Steelhead Populations Research Articles

Using Genetic Data to Determine Origin for Out-Migrating Smolt and Returning Adult Steelhead Trout (Oncorhynchus mykiss) in a Southeast Alaska Drainage.

Oncorhynchus mykiss is a partially migratory salmonid species, and many migratory populations (known as steelhead) have declined in recent decades in the western United States and Canada. Closely related resident populations (known as rainbow trout) may be an effective resource in the recovery of these declining migratory populations. However, the extent to which different populations of resident rainbow trout produce migratory individuals and how likely these individuals are to return as adults to spawn remains unknown. One limitation to answering these questions is the identification of loci that accurately segregate between migratory and resident populations. To address this limitation, we used existing genomic data from a well-studied population of O. mykiss from Southeastern Alaska (Sashin Creek) to identify loci that segregate between phenotypes. We then utilized Double Mismatch Allele-Specific qPCR (DMAS-qPCR) to genotype 233 smolts out-migrating from Sashin Creek and 99 returning adult steelhead trout across a five-year period to determine (a) the origin of out-migrating smolts and returning adults and (b) to quantify the extent to which the resident population contributes to the migratory population. Our results show that 37.3% of out-migrating smolts were produced from resident parents, whereas 19.3% of returning adults had resident parents. Ultimately, these results demonstrate that resident populations of rainbow trout produce migrant offspring that successfully complete their migration and return to spawn, increasing population sizes and likely improving genetic diversity. Therefore, conservation efforts should consider landlocked resident populations for producing smolts when developing recovery plans for migratory steelhead populations.

Genetic divergence and one-way gene flow influence contemporary evolution and ecology of a partially migratory fish.

Recent work has revealed the importance of contemporary evolution in shaping ecological outcomes. In particular, rapid evolutionary divergence between populations has been shown to impact the ecology of populations, communities, and ecosystems. While studies have focused largely on the role of adaptive divergence in generating ecologically important variation among populations, much less is known about the role of gene flow in shaping ecological outcomes. After divergence, populations may continue to interact through gene flow, which may influence evolutionary and ecological processes. Here, we investigate the role of gene flow in shaping the contemporary evolution and ecology of recently diverged populations of anadromous steelhead and resident rainbow trout (Oncorhynchus mykiss). Results show that resident rainbow trout introduced above waterfalls have diverged evolutionarily from downstream anadromous steelhead, which were the source of introductions. However, the movement of fish from above to below the waterfalls has facilitated gene flow, which has reshaped genetic and phenotypic variation in the anadromous source population. In particular, gene flow has led to an increased frequency of residency, which in turn has altered population density, size structure, and sex ratio. This result establishes gene flow as a contemporary evolutionary process that can have important ecological outcomes. From a management perspective, anadromous steelhead are generally regarded as a higher conservation priority than resident rainbow trout, even when found within the same watershed. Our results show that anadromous and resident O. mykiss populations may be connected via gene flow, with important ecological consequences. Such eco-evolutionary processes should be considered when managing recently diverged populations connected by gene flow.

Contrasting patterns of sequence variation in steelhead populations reflect distinct evolutionary processes.

Multiple evolutionary processes influence genome-wide allele frequencies and quantifying effects of genetic drift, and multiple forms of selection remain challenging in natural populations. Here, we investigate variation at major effect loci in contrast to patterns of neutral drift across a wide collection of steelhead (Oncorhynchus mykiss) populations that have declined in abundance due to anthropogenic impacts. Whole-genome resequencing of 74 populations of steelhead revealed genome-wide patterns (~8 million SNPs) consistent with expected neutral population structure. However, allelic variation at major effect loci associated with adult migration timing (chromosome 28: GREB1L/ROCK1) and age at maturity (chromosome 25: SIX6) reflected how selection has acted on phenotypic variation in contrast with neutral structure. Variation at major effect loci was influenced by evolutionary processes with differing signals between the strongly divergent Coastal and Inland lineages, while allele frequencies within and among populations within the Inland lineage have been driven by local natural selection as well as recent anthropogenic influences. Recent anthropogenic effects appeared to have influenced the frequency of major effect alleles including artificial selection for specific traits in hatchery stocks with subsequent gene flow into natural populations. Selection from environmental factors at various scales has also likely influenced variation for major effect alleles. These results reveal evolutionary mechanisms that influence allele frequencies at major effect loci that are critical for conservation of phenotypic traits and life history variation of this protected species.

