Spawning Salmon Density Influences Fruit Production of Salmonberry (Rubus Spectabilis)

Annual spawning migrations by Pacific salmon can provide substantial subsidies to nutrient‐limited freshwater and riparian ecosystems, which can affect the abundance, diversity, and physical characteristics of plant and animal species in these habitats. Here, we provide the first investigation of how salmon subsidies affect reproductive output in plants, focusing on a common riparian shrub, salmonberry (Rubus spectabilis). We studied 14 streams with a range of spawning salmon densities on the central coast of British Columbia, Canada. We determined the effects of chum (Oncorhynchus keta), pink (O. gorbuscha), and total salmon spawning density on the number of fruits per shrub, number of seeds per fruit, fruit weight, and estimated sugar content (° Brix) of salmonberry fruits. We found that the number of fruits per salmonberry shrub increased with increasing salmon density. However, we found no effect of salmon density on the number of seeds per fruit, fruit weight, or sugar content. The effect of salmon density was species‐dependent; the number of fruits per shrub increased with chum salmon density but was not affected by pink salmon density. This could be because chum salmon occur at higher densities and are transferred from water to land at higher rates than pink salmon in our study area. Higher salmonberry fruit production could lead to a larger input of salmonberry fruits to coastal food webs. These results demonstrate how salmon can cross ecological boundaries and influence reproductive output of terrestrial species

Links Between Fluctuations in Sockeye Salmon Abundance and Riparian Forest Productivity Identified by Remote Sensing

Pacific salmon (Oncorhynchus spp.) carcasses can fertilize riparian forests with marine-derived nutrients when populations make their annual return to natal streams to spawn; however, the strength of this cross-system linkage likely varies substantially among years due to the interannual fluctuations in abundance that characterize most salmon populations. Here, we used a 36-yr time series (1984–2019) of satellite imagery and salmon abundance estimates to assess spatiotemporal associations between forest greenness (a measure of plant productivity) and adult sockeye salmon (Oncorhynchus nerka) abundance in the lower Adams River, British Columbia, Canada. The Adams River sockeye population displays a quadrennial pattern of abundance, with a dominant cohort that spawns every four years in numbers that are typically two to three orders of magnitude larger than non-dominant cohorts. We found that variation in forest greenness was consistently explained best by models including dominant cohort year, whereas models lacking an index of salmon abundance were the lowest-ranked. Greenness of riparian vegetation increased by an average of 0.015 NDVI units (approximately 1%) in the summer after a dominant cohort return, and this effect on greenness persisted into the subsequent fall (11–13 months after spawning). The positive association between quadrennial pulses of salmon and riparian greenness occurred in plots both within 30 m of the stream and 95–125 m away from the stream, indicating that the spatial extent of fertilization may occur well beyond areas directly adjacent to the riverbank. These results suggest that forests respond to cyclical variation in salmon abundance and that overwinter storage of marine-derived nutrients within catchments allows plants to capitalize on these nutrients in the following growing season. Continued advances in remote sensing technology will enhance our understanding of cross-system resource linkages and can inform the ecosystem-based management of Pacific salmon

Equivalent roles of marine subsidies and island characteristics in shaping island bird communities

AimSpecies distributions across islands are shaped by dispersal limitations, environmental filters and biotic interactions but the relative influence of each of these processes has rarely been assessed. Here, we examine the relative contributions of island characteristics, marine subsidies, species traits, and species interactions on avian community composition.LocationCentral Coast region of British Columbia, Canada.TaxonTerrestrial breeding birds.MethodsWe observed 3610 individuals of 32 bird species on 89 islands that spanned multiple orders of magnitude in area (0.0002–3 km$^{2}$). We fit a spatially explicit joint species distribution model to estimate the relative contributions of island physical characteristics, island‐specific inputs of marine subsidies, species' traits, and biotic interactions on species distributions. Biogeographic characteristics included island area, isolation, and habitat heterogeneity, while marine influence was represented by forest‐edge soil δ$^{15}$N, wrack biomass, shoreline substrate, and distance to shore. This approach also allowed us to estimate how much variation in distributions resulted from species' biological traits (i.e. body mass, feeding guild, feeding height, and nesting height).ResultsBird species distributions were determined almost equivalently by island biogeographic characteristics (23.5% of variation explained) and marine influence (24.8%). We detected variation in species‐specific responses to both island biogeographic characteristics and marine influence, but no significant effect of any biological trait examined. Additionally, we found evidence that habitat preferences were a more important driver than competitive interactions.Main ConclusionsAlthough most island biogeographic studies focus only on islands' physical characteristics, we found evidence for an equivalent role of marine subsidy in structuring island bird communities. Our study suggests that for small islands, disentangling the effects of island biogeographic characteristics, marine inputs, and biotic interactions is a useful next step in understanding species distributions

