Shade affects magnitude and tactics of juvenile Chinook salmon antipredator behavior in the migration corridor.

Environmental conditions strongly affect antipredator behaviors; however, it is less known how migrating prey adjust antipredator behavior in migration corridors, in part, because active migrants are difficult to observe and study. Migrants are vulnerable and encounter many predators in the corridor, and their propensity to travel towards their destination ties antipredator behavior with movement. We evaluated how environmental risk cues in the migration corridor including in-water habitat structure (present, absent) and overhead shade (sun, shade), and salmon origin (hatchery, wild) affected how juvenile Chinook salmon (Oncorhynchus tshawytscha) reacted to a live predator. We measured how salmon react to predation risk as the difference in time to swim downstream through a 9.1-m long field enclosure with or without a live predatory largemouth bass (Micropterus salmoides). Shade significantly modified the reaction to the predator, and it did so in two ways. First, the magnitude of antipredator behavior was larger in shade compared to direct sun, which suggests salmon perceived shade to be a riskier environment than sun. Second, the escape tactic also varied; salmon slowed down to be cautious in shade and sped up in sun. Structure did not significantly affect behavior and hatchery and wild salmon behaved similarly. Our study suggests that environmental risk cues can shape the magnitude and tactics of how migrants react to predation risk and illustrates how these responses relate to movement with potential to scale up and affect migration patterns.

Comparison of three sampling methods for small-bodied fish in lentic nearshore and open water habitats.

We performed a preliminary evaluation of a mobile sampling platform with adjustable push net and live box (Platform) against two common methods for sampling small-bodied fish (i.e., 10-100 mm) in two distinct lentic habitats. Nearshore (NS) littoral habitat was sampled by Platform and beach seine, and open water (OW) pelagic habitat by Platform and Kodiak trawl. Our goal was to evaluate the Platform's ability to describe fish assemblage structure across habitat types in contrast to common techniques restricted to single habitat types that are less comparable due to gear-specific bias. Platform sample speed had a significant positive effect on recapture efficiency of both nearly neutrally buoyant objects and marked fish. Marked fish recapture efficiencies were similar for Platform in NS and OW, indicating similar efficiency across habitat types. Platform capture efficiency was similar to beach seine and greater than Kodiak trawl. With similar sampling time, the Platform collected more individuals and taxa in NS relative to beach seine and in OW relative to Kodiak trawl. Greater taxa detection by the Platform suggests that it may be effective at detecting species that are numerically rare in specific habitats when compared to these methods. Fish CPUE was significantly greater NS regardless of technique. However, by using the Platform, there is greater confidence that this difference was reliable and not a gear selectivity artifact. Overall, this preliminary study demonstrates the Platform's potential to collect standardized data across NS and OW habitats, track ontogenetic habitat shifts, and detect differences in small-bodied fish taxa richness, relative abundance, and density between NS and OW habitats. Continued experimentation beyond a single reservoir and fish size range is required before consensus can be established regarding the utility of this new push net design.

An escape theory model for directionally moving prey and an experimental test in juvenile Chinook salmon.

Prey evaluate risk and make decisions based on the balance between the costs of predation and those of engaging in antipredator behaviour. Economic escape theory has been valuable in understanding the responses of stationary prey under predation risk; however, current models are not applicable for directionally moving prey. Here we present an extension of existing escape theory that predicts how much predation risk is perceived by directionally moving prey. Perceived risk is measured by the extent antipredator behaviour causes a change in travel speed (the distance to a destination divided by the total time to reach that destination). Cryptic or cautious antipredator behaviour slows travel speed, while prey may also speed up to reduce predator-prey overlap. Next, we applied the sensitization hypothesis to our model, which predicts that prey with more predator experience should engage in more antipredator behaviour, which leads to a larger change in travel speed under predation risk. We then compared the qualitative predictions of our model to the results of a behavioural assay with juvenile Chinook salmon Oncorhynchus tshawytscha that varied in their past predator experience. We timed salmon swimming downstream through a mesh enclosure in the river with and without predator cues present to measure their reaction to a predator. Hatchery salmon had the least predator experience, followed by wild salmon captured upstream (wild-upstream) and wild-salmon captured downstream (wild-downstream). Both wild salmon groups slowed down in response to predator cues, whereas hatchery salmon did not change travel speed. The magnitude of reaction to predator cues by salmon group followed the gradient of previous predator experience, supporting the sensitization hypothesis. Moving animals are conspicuous and vulnerable to predators. Here we provide a novel conceptual framework for understanding how directionally moving prey perceive risk and make antipredator decisions. Our study extends the scope of economic escape theory and improves general understanding of non-lethal effects of predators on moving prey.