Return(s) on investment: Restoration spending in the Columbia River Basin and increased abundance of salmon and steelhead.

The decline in salmon and steelhead populations in the Columbia River Basin has been well documented, as have the decades-long, $9 billion restoration spending efforts by federal and state agencies. These efforts are mainly tied to Endangered Species Act (ESA) mandates for recovery of wild, naturally-spawning threatened or endangered fish species. The impact of these efforts remains poorly understood; many observers, including the federal courts, have long been concerned by the lack of evidence of recovery. Most studies evaluating restoration efforts have examined individual projects for specific species, reaches, or life stages, which limits the ability to make broad inferences at the basin level. There is a need to ask: is there evidence of an overall increase in wild fish abundance associated with the totality of these recovery efforts? To that end, the current study estimates fixed-effects panel regression models of adult returns of four species. Results indicate that restoration spending combined with hatchery production are associated with substantial increases in returning adult fish. Evidence of benefits to wild fish alone, however, require indirect approaches given the commingling of restoration spending with spending on hatchery releases, the impacts of spending on hatchery fish survival, and the density dependence effects of hatchery releases. To accomplish this, the models' predicted adult returns (both hatchery and wild fish) attributed to both spending and hatchery releases are compared to independent estimates of returning hatchery fish based on hatchery survival estimates (smolt-to-adult ratios). The comparison finds the model-predicted levels of adult returns due to spending and hatchery releases do not exceed the survival-rate based estimates for hatcheries alone, so that we are unable to reject the hypothesis of no benefits to wild fish from the restoration spending.

Thiamine Supplementation Improves Survival and Body Condition of Hatchery-Reared Steelhead (Oncorhynchus mykiss) in Oregon.

Early rearing of steelhead (Oncorhynchus mykiss) in Oregon hatcheries is often problematic; fry can become emaciated and die during the period between hatch and first feed. Thiamine (vitamin B1) deficiency has caused early mortality in salmonids; however, the thiamine status of Oregon's steelhead populations is unknown, to date. Of the 26 egg samples from three Oregon hatcheries in 2019, 20 (77%) had thiamine levels < 10 nmol/g, and 13 of those samples (50%) had levels <6.5 nmol/g, suggesting the thiamine deficiency of adult, female steelhead. To investigate if thiamine deficiency was causally related to fry survival, females were injected with buffered thiamine HCl 50 mg/kg prior to spawning; additionally, a subset of eggs were supplemented via bath treatment with thiamine mononitrate (1000 ppm) at spawning. Cumulative fry mortality at 8 weeks post-hatch from thiamine-injected females was 2.9% compared to 13.8% mortality of fry without thiamine supplementation. Fry treated only with the thiamine via bath as eggs had a mortality rate of 6.9%. There were no additional improvements for the survival of fry from injected females that also received a thiamine bath. Furthermore, condition factors were greater in thiamine-supplemented fry than in those that received no thiamine. These data identify thiamine deficiency in Oregon steelhead and suggest supplementation with thiamine can mitigate early rearing mortality.

Predation probabilities and functional responses: How piscivorous waterbirds respond to pulses in fish abundance

