Tracing energy flow in stream food webs using stable isotopes of hydrogen

1. Use of the natural ratios of carbon and nitrogen stable isotopes as tracers of trophic interactions has some clear advantages over alternative methods for food web analyses, yet is limited to situations where organic materials of interest have adequate isotopic separation between potential sources. This constrains the use of natural abundance stable isotope approaches to a subset of ecosystems with biogeochemical conditions favourable to source separation. 2. Recent studies suggest that stable hydrogen isotopes (δD) could provide a robust tracer to distinguish contributions of aquatic and terrestrial production in food webs, but variation in δD of consumers and their organic food sources are poorly known. To explore the utility of the stable hydrogen isotope approach, we examined variation in δD in stream food webs in a forested catchment where variation in δ¹³C has been described previously. 3. Although algal δD varied by taxa and, to a small degree, between sites, we found consistent and clear separation (by an average of 67‰) from terrestrial carbon sources. Environmental conditions known to affect algal δ¹³C, such as water velocity and stream productivity did not greatly influence algal δD, and there was no evidence of seasonal variation. In contrast, algal δ¹³C was strongly affected by environmental factors both within and across sites, was seasonally variable at all sites, and partially overlapped with terrestrial δ¹³C in all streams with catchment areas larger than 10 km². 4. While knowledge of isotopic exchange with water and trophic fractionation of δD for aquatic consumers is limited, consistent source separation in streams suggests that δD may provide a complementary food web tracer to δ¹³C in aquatic food webs. Lack of significant seasonal or spatial variation in δD is a distinct advantage over δ¹³C for applications in many aquatic ecosystems.     

Linking aboveground and belowground food webs through carbon and nitrogen stable isotope analyses

Carbon and nitrogen stable isotope ratios (δ¹³C and δ¹⁵N) have been used for more than two decades in analyses of food web structure. The utility of isotope ratio measurements is based on the observation that consumer δ¹³C values are similar (<1‰ difference) to those of their diet, while consumer δ¹⁵N values are about 3‰ higher than those of their diet. The technique has been applied most often to aquatic and aboveground terrestrial food webs. However, few isotope studies have examined terrestrial food web structure that includes both above- and belowground (detrital) components. Here, we review factors that may influence isotopic signatures of terrestrial consumers in above- and belowground systems. In particular, we emphasize variations in δ¹³C and δ¹⁵N in belowground systems, e.g., enrichment of ¹³C and ¹⁵N in soil organic matter (likely related to soil microbial metabolism). These enrichments should be associated with the high ¹³C (~3‰) enrichment in belowground consumers relative to litter and soil organic matter and with the large variation in δ¹⁵N (~6‰) of the consumers. Because such enrichment and variation are much greater than the trophic enrichment generally used to estimate consumer trophic positions, and because many general predators are considered dependent on energy and material flows from belowground, the isotopic variation in belowground systems should be taken into account in δ¹³C and δ¹⁵N analyses of terrestrial food webs. Meanwhile, by measuring the δ¹³C of key predators, the linkage between above- and belowground systems could be estimated based on observed differences in δ¹³C of primary producers, detritivores and predators. Furthermore, radiocarbon (¹⁴C) measurements will allow the direct estimation of the dependence of predators on the belowground systems.     

A meta‐analysis of the effects of detritus on primary producers and consumers in marine,freshwater, and terrestrial ecosystems

Detritus is a central feature in marine, freshwater, and terrestrial ecosystems. Despite the ubiquity of detritus, ecologists have largely ignored its role in influencing food web structure. We used a meta‐analytic approach to ask three questions about how detritus affects food web structure in a wide variety of ecosystems. First, what is the effect strength of detritus on primary producers, detritivores, herbivores, and predators? Second, what functional role does detritus serve for consumers (energetic, habitat, or both)? Third, how does the effect of detritus on consumers vary between aquatic and terrestrial ecosystems? We found that detritus has strong positive effects on primary producers and consumers in a wide range of ecosystems types. Detritus has a positive direct effect on detritivores by providing both an energetic resource and habitat (refuge from predators). Detritus has equally strong positive effects on herbivores and predators, driven by a positive direct effect of habitat. Detritus has positive effects on consumers in both aquatic and terrestrial ecosystems with 1.7 times stronger effects in terrestrial ecosystems. These results suggest that detritus has strong effects on food‐web structure in a variety of ecosystem types. Even the portion of the food web that is linked most strongly to living plant tissue as its primary energy source is strongly positively affected.     

