Sea to sky: impacts of residual salmon-derived nutrients on estuarine breeding bird communities

Pacific salmon (Oncorhynchus spp.) returning to streams around the North Pacific Rim provide a nutrient subsidy to these ecosystems. While many species of animals feed directly on salmon carcasses each autumn, salmon-derived nutrients can also be stored in coastal habitats throughout the year. The effects of this storage legacy on vertebrates in other seasons are not well understood, especially in estuaries, which can receive a large portion of post-spawning salmon nutrients. We examine the effects of residual salmon-derived nutrients, forest habitats and landscape features on summer breeding birds in estuary forests. We compared models containing environmental variables and combined chum (Oncorhynchus keta) and pink (Oncorhynchus gorbuscha) salmon biomass to test predictions concerning bird density and diversity. We discovered that total bird, insectivore, golden-crowned kinglet and Pacific wren densities and Shannon's diversity in the summer were strongly predicted by salmon biomass in the autumn. For most metrics, this relationship approaches an asymptote beyond 40 000 kg of salmon biomass. Foliage height diversity, watershed catchment area and estuary area were also important predictors of avian communities. Our study suggests that the legacy of salmon nutrients influences breeding bird density and diversity in estuaries that vary across a wide gradient of spawning salmon biomass.     

Uptake of salmon-derived nitrogen by mosses and liverworts in coastal British Columbia

Throughout forested watersheds bordering the North Pacific, anadromous salmon ( Oncorhynchus spp . ) are transferred into coastal forests by numerous predators and scavengers with remnants providing an important nutrient subsidy to riparian zones. The contribution of these nutrients to mosses and liverworts, which are the dominant ground cover of coastal forests, has not been investigated. We examine here δ¹⁵N isotope signatures and foliar percent N of eight moss and liverwort species, as well as moss and liverwort community structure, in multiple habitat blocks that vary in access to salmon nutrients from two watersheds on the central coast of British Columbia. For the most common moss Rhytidiadelphus loreus , we also examine δ¹⁵N values and foliar percent N among salmon carcass micro-sites and along wildlife trails. Overall, δ¹⁵N signatures ranged from 2‰ to 7‰ higher below the falls near the salmon stream than above the small falls that are impassable to salmon, or at the adjacent control watershed that had no salmon. Among micro-sites, δ¹⁵N and %N values were highest near bony carcass remnants from transfer during previous years and high adjacent to wildlife trails indicating spatial heterogeneity in the salmon nutrient pools of these forests. Species richness and prevalence of nitrogen-rich soil indicators were also highest in forests adjacent to the salmon stream. These data suggest an important contribution of salmon-derived nutrients to the non-vascular plants of the riparian zone of old-growth forests throughout the Pacific Rim.     

Salmon species, density and watershed size predict magnitude of marine enrichment in riparian food webs

Resource subsidies across habitat boundaries can structure recipient communities and food webs. In the northern Pacific region, bears Ursus spp. foraging on anadromous salmon Oncorhynchus spp. provide a key link between marine and terrestrial ecosystems, with salmon density, fish size and watershed size as potential predictors of the magnitude of marine subsidy to terrestrial habitats. We use nitrogen and carbon stable isotopes to provide an assessment of the patterns of marine‐enrichment in riparian plants (11 species, 4 guilds) and litter invertebrates (4 guilds) sampled from 27 watersheds in coastal British Columbia, Canada. Watersheds occurred in three geographical regions (Vancouver Island, mainland midcoast and Haida Gwaii) and varied in size, and in biomass (kg m^?1 of spawning length) and species of salmon (chum O. keta, pink O. gorbuscha and coho O. kisutch). δ¹⁵N values in all plant species and invertebrate guilds were positively predicted by total salmon biomass (kg m^?1) and negatively predicted by watershed size. We observed replicated parallel slopes among plant species and invertebrate guilds across the gradient in salmon biomass, with differences in means hypothesized to be due to plant fractionation and animal trophic position. As such, we derived a watershed δ¹⁵N‐index averaged across guilds, and using an information theoretic approach we find that the biomass of chum salmon is a much stronger predictor of the δ¹⁵N‐index than either pink or coho salmon, or the sum biomass of all species. The top linear model contained chum biomass and watershed size. Chum salmon biomass independently predicted δ¹⁵N‐index variation in all three regions of British Columbia. Chum salmon are larger than pink or coho and provide an energetic reward for bears that facilitates carcass transfer, tissue selective foraging, and nutrient distribution by insect scavengers. Analyses of biodiversity and habitat data across many watersheds moves towards a long‐term goal in fisheries ecology to better integrate ecosystem values in salmon conservation.     

