three basins in the blocked area with the greatest amount of predicted habitat were the Powder River, South Fork Payette River, and North Fork Payette River for Chinook salmon, and the Powder River, South Fork Boise River, and South Fork Payette River for steelhead. The modeling approach presented here is complementary to other planning efforts for Chinook salmon and steelhead reintroduction in the blocked area of the Snake River basin, and similar approaches may be useful for reintroduction planning in other systems.
1 Introduction
Habitat loss is a leading cause of declining biodiversity and species abundance worldwide, and anadromous fish species are particularly vulnerable given their unique life histories (; ; ; ). These fish have critical life stages (migration, spawning, incubation and rearing) that occur in freshwater habitats adversely affected by human disturbances such as dam construction, mining, logging, water withdrawal and power generation (; ; ). In western North America, large impassable dams have drastically reduced access to historical habitats of Pacific salmon (Oncorhynchus spp.), contributing to the loss or decline of many populations (; ; ; ). For example, Chinook salmon (O. tshawytscha) are currently restricted from accessing 50–80% of their historical habitat in the Columbia and Snake River basins and in California’s Central Valley (; ; ). Pacific salmon are economically, culturally, and ecologically important, and therefore are the focus of recovery efforts throughout much of their historical range (; ; ; ; ; ).
Salmon recovery efforts are not only critical for biodiversity but also for preserving the heritage and well-being of local Indigenous peoples who have long depended on salmon for cultural, spiritual, and subsistence purposes (; ; ). The loss of salmon has disrupted traditional fishing practices, ceremonies, and diets, resulting in cultural and economic hardships for many Tribal communities (). Thus, in numerous locations, Tribal entities have initiated salmon reintroductions to restore viable populations in historically important areas where they have been absent for decades (; ; ; ). However, salmon remain absent from large swaths of their historical habitat. For example, in the Snake River basin, dams constructed during 1892–1967 eliminated access for Chinook salmon to an estimated 8,249 river kilometers (rkm) of high-quality stream habitat (). This loss adversely affected numerous Tribes including, but not limited to, the Burns Paiute, Fort McDermitt Paiute-Shoshone, Shoshone-Bannock Tribes of the Fort Hall Reservation, and Shoshone-Paiute Tribes of the Duck Valley Reservation (hereafter “Upper Snake River Tribes” [USRT]) who have been forced to exist without salmon for decades (, ; ). In recent years, USRT has initiated efforts to examine salmon reintroduction opportunities throughout the basin, which have focused on assessing historical losses as well as habitat quantity and quality in the blocked area (; ). Because recent information on Pacific salmon distribution and abundance is often lacking in areas where these species have been extirpated, habitat assessments based on environmental variables can be valuable for identifying currently unoccupied reaches with the potential to support reintroduction. Habitat assessments have often been the first step in other salmon reintroductions, as they provide necessary information to inform decisions regarding reintroduction site selection, hatchery fish production, and collection and transport programs to move fish around impassable dams (; , ; ; ).
Field assessments of salmon habitat across large riverscapes are time-intensive, costly, and often impractical, therefore habitat modeling approaches using Geographic Information Systems (GIS) are commonly used as a first step to broadly identify and characterize potential habitat in the reintroduction area (; ). For example, Intrinsic Potential (IP) models have been widely applied across the Pacific Northwest (USA) to assess salmon habitat availability. IP models integrate expert opinion and published literature on species-habitat relationships with geomorphic, climatic, and hydrologic variables from existing spatial datasets to identify and classify stream reaches based on physical habitat potential (e.g., poor, fair, good) (; ; ). Alternatives to IP models include predictions based on empirically derived statistical relationships between habitat variables and species presence or abundance. These relationships can be derived from field sampling datasets (e.g., spawner surveys, beach seines, or electrofishing) and existing habitat data (; ; ). Whereas IP models produce nominal outcomes, empirical models produce continuous probabilities (e.g., 70% probability of salmonid habitat) that are more flexible and information rich. Decisions related to which approach should be used largely depend on the underlying datasets from which models are derived. IP models are a good choice when training datasets are sparsely populated or absent altogether, while empirical models are well suited for large, robust training datasets that allow for statistical analysis and verification. Importantly, the actual distribution and abundance of a species depend on numerous factors beyond the set of attributes often included in habitat models, which are often limited. For example, species distribution and abundance may be affected by unmeasured habitat variables (), prey availability (), inter-and intra-specific interactions (; ; ), or the overall abundance of the species in question (). When species abundance is low, areas that are predicted as “good” habitat may remain unoccupied, even given robust training datasets and high model accuracy (e.g., ). Despite these limitations, habitat models based on empirical data can provide valuable baseline information on habitat quality in currently unoccupied areas.
In this study we developed statistical relationships between habitat variables and empirical data on spring Chinook salmon and summer steelhead (O. mykiss) in the lower Snake River basin, the in-basin region currently accessible to anadromous fish, and projected those relationships to the middle Snake River basin to quantify suitable habitat for these species that are being considered for reintroduction (; ). Our objectives were to (1) identify habitat suitable for two life stages (juvenile rearing and adult spawning) of Chinook salmon and steelhead in the blocked area of the Snake River basin, and (2) develop a management-relevant habitat index using peer-reviewed, publicly available datasets. While this approach was developed to address specific needs related to potential reintroduction in the Snake River basin, these methods provide a useful framework for assessing habitat quantity across large spatial areas of interest where field sampling may be unfeasible.