How predators respond to changes in prey abundance (i.e., functional responses) is foundational to consumer–resource interactions, predator–prey dynamics, and the stability of predator–prey systems. Predation by piscivorous waterbirds on out-migrating juvenile steelhead trout (Oncorhynchus mykiss) is considered a factor affecting the recovery of multiple Endangered Species Act-listed steelhead populations in the Columbia River basin. Waterbird functional responses, however, may vary by predator species and location, with important implications to predator management strategies. We used a 13-year dataset on waterbird abundance across seven breeding colonies (three Caspian tern [Hydroprogne caspia], two double-crested cormorant [Nannopterum auritum], and two California and ring-billed gull [Larus californicus and Larus delawarensis] colonies) and steelhead tag-recovery data (>645,000 tagged and >32,000 recovered steelhead) to quantify weekly predation probabilities and functional responses across waterbird species, colonies, and years. Weekly predation probabilities were highly variable, ranging from 0.01 to 0.30 at tern colonies, 0.01 to 0.20 at cormorant colonies, and 0.03 to 0.13 at gull colonies. Per capita predation probabilities were an order of magnitude higher at inland tern and cormorant colonies relative to estuary colonies of the same species. Terns displayed Type II functional responses across colonies and years, where predation probabilities peaked at low steelhead abundances and declined as steelhead abundance increased (i.e., predator swamping). Cormorants nesting at the large estuary colony (several thousand birds) displayed a Type III functional response, but cormorants nesting at the smaller inland colony (several hundred birds) displayed a Type II response. Consumption probabilities of steelhead by gulls remained consistent across a large range of steelhead availability, suggesting a Type I or a Type III functional response, but a lack of colony abundance data prevented quantifying functional responses. The level of tern predation combined with Type II functional responses indicate possible population-level impacts that could destabilize small or declining prey populations. Conversely, the apparent Type III functional responses of gulls and estuary nesting cormorants are indicative of prey switching behaviors targeted at periods of high steelhead abundance. Our results illustrate the complexity of predator–prey interactions and the importance of quantifying predator- and location-specific functional responses when predicting the efficacy of management strategies to enhance prey populations.

Spatiotemporal distribution of Ceratonova shasta in the lower Columbia River Basin and effects of exposure on survival of juvenile chum salmon Oncorhynchus keta.

In the Columbia River Basin (CRB), USA, anthropogenic factors ranging from dam construction to land use changes have modified riverine flow and temperature regimes and degraded salmon habitat. These factors are directly implicated in native salmon and steelhead (Oncorhynchus species) population declines and also indirectly cause mortality by altering outcomes of ecological interactions. For example, attenuated flows and warmer water temperatures drive increased parasite densities and in turn, overwhelm salmonid resistance thresholds, resulting in high disease and mortality. Outcomes of interactions between the freshwater myxozoan parasite, Ceratonova shasta, and its salmonid hosts (e.g., coho O. kisutch and Chinook salmon O. tshawytscha) are well-described, but less is known about effects on chum salmon O. keta, which have a comparatively brief freshwater residency. The goal of this study was to describe the distribution of C. shasta relative to chum salmon habitat in the CRB and assess its potential to cause mortality in juvenile chum salmon (listed as threatened in the CRB under the U.S. Endangered Species Act). We measured C. shasta densities in water samples collected from chum salmon habitat throughout the lower CRB during the period of juvenile chum salmon outmigration, 2018–2020. In 2019, we exposed caged chum salmon fry from two hatchery stocks at three C. shasta-positive sites to assess infection prevalence and survival. Results demonstrated: (1) C. shasta was detected in spawning streams from which chum salmon have been extirpated but was not detected in contemporary spawning habitat while juvenile chum salmon were present, (2) spatiotemporal overlap occurs between C. shasta and juvenile chum salmon in the Columbia River mainstem, and (3) low densities of C. shasta caused lethal infection in chum salmon fry from both hatchery stocks. Collectively, our results suggest C. shasta may limit recovery of chum salmon now and in the future.

Comparative Susceptibilities of Selected California Chinook Salmon and Steelhead Populations to Isolates of L Genogroup Infectious Hematopoietic Necrosis Virus (IHNV).

