Overwintering distribution, behavior, and survival of adult summer steelhead: Variability among Columbia river populations

Unlike most anadromous salmonids, summer steelhead Oncorhynchus mykiss overwinter in rivers rather than the ocean for 6-10 months prior to spring spawning. Overwintering in rivers may make summer steelhead more vulnerable to harvest and other mortality sources than are other anadromous populations, but there has been little systematic study of this life history strategy. Here, we used a large-scale radiotelemetry study to examine the overwintering behaviors and distributions of 26 summer steelhead stocks within the regulated lower Columbia-Snake River hydrosystem. Over 6 years, we monitored 5,939 fish, of which 3,399 successfully reached spawning tributaries or the upper Columbia River basin and were assigned to specific populations. An estimated 12.4% of fish that reached spawning areas overwintered at least partially within the hydrosystem (annual estimates = 6.8-19.6%), while the remainder overwintered in tributaries. Across all populations, later-arriving fish were more likely to overwinter in the hydrosystem; overwintering percentages ranged from less than 1% for fish tagged in June to over 40% for those tagged in October. Proportionately more interior-basin steelhead (Clearwater, Salmon, and Snake River metapopulations) overwintered in the hydrosystem than did fish from lower-river populations. Steelhead were distributed in mixed-stock assemblages throughout the hydrosystem during winter, usually in reservoirs closest to their home rivers but also in nonnatal tributaries. Overwintering fish moved upstream and downstream between reaches in all months; a nadir occurred in early January and peak egress into spawning tributaries was in March. The estimated survival to tributaries was higher for fish that overwintered in the hydrosystem (82%) than for fish that did not (62%); this difference was largely attributable to low winter harvest rates. Our results suggest that large main-stem habitats, including reservoirs, may be widely used by overwintering summer steelhead. The complex migration behaviors of steelhead indicate both the potential for adaptation and possible susceptibility to future river environment changes.