Variability of <Emphasis Type="Italic">Mya arenaria</Emphasis> growth along an environmental gradient in the Plum Island Sound estuary,Massachusetts, USA

An understanding of the environmental factors that determine how clam growth varies in space and time improves effective mariculture and shellfish management. We examined the importance of temperature, salinity and chlorophyll-a in controlling the spatial pattern of Mya arenaria growth, the commercially important soft-shell clam, in the Plum Island Sound estuary in northeastern Massachusetts, USA. We collected clams (>5.08 cm) monthly during the April to November growing season from which we determined growth rate, maximum size (L-infinity), and time to reach a harvestable size. We also surveyed selected sites along the estuary to estimate the relationship between clam size and weight. We collected environmental data along the estuary, and our data were complemented with data collected and maintained by the Plum Island ecosystems long-term ecological research project. Clams reached harvestable size fastest and had the greatest L-infinity at the most oceanic site (Yacht Club) in the estuary. Clams had the smallest L-infinity and were slowest to reach the harvestable size at the least oceanic site (Railroad Meander). The spatial patterns of clam growth were best explained by a positive distribution of salinity. Salinity significantly accounted for 95 % of the spatial variation of clam growth in the estuary. Snow melt in spring increases freshwater input to the estuary and results in the lowest spring salinity during a year, and this explained the upper estuary limit of clam distribution. IPCC-projected climate change will cause sea-level rise and increasing precipitation in the northeastern USA, which will modify the spatial pattern of salinity in the region’s estuaries. Our research therefore suggests that future management of M. arenaria, an important economic resource for the local economy, should be concerned with the changes of salinity distribution under climate and land-use change.     

Ontogenetic shift to dependence on salmon-derived nutrients in Dolly Varden char from the Iliamna River,Alaska

The upstream migration, spawning, and death of anadromous, semelparous Pacific salmon brings nutrients to terrestrial and aquatic communities around the Pacific Rim. Many fishes use these resources but the relationship between fish body size and the reliance on salmon-derived nutrients might follow one of several patterns related to the onset of egg consumption with body size as fish grow, and possible shifts to alternative prey such as fishes as they grow larger still. In this study, these size-dependent hypotheses of marine subsidy use by resident Dolly Varden, Salvelinus malma, were tested using diet and stable isotope analyses. S. malma did not shift abruptly to a reliance on salmon eggs after they became large enough to eat eggs (i.e., no gape limitation). Rather, fish large enough to eat eggs but < 150 mm showed diets that blended salmon nutrients with aquatic insects, likely because they were spatially segregated from the highest concentration of spawning sockeye salmon, Oncorhynchus nerka. From intermediate through the largest sizes observed (150 to > 600 mm long) S. malma received ca. 80 % of their nutrients from salmon (eggs, flesh, and maggots that had scavenged dead salmon) based on diet analysis and stable isotope ratios despite being large enough to consume fish, as many similarly-sized salmonids do in other ecosystems. The few fish sampled in June, prior to the availability of salmon subsidies, had stable isotope signatures that also reflected heavy (ca. 90 %) reliance on marine sources, likely because they had eaten little since the end of the salmon run the previous fall. This apparent avoidance of piscivory in favor a rich yet pulsed marine subsidy highlights the importance of healthy salmon runs for the sake of not only the salmon but resident fishes that consume them.     