2 Methods
2.1 Project area
The Snake River basin is the largest sub-basin of the Columbia River (in Canada and the United States (U.S.)) and spans six U.S. states including Wyoming, Idaho, Utah, Nevada, Oregon, and Washington (Figure 1; ). The Snake River originates in the Rocky Mountains and flows ~1,674 km downstream to its confluence with the Columbia River near Pasco, Washington (). The watershed covers approximately 280,000 km² and comprises approximately 41% of the total Columbia River basin (). The climate of the basin ranges from semi-arid to alpine and is heavily influenced by the rain shadow of the Cascade Range. Precipitation is highly variable, ranging from 15–25 cm in desert areas, such as the Snake River Plain in southern Idaho and the lower Snake River in southeastern Washington, to over 241 cm in high mountain headwaters such as the Teton Range (; ). Air temperature extremes range from approximately -18 °C in winter to over 38 °C in summer, and high-elevation areas serve as critical cold-water refuges for sensitive fish species as low altitudes warm. Approximately 30% of the watershed is farmland; the Snake River Plain is a global leader in irrigated agriculture with over 3.8 million acres of crops including potatoes, sugar beets, and wheat (). Roughly 50% of the basin consists of semi-arid shrubland and rangeland used primarily for livestock grazing and another 15% is forested (largely managed by the U.S. Forest Service); urban and industrial areas account for only about 1% of the land use, despite housing major cities like Boise and Idaho Falls ().
Figure 1
The Snake River basin supports 35 native fish species including Chinook salmon, sockeye salmon (O. nerka), and steelhead (). These fish undertake an inland journey of over 1,400 km, and the basin comprises approximately 50% of all available cold-water habitat for west coast Pacific salmon (). Shoshone Falls (rkm 990), formed about 14,000 years ago, marks the historical upper limit of fish migration in the Snake River. Hells Canyon Dam (rkm 398) was completed in 1967 () and is owned and operated by Idaho Power. Although Hells Canyon Dam is the most downstream barrier to anadromy on the Snake River, a series of dams constructed by the Bureau of Reclamation in the early 1900s blocked access to major tributaries, including the Boise, Burnt, Malheur, Owyhee, Powder and Payette rivers (). Throughout this work, “lower basin” refers to the Snake River basin from Hells Canyon Dam downstream to the confluence with the Columbia River, and “middle basin” refers to the Snake River from Shoshone Falls to Hells Canyon Dam (Figure 1). Almost half of the middle basin is latitudinally similar to the lower basin, between 44 and 45.5 degrees north latitude. The Owyhee and Bruneau basins are further south than the Payette, Boise, and Malheur basins and do not overlap latitudinally with the lower basin. The middle basin includes 12 damsheds (defined by the watershed area upstream of a major dam) that range in size from 803 km2 (Unity) to 28,156 km2 (Owhyee) (). At the HUC-8 scale, there are 29 basins that range in size from 750 km2 (Middle Snake-Payette) to 8,735 km2 (Bruneau). These two spatial scales overlap, such that some damsheds contain multiple HUC-8 basins and some HUC-8 basins include multiple damsheds.
2.2 Data acquisition
We integrated six datasets for habitat modeling: (1) the National Hydrography Database Plus (NHDPlus, 100 km resolution, https://www.epa.gov/waterdata/nhdplus-national-hydrography-dataset-plus) depicted stream networks (); (2) the NorWeST database (https://research.fs.usda.gov/rmrs/projects/norwest) provided estimated August stream temperatures (); (3) the Variable Infiltration Capacity dataset (VIC, https://research.fs.usda.gov/rmrs/products/dataandtools/archived-western-streamflow-metrics-interior-columbia-river-basin) provided mean summer flow estimates (; ); (4) a dataset stored in Dryad (https://datadryad.org/dataset/doi:10.5061/dryad.d0s7k) contained georeferenced occurrence and rearing abundance records for Chinook salmon and O. mykiss (steelhead are anadromous O. mykiss) throughout the lower Snake River basin (); (5) an environmental DNA (eDNA) dataset (https://www.fs.usda.gov/rds/archive/Catalog/RDS-2018-0010) contained presence-absence surveys for both species (), and (6) StreamNet, a geospatial dataset (https://www.streamnet.org/) provided information about spring Chinook salmon and steelhead spawning locations in the lower Snake River basin (Figure 2) (). StreamNet contained spawning data for Chinook salmon in 11,526 NHDPlus reaches (3,164 presence, 8,362 absence) and spawning data for steelhead in 25,092 reaches (10,734 presence, 14,358 absence).
Figure 2
To identify potential Chinook salmon and steelhead habitat in the blocked area, habitat models were developed and tested in the lower basin and projected to the middle basin. The lower basin contained ~ 57,000 NHDPlus reaches, of which ~33% (18,735) were second-order or larger perennial streams (). The middle basin contained ~67,000 NHDPlus reaches, of which ~21% (13,782) were second-order or larger perennial streams. A comparison of physical attributes of perennial second-order or larger streams revealed lower basin streams were steeper on average and had cooler August stream temperatures than middle basin streams, but summer flow and Strahler stream order were similar (Figure 3). Approximately 80% of intermittent streams in the lower basin were first order compared to ~74% in the middle basin.
