Dynamics of Reciprocal Pulsed Subsidies in Local and Meta-Ecosystems

Temporally variable and reciprocal subsidies between ecosystems are ubiquitous. These spatial flows can generate a suite of direct and indirect effects in local and meta-ecosystems. The focus of most subsidy research, however, has been on the response of consumers in recipient ecosystems to constant subsidies over very short or very long time scales. We derive a meta-ecosystem model to explicitly consider the dynamic feedbacks between local ecosystems coupled through reciprocal pulsed subsidies. We predict oscillating reinforcing and dampening effects of reciprocal pulsed herbivore flows. Maximum reinforcing effects between reciprocal pulsed herbivore flows occur when these flows are in phase with the dynamics of neighboring predators. This prediction is robust to a range of pulse quantities and frequencies. Reciprocal pulsed herbivore subsidies lead to spatial and temporal variability in the strength of trophic cascades in local and meta-ecosystems but these cascading effects are the strongest when reciprocal pulsed subsidies are temporally concentrated. When predators demonstrate a behavioral response to prey abundance, reciprocal pulsed subsidies dampen the strength of local trophic cascades but lead to strong trophic cascades across local ecosystems. The timing of reciprocal pulsed subsidies is a critical component that determines the cascading effects of spatial flows. We show that spatial and temporal variabilities in resources and consumers can have a significant influence on the strength of cascading trophic interactions; therefore, our ability to detect and understand trophic cascades may depend on the scale of inquiry of ecological studies.     

Seasonal variation in the trophic structure of a spatial prey subsidy linking aquatic and terrestrial food webs: adult aquatic insects

Research over the past decade has established spatial resource subsidies as important determinants of food web dynamics. However, most empirical studies have considered the role of subsidies only in terms of magnitude, ignoring an important property of subsidies that may affect their impact in recipient food webs: the trophic structure of the subsidy relative to in situ resources. This may be especially important when subsidies are composed of organisms, as opposed to nutrient subsidies, because the trophic position of subsidy organisms may differ from in situ prey. I explored the relative magnitude and trophic structure of a cross-habitat prey subsidy, adult aquatic insects, in terrestrial habitats along three streams in the south–central United States. Overall, adult aquatic insects contributed more than one-third of potential insect prey abundance and biomass to the terrestrial habitat. This contribution peaked along a permanent spring stream, reaching as high as 94% of abundance and 86% of biomass in winter. Trophic structure of adult aquatic and terrestrial insects differed. Nearly all adult aquatic insects were non-consumers as adults, whereas all but one taxon of terrestrial insects were consumers. Such a difference created a strong relationship between the relative contribution of the prey subsidy and the trophic structure of the prey assemblage: as the proportion of adult aquatic insects increased, the proportion of consumers in the prey assemblage declined. Specific effects varied seasonally and with distance from the stream as the taxonomic composition of the subsidy changed, but general patterns were consistent. These findings show that adult aquatic insect subsidies to riparian food webs not only elevate prey availability, but also alter the trophic structure of the entire winged insect prey assemblage.     

Prey Vulnerability Limits Top-Down Control and Alters Reciprocal Feedbacks in a Subsidized Model Food Web

Resource subsidies increase the productivity of recipient food webs and can affect ecosystem dynamics. Subsidies of prey often support elevated predator biomass which may intensify top-down control and reduce the flow of reciprocal subsidies into adjacent ecosystems. However, top-down control in subsidized food webs may be limited if primary consumers posses morphological or behavioral traits that limit vulnerability to predation. In forested streams, terrestrial prey support high predator biomass creating the potential for strong top-down control, however armored primary consumers often dominate the invertebrate assemblage. Using empirically based simulation models, we tested the response of stream food webs to variations in subsidy magnitude, prey vulnerability, and the presence of two top predators. While terrestrial prey inputs increased predator biomass (+12%), the presence of armored primary consumers inhibited top-down control, and diverted most aquatic energy (∼75%) into the riparian forest through aquatic insect emergence. Food webs without armored invertebrates experienced strong trophic cascades, resulting in higher algal (∼50%) and detrital (∼1600%) biomass, and reduced insect emergence (−90%). These results suggest prey vulnerability can mediate food web responses to subsidies, and that top-down control can be arrested even when predator-invulnerable consumers are uncommon (20%) regardless of the level of subsidy.     