Quantitative Links Between Pacific Salmon and Stream Periphyton

Species’ impacts on primary production can have strong ecological consequences. In freshwater ecosystems, Pacific salmon (Oncorhynchus spp.) may influence stream periphyton through substrate disturbance during spawning and nutrient subsidies from senescent adults. The shape of relationships between the abundance of spawning salmon and stream periphyton, as well as interactions with environmental variables, are incompletely understood and may differ across the geographic range of salmon. We examined these relationships across 24 sockeye salmon (Oncorhynchus nerka) spawning streams in north-central British Columbia, Canada. The influence of salmon abundance and environmental variables (temperature, light, dissolved nutrients, water velocity, watershed size, and invertebrate grazer abundance) on post-spawning periphyton abundance and nitrogen stable isotope signatures, which can indicate the uptake of salmon nitrogen, was evaluated using linear regression models and Akaike Information Criterion. Periphyton nitrogen stable isotope signatures were best described by a positive log-linear relationship with an upstream salmon abundance metric that includes salmon from earlier years. This suggests the presence of a nutrient legacy. In contrast, periphyton abundance was negatively related to the spawning-year salmon density, which likely results from substrate disturbance during spawning, and positively related to dissolved soluble reactive phosphorus prior to spawning, which may indicate phosphorus limitation in the streams. These results suggest that enrichment from salmon nutrients does not always translate into elevated periphyton abundance. This underscores the need to directly assess the outcome of salmon impacts on streams rather than extrapolating from stable isotope evidence for the incorporation of salmon nutrients into food webs.     

Soft-Shell Clam, Mya arenaria, a Convenient Laboratory Animal for Screening Pathogens of Bivalve Mollusks

Attempts to introduce infectious or foreign material into oysters and other bivalve mollusks usually involve force or trauma because of immediate, prolonged adduction of the tightly closing valves. The soft-shell clam, Mya arenaria, is unable to seal its valves completely and relaxes readily, exposing soft tissue and a large siphon. This species is free from fouling organisms and is readily available at all seasons in the New England and mid-Atlantic areas. Suspensions of five strains of Vibrio sp. that cause bacillary necrosis in larval and juvenile bivalve mollusks were injected into the heart, siphon tissue, and the incurrent and excurrent siphon lumina of soft-shell clams. All vibrio strains caused significant mortality, usually within 2 days. Heaviest losses resulted from heart and excurrent siphon injections. No mortality occurred in control clams injected with seawater, broth, Serratia sp., and Escherichia coli. The soft-shell clam appears to be a useful animal for testing the pathogenicity of marine microorganisms for bivalve mollusks.     

Native species vulnerability to introduced predators: testing an inducible defense and a refuge from predation

To manage the impacts of biological invasions, it is important to determine the mechanisms responsible for the effects invasive species have on native populations. When predation by an invader is the mechanism causing declines in a native population, protecting the native species will involve elucidating the factors that affect native vulnerability. To examine those factors, this study measured how a native species responded to an introduced predator, and whether the native response could result in a refuge from predation. Predation by the green crab, Carcinus maenas, has contributed to the decline in numbers of native soft-shell clams, Mya arenaria, and efforts to eradicate crabs have proven futile. We tested how crab foraging affected clam burrowing, and how depth in the sediment affected clam survival. Clams responded to crab foraging by burrowing deeper in the sediment. Clams at shallow depths were more vulnerable to predation by crabs. Results suggest soft-shell clam burrowing is an inducible defense in response to green crab predation because burrowing deeper results in a potential refuge from predation by crabs. For restoring the native clam populations, tents could exclude crabs and protect clams, but when tents must be removed, exposing the clams to cues from foraging crabs should induce the clams to burrow deeper and decrease vulnerability. In general, by exposing potential native prey to cues from introduced predators, we can test how the natives respond, identify whether the response results in a potential refuge, and evaluate the risks to native species survival in invaded communities.     