Figure 3
The Dryad fish survey data were compiled from multiple organizations and included snorkeling and backpack electrofishing records from perennial streams with summer flows <32 m3/sec (). The range of flows included in the dataset resulted in exclusion of mainstem reaches from rearing habitat projections (see section 3.4 below). Some reaches were sampled multiple times per year, or once per year over many years, while other reaches were only sampled once. We eliminated pseudo-replication by grouping fish records by NHDPlus reach, assigned an occurrence value of 1 to a reach if there was an occurrence record during the sampling period and a zero otherwise, and averaged fish abundance records across all sampling occasions within a reach. Abundance summaries were based on 6,189 juvenile Chinook salmon survey records obtained from 455 unique NHDPlus reaches, and 5,182 juvenile steelhead records obtained from 478 NHDPlus reaches (Figure 1). Juvenile Chinook salmon average abundances ranged from 0 to 707 per NHDPlus reach (mean of 31.5) and juvenile steelhead average abundances ranged from 0 to 691 per NHDPlus reach (mean of 53.2). Eighty-six percent of Chinook salmon sample reaches and 84% of steelhead sample reaches were sampled more than once.
We used eDNA data to independently verify the rearing presence-absence habitat models (). The eDNA data were collected by staff representing numerous agencies that have partnered with the National Genomics Center for Wildlife and Fish Conservation (NGC) to create the eDNA Atlas database with coverage throughout the United States. We grouped eDNA records by NHDPlus reach, assigned an occurrence value of 1 to a reach if there was an occurrence record during the sampling period and a 0 otherwise. This resulted in a total of 393 NHDPlus reaches for Chinook salmon and 576 reaches for O. mykiss. Importantly, eDNA sampling was conducted separately from the fish sampling that comprised the Dryad database and thus provided independent verification of the presence-absence models.
The six datasets described here were linked by reach using NHDPlus composite outlet ID after excluding firstorder, intermittent reaches to ensure conservative and reliable habitat estimates (, ). A GIS database was then constructed containing summer flow from June through September [FLOW], August stream temperature [S1], channel slope [SLOPE], fish presence-absence, rearing abundance, spawning presence/absence, and eDNA sampling results. Importantly, VIC and NorWeST modeled datasets contained 1993 to 2011 average summer flow and August stream temperatures that corresponded with decades when the fish survey information was collected (; ).
2.3 Habitat model development
We developed three separate generalized linear models (GLMs) for each species, with spawning (presence/absence), rearing (presence/absence), or rearing abundance (average counts per reach) as response variables. Model selection resulted in a total of six GLMs, which were assembled into a single habitat index model. All analyses, model training, and verification were conducted in the R statistical environment, version 4.5.1 () and ArcGIS Pro Version 3.7 (). The GLMs were constructed (trained) with a 70% randomized draw of NHDPlus reaches that contained data for focal species and life stage. The remaining 30% of reaches were set aside and used to test the predictive performance of the models constructed with the training data.
Binomial logistic regression was used to model spawning and rearing presence-absence for each species () and zero inflated negative binomial regression was used to model rearing abundance (). We tested nine alternative combinations of explanatory variables for each of the six models (Table 1) and used Akaike’s information criterion (AIC) to identify the combination of variables that best explained each response variable (). The full model included SLOPE, S1, log-transformed flow (lnFLOW) to account for differential responses across the range of FLOW values, and quadratic terms for SLOPE and S1, as explanatory variables. Quadratic terms for SLOPE and S1 were evaluated to identify potential nonlinear relationships with habitat. A pseudo R-squared statistic () was generated for each alternative model to quantify explained variability, and leave-one-out cross-validation (LOOCV), which captures absolute error associated with a prediction (), was conducted to confirm model selection results.
Table 1
| Model | Explanatory variables |
|---|---|
| 1 | lnFLOW, SLOPE, S1 |
| 2 | lnFLOW, SLOPE |
| 3 | S1, S12 |
| 4 | lnFLOW, S1 |
| 5 | lnFLOW, SLOPE, SLOPE2, S1, S12 |
| 6 | lnFLOW, S1, S12 |
| 7 | S1 |
| 8 | lnFLOW |
| 9 | SLOPE |
Alternative combinations of explanatory variables that were compared for Chinook salmon and steelhead spawning and rearing models.
2.4 Habitat model verification
The best performing logistic models for rearing and spawning were verified using receiver operating characteristic (ROC) curves to evaluate overall model accuracy based on true positive and true negative rates across a range of probability thresholds (). ROC curves were constructed using the 30% randomized set-aside of NHDPlus reaches from the Dryad dataset, which included 136 set-aside reaches for Chinook salmon rearing, 169 for steelhead rearing, 3,640 for Chinook salmon spawning, and 8,364 for steelhead spawning. For the logistic rearing models, a second ROC curve was generated using the eDNA dataset, which included presence/absence data for Chinook salmon in 393 NHDPlus reaches and presence/absence data for O. mykiss in 576 NHDPlus reaches. The area under the curve (AUC) was calculated based on the ROC curves to summarize and compare overall classification performance for each of the logistic models. The predictive accuracy of each logistic model was evaluated by identifying a probability threshold that balanced sensitivity and specificity of the model projections based on the set-aside data (i.e., omission and commission errors) and then calculating the mean of model sensitivity and specificity [(sensitivity + specificity)/2] at the given probability threshold. The rearing abundance models were verified by calculating Pearson’s correlation coefficient (r) between predicted and observed rearing abundances in 30% set-aside reaches. Additionally, the predictive accuracy of each abundance model was tested based on its ability to predict abundances above versus below the median observed abundance values in the set-aside reaches. Similar to the logistic models, predictive accuracy for the abundance models was evaluated using ROC curves and the mean of model sensitivity and specificity.
