Long‐distance downstream movements by homing adult chinook salmon

Unusually long downstream movements totalling several hundred kilometres to >1100 km were observed during upstream homing migrations of radio‐tagged spring chinook salmon Oncorhynchus tshawytscha in the Columbia and Snake Rivers, U.S.A. Downstream migrants, identified by their repeated ascension and fallback over a series of large hydroelectric dams within the migration corridor, were primarily hatchery‐origin males.     

Non-direct homing behaviours by adult Chinook salmon in a large, multi-stock river system

Two non-direct homing behaviours, overshoot of natal tributaries and temporary non-natal tributary use, were evaluated for 5150 radio-tagged spring–summer Chinook salmon Oncorhynchus tshawytscha from 40 populations in the large Columbia River system. Over 7 years, 2–44% (mean = 15%) of individuals within each group temporarily entered presumed non-natal tributaries. In addition, many Chinook salmon from lower river tributaries initially travelled 3 to >250 km upstream in the main-stem river beyond confluences with presumed natal tributaries before returning to the natal sites ('overshoot'). Both overshoot and temporary tributary use behaviours declined exponentially with increasing distance from the natal tributary. Non-direct homing also increased later in the season as water temperatures rose and was associated with hatchery origin in some cases. The behaviours may reflect a mix of active searching for olfactory cues from natal sites, behavioural thermoregulation and orientation challenges in a large-river migration corridor transformed by dams and reservoirs. While anadromous salmonid homing is generally accurate and precise, these results indicate that route finding can be non-direct, potentially increasing energetic costs and harvest risks during migration.     

Physiological characterization of juvenile Chinook salmon utilizing different habitats during migration through the Columbia River Estuary

Although off-channel habitats in the estuaries of large rivers impart many benefits to fish that rear within them, it is less clear how these habitats benefit migrating anadromous species that utilize these habitats for short periods of time. We evaluated the physiological correlates (nutritional condition, growth, and smoltification) of habitat utilization (main-channel vs. off-channel) by juvenile Chinook salmon Oncorhynchus tshawytscha during emigration. Fish from the off-channel had higher condition factor scores and relative weights than fish from the main-channel throughout the study period. Plasma triglyceride and protein concentrations were significantly different between habitat types and across the sampling period, suggesting that fish utilizing the off-channel habitats were compensating for energy losses associated with emigration as compared to main-channel fish. Growth potential (RNA to DNA ratio) did not vary by habitat or sampling period, presumably due to short residency time. There were no differences in osmoregulatory capacity (gill Na(+), K(+)-ATPase activity) based on habitat type. Our results indicate that short-term off-channel habitat use may mitigate for energy declines incurred during migration, but likely does not impart significant gains in energy stores or growth.     

Return migration of one‐sea‐winter Atlantic salmon in the River Tana

Fifty‐three one‐sea‐winter Atlantic salmon Salmo salar (45–63 cm L _T) were radio‐tagged in the Tana fjord, Barents Sea, in 1995. Thirty‐seven fish (70%) entered the freshwater zone of the River Tana in an average of 3 days after release in the fjord. The migration speeds in the lowest river section below the first riffle area were significantly higher than in the subsequent river section below the second riffle area. Similarly, the observed time spent in the first riffle area was significantly lower than in the next riffle area. The majority of Atlantic salmon entered the river during the hours of high tide and the subsequent ebb tide. In addition, most river entries were recorded around midnight. No effects of river flow on the river entry or migration speed were detected, but the migration speed of Atlantic salmon in both river sections examined was greater at lower temperatures. Twenty‐eight fish (72%) were recaptured in the river, 71% of them with weirs and gillnets, and 29% by rod and line. Over half of the Atlantic salmon (54%) were recaptured within 3 weeks following river entry, and within the first 100 km of the river (56%). The results are discussed in relation to earlier studies on multi‐sea‐winter Atlantic salmon in the River Tana.     