Stream network geomorphology mediates predicted vulnerability of anadromous fish habitat to hydrologic change in southeast Alaska

In rivers supporting Pacific salmon in southeast Alaska, USA, regional trends toward a warmer, wetter climate are predicted to increase mid‐ and late‐21st‐century mean annual flood size by 17% and 28%, respectively. Increased flood size could alter stream habitats used by Pacific salmon for reproduction, with negative consequences for the substantial economic, cultural, and ecosystem services these fish provide. We combined field measurements and model simulations to estimate the potential influence of future flood disturbance on geomorphic processes controlling the quality and extent of coho, chum, and pink salmon spawning habitat in over 800 southeast Alaska watersheds. Spawning habitat responses varied widely across watersheds and among salmon species. Little variation among watersheds in potential spawning habitat change was explained by predicted increases in mean annual flood size. Watershed response diversity was mediated primarily by topographic controls on stream channel confinement, reach‐scale geomorphic associations with spawning habitat preferences, and complexity in the pace and mode of geomorphic channel responses to altered flood size. Potential spawning habitat loss was highest for coho salmon, which spawn over a wide range of geomorphic settings, including steeper, confined stream reaches that are more susceptible to streambed scour during high flows. We estimated that 9–10% and 13–16% of the spawning habitat for coho salmon could be lost by the 2040s and 2080s, respectively, with losses occurring primarily in confined, higher‐gradient streams that provide only moderate‐quality habitat. Estimated effects were lower for pink and chum salmon, which primarily spawn in unconfined floodplain streams. Our results illustrate the importance of accounting for valley and reach‐scale geomorphic features in watershed assessments of climate vulnerability, especially in topographically complex regions. Failure to consider the geomorphic context of stream networks will hamper efforts to understand and mitigate the vulnerability of anadromous fish habitat to climate‐induced hydrologic change.     

Predation on soft-shell clams (Mya arenaria) by the nemertean Cerebratulus lacteus in Atlantic Canada: implications for control measures

The nemertean, Cerebratulus lacteus Verrill (Nemertinea: Heteronemertini), has been identified as an important threat to soft-shell clam (Mya arenariaL.) populations in Atlantic Canada. The biology of this species, however, is still largely unknown. Field and laboratory studies were undertaken in 1998 and 1999 in Prince Edward Island, Canada, to test certain control measures to reduce predation on soft-shell clam populations and to better describe the relationship between C. lacteus and M. arenaria. Field abundance of C. lacteus, M. arenaria and Nereis virens Sars were evaluated in relation to particular habitat modifications that were used as control measures. Sediment manipulations tested were: (1) addition of shells and (2) use of a hydraulic rake. No difference was observed on the abundance of C. lacteus, M. arenaria and N. virens after treatments were applied. In the laboratory, C. lacteus was shown to be an efficient predator of M. arenaria. Clam mortalities reached 100% in the presence of C. lacteus while 0% mortality was observed in its absence. A complementary set of experiments was carried out to see if the sympatric polychaetes N. virens and Glycera dibranchiata Ehlers had any impact on the relationship between C. lacteus and M. arenaria. N. virens showed no impact on C. lacteus predation on clams. The presence of G. dibranchiata significantly reduced the nemertean predation rate. Analysis of clam size selection revealed no significant preference by C. lacteus. Other experimental studies revealed that high predator densities did not impede predation on clams and that C. lacteus preferred soft-shell clams among other commercial bivalve species presented (Mercenaria mercenariaL., Crassostrea virginica Gmelin and Mytilus edulisL.). This study should provide a better understanding of the relationship between C. lacteus and M. arenaria and lead to the development of improved nemertean control measures.     