Species-specific use of allochthonous resources by ground beetles (Carabidae) at a river–land interface

A variety of organisms mediate river–terrestrial linkages through spatial subsidies. However, most empirical studies have classified organisms rather broadly (e.g., by functional group or taxonomic family) and have dismissed species-level linkages at the interface of ecosystems. Here, we show how allochthonous resource use varies among taxonomically similar species of ground beetles (family Carabidae) across seasons (June–September). We investigated seasonal shifts in the distribution of five beetle species and their dietary responses to spatial subsidies (emerging aquatic insects) in a Japanese braided river. Despite their taxonomic closeness, the ground beetles showed species-specific responses to spatial subsidies, and beetle distribution patterns tended to coincide with their diets. Overall, 1–56% of ground beetle diets were derived from aquatic prey. One genus (Bembidion spp.) mainly consumed aquatic prey, while three species fed primarily on terrestrial prey across all seasons. However, one species (Lithochlaenius noguchii) showed shifts in its diet from aquatic to terrestrial prey according to subsidy availability. The observed variation in allochthonous resource use was likely related to species-specific foraging modes, physiological tolerance to dry conditions, and interspecific competition. Our findings suggest that considering species-specific interactions is necessary to fully understand cross-system interactions and recipient food-web dynamics.     

More than a barrier: The complex effects of ecotone vegetation type on terrestrial consumer consumption of an aquatic prey resource

Community structure and dynamics can be influenced by resource transfers between ecosystems, yet little is known about how boundary structure determines both the magnitude of exchanges and their effects on recipient and donor communities. Aquatic and terrestrial ecosystems are often linked by resource fluxes and riparian vegetation is commonly affected by anthropogenic alterations to land use or river hydrological regime. I investigated whether shrubs at the freshwater–terrestrial interface alter the supply, distribution and importance of aquatic prey resources to terrestrial consumers. Shrubs were predicted to alter the larval community composition of aquatic insects and the emergence of winged adults, thus affecting aquatic prey subsidies to terrestrial consumers. In addition, shrubs were hypothesized to alter the microclimatic suitability of the riparian zone for adult aquatic insects, act as a physical barrier to their dispersal and affect terrestrial community composition, particularly the abundance and type of predators that could benefit from the aquatic prey resource. Stable isotope dietary analyses and a survey of shrub‐dominated and open grassland riparian habitats revealed that larval densities of aquatic insects (EPTM: Ephemeroptera, Plecoptera, Trichoptera and Megaloptera) were higher in shrub than grassland habitats; however, reduced emergence and lateral dispersal in shrub areas led to lower densities of adults. The temperature and relative humidity of the riparian zone did not differ between the habitats. Ground‐active terrestrial invertebrate communities had a higher proportion of cursorial spiders in grassland, coinciding with greater abundances of aquatic prey. Aquatic prey contribution to cursorial spider diet matched adult aquatic insect abundances. Overall, riparian shrubs reduced the magnitude, or at least altered the timing, of cross‐ecosystem subsidy supply, distribution and use by consumers through mechanisms operating in both the aquatic and terrestrial ecosystems. Thus, the structure of ecosystem boundaries has complex effects on the strength of biological interactions between adjacent systems.     

Spatial stoichiometry: cross‐ecosystem material flows and their impact on recipient ecosystems and organisms