2.5 Index model
Following model development and verification, spawning and rearing models were projected into the blocked middle basin to identify potential habitat. To make the habitat predictions as accurate as possible, we removed all areas known to have natural blockages to anadromy from the habitat projections (). For each species, we used GLM results to create three binary habitat prediction maps (spawning presence-absence, rearing presence-absence, and rearing abundance) in the middle basin. For the presence-absence models, we converted the predicted probability output for each reach into binary format by selecting a probability threshold that balanced sensitivity and specificity (). Predicted values above the threshold were assigned a 1 and predicted values below the cut-point were assigned to 0. For the rearing density models, we converted the predicted densities into binary format by assigning a 1 to any values above the median observed density and a 0 to any values below the median observed density from the training datasets. We then overlaid the three resulting binary outputs to create a habitat index value from 0 to 3 for each focal species. An index value of 0 indicated that a NHDPlus reach did not contain predicted habitat based on any of the models, whereas a value of 3 indicated that all three binary models predicted habitat in an NHDPlus reach. Intermediate index values (1 or 2) indicated that predicted habitat existed in an NHDPlus reach but was not supported by all three models. Following examination of the index model projections, probability and abundance thresholds for the rearing and spawning models were re-evaluated and adjusted if needed, based on historical information about fish distribution in the middle basin. We summarized predicted index-3 habitat quantity (stream length) in the middle basin at two spatial scales: (1) damshed, and (2) HUC-8 hydrological units, which represent large subbasins inside the project area (Figure 4).
Figure 4
3 Results
3.1 Habitat model development
The final structure of the best performing models varied by species, life stage, and presence/absence versus abundance models (Table 2). In general, the alternative model with the lowest AIC value was selected. The single exception was the Chinook salmon rearing abundance model where the AIC ranking conflicted with LOOCV results. In this case, the simpler model with slightly higher AIC (delta = 7.2) had lower LOOCV, therefore it was selected in lieu of the more complex model. For steelhead rearing presence, the best performing model had only slightly lower AIC (delta = 0.8), but this model was also simpler than the closest competing model and was therefore selected. For the Chinook salmon rearing abundance model and the steelhead rearing presence model, AUC was calculated for the top two competing models. Chinook salmon and steelhead spawning and rearing presence were negatively associated with SLOPE and positively associated with S1 and lnFLOW (Table 3; Supplementary Figure 1). Quadratic terms in the final presence/absence models indicated nonlinear effects of SLOPE and S1 on Chinook salmon rearing and spawning presence and steelhead spawning presence. For the rearing abundance models, the coefficients in the zero-inflation component predicted the probability that fish were absent from the reach, and the coefficients in the count component predicted the abundance at sites where fish were predicted to be present. Chinook salmon rearing abundance at occupied sites was primarily driven by SLOPE, which had a strong negative effect on abundance. Steelhead rearing abundance at occupied sites was positively associated with FLOW and S1 and exhibited unimodal relationships with SLOPE and S1 (Supplementary Figure 1).
Table 2
| Chinook salmon rearing (presence/absence) | ||||||
|---|---|---|---|---|---|---|
| Model | ΔAIC | R2 | LOOCV | AUC | Threshold | Accuracy |
| 5* | 0 | 0.692 | 0.104 | 0.922 | 0.5 | 0.859 |
| 1 | 8.9 | 0.665 | 0.105 | |||
| 2 | 10.6 | 0.657 | 0.107 | |||
| 6 | 29.5 | 0.620 | 0.119 | |||
| 4 | 37.8 | 0.597 | 0.119 | |||
| 8 | 43.2 | 0.579 | 0.125 | |||
| 9 | 76.1 | 0.497 | 0.144 | |||
| 3 | 104.3 | 0.425 | 0.169 | |||
| 7 | 115.7 | 0.386 | 0.170 | |||
| Steelhead rearing (presence/absence) | ||||||
| Model | ΔAIC | R2 | LOOCV | AUC | Threshold | Accuracy |
| 1* | 0 | 0.660 | 0.097 | 0.928 | 0.70 | 0.851 |
| 4 | 0.8 | 0.653 | 0.097 | 0.926 | ||
| 6 | 2.6 | 0.653 | 0.098 | |||
| 5 | 3.8 | 0.660 | 0.099 | |||
| 2 | 28.1 | 0.585 | 0.114 | |||
| 8 | 32.9 | 0.567 | 0.116 | |||
| 7 | 60.7 | 0.491 | 0.136 | |||
| 3 | 62.6 | 0.491 | 0.137 | |||
| 9 | 90.7 | 0.399 | 0.153 | |||
| Chinook salmon rearing (abundance) | ||||||
| Model | ΔAIC | R2 | LOOCV | AUC | Threshold | Accuracy |
| 5 | 0 | 0.603 | 4387.572 | 0.938 | ||
| 1* | 7.2 | 0.584 | 4331.377 | 0.941 | 2 | 0.814 |
| 2 | 14.8 | 0.568 | 4353.705 | |||
| 6 | 78.9 | 0.478 | 4927.045 | |||
| 4 | 79.0 | 0.472 | 4958.561 | |||