Movers and shakers: nutrient subsidies and benthic disturbance predict biofilm biomass and stable isotope signatures in coastal streams

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Immunophenotyping of Mya arenaria neoplastic hemocytes using propidium iodide and a specific monoclonal antibody by flow cytometry

Disseminated neoplasia (DN) is a disorder referred to as hemic neoplasia (HN) in the soft-shell clam Mya arenaria. Traditionally, diagnosis is performed by hematocytology or histology. The intensity of the disease is generally given as the percentage of transformed neoplastic cells out of total number of hemocytes. Flow cytometry techniques have found a field of application in diagnosis of HN with analysis of ploidy. Hemocytes of the soft-shell clams with HN display tetraploid DNA content, as shown by propidium iodide staining. This feature makes difficult HN diagnosis in the soft-shell clam, especially for early stages of the condition, since the percentage of normal circulating cells undergoing mitosis, which also are tetraploid, remains unknown in molluscs. Use of specific monoclonal antibodies in a flow cytometry assay was foreseen as a way to overcome the difficulty. The purpose of this study was to develop a double staining protocol using propidium iodide for hemocyte cycle analysis and the MAb 1E10 for staining of HN cells. Our results showed a correlation between tetraploid and MAb 1E10-stained hemocytes in a single clam with moderate HN. This protocol offers some potential for further investigation of this cell disorder. However, a validation step will be necessary to confirm our preliminary results.     

Effects of Salmon-Derived Nutrients and Habitat Characteristics on Population Densities of Stream-Resident Sculpins

Movement of nutrients across ecosystem boundaries can have important effects on food webs and population dynamics. An example from the North Pacific Rim is the connection between productive marine ecosystems and freshwaters driven by annual spawning migrations of Pacific salmon (Oncorhynchus spp). While a growing body of research has highlighted the importance of both pulsed nutrient subsidies and disturbance by spawning salmon, their effects on population densities of vertebrate consumers have rarely been tested, especially across streams spanning a wide range of natural variation in salmon densities and habitat characteristics. We studied resident freshwater prickly (Cottus asper), and coastrange sculpins (C. aleuticus) in coastal salmon spawning streams to test whether their population densities are affected by spawning densities of pink and chum salmon (O. gorbuscha and O. keta), as well as habitat characteristics. Coastrange sculpins occurred in the highest densities in streams with high densities of spawning pink and chum salmon. They also were more dense in streams with high pH, large watersheds, less area covered by pools, and lower gradients. In contrast, prickly sculpin densities were higher in streams with more large wood and pools, and less canopy cover, but their densities were not correlated with salmon. These results for coastrange sculpins provide evidence of a numerical population response by freshwater fish to increased availability of salmon subsidies in streams. These results demonstrate complex and context-dependent relationships between spawning Pacific salmon and coastal ecosystems and can inform an ecosystem-based approach to their management and conservation.     

Physiological consequences of the salmon louse (Lepeophtheirus salmonis) on juvenile pink salmon (Oncorhynchus gorbuscha): implications for wild salmon ecology and management, and for salmon aquaculture

Pink salmon, Oncorhynchus gorbuscha, are the most abundant wild salmon species and are thought of as an indicator of ecosystem health. The salmon louse, Lepeophtheirus salmonis, is endemic to pink salmon habitat but these ectoparasites have been implicated in reducing local pink salmon populations in the Broughton Archipelago, British Columbia. This allegation arose largely because juvenile pink salmon migrate past commercial open net salmon farms, which are known to incubate the salmon louse. Juvenile pink salmon are thought to be especially sensitive to this ectoparasite because they enter the sea at such a small size (approx. 0.2 g). Here, we describe how 'no effect' thresholds for salmon louse sublethal impacts on juvenile pink salmon were determined using physiological principles. These data were accepted by environmental managers and are being used to minimize the impact of salmon aquaculture on wild pink salmon populations.     