Tracing biogeochemical subsidies from glacier runoff into Alaska's coastal marine food webs

Nearly half of the freshwater discharge into the Gulf of Alaska originates from landscapes draining glacier runoff, but the influence of the influx of riverine organic matter on the trophodynamics of coastal marine food webs is not well understood. We quantified the ecological impact of riverine organic matter subsidies to glacier‐marine habitats by developing a multi‐trophic level Bayesian three‐isotope mixing model. We utilized large gradients in stable (δ¹³C, δ¹⁵N, δ²H) and radiogenic (Δ¹⁴C) isotopes that trace riverine and marine organic matter sources as they are passed from lower to higher trophic levels in glacial‐marine habitats. We also compared isotope ratios between glacial‐marine and more oceanic habitats. Based on isotopic measurements of potential baseline sources, ambient water and tissues of marine consumers, estimates of the riverine organic matter source contribution to upper trophic‐level species including fish and seabirds ranged from 12% to 44%. Variability in resource use among similar taxa corresponded to variation in species distribution and life histories. For example, riverine organic matter assimilation by the glacier‐nesting seabirds Kittlitz's murrelet (Brachyramphus brevirostris) was greater than that of the forest‐nesting marbled murrelet (B. marmoratus). The particulate and dissolved organic carbon in glacial runoff and near surface coastal waters was aged (12100–1500 years BP ¹⁴C‐age) but dissolved inorganic carbon and biota in coastal waters were young (530 years BP ¹⁴C‐age to modern). Thus terrestrial‐derived subsidies in marine food webs were primarily composed of young organic matter sources released from glacier ecosystems and their surrounding watersheds. Stable isotope compositions also revealed a divergence in food web structure between glacial‐marine and oceanic sites. This work demonstrates linkages between terrestrial and marine ecosystems, and facilitates a greater understanding of how climate‐driven changes in freshwater runoff have the potential to alter food web dynamics within coastal marine ecosystems in Alaska.     

Coupling of canopy and understory food webs by ground‐dwelling predators

Understanding food‐web dynamics requires knowing whether species assemblages are compartmentalized into distinct energy channels, and, if so, how these channels are structured in space. We used isotopic analyses to reconstruct the food web of a Kenyan wooded grassland. Insect prey were relatively specialized consumers of either C₃ (trees and shrubs) or C₄ (grasses) plants. Arboreal predators (arthropods and geckos) were also specialized, deriving c. 90% of their diet from C₃‐feeding prey. In contrast, ground‐dwelling predators preyed considerably upon both C₃‐ and C₄‐feeding prey. This asymmetry suggests a gravity‐driven subsidy of the terrestrial predator community, whereby tree‐dwelling prey fall and are consumed by ground‐dwelling predators. Thus, predators in general couple the C₃ and C₄ components of this food web, but ground‐dwelling predators perform this ecosystem function more effectively than tree‐dwelling ones. Although prey subsidies in vertically structured terrestrial habitats have received little attention, they are likely to be common and important to food‐web organization.     