| 8 | 87.6 | 0.450 | 4923.418 | |||
| 9 | 93.2 | 0.441 | 4454.841 | |||
| 3 | 159.3 | 0.320 | 5130.909 | |||
| 7 | 266.7 | 0.050 | 5931.058 | |||
| Steelhead rearing (abundance) | ||||||
| Model | ΔAIC | R2 | LOOCV | AUC | Threshold | Accuracy |
| 5* | 0 | 0.566 | 6400.592 | 0.846 | 13 | 0.702 |
| 6 | 6.3 | 0.546 | 6441.224 | |||
| 4 | 13.4 | 0.529 | 7034.149 | |||
| 1 | 15.0 | 0.533 | 7071.667 | |||
| 2 | 52.5 | 0.470 | 6932.647 | |||
| 8 | 54.0 | 0.456 | 6898.454 | |||
| 3 | 78.4 | 0.419 | 6565.397 | |||
| 7 | 86.6 | 0.395 | 7028.945 | |||
| 9 | 135.0 | 0.293 | 7267.927 | |||
| Chinook salmon spawning (presence/absence) | ||||||
| Model | ΔAIC | R2 | LOOCV | AUC | Threshold | Accuracy |
| 5* | 0 | 0.219 | 0.167 | 0.763 | 0.27 | 0.714 |
| 1 | 74.2 | 0.207 | 0.169 | |||
| 6 | 166.2 | 0.193 | 0.171 | |||
| 2 | 232.6 | 0.182 | 0.173 | |||
| 4 | 254.0 | 0.178 | 0.174 | |||
| 8 | 331.1 | 0.165 | 0.176 | |||
| 9 | 669.3 | 0.109 | 0.184 | |||
| 3 | 1148.4 | 0.026 | 0.196 | |||
| 7 | 1250.7 | 0.007 | 0.198 | |||
| Steelhead spawning (presence/absence) | ||||||
| Model | ΔAIC | R2 | LOOCV | AUC | Threshold | Accuracy |
| 5* | 0 | 0.171 | 0.213 | 0.717 | 0.40 | 0.660 |
| 6 | 83.5 | 0.165 | 0.214 | |||
| 4 | 940.9 | 0.103 | 0.225 | |||
| 1 | 941.8 | 0.103 | 0.225 | |||
| 2 | 1047.2 | 0.095 | 0.226 | |||
| 8 | 1070.0 | 0.093 | 0.226 | |||
| 3 | 1951.5 | 0.026 | 0.240 | |||
| 9 | 2077.3 | 0.016 | 0.242 | |||
| 7 | 2269.5 | 0.001 | 0.245 | |||
Model selection criteria and performance metrics for the models for Chinook salmon and steelhead rearing presence/absence, rearing abundance, and spawning presence/absence.
The top models are indicated with an asterisk. Models were selected based on AIC and LOOCV. If AIC scores were equivalent (<2 point difference) or if AIC results conflicted with LOOCV results, then the simpler model was used for habitat projections. Area under the curve (AUC) was calculated based on receiver operating curves for the best performing models. Thresholds were defined by selecting a value that balanced model sensitivity and specificity for the presence/absence models, and by selecting the median observed value for the abundance models. Accuracy indicates the mean of model sensitivity and specificity [(sensitivity + specificity)/2] at the given threshold value.
Table 3
| Rearing presence/absence models |
|---|
| (1) Chinook salmon presence = -12.55 + (0.097 * lnFLOW) – (80.97 * SLOPE) + (404.09 * SLOPE2) + (1.86 * S1) – (0.07 * S12) |
| (2) Steelhead: Presence = -6.37 + (0.99 * lnFLOW) – (11.41 * SLOPE) + (0.50 * S1) |
| Rearing abundance models |
| (3a) Chinook salmon count model = 6.28 + (0.07 * lnFLOW) - (62.70 * SLOPE) - (0.14 * S1) |
| (3b) Chinook salmon zero inflation model = 5.89 – (1.72 * lnFLOW) + (51.44 * SLOPE) – (0.32 * S1) |
| (4a) Steelhead count model = -5.22 + (0.22 * lnFLOW) + (35.23 * SLOPE) – (464.73 * SLOPE2) + (1.16 * S1) - (0.04 * S12) |
| (4b) Steelhead zero inflation model = -1.43 – (1.35 * lnFLOW) + (13.2 * SLOPE) – (56.06 * SLOPE2) + (1.18 * S1) – (0.08 * S12) |
| Spawning presence/absence models |
| (5) Chinook: -3.09 + (0.45 * lnFLOW) - (16.31 * SLOPE) + (38.30 * SLOPE2) + (0.41 * S1) – (0.02 * S12) |
| (6) Steelhead: = -5.84 + (0.32 * lnFLOW) – (1.45 * SLOPE) + (10.07 * SLOPE2) + (0.94 * S1) – (0.04 * S12) |
Intercepts and coefficients for top models of Chinook salmon and steelhead rearing (presence/absence and abundance) and spawning (presence/absence).
lnFLOW is the log of summer flow (average June – September, 1993 - 2011) of each NHDPlus reach.
S1 is average August water temperature (1993 – 2011) of each NHDPlus reach.
SLOPE is average channel slope (%) of NHDPlus reach.
3.2 Habitat model verification
The Chinook salmon and steelhead rearing presence/absence models produced high AUC values (0.922 and 0.928) and predictive accuracy (0.859 and 0.851) on the 30% set-aside reaches (Chinook salmon N = 136; steelhead N = 169) using 0.50 and 0.70 probability thresholds, respectively (Table 2). The rearing presence/absence models achieved predictive accuracies of 0.731 (Chinook salmon, 0.35 probability threshold) and 0.702 (steelhead, 0.35 probability threshold) with the independent eDNA dataset. Predictions from the Chinook salmon rearing abundance model in the set-aside reaches were moderately correlated with observed values (r = 0.436), and observed values above versus below the median were predicted with AUC of 0.941 and accuracy of 0.841. Predictions from the steelhead rearing abundance model were also moderately correlated with observed values in the set-aside reaches (r = 0.385), with values above versus below the median predicted with AUC of 0.846 and accuracy of 0.702 (Table 2, Figure 5). The Chinook salmon and steelhead spawning presence/absence models had somewhat lower AUC values (0.763 and 0.717) and predictive accuracy (0.714 and 0.660) on the set-aside reaches (Chinook salmon N = 3,640, steelhead N = 7,886) at 0.27 and 0.40 probability thresholds, respectively.