Climate variation is filtered differently among lakes to influence growth of juvenile sockeye salmon in an Alaskan watershed

The physical landscape filters regional climate variation such that the environmental conditions an organism experiences are unique to the characteristics of local habitat features. While it has become apparent that populations may show watershed‐specific responses to changing climate within a geographic region, the population dynamics of Pacific salmon Oncorhynchus spp. suggest that within watershed climate filtering is also important. Growth provides an integrated measure of habitat quality capturing the overall response of individuals to climate as filtered by their habitat and their response to the biological interactions in the ecosystem. We used two different long‐term datasets, scales from returning adults and juvenile length measurements, to assess the response of sockeye salmon O. nerka growth during their juvenile life phase to single and integrated measures of climate within a watershed between 1950 and 2010. Scale growth showed evidence for differences among stocks rearing in different lake habitats within the same Alaska Peninsula watershed. These lakes have substantially different morphometry and showed opposite responses to changes in spring and fall air temperatures. Juvenile length data were also available for one of these stocks and indicated that density effects were relatively weak in contrast to the effects of temperature. While direct measures of juvenile length and measures derived from adult scales showed some similarity in their trends over time, they indicated opposite effects of air temperature. Throughout the range of Pacific salmon, climate change is altering freshwater ecosystems through changes to temperature, precipitation and associated variables. These data suggest that sockeye salmon populations are experiencing climate filtering at a sub‐watershed scale. Maintaining connected, heterogeneous landscapes will therefore likely be important for providing productive habitat for sockeye salmon across a range of climate conditions that they are going to experience under new climate regimes.     

The effects of water depth and density on the growth of a unionid clam*

SUMMARY. 1. Unionid clams from Narrow Lake, Alberta, were collected to quantify the natural variation in growth, to assess the natural variation in abundance, age and size distribution, and growth with water depth in the lake, and to conduct in situ experiments to directly test the effects of water depth (temperature) and clam abundance on clam growth. 2. The unionid clam, Anodonta grandis simpsoniana, showed wide variation in length at a given age. There were no significant differences in growth between clams collected at 1,3, 5, and 7m depths in the lake despite marked differences in water temperature. The wide variation in clam biomass within each depth zone may have masked possible effects of water depth. 3. The effect of water depth and variation in clam density on clam growth was tested directly by stocking clams into small enclosures at densities equivalent to 50, 150, 250, 350 and 450g m-2 (live weight) at each of 1, 3, 5 and 7 m depths in Narrow Lake (each depth and abundance treatment in triplicate). A uniform sandy substrate was used in all enclosures to eliminate any possible effect of substrate type on growth. 4. Mortality was negligible (0.9%) during the experiment. Clam density had no significant effect on clam growth which suggests that clam growth was not food limited in the lake. 5. Clams reared at 7 m grew more slowly than clams reared at 1, 3 and 5 m. Clams reared at 5 m grew more slowly than clams reared at 1 and 3m. Growth of clams reared at 1 and 3m did not differ. These differences in growth were strongly correlated with the measured differences in water temperature between depths. 6. Migration between depths probably accounts for the lack of a depth effect on clams growing in the natural habitat.     

Assessing California's bar-built estuaries using the California Rapid Assessment Method