Thermal exposure of adult Chinook salmon in the Willamette River basin

Radiotelemetry and archival temperature loggers were used to reconstruct the thermal experience of adult spring Chinook salmon (Oncorhynchus tshawytscha) in the highly regulated Willamette River system in Oregon. The study population is threatened and recovery efforts have been hampered by episodically high prespawn mortality that is likely temperature mediated. Over three years, 310 salmon were released with thermal loggers and 68 were recovered in spawning tributaries, primarily at hatchery trapping facilities downstream from high-head dams. More than 190,000 internal body temperature records were collected (mean ~2800 per fish) and associated with 14 main stem and tributary reaches. Most salmon experienced a wide temperature range (minima ~8–10 °C; maxima ~13–22 °C) and 65% encountered potentially stressful conditions (≥18 °C). The warmest salmon temperatures were in lower Willamette River reaches, where some fish exhibited short-duration behavioral thermoregulation. Cumulative temperature exposure, measured by degree days (DD) above 0 °C, varied more than seven-fold among individuals (range=208–1498 DDs) and more than two-fold among sub-basin populations, on average. Overall, ~72% of DDs accrued in tributaries and ~28% were in the Willamette River main stem. DD differences among individuals and populations were related to migration distance, migration duration, and salmon trapping protocols (i.e., extended pre-collection holding in tributaries versus hatchery collection shortly after tributary entry). The combined data provide spatially- and temporally-referenced information on both short-duration stressful temperature exposure and the biologically important total exposure. Thermal exposure in this population complex proximately influences adult salmon physiology, maturation, and disease processes and ultimately affects prespawn mortality and fitness. The results should help managers develop more effective salmon recovery plans in basins with marginal thermal conditions.     

Confirmation of the presence and use of sandy beach surf-zones by juvenile Chinook salmon

Migration patterns and habitat use of sub-yearling Chinook salmon during initial ocean entrance is poorly understood. Twenty-five years ago, sub-yearling Chinook salmon were hypothesized to stay close to shore (<5 km). To test this hypothesis we sampled the surf-zone of a southern Oregon dissipative sandy beach throughout the summer of 2006 (06/07–09/29) using a beach seine in 1 m of water depth. We caught 48 sub-yearlings over six dates (07/22 to 09/01). Mean standard length of Chinook salmon caught in the surf-zone increased from 9.1 ± 0.6 (07/22/06) to 11.6 ± 0.7 cm (09/01/06), suggesting a mean increase of 0.6 mm in standard length (S.L.) per day. Early in the summer, smaller fish fed mostly on amphipods. Later in the summer, larger juveniles fed primarily on larval and juvenile fish. All prey items were common in the surf-zone. Juveniles appear to migrate from the estuary to the surf-zone where they feed on the local zooplankton for up to two summer months before migrating offshore.     

Watershed‐scale climate influences productivity of Chinook salmon populations across southcentral Alaska

The ecosystems supporting Pacific salmon (Oncorhynchus spp.) are changing rapidly as a result of climate change and habitat alteration. Understanding how—and how consistently—salmon populations respond to changes at regional and watershed scales has major implications for fisheries management and habitat conservation. Chinook salmon (O. tshawytscha) populations across Alaska have declined over the past decade, resulting in fisheries closures and prolonged impacts to local communities. These declines are associated with large‐scale climate drivers, but uncertainty remains about the role of local conditions (e.g., precipitation, streamflow, and stream temperature) that vary among the watersheds where salmon spawn and rear. We estimated the effects of these and other environmental indicators on the productivity of 15 Chinook salmon populations in the Cook Inlet basin, southcentral Alaska, using a hierarchical Bayesian stock‐recruitment model. Salmon spawning during 2003–2007 produced 57% fewer recruits than the previous long‐term average, leading to declines in adult returns beginning in 2008. These declines were explained in part by density dependence, with reduced population productivity following years of high spawning abundance. Across all populations, productivity declined with increased precipitation during the fall spawning and early incubation period and increased with above‐average precipitation during juvenile rearing. Above‐average stream temperatures during spawning and rearing had variable effects, with negative relationships in many warmer streams and positive relationships in some colder streams. Productivity was also associated with regional indices of streamflow and ocean conditions, with high variability among populations. The cumulative effects of adverse conditions in freshwater, including high spawning abundance, heavy fall rains, and hot, dry summers may have contributed to the recent population declines across the region. Identifying both coherent and differential responses to environmental change underscores the importance of targeted, watershed‐specific monitoring and conservation efforts for maintaining resilient salmon runs in a warming world.     