Predator and detritivore niche width helps to explain biocomplexity of experimental detritus-based food webs in four aquatic and terrestrial ecosystems

In the study of food webs, the existence and explanation of recurring patterns, such as the scale invariance of linkage density, predator–prey ratios and mean chain length, constitute long-standing issues. Our study focused on litter-associated food webs and explored the influence of detritivore and predator niche width (as δ¹³C range) on web topological structure. To compare patterns within and between aquatic and terrestrial ecosystems and take account of intra-habitat variability, we constructed 42 macroinvertebrate patch-scale webs in four different habitats (lake, lagoon, beech forest and cornfield), using an experimental approach with litterbags. The results suggest that although web differences exist between ecosystems, patterns are more similar within than between aquatic and terrestrial web types. In accordance with optimal foraging theory, we found that the niche width of predators and prey increased with the number of predators and prey taxa as a proportion of total taxa in the community. The tendency was more marked in terrestrial ecosystems and can be explained by a lower per capita food level than in aquatic ecosystems, particularly evident for predators. In accordance with these results, the number of links increased with the number of species but with a significantly sharper regression slope for terrestrial ecosystems. As a consequence, linkage density, which was found to be directly correlated to niche width, increased with the total number of species in terrestrial webs, whereas it did not change significantly in aquatic ones, where connectance scaled negatively with the total number of species. In both types of ecosystem, web robustness to rare species removal increased with connectance and the niche width of predators. In conclusion, although limited to litter-associated macroinvertebrate assemblages, this study highlights structural differences and similarities between aquatic and terrestrial detrital webs, providing field evidence of the central role of niche width in determining the structure of detritus-based food webs and posing foraging optimisation constraints as a general mechanistic explanation of food web complexity differences within and between ecosystem types.     

Trophic structure and major trophic links in conventional versus organic farming systems as indicated by carbon stable isotope ratios of fatty acids

Using bulk tissue and fatty acid ¹³C analysis we investigated major trophic pathways from soil microorganisms to microbial consumers to predators in conventional versus organic farming systems planted for the first time with maize. Organic farming led to an increase in microbial biomass in particular that of fungi as indicated by phospholipid fatty acids (PLFAs). Microbial PLFAs reflected the conversion from C₃ to C₄ plants by a shift in δ¹³C of 2‰, whereas the isotopic signal in fatty acids (FAs) of Collembola was much more pronounced. In the euedaphic Protaphorura fimata the δ¹³C values in maize fields exceeded that in C₃ (soybean) fields by up to 10‰, indicating a close relationship between diet and vegetation cover. In the epedaphic Orchesella villosaδ¹³C values shifted by 4‰, suggesting a wider food spectrum including carbon of former C₃ crop residues. Differences in δ¹³C of corresponding FAs in consumers and resources were assessed to assign food web links. P. fimata was suggested as root and fungal feeder in soybean fields, fungal feeder in conventional and leaf consumer in organically managed maize fields. O. villosa likely fed on root and bacteria under soybean, and bacteria and fungi under maize. Comparison of δ¹³C values in FAs of the cursorial spider Pardosaagrestis and O. villosa implied the latter as important prey species in soybean fields. In contrast, the web‐building spider Mangora acalypha showed no predator–prey relationship with Collembola. The determination of δ¹³C values in trophic biomarker FAs allowed detailed insight into the structure of the decomposer food web and identified diet‐shifts in both consumers at the base of the food web and in top predators in organic versus conventional agricultural systems. The results indicate changes in major trophic links and therefore carbon flux through the food web by conversion of conventional into organic farming systems.     

Nitrogen isotopes in otoliths reconstruct ancient trophic position

The ratio of ¹⁵N/¹⁴N (δ¹⁵N) from consumer and prey tissue is commonly used in ecological studies to determine trophic level, food web structure, and mean trophic level in aquatic ecosystems. There is a predictable positive relationship between the δ¹⁵N values in tissue and trophic level, caused by the bioaccumulation of ¹⁵N in tissues of consumers with each step up the food chain. Reconstructing trophic structure or food chain length over time may provide resource managers with insights about ecosystem biodiversity and resilience. Yet, in many marine systems the absence of baseline information before anthropogenic disturbances makes comparative studies addressing ecosystem responses extremely difficult. Here we attempt to retrospectively reconstruct trophic position in four species of fish from the upper Gulf of California, Mexico before perturbations such as overfishing or the damming of the Colorado River. We first validated if otolith δ¹⁵N approximates the δ¹⁵N observed in fish tissue. We then used the δ¹⁵N encapsulated in ancient fish otoliths that are between 1,000 and 5,500 years old to define the food web structure. Our results suggested that δ¹⁵N in otoliths has slightly more positive δ¹⁵N than soft tissue. The δ¹⁵N values from ancient otoliths appropriately defined the fishes’ relative trophic position. We found significant differences in δ¹⁵N between functional groups, apex predator versus intermediate predators. Juveniles and adult fishes displayed trophic separation between functional groups. Our findings advocate the application of δ¹⁵N analysis of prehistoric otoliths for establishing pre-disturbance ecological benchmarks.     