Simple SummaryChinook salmon in California conservation hatcheries have suffered disease outbreaks and significant mortality due to the fish virus infectious hematopoietic necrosis virus (IHNV) since at least the 1940s. Although steelhead trout in California are also occasionally infected with the same virus they do not typically experience disease. In this study the susceptibility of California Chinook salmon and steelhead trout was defined in controlled experiments that exposed groups of juvenile fish to California IHNV strains (L genogroup). The results confirmed high mortality among Chinook salmon but very low mortality of steelhead trout, despite identical virus exposures. Tests of varying conditions found increasing mortality of Chinook salmon with increasing virus dose, but reduced mortality at higher temperatures or with increased age or size of fish. Among Chinook salmon that survived virus exposure the persistence of virus was detected in one fish 8 months after virus exposure. These findings demonstrate that L genogroup IHNV in California has host specificity for Chinook salmon, and provide an understanding of factors that contribute to disease epidemics in California hatchery Chinook salmon.Salmonid species demonstrate varied susceptibility to the viral pathogen infectious hematopoietic necrosis virus (IHNV). In California conservation hatcheries, juvenile Chinook salmon (Oncorhynchus tshawytscha) have experienced disease outbreaks due to L genogroup IHNV since the 1940s, while indigenous steelhead (anadromous O. mykiss) appear relatively resistant. To characterize factors contributing to the losses of California salmonid fish due to IHNV, three populations of Chinook salmon and two populations of steelhead native to California watersheds were compared in controlled waterborne challenges with California L genogroup IHNV isolates at viral doses of 104–106 pfu mL−1. Chinook salmon fry were moderately to highly susceptible (CPM = 47–87%) when exposed to subgroup LI and LII IHNV. Susceptibility to mortality decreased with increasing age and also with a higher temperature. Mortality for steelhead fry exposed to two IHNV isolates was low (CPM = 1.3–33%). There was little intraspecies variation in susceptibility among populations of Chinook salmon and no differences in virulence between viruses strains. Viral persistence was demonstrated by the isolation of low levels of infectious IHNV from the skin of two juvenile Chinook salmon at 215 d post exposure. The persistence of the virus among Chinook salmon used for stocking into Lake Oroville may be an explanation for the severe epidemics of IHN at the Feather River hatchery in 1998–2002.

Size and Distribution of Parr Produced from Natural‐ and Hatchery‐Origin Steelhead Spawning Naturally in a Small Pacific Northwest Coastal Stream

AbstractRecent studies suggest that steelhead Oncorhynchus mykiss produced from local hatchery‐origin (HOR) adults have lower lifetime fitness than their natural‐origin (NOR) counterparts. To increase our understanding of this pattern, we compared age‐1 parr size and distribution produced by a local integrated population of HOR and NOR adults spawning naturally across a range of environmental conditions. Across 8 years, we used genetic parentage assignments and field measurements of parr in conjunction with creek temperature and flow data to find small, nonbiologically significant differences in size between parr of different parent origins. This suggests that parr produced by HOR adults are acquiring enough resources to grow at a rate similar to that of parr produced by NOR adults. In contrast to origin, we found strong positive associations between mean size of parr and annual mean water temperature and summer flow. Additionally, the distribution of parr was similar and HOR‐produced parr were not skewed relative to the location of the hatchery. Parr occupying the full extent of accessible and suitable habitat might be a positive outcome for hatchery programs seeking to supplement an existing population or replace an extirpated population. However, these results also imply that HOR fish could be competing with NOR fish for food and space, which might be less desirable if the goal of the hatchery program is harvest and not supplementation. Our results highlight the importance of clearly articulated goals for the hatchery program, as the observed pattern could be deemed a benefit or a risk depending on the hatchery's purpose. Lastly, projections for higher water temperatures and reduced summer flows, when considered in the context of the correlations we observed for size of parr, imply that coastal steelhead populations could start experiencing negative climate impacts with respect to steelhead parr growth metrics during the summer in the next few decades.

Counting the Parts to Understand the Whole: Rethinking Monitoring of Steelhead in California’s Central Valley