Cross‐ecosystem material flows, in the form of inorganic nutrients, detritus and organisms, spatially connect ecosystems and impact food web dynamics. To date research on material flows has focused on the impact of the quantity of these flows and largely ignored their elemental composition, or quality. However, the ratios of elements like carbon, nitrogen and phosphorus can influence the impact material flows have on food web interactions through stoichiometric mismatches between resources and consumers. The type and movement of materials likely vary in their ability to change stoichiometric constraints within the recipient ecosystem and materials may undergo changes in their own stoichiometry during transport. In this literature review we evaluate the importance of cross‐ecosystem material flows within the framework of ecological stoichiometry. We explore how movement in space and time impacts the stoichiometry of material flow, as these transformations are essential to consider when assessing the ability of these flows to impact food web productivity and ecosystem functioning. Our review suggests that stoichiometry of cross‐ecosystem material flows are highly dynamic and undergo changes during transport across the landscape or from human influence. These material flows can impact recipient organisms if they change stoichiometry of the abiotic medium, or provide resources that have a different stoichiometry to in situ resources. They might also alter consumer excretion rates, in turn altering the availability of nutrients in the recipient ecosystem. These alterations in stoichiometric constraints of recipient organisms can have cascading trophic effects and shape food web dynamics. We highlight significant gaps in the literature and suggest new avenues for research that explore how cross‐ecosystem material flows impact recipient ecosystems when considering differences in stoichiometric quality, their movement through the landscape and across ecosystem boundaries, and the nutritional constraints of the recipient organisms.     

Resource subsidies between stream and terrestrial ecosystems under global change

Streams and adjacent terrestrial ecosystems are characterized by permeable boundaries that are crossed by resource subsidies. Although the importance of these subsidies for riverine ecosystems is increasingly recognized, little is known about how they may be influenced by global environmental change. Drawing from available evidence, in this review we propose a conceptual framework to evaluate the effects of global change on the quality and spatiotemporal dynamics of stream–terrestrial subsidies. We illustrate how changes to hydrological and temperature regimes, atmospheric CO₂ concentration, land use and the distribution of nonindigenous species can influence subsidy fluxes by affecting the biology and ecology of donor and recipient systems and the physical characteristics of stream–riparian boundaries. Climate‐driven changes in the physiology and phenology of organisms with complex life cycles will influence their development time, body size and emergence patterns, with consequences for adjacent terrestrial consumers. Also, novel species interactions can modify subsidy dynamics via complex bottom‐up and top‐down effects. Given the seasonality and pulsed nature of subsidies, alterations of the temporal and spatial synchrony of resource availability to consumers across ecosystems are likely to result in ecological mismatches that can scale up from individual responses, to communities, to ecosystems. Similarly, altered hydrology, temperature, CO₂ concentration and land use will modify the recruitment and quality of riparian vegetation, the timing of leaf abscission and the establishment of invasive riparian species. Along with morphological changes to stream–terrestrial boundaries, these will alter the use and fluxes of allochthonous subsidies associated with stream ecosystems. Future research should aim to understand how subsidy dynamics will be affected by key drivers of global change, including agricultural intensification, increasing water use and biotic homogenization. Our conceptual framework based on the match–mismatch between donor and recipient organisms may facilitate understanding of the multiple effects of global change and aid in the development of future research questions.     

Consumer-specific responses to riverine subsidy pulses in a riparian arthropod assemblage

1. Aquatic resource fluxes from streams can provide significant subsidies for riparian consumers. Because aquatic resource fluxes can be highly variable in space and time, the subsidy efficiency (i.e. transfer to the recipient food web) is controlled by the short‐term aggregative response of riparian consumers. 2. Field manipulations of stream‐derived invertebrate prey subsidies were used to examine specific aggregative responses of ground‐dwelling arthropods to riverine subsidy pulses in a braided‐river (Tagliamento River, NE Italy). Subsidy manipulation comprised short‐term reductions of natural stream‐derived subsidies and increased subsidies of stream‐derived invertebrate prey during four seasons. 3. We hypothesised that specific aggregative responses of riparian arthropods depend on their specialisation on aquatic insects which was inferred from stable isotope analysis. Natural riverine subsidy sources including aquatic insect emergence and surface‐drifting organisms were quantified. 4. Arthropods responded significantly with a reduction in abundance by 51%, at reduced subsidies and an increase by 110% at increased subsidies, when averaged over all seasons. Different arthropod taxa responded differently to subsidy manipulations in relation to their specialisation on aquatic subsidies: ground beetles with a diet consisting predominantly of aquatic insects responded only to subsidy reductions, indicating that their local abundance was not limited by natural stream‐derived subsidies; lycosid spiders with a partly aquatic diet showed no significant response; and ants, although relying on a terrestrial diet, responded positively to added stream‐derived invertebrate prey, indicating that stranding of surface‐drifting terrestrial invertebrates represented an important subsidy pathway. 5. Ground beetles and lycosid spiders were seasonally separated in their use of aquatic subsidies. Results indicate that the life‐history characteristics of riparian consumers can control the subsidy efficiency for the recipient community. By the effective uptake of pulsed riverine‐derived subsidies, riparian arthropods can enhance the transfer of riverine food sources to the riparian food web.     