Scale-dependent effects of marine subsidies on the island biogeographic patterns of plants

Although species richness can be determined by different mechanisms at different spatial scales, the role of scale in the effects of marine inputs on island biogeography has not been studied explicitly. Here, we evaluated the potential influence of island characteristics and marine inputs (seaweed wrack biomass and marine-derived nitrogen in the soil) on plant species richness at both a local (plot) and regional (island) scale on 92 islands in British Columbia, Canada. We found that the effects of subsidies on species richness depend strongly on spatial scale. Despite detecting no effects of marine subsidies at the island scale, we found that as plot level subsidies increased, species richness decreased; plots with more marine-derived nitrogen in the soil hosted fewer plant species. We found no effect of seaweed wrack at either scale. To identify potential mechanisms underlying the decrease in diversity, we fit a spatially explicit joint species distribution model to evaluate species level responses to marine subsidies and effects of biotic interactions among species. We found mixed evidence for competition for both light and nutrients, and cannot rule out an alternative mechanism; the observed decrease in species richness may be due to disturbances associated with animal-mediated nutrient deposits, particularly those from North American river otters (Lontra canadensis). By evaluating the scale-dependent effects of marine subsidies on island biogeographic patterns of plants and revealing likely mechanisms that act on community composition, we provide novel insights on the scale dependence of a fundamental ecological theory, and on the rarely examined links between marine and terrestrial ecosystems often bridged by animal vectors

Biogeographic features mediate marine subsidies to island food webs

Although marine subsidies often enrich terrestrial ecosystems, their influence is known to be context-dependent. Additionally, the multitrophic impact of marine subsidies has not been traced through food webs across physically diverse islands. Here, we test predictions about how island characteristics can affect marine enrichment of food web constituents and how nutrients flow through island food webs. To evaluate enrichment and trace marine nutrients across food webs, we used stable isotopes of soil, flora, and fauna (n = 4752 samples) collected from 97 islands in British Columbia, Canada. Island area was the strongest predictor of enrichment across taxa; we found that samples were more 15N-rich on smaller islands. Enrichment declined with distance from shore but less so on small islands, implying a higher per-unit-area subsidy effect. These area and distance-to-shore effects were taxon-specific, and nearly twice as strong in basal food web groups. We also found that increases in δ15N correlated with increases in %N in basal trophic groups, as well as in songbirds, implying biologically relevant uptake of a potentially limiting nutrient. Path analysis demonstrated that subsidies in soil flow through plants and detritivores, and into upper-level consumers. Our results reveal an interplay between island biogeography and marine subsidies in shaping island food webs through bottom-up processes

Marine subsidies mediate patterns in avian island biogeography

The classical theory of island biogeography, which predicts species richness using island area and isolation, has been expanded to include contributions from marine subsidies, i.e. subsidized island biogeography (SIB) theory. We tested the effects of marine subsidies on species diversity and population density on productive temperate islands, evaluating SIB predictions previously untested at comparable scales and subsidy levels. We found that the diversity of terrestrial breeding bird communities on 91 small islands (approx. 0.0001-3 km2) along the Central Coast of British Columbia, Canada were correlated most strongly with island area, but also with marine subsidies. Species richness increased and population density decreased with island area, but isolation had no measurable influence. Species richness was negatively correlated with marine subsidy, measured as forest-edge soil δ15N. Density, however, was higher on islands with higher marine subsidy, and a negative interaction between area and subsidy indicates that this effect is stronger on smaller islands, offering some support for SIB. Our study emphasizes how subsidies from the sea can shape diversity patterns on islands and can even exceed the importance of isolation in determining species richness and densities of terrestrial biota