Steelhead (Oncorhynchus mykiss expressing an anadromous life history) in the Sacramento and San Joaquin rivers and their tributaries in California’s Central Valley (CCV) belong to a Distinct Population Segment (DPS) that is listed as threatened under the US Endangered Species Act. Although contemporary management and recovery plans include numerous planned and ongoing efforts seeking to aid in DPS recovery—such as gravel augmentation, manipulation of spring flows, and restoration of rearing and spawning habitat—a paucity of data precludes the possibility of evaluating the effect of these actions on populations of Steelhead in CCV streams. Knowledge gaps relating to historic and current abundance, population-specific ratios of resident and anadromous life-history expression, and the influence of hatchery-reared fish remain largely unaddressed. This is partly a result of aspects of Steelhead biology that make them difficult to monitor, including the multitude of factors that contribute to the expression of anadromy, polymorphic populations, and migration periods that coincide with challenging field conditions. However, these gaps in understanding are also partly the result of an institutional focus on Chinook Salmon (Oncorhynchus tshawytscha) and a pervasive notion that actions benefiting Chinook populations will also benefit Steelhead populations. To evaluate these gaps and to suggest approaches for assessing DPS recovery actions, we review available data and existing monitoring efforts, and consider the actions necessary to inform the development of targeted O. mykiss monitoring programs. Current management and recovery goals focus on abundance estimates of Steelhead only, yet current monitoring is insufficient for reliable estimates. We argue that a reallocation of monitoring resources to better understand the interaction between resident O. mykiss and Steelhead would provide better data to estimate the vital rates needed to evaluate the effects of recovery actions.

Counting the Parts to Understand the Whole: Rethinking Monitoring of Steelhead in California’s Central Valley

Steelhead (Oncorhynchus mykiss expressing an anadromous life history) in the Sacramento and San Joaquin rivers and their tributaries in California’s Central Valley (CCV) belong to a Distinct Population Segment (DPS) that is listed as threatened under the US Endangered Species Act. Although contemporary management and recovery plans include numerous planned and ongoing efforts seeking to aid in DPS recovery—such as gravel augmentation, manipulation of spring flows, and restoration of rearing and spawning habitat—a paucity of data precludes the possibility of evaluating the effect of these actions on populations of Steelhead in CCV streams. Knowledge gaps relating to historic and current abundance, population-specific ratios of resident and anadromous life-history expression, and the influence of hatchery-reared fish remain largely unaddressed. This is partly a result of aspects of Steelhead biology that make them difficult to monitor, including the multitude of factors that contribute to the expression of anadromy, polymorphic populations, and migration periods that coincide with challenging field conditions. However, these gaps in understanding are also partly the result of an institutional focus on Chinook Salmon (Oncorhynchus tshawytscha) and a pervasive notion that actions benefiting Chinook populations will also benefit Steelhead populations. To evaluate these gaps and to suggest approaches for assessing DPS recovery actions, we review available data and existing monitoring efforts, and consider the actions necessary to inform the development of targeted O. mykiss monitoring programs. Current management and recovery goals focus on abundance estimates of Steelhead only, yet current monitoring is insufficient for reliable estimates. We argue that a reallocation of monitoring resources to better understand the interaction between resident O. mykiss and Steelhead would provide better data to estimate the vital rates needed to evaluate the effects of recovery actions.

Influence of Steelhead Trout Stocking Size and Location on Emigration and Adult Returns

AbstractEffective stocking programs are an essential part of maintaining most high‐quality tributary steelhead Oncorhynchus mykiss fisheries in the Great Lakes, requiring sufficient numbers of stocked fish to survive, imprint, and return as adults. New York’s Lake Erie tributaries have long supported popular and economically important recreational steelhead fisheries, attracting anglers from across the country. Recent research has illustrated New York’s dependence on “stray” steelhead from other jurisdictions, making New York fisheries vulnerable to other agencies’ management decisions. We hypothesized that the limited contribution by New York‐stocked steelhead was attributable to poor poststocking survival and imprinting due to the smaller stocking size relative to other Lake Erie jurisdictions. We took an experimental approach to describe how the combination of stocking size and within‐stream stocking location interact to influence emigration and adult returns and identify mechanisms that may be driving these relationships, with the goal of identifying stocking strategies that improve adult returns. Stocked steelhead &gt;150 mm total length emigrated from the stream by the end of May; small‐size steelhead tended to remain in the stream and exhibited upstream migration. The residual population of steelhead lost body condition and likely experienced high mortality. Adult returns from large‐size fish that were stocked 6.4 km upstream were 4.1 times greater than small‐size fish stocked upstream and six times greater than fish stocked near the lake confluence (0.8 km upstream). We conclude that many of the steelhead stocked by New York are too small to imprint and emigrate, instead remaining as stream residents that experience poor survival and ultimately make minimal contributions to the adult fishery. Increases in the stocking size of New York steelhead coupled with stocking well upstream of the lake confluence would result in increased adult returns of New York‐stocked fish to their streams of origin and improved fishery performance and resiliency.