Unnatural selection of salmon life histories in a modified riverscape.

Altered river flows and fragmented habitats often simplify riverine communities and favor non-native fishes, but their influence on life-history expression and survival is less clear. Here, we quantified the expression and ultimate success of diverse salmon emigration behaviors in an anthropogenically altered California river system. We analyzed two decades of Chinook salmon monitoring data to explore the influence of regulated flows on juvenile emigration phenology, abundance, and recruitment. We then followed seven cohorts into adulthood using otolith (ear stone) chemical archives to identify patterns in time- and size-selective mortality along the migratory corridor. Suppressed winter flow cues were associated with delayed emigration timing, particularly in warm, dry years, which was also when selection against late migrants was the most extreme. Lower, less variable flows were also associated with reduced juvenile and adult production, highlighting the importance of streamflow for cohort success in these southernmost populations. While most juveniles emigrated from the natal stream as fry or smolts, the survivors were dominated by the rare few that left at intermediate sizes and times, coinciding with managed flows released before extreme summer temperatures. The consistent selection against early (small) and late (large) migrants counters prevailing ecological theory that predicts different traits to be favored under varying environmental conditions. Yet, even with this weakened portfolio, maintaining a broad distribution in migration traits still increased adult production and reduced variance. In years exhibiting large fry pulses, even marginal increases in their survival would have significantly boosted recruitment. However, management actions favoring any single phenotype could have negative evolutionary and demographic consequences, potentially reducing adaptability and population stability. To recover fish populations and support viable fisheries in a warming and increasingly unpredictable climate, coordinating flow and habitat management within and among watersheds will be critical to balance trait optimization versus diversification.

State-dependent life history models in a changing (and regulated) environment: steelhead in the California Central Valley.

We use a state dependent life history model to predict the life history strategies of female steelhead trout (Oncorhynchus mykiss) in altered environments. As a case study of a broadly applicable approach, we applied this model to the American and Mokelumne Rivers in central California, where steelhead are listed as threatened. Both rivers have been drastically altered, with highly regulated flows and translocations that may have diluted local adaptation. Nevertheless, evolutionary optimization models could successfully predict the life history displayed by fish on the American River (all anadromous, with young smolts) and on the Mokelumne River (a mix of anadromy and residency). The similar fitness of the two strategies for the Mokelumne suggested that a mixed strategy could be favored in a variable environment. We advance the management utility of this framework by explicitly modeling growth as a function of environmental conditions and using sensitivity analyses to predict likely evolutionary endpoints under changed environments. We conclude that the greatest management concern with respect to preserving anadromy is reduced survival of emigrating smolts, although large changes in freshwater survival or growth rates are potentially also important. We also demonstrate the importance of considering asymptotic size along with maximum growth rate.

Salmon, wildlife, and wine: marine-derived nutrients in human-dominated ecosystems of central California.

Pacific salmon transfer large quantities of marine-derived nutrients to adjacent forest ecosystems with profound effects on plant and wildlife production. We investigated this process for two highly modified California wine country rivers, one with consistent salmon runs (Mokelumne River) and one without (Calaveras River). Mokelumne River Chinook salmon transported biomass and N comparable to Pacific Northwest salmon streams. Calaveras River levels were much less. Scavenger numbers correlated with salmon carcass counts over time on the Mokelumne River but not the Calaveras River. Likewise, salmon carcasses were consumed significantly faster on the Mokelumne River. Native riparian vegetation as well as cultivated wine grapes adjacent to Mokelumne River spawning sites received 18-25% of foliar N from marine sources, significantly higher than vegetation along the Calaveras River. These data suggest that robust salmon runs continue to provide important ecological services with high economic value, even in impaired watersheds. Loss of Pacific salmon can not only negatively affect stream and riparian ecosystem function, but can also affect local economies where agriculture and salmon streams coexist.