Does the strength of cross‐ecosystem trophic cascades vary with ecosystem size? A test using a natural microcosm

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Marine resource flows to terrestrial arthropod predators on a temperate island: the role of subsidies between systems of similar productivity

Marine-terrestrial resource flows can subsidies recipient consumers at various trophic levels. Theory suggests that the importance of such spatial subsidies depends on the productivity gradient between adjacent systems; however, the empirical data required to test this assumption are scarce. Most studies of marine-terrestrial subsidies have been performed in arid coastal habitats of low productivity surrounded by productive ocean waters. We examined the importance of marine resource inputs for terrestrial consumers on a temperate, productive forest island surrounded by a marine system of similar productivity. The importance of marine resources for the dominant arthropod consumers was estimated using stable isotopes and linear mixing models. We compared isotopic signatures of spiders and ants captured along a gradient from shore to inland to estimate how far marine-derived energy penetrates the island. We evaluated the distribution of ground-dwelling arthropods using pitfall-trap transects extending from the supratidal-forest boundary to the middle of the island. The contribution of marine-derived energy assimilated by arthropod consumers differed both among taxa and location. Marine-derived resources contributed >80% to the assimilated C of intertidal spiders and 5-10% for spiders at the forest edge and further inland. Ants assimilated 20% of their C from marine-derived resources and this proportion was not affected by distance from shore. Spiders, ants, and all arthropods combined exhibited no spatial aggregation towards the shore. Our results indicate that on temperate islands marine-terrestrial subsidies might be predominantly an edge effect, confined to intertidal consumers. Mobile consumers that opportunistically forage in intertidal habitats play an important role in transferring marine-derived energy further inland. This suggests that the importance of the productivity gradient for spatial subsidies can be modified by the mobility traits of the recipient consumers and their degree of specialization on the interface habitat.     

Seasonal subsidy stabilizes food web dynamics: Balance in a heterogeneous landscape

Resource subsidies from external habitats can substantially affect the food web dynamics of local habitats. In this paper, we explore a mathematical model that is tailored for a stream food web, studied by Nakano and colleagues, in which consumers, in situ prey and subsidies all show seasonal fluctuation. The model reveals that the food web dynamics are stabilized if subsidies increase in summer when in situ productivity is low. Consumer dynamics are stabilized because subsidies complement seasonal resource deficiency. In situ prey dynamics are stabilized because subsidies indirectly balance the predation pressure by consumers, with seasonal change in prey carrying capacity. In summer when prey carrying capacity is low, seasonally abundant subsidies indirectly decrease predation pressure, whereas in winter, with high prey carrying capacity, scarce subsidies increase the predation pressure. Our results suggest that temporal productivity differences between spatially linked habitats are important to promote the stability of food web dynamics in a landscape context.     

Effects of an invasive plant transcend ecosystem boundaries through a dragonfly-mediated trophic pathway

Trophic interactions can strongly influence the structure and function of terrestrial and aquatic communities through top-down and bottom-up processes. Species with life stages in both terrestrial and aquatic systems may be particularly likely to link the effects of trophic interactions across ecosystem boundaries. Using experimental wetlands planted with purple loosestrife (Lythrum salicaria), we tested the degree to which the bottom-up effects of floral density of this invasive plant could trigger a chain of interactions, changing the behavior of terrestrial flying insect prey and predators and ultimately cascading through top-down interactions to alter lower trophic levels in the aquatic community. The results of our experiment support the linkage of terrestrial and aquatic food webs through this hypothesized pathway, with high loosestrife floral density treatments attracting high levels of visiting insect pollinators and predatory adult dragonflies. High floral densities were also associated with increased adult dragonfly oviposition and subsequently high larval dragonfly abundance in the aquatic community. Finally, high-flower treatments were coupled with changes in zooplankton species richness and shifts in the composition of zooplankton communities. Through changes in animal behavior and trophic interactions in terrestrial and aquatic systems, this work illustrates the broad and potentially cryptic effects of invasive species, and provides additional compelling motivation for ecologists to conduct investigations that cross traditional ecosystem boundaries.     