Figure 5
3.3 Habitat model projections
Model projections in the middle basin produced distinct patterns for each species and life stage (Supplementary Figures 2, 3). The Chinook salmon rearing presence/absence model produced higher presence probabilities in large tributaries with low channel gradients. The steelhead rearing presence/absence model produced higher model probabilities throughout much of the basin, except for the smallest tributaries at higher elevations. Projected rearing abundance patterns in the middle basin also differed between the two focal species. Chinook salmon rearing densities were predicted to be high in some large and intermediate sized tributaries, but not in smaller tributaries. Steelhead rearing densities were predicted to be high in a wider range of large, intermediate, and small tributaries, but in general, upper elevation tributaries on the eastern edge had low predicted abundance, consistent with the rearing presence/absence projections. Predicted spawning habitat for Chinook salmon generally included large and intermediate tributaries, but smaller tributaries in the northern and eastern portions of the basin also had high predicted probabilities. Predicted steelhead spawning habitat was widespread, but the highest predicted probabilities were in small to intermediate sized tributaries at relatively high elevations.
3.4 Index models and habitat summaries
Based on the initial probability and abundance thresholds for each of the presence and abundance models (Table 2), predictions for index-3 habitat values totaled 1,185 km for Chinook salmon (9.1% of perennial, >1st order streams in the basin), and 5,197 km for steelhead (39% of perennial, >1st order streams in the basin). For Chinook salmon, projected index-3 habitat based on initial threshold values did not include habitat in the upper Malheur and Bruneau river basins. These basins historically produced Chinook salmon, and there is no evidence that habitat capable of supporting Chinook salmon has disappeared. Because the index was highly sensitive to small changes in the threshold value for Chinook salmon rearing presence/absence, the threshold for this model was adjusted from 0.50 to 0.35 to generate the index. This adjustment resulted in 2,887 km of predicted Chinook salmon habitat (<32 m3/sec; 22% of the stream network) and included all major basins that historically produced Chinook salmon (Figure 6). Projected index-3 habitat for steelhead initially included more than half of the middle basin, but these relatively indiscriminate results were minimally informative for reintroduction planning. The steelhead index was highly sensitive to the spawning habitat probability threshold; therefore, we increased the threshold from 0.40 to 0.55, resulting in 2,434 km of predicted habitat (Figure 7; 18% of the stream network).
Figure 6
Figure 7
The two habitat index models produced similar patterns for each focal species by damshed, with the most abundant predicted Chinook salmon and steelhead habitat in the Black Canyon diversion, Brownlee, and Boise River diversion damsheds and least abundant habitat in the Unity, Warm Springs, and Agency Valley damsheds (Figure 8; Table 4). Predicted habitat by HUC-8 differed slightly between the two focal species. For Chinook salmon, the Powder, South Fork Payette, and North Fork Payette basins contained the most predicted habitat, whereas for steelhead, the Powder, South Fork Boise, and South Fork Payette river basins contained the largest amounts of predicted habitat (Figure 8; Table 5).
Figure 8
Table 4
| DSpecies | Damshed | DRKM of index-3 habitat |
|---|---|---|
| Chinook salmon | Agency Valley | 36.1 |
| Black Canyon Diversion | 761.6 | |
| Boise River Diversion | 572.3 | |
| Brownlee | 594.7 | |
| Bruneau | 183.2 | |
| CJ Strike | 174.8 | |
| Hells Canyon | 81.4 | |
| Owyhee | 307.8 | |
| Thief Valley | 125.3 | |
| Unity | 14.1 | |
| Warm Springs | 35.5 | |
| Total = | 2886.8 | |
| Steelhead | Agency Valley | 38.4 |
| Black Canyon Diversion | 697.7 | |
| Boise River Diversion | 594.3 | |
| Brownlee | 408.7 | |
| Bruneau | 111.9 | |
| CJ Strike | 127.3 | |
| Hells Canyon | 123.2 | |
| Owyhee | 166.3 | |
| Thief Valley | 124.1 | |
| Unity | 11.5 | |
| Warm Springs | 30.7 | |
| Total = | 2434.1 |
River kilometers (RKM) of index-3 Chinook salmon and steelhead habitat at the damshed scale in the middle Snake River basin.
Bold text identifies the three damsheds (per species) with the largest amount of predicted habitat.
Table 5
| Species | HUC-8 | RKM of index-3 habitat |
|---|---|---|
| Chinook salmon | Boise-Mores | 51.6 |
| Brownlee | 81.4 | |
| Bruneau | 183.2 | |
| Burnt OR | 111.5 | |
| Crooked-Rattlesnake | 15.0 | |
| Lower Boise | 85.4 | |
| Lower Malheur | 22.1 | |
| Lower Owyhee | 3.0 | |
| MF Payette | 109.8 | |
| Middle Owyhee | 3.6 | |
| Middle Snake-Payette | 2.1 | |
| NF-MF Boise | 246.1 | |
| NF Payette | 278.4 | |
| Payette | 87.5 | |
| Powder | 293.7 | |
| Salmon Falls | 161.5 | |
| SF Boise | 270.0 | |
| SF Owyhee | 83.1 | |
| SF Payette | 285.9 | |
| Upper Malheur | 71.5 | |
| Upper Owyhee | 203.1 | |
| Upper Snake-Rock | 13.2 | |
| Weiser | 202.9 | |
| Willow | 21.1 | |
| Total = | 2886.8 | |
| Steelhead | Boise-Mores | 57.6 |
| Brownlee | 123.2 | |
| Bruneau | 111.9 | |
| Burnt OR | 50.5 | |
| Lower Boise | 76.4 | |
| MF Payette | 123.1 | |
| NF-MF Boise | 233.6 | |
| NF Payette | 255.8 | |
| Payette | 55.4 | |
| Powder | 301.6 | |
| Salmon Falls | 109.5 | |
| SF Boise | 298.4 | |
| SF Owyhee | 23.1 | |
| SF Payette | 263.4 | |
| Upper Malheur | 69.1 | |
| Upper Owyhee | 143.2 | |
| Upper Snake-Rock | 17.8 | |
| Weiser | 120.1 | |
| Willow | 0.233 | |
| Total = | 2434.1 |
River kilometers (RKM) of index-3 Chinook salmon and steelhead habitat at the HUC-8 basin scale in the middle Snake River basin.
