Resources from another place and time: responses to pulses in a spatially subsidized system

As the theoretical bases for the dynamics of spatially subsidized communities emerge, ecologists question whether spatially subsidized communities exhibit similar structure or dynamics to communities that receive strongly pulsed resources. In both cases, communities may be structured by responses to resources that are potentially absent at any given point in time (pulsed communities) or space (subsidized communities), even if pulsed resources are part of the in situ productivity of the system or the subsidies arrive as a relatively constant input from a nearby system. The potential for significant spatial or temporal resource limitation, therefore, may be a key factor influencing in similar ways the persistence of populations, the structure and dynamics of communities, and the evolution of specific life history traits. In most complex systems, however, multiple resources may arrive for various trophic entities at various points in time and from various points in space, and thus it may be difficult to separate or compare the dynamics of spatially subsidized and pulsed systems. In this paper, we explore the effects of interactions between pulses and subsidies in plant and animal populations and communities on highly pulsed and variably subsidized islands in the Gulf of California. While many of the plant and animal communities on the unsubsidized islands in this system respond to pulses of rain in classic ways, responses to these rain pulses on islands subsidized by seabird guano or other marine resources are quite different and variable, and depend on a combination of life history characteristics, physiology, competitive interactions, and trophic relationships. These variable responses to rain pulses then translate into large differences in dynamics and community structure of subsidized vs. unsubsidized islands. Indeed, most systems experience both temporal pulses and spatial subsidies. When considered in tandem, complementary or synergistic effects of the multiple, temporally and spatially variable resources may emerge that help explain complex food web structure and dynamics.     

Reciprocal subsidies between freshwater and terrestrial ecosystems structure consumer resource dynamics

Cross-ecosystem movements of material and energy, particularly reciprocal resource fluxes across the freshwater-land interface, have received major attention. Freshwater ecosystems may receive higher amounts of subsidies (i.e., resources produced outside the focal ecosystem) than terrestrial ecosystems, potentially leading to increased secondary production in freshwaters. Here we used a meta-analytic approach to quantify the magnitude and direction of subsidy inputs across the freshwater-land interface and to determine subsequent responses in recipient animals. Terrestrial and freshwater ecosystems differed in the magnitude of subsidies they received, with aquatic ecosystems generally receiving higher subsidies than terrestrial ecosystems. Surprisingly, and despite the large discrepancy in magnitude, the contribution of these subsidies to animal carbon inferred from stable isotope composition did not differ between freshwater and terrestrial ecosystems, likely due to the differences in subsidy quality. The contribution of allochthonous subsidies was highest to primary consumers and predators, suggesting that bottom-up and top-down effects may be affected considerably by the input of allochthonous resources. Future work on subsidies will profit from a food web dynamic approach including indirect trophic interactions and propagating effects.     

Predators, prey, and transient states in the assembly of spatially structured communities

Ecological theory suggests that both dispersal limitation and resource limitation can exert strong effects on community assembly. However, empirical studies of community assembly have focused almost exclusively on communities with a single trophic level. Thus, little is known about the combined effects of dispersal and resource limitation on assembly of communities with multiple trophic levels. We performed a landscape-scale experiment using spatially arranged mesocosms to study effects of dispersal and resource limitation on the assembly dynamics of aquatic invertebrate communities with two trophic levels. We found that interplay between dispersal and resource limitation regulated the assembly of predator and prey trophic levels in these pond communities. Early in assembly, predators and prey were strongly dispersal limited, and resource (i.e., prey) availability did not influence predator colonization. Later in assembly, after predators colonized, resource limitation was the strongest driver of predator abundance, and dispersal limitation played a negligible role. Thus, habitat isolation affected predators directly by reducing predator colonization rate, and indirectly through the effect of distance on prey availability. Dispersal and resource limitation of predators resulted in a transient period in which predators were absent or rare in isolated habitats. This period may be important for understanding population dynamics of vulnerable prey species. Our findings demonstrate that dispersal and resource limitation can jointly regulate assembly dynamics in multi-trophic systems. They also highlight the need to develop a temporal picture of the assembly process in multi-trophic communities because the availability and spatial distribution of limiting resources (i.e., prey) and the distribution of predators can shift radically over time.     