Variable littoral‐pelagic coupling as a food‐web response to seasonal changes in pelagic primary production

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Food Web Structure and Basal Resource Utilization along a Tropical Island Stream Continuum, Puerto Rico

Tropical stream food webs are thought to be based primarily on terrestrial resources (leaf litter) in small forested headwater streams and algal resources in larger, wider streams. In tropical island streams, the dominant consumers are often omnivorous freshwater shrimps that consume algae, leaf litter, insects, and other shrimps. We used stable isotope analysis to examine (1) the relative importance of terrestrial and algal‐based food resources to shrimps and other consumers and determine (2) if the relative importance of these food resources changed along the stream continuum. We examined δ¹⁵N and δ¹³C signatures of leaves, algae, macrophytes, biofilm, insects, snails, fishes, and shrimps at three sites (300, 90, and 10 m elev.) along the Río Espíritu Santo, which drains the Caribbean National Forest, Puerto Rico. Isotope signatures of basal resources were distinct at all sites. Results of two‐source δ¹³C mixing models suggest that shrimps relied more on algal‐based carbon resources than terrestrially derived resources at all three sites along the continuum. This study supports other recent findings in tropical streams, demonstrating that algal‐based resources are very important to stream consumers, even in small forested headwater streams. This study also demonstrates the importance of doing assimilation‐based analysis (i.e., stable isotope or trophic basis of production) when studying food webs.     

Lake size and fish diversity determine resource use and trophic position of a top predator in high‐latitude lakes

Prey preference of top predators and energy flow across habitat boundaries are of fundamental importance for structure and function of aquatic and terrestrial ecosystems, as they may have strong effects on production, species diversity, and food‐web stability. In lakes, littoral and pelagic food‐web compartments are typically coupled and controlled by generalist fish top predators. However, the extent and determinants of such coupling remains a topical area of ecological research and is largely unknown in oligotrophic high‐latitude lakes. We analyzed food‐web structure and resource use by a generalist top predator, the Arctic charr Salvelinus alpinus (L.), in 17 oligotrophic subarctic lakes covering a marked gradient in size (0.5–1084 km²) and fish species richness (2–13 species). We expected top predators to shift from littoral to pelagic energy sources with increasing lake size, as the availability of pelagic prey resources and the competition for littoral prey are both likely to be higher in large lakes with multispecies fish communities. We also expected top predators to occupy a higher trophic position in lakes with greater fish species richness due to potential substitution of intermediate consumers (prey fish) and increased piscivory by top predators. Based on stable carbon and nitrogen isotope analyses, the mean reliance of Arctic charr on littoral energy sources showed a significant negative relationship with lake surface area, whereas the mean trophic position of Arctic charr, reflecting the lake food‐chain length, increased with fish species richness. These results were supported by stomach contents data demonstrating a shift of Arctic charr from an invertebrate‐dominated diet to piscivory on pelagic fish. Our study highlights that, because they determine the main energy source (littoral vs. pelagic) and the trophic position of generalist top predators, ecosystem size and fish diversity are particularly important factors influencing function and structure of food webs in high‐latitude lakes.     