Bold text identifies the three HUC-8 units (per species) with the largest amount of predicted habitat.
Mainstem reaches with mean summer flows that exceeded 32 m3/sec were excluded from the habitat index projections because these flow rates fell outside the calibration range of the rearing models. However, these flows were within the calibration range of the spawning habitat model that was built with StreamNet data, and therefore spawning habitat projections could be made for the 1,381 km of mainstem habitat that exceeded 32 m3/sec. Projections from spawning models in these mainstem reaches identified 494 km of predicted Chinook salmon spawning habitat and 138 km of predicted steelhead spawning habitat. Adding predicted mainstem spawning habitat (FLOW >32 m3/sec) to the index-3 habitat predictions increased estimates to 3,381 km of predicted Chinook salmon habitat and 2,572 km of predicted steelhead habitat.
4 Discussion
In this study, a rich set of publicly available datasets, combined with a simple, flexible modeling framework were used to evaluate habitat availability for critical freshwater life stages of Chinook salmon and steelhead in the blocked area of the Snake River basin. The goal of this effort was to generate information useful for advancing salmon reintroduction efforts aimed at ecological and cultural restoration in the basin. Our results indicate that a large amount of habitat is currently available within the blocked area to support juvenile and adult life stages of Chinook salmon and steelhead. Model estimates suggest that at least 2,887 rkm of suitable habitat exists for Chinook salmon and 2,434 rkm of habitat exists for steelhead. The largest habitat quantities were identified in the Powder River, North Fork Payette River, and South Fork Payette River damsheds for Chinook salmon and in the Powder River, South Fork Boise River, and North Fork Payette River damsheds for steelhead. The predicted probability and abundance thresholds used to generate these habitat estimates can be adjusted to be more or less conservative depending on the needs of managers and could be used in conjunction with other assessments (e.g., IP modeling, field surveys) to provide additional lines of evidence for habitat availability. Moreover, the framework provided here demonstrates the utility of publicly available data for evaluating habitat availability for Pacific salmon and other anadromous species in areas that are currently inaccessible.
The blocked area of the Snake River basin represents a vast expanse of ancestral salmon habitat—once biologically rich, culturally vital, and ecologically dynamic (). Restoring access to the blocked area of the Snake River basin, which contains a large amount of the existing coldwater habitat in the Columbia River basin (), has the potential to significantly benefit salmon and steelhead recovery efforts in the region by providing opportunities to increase abundance, productivity, spatial structure, and diversity, the four characteristics that influence the biological viability of salmon populations (). For USRT, this region is not only a homeland but a place shaped by and deeply connected to anadromous salmonids (). The long-standing loss of salmon and steelhead due to dam construction, habitat degradation, and disconnection from the ocean has had profound ecological, cultural, and economic consequences (; ; ). As momentum grows around reintroducing salmon into historical habitats blocked by dams, habitat modeling provides a foundational tool that can be used to guide restoration efforts and support the sovereign rights, health, and heritage of Tribes and other key stakeholders (; ; ).
Our estimates of habitat availability for Chinook salmon (2,287 km) are notably lower (~72% reduction) than those produced during an earlier study (8,249 km; ). For that study, an IP modeling approach was used to estimate historical habitat losses under the assumption that all habitat in the basin was pristine prior to European colonization and development. The IP model incorporated generic parameters that included bankfull channel width, channel gradient, and valley-width-ratio (). For our approach, we included several elements that were intended to refine habitat prediction so the differences in habitat quantity estimates between the two studies are not surprising. Notably, we omitted intermittent, 1st-order streams, and stream reaches located upstream of impassable fish barriers to eliminate habitat that would not have been historically accessible to anadromous fish species. We also explicitly evaluated parameters that are biologically important for Pacific salmon species including summer streamflow, summer water temperature, and channel slope. Additionally, our index estimates are based on empirically derived probability and abundance thresholds for salmon that may be more stringent than the criteria used for previous IP modeling efforts. Although there are notable differences between results from the two studies, they collectively demonstrate the existence of substantial habitat suitable for supporting Chinook salmon and steelhead in the blocked area of the Snake River basin. It is important to note that numerous ecological factors influence how Chinook salmon and steelhead distribute within available habitat (; ; ; ; ). Thus, all reaches that were identified as suitable may not actually be used by salmon and steelhead if reintroduction is implemented in the blocked area. Once salmon and steelhead are present within specific watersheds, additional research and monitoring would be required to identify priority areas based on realized habitat use patterns.
The modeling approach presented here may be applicable for quantifying habitat availability in other systems that currently lack anadromous fish, particularly in spatially expansive areas where comprehensive field surveys may be impractical. This approach effectively identified priority habitats for two anadromous fish species based on habitat variables important for both juvenile and adult life stages in a study area that spanned multiple states and included 29 basins that individually ranged in size from 750 to 8,735 km2. Results generated from this modeling effort provided overall estimates of habitat availability within the study area and identified (to the reach level) potential habitat for both adult spawning and juvenile rearing (Figures 6, 7). This establishes a critical baseline that will benefit future studies by eliminating a large amount of unsuitable habitat from consideration in an expansive area of interest, thereby allowing efforts to focus on high priority reaches that are most likely to support anadromous fish populations.