Outmigration survival of a threatened steelhead population through a tidal estuary

Juvenile steelhead (Oncorhynchus mykiss) are exposed to numerous threats in heterogeneous, estuarine environments, yet understanding of survival patterns and processes during this migratory stage is often limited by studies that use surrogate species or are restricted in duration and spatial specificity. Lack of detailed survival information in this critical migratory stage limits the effectiveness of management to maintain juvenile life history diversity in threatened populations. We used acoustic telemetry with multistate mark–recapture models to investigate survival patterns during a key stage of the juvenile emigration of anadromous steelhead through the Sacramento–San Joaquin River Delta of California, United States, over multiple years, including three drought years. Survival was highly variable both within and among the six years of the study; estimated total survival through the Delta ranged from 0.06 (May 2014) to 0.69 (March 2011). Survival in the upstream reaches was associated with river discharge into the Delta, while survival through the lower reaches was associated with migration route. The lack of a single factor associated with survival in all reaches counteracts preconceived ideas of survival processes. Hydrodynamic manipulation and habitat improvements are recommended to support this anadromous population in a changing climate.

Overwintering Distribution and Postspawn Survival of Steelhead in the Upper Columbia River Basin

AbstractAdult summer‐run steelheadOncorhynchus mykissoverwinter in freshwater for several months prior to spawning. In systems supporting mixed populations of fisheries and conservation importance, understanding the distribution and survival of pre‐ and postspawn fish is necessary for informed management. The upper Columbia River supports hatchery‐origin components valued by anglers, natural‐origin components of conservation concern, and temporary strays from downstream populations. We used radiotelemetry and PIT telemetry to monitor the behavior, distribution, and survival of adult steelhead during fall, overwintering, spawning, and postspawn periods, with a focus on use of the main stem versus four major tributaries. Adult steelhead (N = 807) were tagged at Priest Rapids Dam in 2015 and 2016. One‐fifth of steelhead fell back below Priest Rapids Dam and did not reascend. A slight majority of tagged steelhead that overwintered upstream of Priest Rapids Dam did so in main‐stem reservoirs (54%;N = 548). Overwintering in the main‐stem Columbia River was more likely for later‐arriving steelhead and was concentrated in the upstream‐most reservoir. Winter tributary use was highest in the Wenatchee (26%; 2016) and Methow (18%; 2015) rivers, whereas no steelhead overwintered in the Entiat River. Harvest of hatchery‐origin steelhead was 18% in 2015 and was near zero in 2016, when the fishery was suspended due to low adult returns. After accounting for reported harvest, annual overwinter survival did not differ between main‐stem and tributary habitats, and relatively low adjusted survival of hatchery‐origin steelhead in 2015 suggested unreported harvest. In contrast to low iteroparity rates (&lt;3%), the majority of postspawn steelhead (56.5%) exited tributaries as kelts; kelt survival to Bonneville Dam was 65% in 2016 and 23% in 2017. Collectively, the results highlight the importance of understanding patterns of habitat use and mortality in steelhead populations when managers are faced with balancing harvest and conservation goals.

The Impacts of Dam Construction and Removal on the Genetics of Recovering Steelhead (Oncorhynchus mykiss) Populations across the Elwha River Watershed

Dam construction and longitudinal river habitat fragmentation disrupt important life histories and movement of aquatic species. This is especially true for Oncorhynchus mykiss that exhibits both migratory (steelhead) and non-migratory (resident rainbow) forms. While the negative effects of dams on salmonids have been extensively documented, few studies have had the opportunity to compare population genetic diversity and structure prior to and following dam removal. Here we examine the impacts of the removal of two dams on the Elwha River on the population genetics of O. mykiss. Genetic data were produced from >1200 samples collected prior to dam removal from both life history forms, and post-dam removal from steelhead. We identified three genetic clusters prior to dam removal primarily explained by isolation due to dams and natural barriers. Following dam removal, genetic structure decreased and admixture increased. Despite large O. mykiss population declines after dam construction, we did not detect shifts in population genetic diversity or allele frequencies of loci putatively involved in migratory phenotypic variation. Steelhead descendants from formerly below and above dammed populations recolonized the river rapidly after dam removal, suggesting that dam construction did not significantly reduce genetic diversity underlying O. mykiss life history strategies. These results have significant evolutionary implications for the conservation of migratory adaptive potential in O. mykiss populations above current anthropogenic barriers.