Broad sampling and diverse biomarkers allow characterization of nearshore particulate organic matter

Suspended particulate organic matter (POM) is a primary food source for benthic and pelagic consumers in aquatic and marine ecosystems. POM is potentially composed of many sources including phytoplankton, bacteria, zooplankton and macrophyte (seaweed and seagrass) and terrestrial detritus. The relative importance of these sources to POM consumers is debated, in large part due to differing interpretations of stable isotope and fatty acid biomarkers. We investigated POM composition in a nearshore marine ecosystem using multiple methods including visual quantification of living and detrital components, multiple stable isotope (MSI) and fatty acid (FA) analyses. Sampling was conducted at multiple temporal and spatial scales to 1) determine the range of variability in POM biomarkers, 2) quantitatively evaluate δ¹³C, δ¹⁵N, δ³⁴S and FA biomarkers with proportional abundance of putative sources and 3) determine the availability of phytoplankton, macrophytes and terrestrial carbon in nearshore POM. Variation of total FA concentration and proportions, and δ¹³C and δ³⁴S were strongly correlated to phytoplankton abundance, at tidal and seasonal timescales. Using multivariate multiple regressions, MSI and FA explained 59.6% and 89.7% of the variation in POM composition, respectively. As phytoplankton abundance increased, total FA concentration increased concurrent to δ¹³C and δ³⁴S enrichment. In high detritus samples, bacterial FA and saturated FA were proportionally higher, corresponding to depletion of δ¹³C and δ³⁴S and enrichment of δ¹⁵N. We identify MSI and FA biomarkers that are good predictors of diatom, dinoflagellate and detrital contributions to the POM. The results of this multi‐scale study show that POM composition is highly dynamic and largely driven by phytoplankton abundance, with minor contributions from terrestrial or macrophyte subsidies. This quantitative approach provides novel and critical empirical information linking POM compositional dynamics to specific biomarkers that are commonly used for tracking energy subsidies and biogeochemical cycling in aquatic ecosystems.     

The linkage between riparian predators and aquatic insects across a stream-resource spectrum

1. Spatial subsidies, defined as the flow of energy, nutrients, organisms or pollutants from one habitat to another, have been shown to affect the food–web dynamics in a wide range of ecosystems. An important subsidy to riparian communities is the contribution of adult stream insects to terrestrial predators such as birds, bats and lizards, but also invertebrates including ground and web‐building spiders. 2. We surveyed 37 first‐ and second‐order forest streams across differing environmental gradients in the Central South Island, New Zealand, to investigate the relationship between potential aquatic prey subsidies and predatory riparian arachnids. We anticipated that stream‐insect biomass would be positively associated with riparian arachnids, as a result of emergent adult aquatic insect subsidies to the adjacent habitat. 3. We confirmed positive associations between stream‐insect biomass as a predictor variable and riparian arachnid biomass (R² = 0.42, F_1,34 = 25.2, P < 0.001) and web densities (R² = 0.45, F_1,14 = 11.5, P < 0.01) respectively as dependent variables after adjusting for the confounding effects of environmental variables. Hierarchical partitioning confirmed the importance of stream insect biomass as a statistically significant contributor to the total explained variance in analyses calculated for arachnid biomass, abundance and web density. 4. A concurrent survey of spider‐web density along 20‐m transects from the stream edge into the forest indicated a strong decline in web‐building spider density moving away from the stream (R² = 0.41, F_1,158 = 109, P < 0.001), with stream‐insect biomass as a significant covariate (F_1,149 = 17.7, P < 0.001). 5. Our results suggest that productivity gradients present in the donor system affect the magnitude of the interaction between adjacent habitats. Productivity gradients may lead to increased reciprocal subsidies through a positive feedback loop involving the predation of spiders and other predatory terrestrial invertebrates by aquatic predators. However, terrestrial insectivores such as birds, bats and lizards that are not readily used as prey by aquatic predators may circumvent the feedback cycle by consuming a large proportion of emergent aquatic‐insect biomass. This may lead to asymmetry in the strength of food–web linkages between aquatic and terrestrial habitats.     