Microsatellite variation reveals weak genetic structure and retention of genetic variability in threatened Chinook salmon (<Emphasis Type="Italic">Oncorhynchus tshawytscha</Emphasis>) within a Snake River watershed

Pacific salmon (Oncorhynchus spp.) have been central to the development of management concepts associated with evolutionarily significant units (ESUs), yet there are still relatively few studies of genetic diversity within threatened and endangered ESUs for salmon or other species. We analyzed genetic variation at 10 microsatellite loci to evaluate spatial population structure and genetic variability in indigenous Chinook salmon (Oncorhynchus tshawytscha) across a large wilderness basin within a Snake River ESU. Despite dramatic 20th century declines in abundance, these populations retained robust levels of genetic variability. No significant genetic bottlenecks were found, although the bottleneck metric (M ratio) was significantly correlated with average population size and variability. Weak but significant genetic structure existed among tributaries despite evidence of high levels of gene flow, with the strongest genetic differentiation mirroring the physical segregation of fish from two sub-basins. Despite the more recent colonization of one sub-basin and differences between sub-basins in the natural level of fragmentation, gene diversity and genetic differentiation were similar between sub-basins. Various factors, such as the (unknown) genetic contribution of precocial males, genetic compensation, lack of hatchery influence, and high levels of current gene flow may have contributed to the persistence of genetic variability in this system in spite of historical declines. This unique study of indigenous Chinook salmon underscores the importance of maintaining natural populations in interconnected and complex habitats to minimize losses of genetic diversity within ESUs.     

Olfactory gene expression in migrating adult sockeye salmon Oncorhynchus nerka

Expression of 12 olfactory genes was analysed in adult sockeye salmon Oncorhynchus nerka nearing spawning grounds and O. nerka that had strayed from their natal migration route. Variation was found in six of these genes, all of which were olfc olfactory receptors and had lower expression levels in salmon nearing spawning grounds. The results may reflect decreased sensitivity to natal water olfactory cues as these fish are no longer seeking the correct migratory route. The expression of olfactory genes during the olfactory‐mediated spawning migration of Pacific salmon Oncorhynchus spp. is largely unexplored and these findings demonstrate a link between migratory behaviours and olfactory plasticity that provides a basis for future molecular research on salmon homing.     

Linking landscape-level change to habitat quality: an evaluation of restoration actions on the freshwater habitat of spring-run Chinook salmon

1. Conservation planning is often hampered by the lack of causal quantitative links between landscape characteristics, restoration actions and habitat conditions that impact the status of imperilled species. Here we present a first step toward linking actions on the landscape to the population status of endangered stream‐type Chinook salmon (Oncorhynchus tshawytscha). 2. We developed relationships between land use, landscape characteristics and freshwater habitat of spring Chinook salmon in the Wenatchee River basin. Available data allowed us to find relationships that described water temperatures at several life stages (prespawning, egg incubation and summer rearing) and substratum characteristics, including fine sediments, cobble and embeddedness. Predictors included altitude, gradient, mean annual precipitation, total and riparian forest cover, road density, impervious surface and alluvium. We used a model averaging approach to account for parameter and model selection uncertainty. Key predictors were total forest cover and impervious surface area for prespawning and summer rearing temperatures; precipitation and stream gradients were important predictors of the percent of fine sediments in stream substrata. 3. We estimated habitat conditions using these relationships in three alternative landscape scenarios: historical, no restoration and one that included a set of restoration actions from local conservation planning. We found that prespawning and summer temperatures were estimated to be slightly higher historically relative to current conditions in dry sparsely forested areas, but lower in some important Chinook salmon spawning and rearing areas and lower in those locations under the restoration scenario. Fine sediments were lower in the historical scenario and were reduced as a consequence of restoration actions in two areas currently unoccupied by Chinook salmon that contain reaches with some potential for high quality spawning and rearing. Cobble and embeddedness in general were predicted to be higher historically and changed little as a result of restoration actions relative to current conditions. 4. This modelling framework converts suites of restoration actions into changes in habitat condition, thereby enabling restoration planners to evaluate alternative combinations of proposed actions. It also provides inputs to models linking habitat conditions to population status. This approach represents a first step in estimating impacts of restoration strategies, and can provide key information for conservation managers and planners.     