Bar-built estuaries are generally found at the mouths of smaller watersheds with seasonal precipitation, episodic streamflow and seasonal swell dynamics. Low streamflows and constructive wave forces form a sand bar at the mouth isolating the stream from the ocean, creating a ponded lagoon, and inundating the surrounding marsh plain. Bar-built estuaries are wide spread in California comprising over 50 percent of California's more than 500 estuaries. By connecting terrestrial, freshwater, and marine realms bar-built estuaries are complex and dynamic providing great habitat and ecosystem services. California has suffered some of the highest losses of wetland habitats, and the wetland habitats of bar-built estuaries continue to be threatened by further development, pollution, and climate related changes including diminished streamflows and sea level rise. Given this loss and threat we developed a California Rapid Assessment Method (CRAM) to assess the condition of California's bar-built estuaries. CRAM uses visual indicators to accurately reflect current wetland condition with regards to buffer habitat, hydrology, physical complexity, plant diversity and structure, and landscape influences. Here we validate this method by comparing results of CRAM for bar-built estuaries to other accepted measures of wetland condition that we simultaneously collected with CRAM including vegetative surveys, water nutrient levels, and GIS landscape scale measures of stress for 32 sites throughout California. CRAM correlated well with each of these three independent methods of assessing condition. Notably, the Environmental Monitoring and Assessment Program (EMAP) number of natives metric significantly correlated with CRAM Index and all Attribute scores. The strong correlations of CRAM to nutrient levels is particularly important considering the documentation of the negative impact of nutrients on fish populations, the use of bar-built estuaries by juveniles of commercially important species, and the nursery role of bar-built estuaries for maintaining imperiled populations of species such as steelhead. GIS measured percent impervious, percent agriculture, and percent dams all correlated well with expected CRAM Attribute scores at appropriate watershed scales relative to the area of inference for each CRAM metric. Further, CRAM for bar-built estuaries works well throughout California's diversity of environmental conditions and regardless of geography, timing, or whether the bar was open or closed during the survey. We hope that the availability of CRAM combined with available data repositories will enable local, state, and federal decision makers to better manage, restore, and conserve valuable bar-built estuaries in the face of continual threats like development, drought, and sea level rise.     

Reduction of attractiveness to the sea star Asterias forbesi (Desor) by the clam Mercenaria mercenaria (Linnaeus)

Both in field and laboratory choice tests, the sea star, Asterias forbesi (Desor), was attracted to distant upstream clams, Mercenaria mercenaria (Linnaeus). Clams exposed to upstream sea stars were chosen less frequently by downstream sea stars than clams without sea stars upstream. Sea stars neither attracted nor repelled downstream conspecifics.When clams were exposed to upstream sea stars, their oxygen consumption decreased, as did their pumping rate and activity (as measured by number of visible siphons). The former may result from one or both of the latter.It is concluded that clam and sea star sense each other over a distance by chemical cues. The response of the clam is a general lowering of activity which may result in decreased attractiveness to sea star predators. This response may serve as a defensive measure against distance detection by Asterias forbesi.     

Ecosystem services transcend boundaries: estuaries provide resource subsidies and influence functional diversity in coastal benthic communities

Background

Estuaries are highly productive ecosystems that can export organic matter to coastal seas (the ‘outwelling hypothesis’). However the role of this food resource subsidy on coastal ecosystem functioning has not been examined.

Methodology/Principal Findings

We investigated the influence of estuarine primary production as a resource subsidy and the influence of estuaries on biodiversity and ecosystem functioning in coastal mollusk-dominated sediment communities. Stable isotope values (δ¹³C, δ¹⁵N) demonstrated that estuarine primary production was exported to the adjacent coast and contributed to secondary production up to 4 km from the estuary mouth. Further, isotope signatures of suspension feeding bivalves on the adjacent coast (Dosinia subrosea) closely mirrored the isotope values of the dominant bivalves inside the estuaries (Austrovenus stutchburyi), indicating utilization of similar organic matter sources. However, the food subsidies varied between estuaries; with estuarine suspended particulate organic matter (SPOM) dominant at Tairua estuary, while seagrass and fringing vegetation detritus was proportionately more important at Whangapoua estuary, with lesser contributions of estuarine SPOM. Distance from the estuary mouth and the size and density of large bivalves (Dosinia spp.) had a significant influence on the composition of biological traits in the coastal macrobenthic communities, signaling the potential influence of these spatial subsidies on ecosystem functioning.

Conclusions/Significance

Our study demonstrated that the locations where ecosystem services like productivity are generated are not necessarily where the services are utilized. Further, we identified indirect positive effects of the nutrient subsidies on biodiversity (the estuarine subsidies influenced the bivalves, which in turn affected the diversity and functional trait composition of the coastal sediment macrofaunal communities). These findings highlight the importance of integrative ecosystem-based management that maintains the connectivity of estuarine and coastal ecosystems.