Stable isotope analyses on archived fish scales reveal the long‐term effect of nitrogen loads on carbon cycling in rivers

Stable isotope analysis of organic matter in sediment records has long been used to track historical changes in productivity and carbon cycling in marine and lacustrine ecosystems. While flow dynamics preclude stratigraphic measurements of riverine sediments, such retrospective analysis is important for understanding biogeochemical cycling in running waters. Unique collections of riverine fish scales were used to analyse δ¹⁵N and δ¹³C variations in the food web of two European rivers that experience different degrees of anthropogenic pressure. Over the past four decades, dissolved inorganic N loading remained low and constant in the Teno River (70°N, Finland); in contrast, N loading increased fourfold in the Scorff River (47°N, France) over the same period. Archived scales of Atlantic salmon parr, a riverine life‐stage that feeds on aquatic invertebrates, revealed high δ¹⁵N values in the Scorff River reflecting anthropogenic N inputs to that riverine environment. A strong correlation between dissolved inorganic N loads and δ¹³C values in fish scales was observed in the Scorff River, whereas no trend was found in the Teno River. This result suggests that anthropogenic N‐nutrients enhanced atmospheric C uptake by primary producers and its transfer to fish. Our results illustrate for the first time that, as for lakes and marine ecosystems, historical changes in anthropogenic N loading can affect C cycling in riverine food webs, and confirm the long‐term interactions between N and C biogeochemical cycles in running waters.     

Ontogenetic and individual diet variation in amphibian larvae across an environmental gradient

1. Variation among individuals within size or age classes can have profound effects on community dynamics and food‐web structure. We investigated the potential influence of habitat disturbance on intrapopulation niche variation. 2. Amphibians occupy a range of lentic habitats from short‐hydroperiod intermittent ponds to long‐hydroperiod permanent ponds. We quantified ontogenetic diet variation and individual specialisation in wood frog tadpoles (Lithobates sylvaticus) and blue‐spotted salamander larvae (Ambystoma laterale) to investigate the influence of hydroperiod on population niche width across a natural hydroperiod gradient using stable isotope and gut content analyses. In one of the few tests using larval forms, we tested the niche variation hypothesis, which predicts that populations with larger niche widths also have increased individual variation. 3. Our results support the niche variation hypothesis, indicating that more generalised populations exhibit higher within‐individual diet variation. We report gradual changes in the relative importance of diet items, decreased dietary overlap and increased trophic position in L. sylvaticus throughout development. A. laterale became more enriched in δ¹³C and increased in δ¹⁵N throughout its larval period. We did not find a relationship between hydroperiod and niche parameters, indicating that niches are conserved across heterogeneous habitats. In contrast to most documented cases, we estimated low levels of individual specialisation in amphibian larvae. 4. Amphibians are an important link between aquatic and terrestrial ecosystems, whereby diet shifts can influence food‐web structure by altering energy flow pathways and the trophic position of higher consumers, ultimately changing food‐chain length.     

Temporal and spatial trends in marine carbon isotopes in the Arctic Ocean and implications for food web studies

The Arctic is undergoing unprecedented environmental change. Rapid warming, decline in sea ice extent, increase in riverine input, ocean acidification and changes in primary productivity are creating a crucible for multiple concurrent environmental stressors, with unknown consequences for the entire arctic ecosystem. Here, we synthesized 30 years of data on the stable carbon isotope (δ¹³C) signatures in dissolved inorganic carbon (δ¹³C‐DIC; 1977–2014), marine and riverine particulate organic carbon (δ¹³C‐POC; 1986–2013) and tissues of marine mammals in the Arctic. δ¹³C values in consumers can change as a result of environmentally driven variation in the δ¹³C values at the base of the food web or alteration in the trophic structure, thus providing a method to assess the sensitivity of food webs to environmental change. Our synthesis reveals a spatially heterogeneous and temporally evolving δ¹³C baseline, with spatial gradients in the δ¹³C‐POC values between arctic shelves and arctic basins likely driven by differences in productivity and riverine and coastal influence. We report a decline in δ¹³C‐DIC values (?0.011‰ per year) in the Arctic, reflecting increasing anthropogenic carbon dioxide (CO₂) in the Arctic Ocean (i.e. Suess effect), which is larger than predicted. The larger decline in δ¹³C‐POC values and δ¹³C in arctic marine mammals reflects the anthropogenic CO₂ signal as well as the influence of a changing arctic environment. Combining the influence of changing sea ice conditions and isotopic fractionation by phytoplankton, we explain the decadal decline in δ¹³C‐POC values in the Arctic Ocean and partially explain the δ¹³C values in marine mammals with consideration of time‐varying integration of δ¹³C values. The response of the arctic ecosystem to ongoing environmental change is stronger than we would predict theoretically, which has tremendous implications for the study of food webs in the rapidly changing Arctic Ocean.     