Marine subsidies mediate patterns in avian island biogeography

The classical theory of island biogeography, which predicts species richness using island area and isolation, has been expanded to include contributions from marine subsidies, i.e. subsidized island biogeography (SIB) theory. We tested the effects of marine subsidies on species diversity and population density on productive temperate islands, evaluating SIB predictions previously untested at comparable scales and subsidy levels. We found that the diversity of terrestrial breeding bird communities on 91 small islands (approx. 0.0001-3 km2) along the Central Coast of British Columbia, Canada were correlated most strongly with island area, but also with marine subsidies. Species richness increased and population density decreased with island area, but isolation had no measurable influence. Species richness was negatively correlated with marine subsidy, measured as forest-edge soil δ15N. Density, however, was higher on islands with higher marine subsidy, and a negative interaction between area and subsidy indicates that this effect is stronger on smaller islands, offering some support for SIB. Our study emphasizes how subsidies from the sea can shape diversity patterns on islands and can even exceed the importance of isolation in determining species richness and densities of terrestrial biota

Scale-dependent effects of marine subsidies on the island biogeographic patterns of plants

Although species richness can be determined by different mechanisms at different spatial scales, the role of scale in the effects of marine inputs on island biogeography has not been studied explicitly. Here, we evaluated the potential influence of island characteristics and marine inputs (seaweed wrack biomass and marine-derived nitrogen in the soil) on plant species richness at both a local (plot) and regional (island) scale on 92 islands in British Columbia, Canada. We found that the effects of subsidies on species richness depend strongly on spatial scale. Despite detecting no effects of marine subsidies at the island scale, we found that as plot level subsidies increased, species richness decreased; plots with more marine-derived nitrogen in the soil hosted fewer plant species. We found no effect of seaweed wrack at either scale. To identify potential mechanisms underlying the decrease in diversity, we fit a spatially explicit joint species distribution model to evaluate species level responses to marine subsidies and effects of biotic interactions among species. We found mixed evidence for competition for both light and nutrients, and cannot rule out an alternative mechanism; the observed decrease in species richness may be due to disturbances associated with animal-mediated nutrient deposits, particularly those from North American river otters (Lontra canadensis). By evaluating the scale-dependent effects of marine subsidies on island biogeographic patterns of plants and revealing likely mechanisms that act on community composition, we provide novel insights on the scale dependence of a fundamental ecological theory, and on the rarely examined links between marine and terrestrial ecosystems often bridged by animal vectors

Biogeographic features mediate marine subsidies to island food webs

Abstract Although marine subsidies often enrich terrestrial ecosystems, their influence is known to be context‐dependent. Additionally, the multitrophic impact of marine subsidies has not been traced through food webs across physically diverse islands. Here, we test predictions about how island characteristics can affect marine enrichment of food web constituents and how nutrients flow through island food webs. To evaluate enrichment and trace marine nutrients across food webs, we used stable isotopes of soil, flora, and fauna (n = 4752 samples) collected from 97 islands in British Columbia, Canada. Island area was the strongest predictor of enrichment across taxa; we found that samples were more 15N‐rich on smaller islands. Enrichment declined with distance from shore but less so on small islands, implying a higher per‐unit‐area subsidy effect. These area and distance‐to‐shore effects were taxon‐specific, and nearly twice as strong in basal food web groups. We also found that increases in δ15N correlated with increases in %N in basal trophic groups, as well as in songbirds, implying biologically relevant uptake of a potentially limiting nutrient. Path analysis demonstrated that subsidies in soil flow through plants and detritivores, and into upper‐level consumers. Our results reveal an interplay between island biogeography and marine subsidies in shaping island food webs through bottom‐up processes