Thermal biology and bioenergetics of different upriver migration strategies in a stock of summer-run Chinook salmon

By combining biotelemetry with animal-borne thermal loggers, we re-created the thermal histories of 21 summer-run Chinook salmon (Oncorhynchus tshawytscha) migrating in the Puntledge River, a hydropower impacted river system on Vancouver Island, British Columbia, Canada. Daily maximum water temperatures in the Puntledge River during the summer-run adult Chinook salmon migration and residency period frequently exceeded 21 °C, a value that has been observed to elicit behavioral thermoregulation in other Chinook salmon populations. We therefore compared river temperatures to body temperatures of 16 fish that migrated through the river to understand if cool-water refuge was available and being used by migrants. In addition, we used thermal histories from fish and thermal loggers distributed in the river to model the effect of thermal habitat on energy density using a bioenergetics model. In general, we found no evidence that cool-water refuge existed in the river, suggesting that there is no opportunity for fish to behaviorally thermoregulate during upriver migration through the regulated portion of the river. Of the thermal histories used in the bioenergetics model, fish that reached an upstream lake were able to access cooler, deeper waters, which would have reduced energy consumption compared to fish that only spent time in the warmer river. Consequently, the Puntledge River water temperatures are likely approaching and in some cases exceeding the thermal limits of the summer-run Chinook salmon during the spawning migration. Further warming may cause more declines in the stock.     

Population structure of lake‐type and river‐type sockeye salmon in transboundary rivers of northern British Columbia

The population structure of 'lake‐type' and 'river‐type' sockeye salmon Oncorhynchus nerka , primarily in transboundary rivers in northern British Columbia, was examined with a survey of microsatellite variation. Variation at 14 microsatellite loci was surveyed from c . 3000 lake‐type and 3200 river‐type sockeye salmon from 47 populations in six river drainages in British Columbia. The mean F _ST for the 14 microsatellite loci and 47 populations was 0·068, and 0·034 over all river‐type populations. River‐type sockeye salmon were more genetically diverse than lake‐type sockeye salmon, with expected heterozygosity of river‐type sockeye salmon 0·72 and with an average 12·7 alleles observed per locus, whereas expected heterozygosity of lake‐type sockeye salmon was 0·65 with and average 10·5 alleles observed per locus. River drainage of origin was a significant unit of population structure. There was clear evidence of genetic differentiation among river‐type populations of sockeye salmon from different drainages over a broad geographic range in British Columbia.     

Distribution of the invasive New Zealand mudsnail (Potamopyrgus antipodarum) in the Columbia River Estuary and its first recorded occurrence in the diet of juvenile Chinook salmon (Oncorhynchus tshawytscha)

Estuaries play an important role as nurseries and migration corridors for Chinook salmon and other fishes. The invasive New Zealand mudsnail, Potamopyrgus antipodarum (Gray, 1843), has been noted in the Columbia River Estuary and other estuaries in the western USA, yet no studies have addressed the estuarine impacts of this invader. Our data show P. antipodarum is currently found in five peripheral bays and many tributaries of the Columbia River Estuary, where it can constitute a major portion of the benthic invertebrate biomass and where it co-occurs with native amphipod species. We review the history of the P. antipodarum invasion in the Columbia River Estuary and discuss potential impacts on estuarine food webs. We also report the first occurrence of P. antipodarum in the diet of juvenile Chinook salmon from the Columbia River Estuary. Although present in Chinook diets at very low frequencies, our observations of P. antipodarum in Chinook gut contents may represent early stages of food web change due to the establishment of dense estuarine snail populations. Additional research is needed to determine the effects of P. antipodarum on benthic resources, native benthic invertebrates, and benthic predators. We encourage biologists working in western USA estuaries to be alert to the possibility of encountering P. antipodarum in benthic habitats and predator diets.

A latitudinal cline in the Chinook salmon (Oncorhynchus tshawytscha) Clock gene: evidence for selection on PolyQ length variants

A critical seasonal event for anadromous Chinook salmon (Oncorhynchus tshawytscha) is the time at which adults migrate from the ocean to breed in freshwater. We investigated whether allelic variation at the circadian rhythm genes, OtsClock1a and OtsClock1b, underlies genetic control of migration timing among 42 populations in North America. We identified eight length variants of the functionally important polyglutamine repeat motif (PolyQ) of OtsClock1b while OtsClock1a PolyQ was highly conserved. We found evidence of a latitudinal cline in average allele length and frequency of the two most common OtsClock1b alleles. The shorter 335 bp allele increases in frequency with decreasing latitude while the longer 359 bp allele increases in frequency at higher latitudes. Comparison to 13 microsatellite loci showed that 335 and 359 bp deviate significantly from neutral expectations. Furthermore, a hierarchical gene diversity analysis based on OtsClock1b PolyQ variation revealed that run timing explains 40.9 per cent of the overall genetic variance among populations. By contrast, an analysis based on 13 microsatellite loci showed that run timing explains only 13.2 per cent of the overall genetic variance. Our findings suggest that length polymorphisms in OtsClock1b PolyQ may be maintained by selection and reflect an adaptation to ecological factors correlated with latitude, such as the seasonally changing day length.     