Genetic interactions among native and introduced stocks of Oncorhynchus mykiss in the upper Willamette River, Oregon

First recognized as threatened with extinction in 1999, native winter steelhead Oncorhynchus mykiss from Oregon’s Upper Willamette River (UWR) declined to a record-low 543 adult fish in 2017. This anadromous species has been seriously affected by habitat loss caused by impassable dams, intense predation from pinnipeds, and water pollution. Genetic and ecological risks posed by non-native hatchery steelhead may also limit recovery. Here we used 15 microsatellite markers to investigate the genetic composition of naturally-produced juvenile O. mykiss, collected from the most downstream UWR location and other sites throughout the upper basin. Our results, based on 1012 individuals, identified some natural production by summer steelhead released by ongoing hatchery programs, but even greater influence from discontinued non-native winter steelhead hatchery programs. Genetic influence from hatchery programs was nearly absent in above-dam habitats, but concentrated in below-dam habitats and rivers stocked with hatchery steelhead. Although informative, our microsatellite data lacked sufficient power to assign individual fish to hybrid classes, and we urge caution when interpreting similar results from few loci. Taken together, our findings suggest that fish sorting facilities can assist with the conservation of steelhead populations reintroduced above dams, and that hatchery programs can have a lasting genetic influence over naturally-spawning populations, even after stocking has ceased. To conserve the genetic integrity of native UWR steelhead, we recommend that managers consider the lineage of steelhead used for reintroductions, continue to implement policies that limit genetic risks from existing hatchery programs and explore ways to alleviate persistent risks from discontinued programs.

Differences in growth and condition of juvenile Oncorhynchus mykiss related to sex and a migration-associated genomic region

The expression of anadromy in partially migratory salmonid populations is influenced by sex-specific interactions among an individual’s genotype, condition, and environment, but genotype–phenotype relationships prior to the expression of migratory type are poorly understood. We examined whether juvenile growth and condition differed with respect to sex and a migration-associated genomic region (Omy05) in a coastal California population of steelhead and rainbow trout (Oncorhynchus mykiss). Sex and Omy05 genotype had additive effects on annual growth from age 0+ to age 1+ (∼6–18 months old), with higher growth in males and individuals with copies of the rearranged–resident haplotype. Condition at age 1+ increased with the number of copies of the rearranged–resident haplotype, with similar support for additive or interactive effects with sex; additive effects suggested that genotype differences occurred in both sexes and also that condition was slightly higher in males, while interactive effects suggested that the genotype differences occurred only in males. Our results indicate that phenotypic differences related to sex and Omy05 genotype occur well in advance of anadromous migration or freshwater maturation in this southern O. mykiss population.

Parentage-based tagging improves escapement estimates for ESA-listed adult Chinook salmon and steelhead in the Snake River basin

Parentage-based tagging (PBT) is a nonlethal, genetic tagging method that has been successfully applied in hatchery-supplemented populations to manage hatchery brood stock and monitor hatchery harvest and straying rates. We show that PBT can also improve the accuracy of escapement estimates by significantly reducing the number of hatchery-origin fish falsely classified as natural-origin. Unlike conventional abundance estimates, which use physical marks and tags to distinguish hatchery individuals from their wild counterparts, PBT identifies origin independent of physical form. We applied PBT to populations of Chinook salmon (Oncorhynchus tshawytscha) and steelhead (Oncorhynchus mykiss), which are classified as Threatened under the Endangered Species Act and subject to extensive hatchery supplementation efforts. For spawn years 2014–2018, 16 511 adipose-intact Chinook salmon and 21 953 adipose-intact steelhead were sampled, and PBT identified 19.6% of returning Chinook salmon and 8.3% of steelhead were of hatchery-origin, despite having no physical or mechanical marks. The 90% confidence intervals for escapement estimates of natural-origin Chinook salmon and steelhead made with and without corrections using PBT were nonoverlapping for nine of ten comparisons, indicating that failing to account for unmarked, untagged hatchery-origin fish would result in a significant overestimation of natural abundance.