Environmental impact propagated by cross-system subsidy: chronic stream pollution controls riparian spider populations

Resource subsidies between habitats are common and create the potential for the propagation of environmental impacts across system boundaries. However, recent understanding of the potential for subsidy-mediated cross-system impact propagations is limited and primarily based on passive flows of nutrients and detritus or short-term effects. Here, we assess the effects of sustained alterations in aquatic insect emergence (active subsidy pathway), due to chronic stream pollution, for riparian spiders. The sustained reduction in aquatic insect densities at the polluted reaches resulted in a marked decline in web spider population density and a shift in spider community composition. Our results provide the first evidence that stream pollution can control populations and community structure of terrestrial predators via sustained alterations in aquatic subsidies, emphasizing the role of subtle trophic linkages in the transmission of environmental impacts across ecosystem boundaries.     

Anthropogenic subsidies alter stream consumer-resource stoichiometry, biodiversity, and food chains.

Urbanization is dramatically changing nutrient and organic matter regimes in streams, yet the community and ecosystem implications often remain obscure. We assessed the consequences of sewage-derived particulate organic matter (SDPOM) for invertebrate community structure and function in a headwater stream. Using stable isotope analyses, we found assimilation of organic SDPOM to double community secondary production, and stoichiometric analyses revealed SDPOM enriched in phosphorus (P) to foster putatively fast-growing, P-rich consumers in the subsidized reach. This altered consumer-resource stoichiometry impacted both community structure and nutrient fluxes through the invertebrate community. Community structure shifted toward significantly reduced diversity and evenness in the subsidized reach and consequently toward shorter food chains. Our integration of ecological stoichiometry with stable isotope analyses and food web ecology expands the previous focus of traditional ecotoxicology and ecophysiology to an ecosystem-level appreciation of pollutant ecology.     

Trophic segregation between sexes in the Black Skimmer revealed through the analysis of stable isotopes

The degree of individual or gender variation when exploiting food resources is an important aspect in the study of foraging ecology within a population. Previous information on non-breeding skimmers obtained through conventional methodologies suggested sex-related differences in prey species. In this study, stable isotope techniques were used to investigate the intraspecific segregation in diet and foraging habits of the Black Skimmer (Rynchops niger intercedens) at Mar Chiquita Coastal Lagoon (37°40′S, 57°22′W), Argentina. These results were compared with contemporary data on the trophic composition obtained by conventional methodologies. Blood samples were taken from birds captured with mist-nets during their non-breeding season. The isotopic signatures of skimmers showed a diet mainly composed of marine prey with some degree of estuarine fish intake. When comparing diet between sexes, males showed enrichment in ¹⁵N compared to females, while no differences were observed in ¹³C. The use of mixing models revealed differences in the relative composition of prey in the diet of male and female skimmers. This study highlights stable isotope analysis as a valuable tool to test inter-individual differences and sexual segregation in trophic ecology of Black Skimmers as compared to conventional methodologies. The results show a trophic segregation in the Black Skimmer during the non-breeding season that can be explained by differences in prey species and larger prey sizes of male skimmers. Our findings have significant implications for conservation since any environmental change occurring at wintering areas might have profound effects on several avian life-history traits, and could be different for males and females due to trophic segregation.     

Landscape of fear influences the relative importance of consumptive and nonconsumptive predator effects

Predators can initiate trophic cascades by consuming and/or scaring their prey. Although both forms of predator effect can increase the overall abundance of prey's resources, nonconsumptive effects may be more important to the spatial and temporal distribution of resources because predation risk often determines where and when prey choose to forage. Our experiment characterized temporal and spatial variation in the strength of consumptive and nonconsumptive predator effects in a rocky intertidal food chain consisting of the predatory green crab (Carcinus maenas), an intermediate consumer (the dogwhelk, Nucella lapillus), and barnacles (Semibalanus balanoides) as a resource. We tracked the survival of individual barnacles through time to map the strength of predator effects in experimental communities. These maps revealed striking spatiotemporal patterns in Nucella foraging behavior in response to each predator effect. However, only the nonconsumptive effect of green crabs produced strong spatial patterns in barnacle survivorship. Predation risk may play a pivotal role in determining the small-scale distribution patterns of this important rocky intertidal foundation species. We suggest that the effects of predation risk on individual foraging behavior may scale up to shape community structure and dynamics at a landscape level.     