A key challenge for assessing habitat availability in currently unoccupied areas is the lack of contemporary occurrence data. Our framework addresses this challenge by (1) developing and validating models based on salmon presence and abundance records in a geographical area that is adjacent and latitudinally similar to the focal area and that lies within the same basin, and (2) projecting probability and abundance estimates into the unoccupied focal area. The present study benefitted from robust fish presence, abundance, and eDNA datasets in the lower Snake River basin, publicly available information on physical stream attributes, and apparent similarities between the lower basin and middle basin based on several habitat parameters (Figure 3). Combining the results of three complementary model outputs (rearing presence/absence, rearing abundance, and spawning presence/absence) strengthens inference and may be particularly valuable to managers, as it identifies habitat that is suitable for multiple life stages. However, we also identified several limitations of this approach. For example, our analysis relied on modeled temperature and flow data produced during 1993-2011, and current and future climate conditions may shift the distribution and abundance of suitable habitat in the focal area. Additionally, this modeling framework assumes that fish in the lower basin are biologically representative of fish that were historically present in the middle basin. Additionally, the index model outputs provided here are sensitive to the selection of thresholds that were used to convert continuous probability and abundance predictions into binary habitat maps. Ultimately, threshold choices require judgement based on biological and historical knowledge of the region and the species of interest.
5 Conclusion
In summary, this study provides new information on the quantity of suitable habitat for juvenile and adult life stages of Chinook salmon and steelhead in the blocked area of the Snake River basin while also creating a template that can be useful in other large systems where ecological restoration is of interest. Given the scale of the basin and the complexities associated with reintroducing salmon across this vast, fragmented landscape, prioritization will likely be an essential part of the planning process. Results from this study can be used to support ranking processes focused on prioritizing watersheds or stream segments based on ecological potential, restoration needs, cultural significance, and logistic feasibility. This prioritization can be used to help identify areas where probability of success is greatest for early reintroduction efforts, pilot studies, cultural programs and habitat investments. It also creates a transparent, data-driven approach that can be used to allocate funding and effort toward areas where ecological and cultural returns on investment are highest. These efficiencies and project transparencies will be critical for effective stakeholder buy-in and long-term success.
Statements
Data availability statement
Publicly available datasets were analyzed in this study. This data can be found here: Multiple sources, described and cited in the text.
Author contributions
JRH: Conceptualization, Data curation, Formal analysis, Investigation, Methodology, Validation, Visualization, Writing – original draft, Writing – review & editing. JMH: Writing – original draft, Writing – review & editing. DD: Funding acquisition, Writing – review & editing. CC: Formal analysis, Methodology, Visualization, Writing – review & editing. TK: Supervision, Writing – review & editing.
Funding
The author(s) declared that financial support was received for this work and/or its publication. The author(s) declare that financial support was received for this work and/or its publication. Funding for this project was from a grant provided by the Upper Snake River Tribes via a grant, contract number A20AV00822, provided byfrom the U.S. Department of Indian Affairs Hydro-Compliance Program. (Contract Number A20AV00822).
Acknowledgments
The authors would like to thank the following contributors: Dan Isaak and Aimee Fullerton for their input into modeling approaches; Scott Hauser for his insights into salmon and steelhead habitat in the Snake River basin; Steve Waste for supporting the project and coordinating meetings to discuss research and modeling approaches; James White for reviewing the draft manuscript and providing additional insights regarding habitat modeling approaches; Two reviewers whose efforts improved the final document; Any use of trade, firm, or product names is for descriptive purposes only and does not imply endorsement by the U.S. Government.
Conflict of interest
The author(s) declared that this work was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
Generative AI statement
The author(s) declared that generative AI was not used in the creation of this manuscript.
Any alternative text (alt text) provided alongside figures in this article has been generated by Frontiers with the support of artificial intelligence and reasonable efforts have been made to ensure accuracy, including review by the authors wherever possible. If you identify any issues, please contact us.
Publisher’s note
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Supplementary material
The Supplementary Material for this article can be found online at: https://www.frontiersin.org/articles/10.3389/fevo.2026.1924685/full#supplementary-material
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Summary
Keywords
rearing habitat, reintroduction, salmon, spawning habitat, steelhead
Citation
Hatten JR, Hardiman JM, Daw DA, Couch CE and Kock TJ (2026) Estimating habitat availability for Chinook salmon (Oncorhynchus tshawytscha) and steelhead (O. mykiss) to inform reintroduction planning in the middle Snake River basin, USA. Front. Ecol. Evol. 14:1924685. doi: 10.3389/fevo.2026.1924685
Received
30 June 2026
Revised
04 August 2026
Accepted
24 August 2026
Published
11 September 2026
Volume
14 - 2026
Edited by
Nicholas A. Som, California Polytechnic University, United States
Reviewed by
Tinyiko Cavin Shivambu, University of South Africa, South Africa
Carlos Polivka, Forest Service (USDA), United States
Updates
Copyright
This work is authored by James R. Hatten, Jill M. Hardiman, Claire E. Couch, Tobias J. Kock on behalf of the U.S. Government and as regards Dr. Hatten, Dr. Hardiman, Dr. Couch, Dr. Kock and the U.S. Government, is not subject to copyright protection in the United States. Foreign and other copyrights may apply.
This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) or copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.
*Correspondence: Tobias J. Kock, tkock@usgs.gov
Disclaimer
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