Understanding Life History Diversity of a Wild Steelhead Population and Managing for Resiliency

AbstractDiversity in steelhead Oncorhynchus mykiss (anadromous Rainbow Trout) is often characterized by varying time spent in freshwater and saltwater environments. However, ontogenetic shifts are not often highlighted as an important diversity characteristic in salmonids. Larger water bodies can play a key role in population stability by buffering limited resources in smaller tributary systems. Based on freshwater and saltwater ages of adult steelhead in Fish Creek, Idaho, there were 12 different life history trajectories observed. Juvenile abundance varied from 12,083 (SE = 1,225.7) to 62,463 (SE = 1,753.8), with ages ranging from young of the year (age 0) to 4 years. Adult abundance fluctuated from 17 (SE &lt; 0.1) to 499 (SE = 67.1), and total age of adults varied from 3 to 7 years. Diversity was most evident in juvenile movement and rearing strategies. We observed four types of movement and rearing pathways in which juveniles either directly migrated to the ocean from Fish Creek or resided for up to three winters in main‐stem river habitat in the Clearwater River basin prior to ocean migration. Most juveniles emigrated in the fall at age 2, and most ocean‐emigrating smolts were age 3. Subsequently, most juveniles resided for at least one additional winter period before ocean migration. Juvenile diversity was also reflected in adult ages; the mean proportion that spent 3 years in freshwater was 0.55 (SE = 0.03), and the mean proportion that spent 2 years in freshwater was 0.41 (SE = 0.03). Adult sex ratio was female biased (mean ± SE = 0.66 ± 0.02), with a higher proportion of females (0.89 ± 0.03) than males (0.65 ± 0.05) residing in the ocean for at least 2 years. Density‐dependent mechanisms were evident in juvenile rearing habitat in Fish Creek for cohorts of high female abundance. Population productivity potentials can be met when tributary and main‐stem rearing habitat can be utilized, and this will be essential in attaining recovery goals.

Distribution of genetic variation underlying adult migration timing in steelhead of the Columbia River basin.

Fish migrations are energetically costly, especially when moving between freshwater and saltwater, but are a viable strategy for Pacific salmon and trout (Oncorhynchus spp.) due to the advantageous resources available at various life stages. Anadromous steelhead (O. mykiss) migrate vast distances and exhibit variation for adult migration phenotypes that have a genetic basis at candidate genes known as greb1L and rock1. We examined the distribution of genetic variation at 13 candidate markers spanning greb1L, intergenic, and rock1 regions versus 226 neutral markers for 113 populations (n = 9,471) of steelhead from inland and coastal lineages in the Columbia River. Patterns of population structure with neutral markers reflected genetic similarity by geographic region as demonstrated in previous studies, but candidate markers clustered populations by genetic variation associated with adult migration timing. Mature alleles for late migration had the highest frequency overall in steelhead populations throughout the Columbia River, with only 9 of 113 populations that had a higher frequency of premature alleles for early migration. While a single haplotype block was evident for the coastal lineage, we identified multiple haplotype blocks for the inland lineage. The inland lineage had one haplotype block that corresponded to candidate markers within the greb1L gene and immediately upstream in the intergenic region, and the second block only contained candidate markers from the intergenic region. Haplotype frequencies had similar patterns of geographic distribution as single markers, but there were distinct differences in frequency between the two haplotype blocks for the inland lineage. This may represent multiple recombination events that differed between lineages where phenotypic differences exist between freshwater entry versus arrival timing as indicated by Micheletti et al. (2018a). Redundancy analyses were used to model environmental effects on allelic frequencies of candidate markers, and significant variables were migration distance, temperature, isothermality, and annual precipitation. This study improves our understanding of the spatial distribution of genetic variation underlying adult migration timing in steelhead as well as associated environmental factors and has direct conservation and management implications.