Temporal patterns in adult salmon migration timing across southeast Alaska

Pacific salmon migration timing can drive population productivity, ecosystem dynamics, and human harvest. Nevertheless, little is known about long‐term variation in salmon migration timing for multiple species across broad regions. We used long‐term data for five Pacific salmon species throughout rapidly warming southeast Alaska to describe long‐term changes in salmon migration timing, interannual phenological synchrony, relationships between climatic variation and migratory timing, and to test whether long‐term changes in migration timing are related to glaciation in headwater streams. Temporal changes in the median date of salmon migration timing varied widely across species. Most sockeye populations are migrating later over time (11 of 14), but pink, chum, and especially coho populations are migrating earlier than they did historically (16 of 19 combined). Temporal trends in duration and interannual variation in migration timing were highly variable across species and populations. The greatest temporal shifts in the median date of migration timing were correlated with decreases in the duration of migration timing, suggestive of a loss of phenotypic variation due to natural selection. Pairwise interannual correlations in migration timing varied widely but were generally positive, providing evidence for weak region‐wide phenological synchrony. This synchrony is likely a function of climatic variation, as interannual variation in migration timing was related to climatic phenomenon operating at large‐ (Pacific decadal oscillation), moderate‐ (sea surface temperature), and local‐scales (precipitation). Surprisingly, the presence or the absence of glaciers within a watershed was unrelated to long‐term shifts in phenology. Overall, there was extensive heterogeneity in long‐term patterns of migration timing throughout this climatically and geographically complex region, highlighting that future climatic change will likely have widely divergent impacts on salmon migration timing. Although salmon phenological diversity will complicate future predictions of migration timing, this variation likely acts as a major contributor to population and ecosystem resiliency in southeast Alaska.     

The influence of hydrology and waterway distance on population structure of Chinook salmon Oncorhynchus tshawytscha in a large river

Adult Chinook salmon Oncorhynchus tshawytscha navigate in river systems using olfactory cues that may be influenced by hydrologic factors such as flow and the number, size and spatial distribution of tributaries. Thus, river hydrology may influence both homing success and the level of straying (gene flow), which in turn influences population structure. In this study, two methods of multivariate analysis were used to examine the extent to which four indicators of hydrology and waterway distance explained population structure of O. tshawytscha in the Yukon River. A partial Mantel test showed that the indicators of hydrology were positively associated with broad‐scale (Yukon basin) population structure, when controlling for the influence of waterway distance. Multivariate multiple regression showed that waterway distance, supplemented with the number and flow of major drainage basins, explained more variation in broad‐scale population structure than any single indicator. At an intermediate spatial scale, indicators of hydrology did not appear to influence population structure after accounting for waterway distance. These results suggest that habitat changes in the Yukon River, which alter hydrology, may influence the basin‐wide pattern of population structure in O. tshawytscha. Further research is warranted on the role of hydrology in concert with waterway distance in influencing population structure in Pacific salmon.     

A mobile sex‐determining region,male‐specific haplotypes and rearing environment influence age at maturity in Chinook salmon

Variation in age at maturity is an important contributor to life history and demographic variation within and among species. The optimal age at maturity can vary by sex, and the ability of each sex to evolve towards its fitness optimum depends on the genetic architecture of maturation. Using GWAS of RAD sequencing data, we show that age at maturity in Chinook salmon exhibits sex‐specific genetic architecture, with age at maturity in males influenced by large (up to 20 Mb) male‐specific haplotypes. These regions showed no such effect in females. We also provide evidence for translocation of the sex‐determining gene between two different chromosomes. This has important implications for sexually antagonistic selection, particularly that sex linkage of adaptive genes may differ within and among populations based on chromosomal location of the sex‐determining gene. Our findings will facilitate research into the genetic causes of shifting demography in Chinook salmon as well as a better understanding of sex determination in this species and Pacific salmon in general.