Tangled webs: reciprocal flows of invertebrate prey link streams and riparian zones

1. Streams and their adjacent riparian zones are closely linked by reciprocal flows of invertebrate prey. We review characteristics of these prey subsidies and their strong direct and indirect effects on consumers and recipient food webs. 2. Fluxes of terrestrial invertebrates to streams can provide up to half the annual energy budget for drift‐feeding fishes such as salmonids, despite the fact that input occurs principally in summer. Inputs appear highest from closed‐canopy riparian zones with deciduous vegetation and vary markedly with invertebrate phenology and weather. Two field experiments that manipulated this prey subsidy showed that it affected both foraging and local abundance of stream fishes. 3. Emergence of adult insects from streams can constitute a substantial export of benthic production to riparian consumers such as birds, bats, lizards, and spiders, and contributes 25–100% of the energy or carbon to such species. Emergence typically peaks in early summer in the temperate zone, but also provides a low‐level flux from autumn to spring in ice‐free streams. This flux varies with in‐stream productivity, and declines exponentially with distance from the stream edge. Some predators aggregate near streams and forage on these prey during periods of peak emergence, whereas others rely on the lower subsidy from autumn through spring when terrestrial prey are scarce. Several field experiments that manipulated this subsidy showed that it affected the short‐term behaviour, growth, and abundance of terrestrial consumers. 4. Reciprocal prey subsidies also have important indirect effects on both stream and riparian food webs. Theory predicts that allochthonous prey should increase density of subsidised predators, thereby increasing predation on in situ prey and causing a negative indirect effect via apparent competition. However, short‐term experiments have produced either positive or negative indirect effects. These contrasting results may be due to characteristics of the subsidies and individual consumers, but could also result from differences in experimental designs. 5. New study approaches are needed to better determine the direct and indirect effects of reciprocal prey subsidies. Experiments coupled with comparative research will be required to measure their effects on individual consumer fitness and population demographics. Future work should investigate whether reciprocal prey fluxes stabilise linked stream–riparian ecosystems, explore how landscape context affects the magnitude and importance of subsidies, and determine how impacts of human disturbance can propagate between streams and riparian zones via these trophic linkages. Study of these reciprocal connections is helping to define a more holistic perspective of catchments, and has the potential to shape new directions for ecology in general.     

Differential support of lake food webs by three types of terrestrial organic carbon

Organic carbon inputs from outside of ecosystem boundaries potentially subsidize recipient food webs. Four whole-lake additions of dissolved inorganic ¹³C were made to reveal the pathways of subsidies to lakes from terrestrial dissolved organic carbon (t-DOC), terrestrial particulate organic carbon (t-POC) and terrestrial prey items. Terrestrial DOC, the largest input, was a major subsidy of pelagic bacterial respiration, but little of this bacterial C was passed up the food web. Zooplankton received <2% of their C from the t-DOC to bacteria pathway. Terrestrial POC significantly subsidized the production of both zooplankton and benthic invertebrates, and was passed up the food web to Chaoborus and fishes. This route supplied 33–73% of carbon flow to zooplankton and 20–50% to fishes in non-fertilized lakes. Terrestrial prey, by far the smallest input, provided some fishes with >20% of their carbon. The results show that impacts of cross-ecosystem subsidies depend on characteristics of the imported material, the route of entry into the food web, the types of consumers present, and the productivity of the recipient system.     