Opportunistic top predators partition food resources in a tropical freshwater ecosystem

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Linking Land and Sea: Different Pathways for Marine Subsidies

Nutrients and energy derived from marine autotrophs subsidize shore ecosystems, increasing productivity and affecting food web dynamics and structure. In this study we examined how the inland reach of such inflow effects depends on vectors carrying the marine inflow inland and on landscape structure. We used stable isotopes of carbon and nitrogen to examine the roles of arthropod vectors in carrying marine-derived nutrients inland in two very different shore ecosystems: shore meadows in Sweden with marine inflows of algae and emerging chironomid midges; and sandy beaches and shore dunes in south-western Australia with marine inflows of algae and seagrass. In a colonization experiment, we found that deposited wrack on the beach is quickly colonized by both grazers and predators. However, in both systems we found a larger inland reach of the marine subsidy than could be accounted for by deposited macrophytes on shores alone, and that dipterans and spiders potentially functioned as vectors for the inflow. Our results indicate that marine inflows are important for near-shore terrestrial ecosystems well above the water’s edge, and that this effect is largely due to arthropod vectors (mainly dipterans and spiders) in both low-productivity sandy beach ecosystems at the Indian Ocean coast of Australia, and more productive shore meadows on the Baltic Sea coast of Sweden. Our findings also suggest that the type of vector transporting marine material inland may be as important as the productivity contrast between ecosystems for explaining the degree of marine influence on the terrestrial system.     

Ecosystem Linkages Between Lakes and the Surrounding Terrestrial Landscape in Northeast Iceland

Despite a recent emphasis on understanding cross-habitat interactions, few studies have examined the ecological linkages between lakes and surrounding terrestrial habitats. The current paradigm of land–lake interactions is typically unidirectional: the view is that nutrients and matter are transported downslope from the surrounding watershed to their ultimate lacustrine destination. Emergent aquatic insects, which spend their larval stages in lake sediments and emerge as adults to mate over land, can act as vectors of material, energy and nutrients from aquatic to terrestrial habitats. In this study, we document a gradient of midge (Diptera: Chironomidae) infall rates into terrestrial habitats (measured as g dw midges m^?2 d^?1) surrounding eight lakes in Northern Iceland (≈66°N latitude). Lakes ranged from having virtually no midge infall (for example, Helluvaðstjörn, 0.03 g m^?2 d^?1) to extreme levels (for example, Mývatn, 19 g m^?2 d^?1) with abundances of midges decreasing logarithmically with distance from shore. Annual midge input rates are estimated as high as 1200–2500 kg midges ha^?1 y^?1. As midges are approximately 9.2% total N, this can result in a significant fertilization effect of terrestrial habitats with consequences for plant quality and community structure. In addition, we used naturally-occurring δ¹³C and δ¹⁵N isotopes to examine food web structure and diet sources of terrestrial arthropod consumers surrounding lakes with differing amounts of midge input. Terrestrial arthropods showed increased utilization of aquatic-derived (that is, midge) C relative to terrestrial sources as midge infall increased. This pattern was particularly pronounced for predators, such as spiders and opiliones, and some detritivores (Collembola). These findings suggest that, despite being largely ignored, aquatic-to-terrestrial linkages can be large and midges can fuel terrestrial communities by directly serving as resources for predators and decomposers.