Intertidal clams exhibit population synchrony across spatial and temporal scales.

Long-term datasets can be particularly useful for parsing out factors influencing populations, yet few studies have utilized continuous datasets to quantify population dynamics in bivalve molluscs. We used dynamic factor analysis on a clam biomass dataset spanning 28 yr and five distinct regions in the southern Salish Sea to determine (1) if native intertidal clam populations exhibit synchrony and (2) what environmental covariates may be correlated with these population trends. Once covariates were accounted for, the model with the most data support included three predominant trends to describe multidecadal change in clam biomass. Intraspecific synchrony was highest among Saxidomus gigantea and Leukoma staminea populations, with no clear evidence of covariance in Clinocardium nuttallii. Specifically, we quantified a pronounced decadal decline in L. staminea and an increase in S. gigantea biomass on most beaches. No beaches showed synchrony in trends across all three species, indicating that species-specific trends (regardless of location) were more common than beach-specific trends (regardless of species). Seven environmental covariates were evaluated in their capacity to explain variability in annual mean biomass. Of these, the North Pacific Gyre Oscillation lagged 4 yr prior to the observation year was most supported by the data in the best fitting model, implying that 4 yr old clam biomass is partially determined by oceanographic processes affecting larval clams. Although results suggest large-scale density-independent factors play a role in venerid clam population dynamics, it is also likely local factors account for variability not explained by our model.

Diel horizontal migration in streams: juvenile fish exploit spatial heterogeneity in thermal and trophic resources.

Vertical heterogeneity in the physical characteristics of lakes and oceans is ecologically salient and exploited by a wide range of taxa through diel vertical migration to enhance their growth and survival. Whether analogous behaviors exploit horizontal habitat heterogeneity in streams is largely unknown. We investigated fish movement behavior at daily timescales to explore how individuals integrated across spatial variation in food abundance and water temperature. Juvenile coho salmon made feeding forays into cold habitats with abundant food, and then moved long distances (350-1300 m) to warmer habitats that accelerated their metabolism and increased their assimilative capacity. This behavioral thermoregulation enabled fish to mitigate trade-offs between trophic and thermal resources by exploiting thermal heterogeneity. Fish that exploited thermal heterogeneity grew at substantially faster rates than did individuals that assumed other behaviors. Our results provide empirical support for the importance of thermal diversity in lotic systems, and emphasize the importance of considering interactions between animal behavior and habitat heterogeneity when managing and restoring ecosystems.

Temperature-associated population diversity in salmon confers benefits to mobile consumers.

Habitat heterogeneity can generate intraspecific diversity through local adaptation of populations. While it is becoming increasingly clear that population diversity can increase stability in species abundance, less is known about how population diversity can benefit consumers that can integrate across population diversity in their prey. Here we demonstrate cascading effects of thermal heterogeneity on trout-salmon interactions in streams where rainbow trout rely heavily on the seasonal availability of anadromous salmon eggs. Water temperature in an Alaskan stream varied spatially from 5 degrees C to 17.5 degrees C, and spawning sockeye salmon showed population differentiation associated with this thermal heterogeneity. Individuals that spawned early in cool regions of the 5 km long stream were genetically differentiated from those spawning in warmer regions later in the season. Sockeye salmon spawning generates a pulsed resource subsidy that supports the majority of seasonal growth in stream-dwelling rainbow trout. The spatial and temporal structuring of sockeye salmon spawn timing in our focal stream extended the duration of the pulsed subsidy compared to a thermally homogeneous stream with a single population of salmon. Further, rainbow trout adopted movement strategies that exploited the multiple pulses of egg subsidies in the thermally heterogeneous stream. Fish that moved to track the resource pulse grew at rates about 2.5 times higher than those that remained stationary or trout in the reference stream with a single seasonal pulse of eggs. Our results demonstrate that habitat heterogeneity can have important effects on the population diversity of dominant species, and in turn, influence their value to species that prey upon them. Therefore, habitat homogenization may have farther-reaching ecological effects than previously considered.

Thermal heterogeneity mediates the effects of pulsed subsidies across a landscape.

Spatial and temporal heterogeneity interact to make the foraging rates of individuals more variable than expected from models assuming that spatial and temporal dimensions of habitat conditions operate independently. For example, trophic resource pulses produce temporal patches of prey superabundance, yet little is known about how spatial heterogeneity in habitat conditions mediates the ability of consumers to exploit these high-quality foraging opportunities. We studied how spatial variation in water temperature regulates the potential for juvenile coho salmon to exploit a seasonal pulsed subsidy of eggs produced by anadromous sockeye salmon. Streams within the Wood River watershed, southwestern Alaska, USA, ranged in mean summer temperature from 3.6 degrees to 14.5 degrees C. Growth of juvenile coho prior to the arrival of the seasonal egg subsidy was positively related to water temperature among streams. An in situ experiment combined with field samples of diets revealed a size threshold for egg consumption; only individuals longer than approximately 70 mm could consume eggs due to gape limitation of smaller individuals. A bioenergetics simulation demonstrated that water temperature regulated whether age-0 coho salmon could grow large enough to exceed the size threshold for egg consumption. Coho salmon that consumed eggs had energy rations that were five times higher than fish that did not consume eggs, resulting in a positive feedback of water temperature on their integrated seasonal growth. Across this landscape, heterogeneity in water temperature mediates individual- and population-level responses to seasonally available resource pulses. Our study illustrates that ecological mechanisms, such as size-based foraging asymmetries, can magnify the effects of climate change compared to predictions based on physiology alone.

Habitat saturation drives thresholds in stream subsidies.

Understanding how abundance regulates the effects of organisms on their ecosystems remains a critical goal of ecology, especially for understanding inter-ecosystem transfers of energy and nutrients. Here we examined how territoriality and nest-digging by anadromous salmon mediate trophic subsidies to stream fishes. Salmon eggs become available for consumption primarily by the digging of salmon that superimpose their nests on previous nests. An individual-based model of spawning salmon predicted that territoriality and habitat saturation produce a nonlinear effect of salmon density on numbers of available eggs to resident predators. Field studies in Alaskan streams found that higher densities of salmon produce disproportionately more eggs in stream drift and in diets of resident fishes (Arctic grayling and rainbow trout). Bioenergetics model simulations indicated that these subsidies produce substantially enhanced growth rates of trout. These results demonstrate that small changes in salmon abundance can drive large changes in subsidies to stream food webs. Thus, the ecological consequences of population declines of keystone species, such as salmon, will be exacerbated when behavior generates nonlinear impacts.