Reducing dissolved phosphorus loading to the Salton Sea with aluminum sulfate

Terrestrial support of aquatic food webs is becoming well established in the science of ecology. However, while terrestrial subsidies of energy have been shown to exert strong effects on aquatic food webs, it is less clear how variations in these subsidies, via natural or anthropogenic factors, will affect recipient ecosystems. To assess the influence of variations in terrestrial subsidies on an aquatic food web, we manipulated leaf-litter inputs in artificial ponds. Decreasing litter inputs did not affect any of the response variables in artificial ponds. This may be because the minimal amount of terrestrial carbon present combined with autochthonous production was enough to sustain the food web and/or the food web was altered in ways not detected by the experimental design. However, increasing leaf-litter inputs increased the percent survival and developmental rate of larval wood frogs (Rana sylvatica). Conversely, increasing litter input appeared to have no influence on zooplankton or salamander larvae. Increasing litter inputs also increased the dissolved organic carbon content and decreased the percent saturation of dissolved oxygen in artificial ponds. As system respiration in aquatic systems is frequently dominated by microbial respiration, we hypothesize that the effects of increasing litter input on wood frogs were the result of an increase food resources (i.e., microbes) for tadpoles. The lack of a response by salamander larvae and zooplankton may be due to the densities of zooplankton in tanks providing enough food for salamanders in all treatments, variation among specific zooplankton species in their ability to exploit these resources and transfer energy to salamanders, or omnivory among zooplankton offsetting the affects of leaf-litter inputs. Additional work is needed to determine the influence of litter inputs on zooplankton and salamanders in this community. These data demonstrate that variations in leaf-litter inputs can influence food web structure; however, the importance of these variations will likely be dependent upon the trophic position of various consumers. Handling editor: J. Cole     

Context dependency of animal resource subsidies

The transport of resource subsidies by animals has been documented across a range of species and ecosystems. Although many of these studies have shown that animal resource subsidies can have significant effects on nutrient cycling, ecosystem productivity, and food‐web structure, there is a great deal of variability in the occurrence and strength of these effects. Here we propose a conceptual framework for understanding the context dependency of animal resource subsidies, and for developing and testing predictions about the effects of animal subsidies over space and time. We propose a general framework, in which abiotic characteristics and animal vector characteristics from the donor ecosystem interact to determine the quantity, quality, timing, and duration (QQTD) of an animal input. The animal input is translated through the lens of recipient ecosystem characteristics, which include both abiotic and consumer characteristics, to yield the QQTD of the subsidy. The translated subsidy influences recipient ecosystem dynamics through effects on both trophic structure and ecosystem function, which may both influence the recipient ecosystem's response to further inputs and feed back to influence the donor ecosystem. We present a review of research on animal resource subsidies across ecosystem boundaries, placed within the context of this framework, and we discuss how the QQTD of resource subsidies can influence trophic structure and ecosystem function in recipient ecosystems. We explore the importance of understanding context dependency of animal resource subsidies in increasingly altered ecosystems, in which the characteristics of both animal vectors and donor and recipient ecosystems may be changing rapidly. Finally, we make recommendations for future research on animal resource subsidies, and resource subsidies in general, that will increase our understanding and predictive capacity about their ecosystem effects.     

Increasing donor ecosystem productivity decreases terrestrial consumer reliance on a stream resource subsidy

Because nutrient enrichment can increase ecosystem productivity, it may enhance resource flows to adjacent ecosystems as organisms cross ecosystem boundaries and subsidize predators in recipient ecosystems. Here, we quantified the biomass and abundance of aquatic emergence and terrestrial spiders in a reference and treatment stream that had been continuously enriched with nitrogen and phosphorus for 5 years. Because we previously showed that enrichment increased secondary production of stream consumers, we predicted that aquatic emergence flux would be higher in the treatment stream, subsequently increasing the biomass and abundance of terrestrial spiders. Those increases were predicted to be greatest for spiders specializing on aquatic emergence subsidies (e.g., Tetragnathidae). By adding a ¹⁵N stable isotope tracer to both streams, we also quantified nitrogen flow from the stream into the riparian community. Emergence biomass, but not abundance, was higher in the treatment stream. The average body size of emerging adult insects and the relative dominance of Trichoptera adults were also greater in the treatment stream. However, spider biomass did not differ between streams. Spiders also exhibited substantially lower reliance on aquatic emergence nitrogen in the treatment stream. This reduced reliance likely resulted from shifts in the body size distributions and community composition of insect emergence that may have altered predator consumption efficiency in the treatment stream. Despite nutrient enrichment approximately doubling stream productivity and associated cross-ecosystem resource flows, the response of terrestrial predators depended more on the resource subsidy’s characteristics that affected the predator’s ability to capitalize on such increases.     

