Salmon nutrients are associated with the phylogenetic dispersion of riparian flowering-plant assemblages.

A signature of nonrandom phylogenetic community structure has been interpreted as indicating community assembly processes. Significant clustering within the phylogenetic structure of a community can be caused by habitat filtering due to low nutrient availability. Nutrient limitation in temperate Pacific coastal rainforests can be alleviated to some extent by marine nutrient subsidies introduced by migrating salmon, which leave a quantitative signature on the makeup of plant communities near spawning streams. Thus, nutrient-mediated habitat filtering could be reduced by salmon nutrients. Here, we ask how salmon abundance affects the phylogenetic structure of riparian flowering plant assemblages across 50 watersheds in the Great Bear Rainforest of British Columbia, Canada. Based on a regional pool of 60 plant species, we found that assemblages become more phylogenetically dispersed and species poor adjacent to streams with higher salmon spawning density. In contrast, increased phylogenetic clumping and species richness was seen in sites with low salmon density, with steeper slopes, further from the stream edge, and within smaller watersheds. These observations are all consistent with abiotic habitat filtering and biotic competitive exclusion acting together across local and landscape-scale gradients in nutrient availability to structure assembly of riparian flowering plants. In this case, rich salmon nutrients appear to release riparian flowering-plant assemblages from the confines of a low-nutrient habitat filter that drives phylogenetic clustering.

Salmon subsidize an escape from a size spectrum.

A general rule in ecology is that the abundance of species or individuals in communities sharing a common energy source decreases with increasing body size. However, external energy inputs in the form of resource subsidies can modify this size spectrum relationship. Here, we provide the first test of how a marine resource subsidy can affect size spectra of terrestrial communities, based on energy derived from Pacific salmon carcasses affecting a forest soil community beside streams in western Canada. Using both species-based and individual approaches, we found size structuring in this forest soil community, and transient community-wide doubling of standing biomass in response to energy pulses from Pacific salmon carcasses. One group of species were clear outliers in the middle of the size spectrum relationship: larval calliphorid and dryomyzid flies, which specialize on salmon carcasses, and which showed a tenfold increase in biomass in their size class when salmon were available. Thus, salmon subsidize their escape from the size spectrum. These results suggest that using a size-based perspective of resource subsidies can provide new insights into the structure and functioning of food webs.

Marine subsidy promotes spatial and dietary niche variation in an omnivore, the Keen's mouse (Peromyscus keeni).

Marine-derived resource subsidies can generate intrapopulation variation in the behaviors and diets of terrestrial consumers. How omnivores respond, given their multiple trophic interactions, is not well understood. We sampled mice (Peromyscus keeni) and their food sources at five sites on three islands of the Central Coast of British Columbia, Canada, to test predictions regarding variation in the spatial behavior and consumption of marine-subsidized foods among individuals. About 50% of detections (n = 27 recaptures) occurred at traps closest to shoreline (25 m), with capture frequencies declining significantly inland (up to 200 m). Stable isotope signatures (δ 13C and δ 15N), particularly δ 15N, in plant foods, forest arthropod prey, and mouse feces were significantly enriched near shorelines compared with inland, while δ 13C patterns were more variable. Bayesian isotope mixing models applied to isotope values in mouse hair indicated that over one-third (35-37%) of diet was comprised of beach-dwelling arthropods, a marine-derived food source. Males were more abundant near the shoreline than females and consumed more marine-derived prey, regardless of reproductive status or availability of other food sources. Our results identify how multiple pathways of marine nutrient transfer can subsidize terrestrial omnivores and how subsets of recipient populations can show variation in spatial and dietary response.

Salmon abundance and patterns of forest greenness as measured by satellite imagery.

Linkages across ecosystems can shape productivity. Salmon carcasses are exemplary of cross-system linkages, because they can fertilize riparian vegetation and shape patterns of terrestrial biodiversity. Detection of salmon fertilization effects has been confined to field-based studies that are limited in scale. Here we use satellite images to quantify the effects of salmon on greenness of riparian vegetation. We measure tree greenness across spatial and temporal gradients of salmon fertilization effects in two regions. In the first case study, we find that deciduous trees are greener in years following large salmon spawning events, and that the magnitude of this effect was related to the specific abundance of spawning salmon. In the second case study we compare greenness of mixed evergreen and deciduous forests across different watersheds that have different salmon spawning densities. We found greenness was related positively to salmon spawning density near streams with high evergreen cover and flat stream banks. These findings suggest that the effect of salmon carcasses on riparian vegetation may be detectable from space. Further work on this approach, especially with high spatial, temporal and spectral data, may allow estimation of the spatial extent of nutrient enrichment from salmon carcasses and aid ecosystem-based management to protect important ecosystem linkages.

Salmon-derived nitrogen in terrestrial invertebrates from coniferous forests of the Pacific Northwest.

BACKGROUND: Bi-directional flow of nutrients between marine and terrestrial ecosystems can provide essential resources that structure communities in transitional habitats. On the Pacific coast of North America, anadromous salmon (Oncorhynchus spp.) constitute a dominant nutrient subsidy to aquatic habitats and riparian vegetation, although the contribution to terrestrial habitats is not well established. We use a dual isotope approach of delta15N and delta13C to test for the contribution of salmon nutrients to multiple trophic levels of litter-based terrestrial invertebrates below and above waterfalls that act as a barrier to salmon migration on two watersheds in coastal British Columbia. RESULTS: Invertebrates varied predictably in delta15N with enrichment of 3-8 per thousand below the falls compared with above the falls in all trophic groups on both watersheds. We observed increasing delta15N levels in our invertebrate groups with increasing consumption of dietary protein. Invertebrates varied in delta13C but did not always vary predictably with trophic level or habitat. From 19.4 to 71.5% of invertebrate total nitrogen was originally derived from salmon depending on taxa, watershed, and degree of fractionation from the source. CONCLUSIONS: Enrichment of delta15N in the invertebrate community below the falls in conjunction with the absence of delta13C enrichment suggests that enrichment in delta15N occurs primarily through salmon-derived nitrogen subsidies to litter, soil and vegetation N pools rather than from direct consumption of salmon tissue or salmon tissue consumers. Salmon nutrient subsidies to terrestrial habitats may result in shifts in invertebrate community structure, with subsequent implications for higher vertebrate consumers, particularly the passerines.

Impacts of salmon on riparian plant diversity.

The study of natural gradients in nutrient subsidies between ecosystems allows for predictions of how changes in one system can affect biodiversity in another. We performed a large-scale empirical test of the role of Pacific salmon (Oncorhynchus spp.) in structuring riparian plant communities. A comparison of 50 watersheds in the remote Great Bear Rainforest of British Columbia's central coast in Canada shows that salmon influence nutrient loading to plants,shifting plant communities toward nutrient-rich species, which in turn decreases plant diversity.These effects are mediated by interactions between salmon density and the physical characteristics of watersheds. Predicting how salmon affect terrestrial ecosystems is central to conservation plans that aim to better integrate ecosystem values into resource management.