The relationships among behavior, environment, and migration success in anadromous fishes are poorly understood. We monitored migration behavior at eight Columbia and Snake river dams for 18 286 adult Chinook salmon (Oncorhynchus tshawytscha) and steelhead (sea-run Oncorhynchus mykiss) over 7 years using radiotelemetry. When statistically controlling for variation in flow, temperature, fisheries take, and other environmental variables, we observed that unsuccessful individuals — those not observed to reach spawning areas — had longer passage times at nearly all dams than fish that eventually reached tributaries. In many cases, times were also longer for unsuccessful adults passing through a multiple-dam reach. Four ecological mechanisms may have contributed to these patterns: (i) environmental factors not accounted for in the analyses; (ii) inefficient responses by some fish to passage conditions at dams that resulted in slowed passage, energetic depletion, and unsuccessful migration; (iii) ongoing selection for traits needed to pass obstructions; and (or) (iv) passage rate was not directly linked to migration success, but rather, both resulted from relatively poor phenotypic condition upon river entry in unsuccessful migrants. Overall, these results illustrate the need for a mechanistic understanding of the factors that influence migration success and the need for fitness-based criteria to assess the effects of dams on anadromous fishes.

We assessed upstream migration rates of more than 12,000 radio-tagged adult Chinook salmon Oncorhynchus tshawytscha and steelhead O. mykiss past a series of dams and reservoirs on the Columbia and Snake rivers. Most fish passed each dam in less than 2 d. Migration behavior in reservoirs and through multiple dam-reservoir reaches varied within and between years and between species. Within years, spring-summer Chinook salmon migrated more rapidly as water temperature and date of migration increased; between years, spring-summer Chinook salmon migrated fastest in low-discharge years. Steelhead migrations slowed dramatically when summer water temperatures peaked within each year, then increased as rivers cooled in fall. Mean summer temperatures explained more between-year variation in steelhead passage rates than did differences in discharge. Fall Chinook salmon migration rates also slowed during periods of warm water. Protracted passage times within the hydrosystem were most likely for fish from all runs that fell back over and reascended dams and for steelhead that sought thermal refugia by straying temporarily into coldwater tributaries.

An understanding of the migration timing patterns of Pacific salmon Oncorhynchus spp. and steelhead O. mykiss is important for managing complex mixed-stock fisheries and preserving genetic and life history diversity. We examined adult return timing for 3,317 radio-tagged fish from 38 stocks of Columbia River basin spring-summer Chinook salmon O. tshawytscha over 5 years. Stock composition varied widely within and between years depending on the strength of influential populations. Most individual stocks migrated at similar times each year relative to overall runs, supporting the hypotheses that run timing is predictable, is at least partially due to genetic adaptation, and can be used to differentiate between some conspecific populations. Arrival timing of both aggregated radio-tagged stocks and annual runs was strongly correlated with river discharge; stocks arrived earlier at Bonneville Dam and at upstream dams in years with low discharge. Migration timing analyses identified many between-stock and between-year differences in anadromous salmonid return behavior and should aid managers interested in protection and recovery of evolutionarily significant populations.

Accurate estimates of escapement by adult anadromous salmonids are difficult, especially in large, multistock river systems. We used radiotelemetry and a fishery reward program to calculate escapement, harvest, and unaccounted for loss rates for 10 498 adult chinook salmon (Oncorhynchus tshawytscha) and 5324 steelhead (Oncorhynchus mykiss) during six return years in the Columbia River basin. Mean annual escapements to spawning sites, hatcheries, or the upper bounds of the monitored hydrosystem were 73.4% (spring–summer chinook salmon), 61.3% (fall chinook salmon), and 62.6% (steelhead). Mean reported harvest rates were 8.7% (spring–summer chinook), 22.0% (fall chinook), and 15.1% (steelhead) within the mainstem hydrosystem and 5.9%, 3.4%, and 5.7%, respectively, in lower hydrosystem tributaries. On average, 12%–17% of each run had unknown fates in the mainstem hydrosystem. Escapement, harvest, and loss varied significantly between runs and years, within runs between known-origin subbasin stocks, and between interdam river reaches. Multiyear quantitative assessments like this can reduce uncertainty, clarify inter- and intra-annual variability, and help managers better evaluate fisheries, identify conservation priorities, and help protect evolutionarily significant populations.