The dynamics of spatially coupled food webs

The dynamics of ecological systems include a bewildering number of biotic interactions that unfold over a vast range of spatial scales. Here, employing simple and general empirical arguments concerning the nature of movement, trophic position and behaviour we outline a general theory concerning the role of space and food web structure on food web stability. We argue that consumers link food webs in space and that this spatial structure combined with relatively rapid behavioural responses by consumers can strongly influence the dynamics of food webs. Employing simple spatially implicit food web models, we show that large mobile consumers are inordinately important in determining the stability, or lack of it, in ecosystems. More specifically, this theory suggests that mobile higher order organisms are potent stabilizers when embedded in a variable, and expansive spatial structure. However, when space is compressed and higher order consumers strongly couple local habitats then mobile consumers can have an inordinate destabilizing effect. Preliminary empirical arguments show consistency with this general theory.     

Ecosystem linkages revealed by experimental lake-derived isotope signal in heathland food webs

Cross-ecosystem movement of nutrients and biomass can have important effects on recipient systems. Emerging aquatic insects are subsidies to terrestrial ecosystems and can influence foodweb interactions in riparian systems. In a 2-year field experiment, we simulated aquatic insect deposition by adding adult midge carcasses (150?g dry mass m^?2 year^?1) to 1-m² heathland plots at a site with low natural midge deposition. We established four levels of midge-addition treatments and measured stable isotopes (??¹³C and ??¹⁵N) in plants and arthropods within each treatment. We used a multiple-source isotope Bayesian mixing model to estimate the terrestrial versus aquatic contribution to the diets of arthropods. Aquatic resources were incorporated into plant, detritivore, and predator biomass. Detritivorous Collembola showed the greatest difference in isotope values (+3??? ??¹⁵N and +4??? ??¹³C) between midge-addition and reference treatments. Isotope values of small spiders followed the same trend of enrichment as Collembola while other arthropods (mites and large spiders) were only enriched after 2?years of midge addition. Although predator diets did not change, they became isotopically enriched via their likely prey (Collembola). Plants also had elevated ??¹⁵N (+1???) in midge-addition treatments. The time required and amount of midge-derived C and N detected varied and depended on trophic position. Midge-derived nutrients were no longer present in arthropod biomass in the year following midge addition. Aquatic insect carcasses can be rapidly incorporated into terrestrial food webs in nearshore habitats, and repeated inputs can be detected at multiple trophic levels, thus highlighting the importance of the detrital pathway for aquatic to terrestrial cross-ecosystem subsidies.     

Trophic interception: how a boundary-foraging organism influences cross-ecosystem fluxes

The rate at which subsidies move between habitats is a function not only of those factors that facilitate such transfers, but also of factors that limit or restrict the movement of subsidies. The interruption or redirection of subsidies by organisms foraging at the boundary between habitats, or trophic interception, has the potential to substantially restructure the food webs of recipient habitats. In this study we describe how a waterstrider, Aquarius remigis , limits the transfer of a subsidy across the land-water boundary. Prey interception varied with the type of stream habitat; on average, waterstriders in isolated pools intercepted 71.8% of experimental prey inputs of all sizes compared with 21.5% in connected pools and 0.8% in riffles. Across all stream habitat types, waterstriders consumed 43.2% of the experimental inputs of the smallest prey used, the prey size class most similar to natural inputs in our study area. We estimate that foraging waterstriders may intercept 0.3–1.2 kg of terrestrial prey subsidy for every 100 m of stream channel during three summer months, resources that could otherwise support 13–58 young-of-the-year salmonids. In controlled trials, waterstriders significantly altered the amount of terrestrial prey directly consumed by fish, while fish also altered waterstrider interception of those prey. Interestingly, when waterstriders and fish were present together, more terrestrial prey were lost to the streambed than when either or both of these consumers were absent, making this resource available to benthic detritivores, and facilitating the direct incorporation of terrestrial nutrients into stream detrital webs. Overall, we demonstrate that organisms that forage at habitat boundaries can control the quantity and quality of subsidies arriving in recipient habitats, potentially altering food web structures within those habitats.