26Fishways designed for salmonids often restrict passage by non-salmonids and effective tools are 27 needed both to identify passage problems for non-target species and to inform remediation 28 planning. In this case study, we used migration histories from 2170 radio-tagged adult Pacific 29 lamprey (Entosphenus tridentatus) to identify locations of poor passage ("bottlenecks") at a 30 large, multi-fishway dam. Over ten years, 49% of tagged lamprey that entered fishways failed to 31 pass the dam. Models accounting for repeated attempts by individual lamprey indicated 32 successful passage strongly depended on attempted passage route. Success also varied with time 33 of fishway entry, water temperature, and lamprey body size. Most failed passage attempts 34 terminated in lower fishway segments, but extensive seasonal shifts in bottleneck locations were 35 detected. Ranking metrics helped prioritize bottlenecks and identified sites where structural or 36 operational modifications should improve lamprey passage. Our integration of spatially-37 intensive monitoring with novel analytical techniques was critical to understanding the complex 38 relationships between fishway features, environmental variation, and lamprey behavior. The 39 prioritization framework can be applied to a wide range of fish passage assessments. 40 41 42

Upstream migration rates were assessed for 1801 radio-tagged adult spring-summer Chinook salmon Oncorhynchus tshawytscha through 12 unimpounded river reaches in the Columbia River basin from 1997 to 2002. Reaches were 36 to 241 km long (mean ¼ 130 km) and included sections of the large Columbia and Snake Rivers and smaller free-flowing tributaries. Median Chinook salmon migration rates ranged from <10 km day À1 in the Deschutes and Clearwater Rivers to >35 km day À1 in the Columbia and Snake Rivers. Using multivariate analyses, migration date explained the most variance in Chinook salmon migration rates while river discharge, migration year and migration reach were secondary. Both within and between years, Chinook salmon migrated more rapidly as migration date increased and more slowly when discharge was high. Arrival at high elevation spawning grounds at appropriate times and increased metabolic activity and reproductive maturation may explain the greater power of migration date, relative to river discharge, in predicting migration rates of Columbia basin spring-summer Chinook salmon. # 2004 The Fisheries Society of the British Isles

Rising river temperatures in western North America have increased the energetic costs of migration and the risk of premature mortality in many Pacific salmon (Oncorhynchus spp.) populations. Predicting and managing risks for these populations requires data on acute and cumulative thermal exposure, the spatio-temporal distribution of adverse conditions, and the potentially mitigating effects of cool-water refuges. In this study, we paired radiotelemetry with archival temperature loggers to construct continuous, spatially-explicit thermal histories for 212 adult Chinook salmon (O. tshawytscha) and 200 adult steelhead (O. mykiss). The fish amassed ~500,000 temperature records (30-min intervals) while migrating through 470 kilometers of the Columbia and Snake rivers en route to spawning sites in Idaho, Oregon, and Washington. Spring- and most summer-run Chinook salmon migrated before river temperatures reached annual highs; their body temperatures closely matched ambient temperatures and most had thermal maxima in the lower Snake River. In contrast, many individual fall-run Chinook salmon and most steelhead had maxima near thermal tolerance limits (20–22 °C) in the lower Columbia River. High temperatures elicited extensive use of thermal refuges near tributary confluences, where body temperatures were ~2–10 °C cooler than the adjacent migration corridor. Many steelhead used refuges for weeks or more whereas salmon use was typically hours to days, reflecting differences in spawn timing. Almost no refuge use was detected in a ~260-km reach where a thermal migration barrier may more frequently develop in future warmer years. Within population, cumulative thermal exposure was strongly positively correlated (0.88 ≤ r ≤ 0.98) with migration duration and inconsistently associated (-0.28 ≤ r ≤ 0.09) with migration date. All four populations have likely experienced historically high mean and maximum temperatures in recent years. Expected responses include population-specific shifts in migration phenology, increased reliance on patchily-distributed thermal refuges, and natural selection favoring temperature-tolerant phenotypes.

Optimization of fishways to pass multiple species is challenging because life history, swimming ability, and behavior often differ among species. For example, high fishway water velocities designed to attract adult Pacific salmon Oncorhynchus spp. at Columbia River dams inhibit fishway entrance and passage success of adult Pacific lampreys Lampetra tridentata, a species of conservation concern. We tested whether reduced water velocities (∼1.2 m/s, 0.15 m of head) at Bonneville Dam fishway openings improved entrance efficiency and other passage metrics for radio‐tagged Pacific lampreys compared with control velocities (>1.98 m/s, 0.46 m of head) and near‐zero (“standby”) velocities. Lamprey entrance efficiencies were significantly higher in the reduced‐velocity treatment (26–29%) than in the control (13–20%) or standby (5–9%) treatment. In some years, significantly more Pacific lampreys passed through fishway collection channels and transition pools and reached the fish ladder during reduced‐velocity treatment conditions, indicating that benefits extended beyond fishway entrances. However, overall passage efficiency at the dam was relatively unchanged, suggesting that additional passage bottlenecks for Pacific lampreys exist upstream from fishway entrances. The experiment demonstrated how operational changes can improve passage performance and how exploiting behavioral differences among species can improve multispecies management.