Role of the fish Astyanax aeneus (Characidae) as a keystone nutrient recycler in low-nutrient neotropical streams

Nutrient recycling by animals is a potentially important biogeochemical process in both terrestrial and aquatic ecosystems. Stoichiometric traits of individual species may result in some taxa playing disproportionately important roles in the recycling of nutrients relative to their biomass, acting as keystone nutrient recyclers. We examined factors controlling the relative contribution of 12 Neotropical fish species to nutrient recycling in four streams spanning a range of phosphorus (P) levels. In high-P conditions (135 microg/L soluble reactive phosphorus, SRP), most species fed on P-enriched diets and P excretion rates were high across species. In low-P conditions (3 microg/L SRP), aquatic food resources were depleted in P, and species with higher body P content showed low rates of P recycling. However, fishes that were subsidized by terrestrial inputs were decoupled from aquatic P availability and therefore excreted P at disproportionately high rates. One of these species, Astyanax aeneus (Characidae), represented 12% of the total population and 18% of the total biomass of the fish assemblage in our focal low-P study stream but had P excretion rates > 10-fold higher than other abundant fishes. As a result, we estimated that P excretion by A. aeneus accounted for 90% of the P recycled by this fish assemblage and also supplied approximately 90% of the stream P demand in this P-limited ecosystem. Nitrogen excretion rates showed little variation among species, and the contribution of a given species to ecosystem N recycling was largely dependent upon the total biomass of that species. Because of the high variability in P excretion rates among fish species, ecosystem-level P recycling could be particularly sensitive to changes in fish community structure in P-limited systems.     

Diet of young Atlantic bluefin tuna (<Emphasis Type="Italic">Thunnus thynnus</Emphasis>) in eastern and western Atlantic foraging grounds

Atlantic bluefin tuna (Thunnus thynnus) are highly migratory predators whose abundance, distribution, and somatic condition have changed over the past decades. Prey community composition and abundance have also varied in several foraging grounds. To better understand underlying food webs and regional energy sources, we performed stomach content and stable isotope analyses on mainly juvenile (60–150 cm curved fork length) bluefin tuna captured in foraging grounds in the western (Mid-Atlantic Bight) and eastern (Bay of Biscay) Atlantic Ocean. In the Mid-Atlantic Bight, bluefin tuna diet was mainly sand lance (Ammodytes spp., 29% prey weight), consistent with historic findings. In the Bay of Biscay, krill (Meganyctiphanes norvegica) and anchovy (Engraulis encrasicolus) made up 39% prey weight, with relative consumption of each reflecting annual changes in prey abundance. Consumption of anchovies apparently declined after the local collapse of this prey resource. In both regions, stable isotope analysis results showed that juvenile bluefin tuna fed at a lower trophic position than indicated by stomach content analysis. In the Mid-Atlantic Bight, stable isotope analyses suggested that >30% of the diet was prey from lower trophic levels that composed <10% of the prey weights based upon traditional stomach content analyses. Trophic position was similar to juvenile fish sampled in the NW Atlantic but lower than juveniles sampled in the Mediterranean Sea in previous studies. Our findings indicate that juvenile bluefin tuna targeted a relatively small range of prey species and regional foraging patterns remained consistent over time in the Mid-Atlantic Bight but changed in relation to local prey availability in the Bay of Biscay.     

Aquatic and terrestrial organic matter in the diet of stream consumers: implications for mercury bioaccumulation

The relative contribution of aquatic vs. terrestrial organic matter to the diet of consumers in fluvial environments and its effects on bioaccumulation of contaminants such as mercury (Hg) remain poorly understood. We used stable isotopes of carbon and nitrogen in a gradient approach (consumer isotope ratio vs. periphyton isotope ratio) across temperate streams that range in their pH to assess consumer reliance on aquatic (periphyton) vs. terrestrial (riparian vegetation) organic matter, and whether Hg concentrations in fish and their prey were related to these energy sources. Taxa varied in their use of the two sources, with grazing mayflies (Heptageniidae), predatory stoneflies (Perlidae), one species of water strider (Metrobates hesperius), and the fish blacknose dace (Rhinichthys atratulus) showing strong connections to aquatic sources, while Aquarius remigis water striders and brook trout (Salvelinus fontinalis) showed a weak link to in-stream production. The aquatic food source for consumers, periphyton, had higher Hg concentrations in low-pH waters, and pH was a much better predictor of Hg in predatory invertebrates that relied mainly on this food source vs. those that used terrestrial C. These findings suggest that stream biota relying mainly on dietary inputs from the riparian zone will be partially insulated from the effects of water chemistry on Hg availability. This has implications for the development of a whole-system understanding of nutrient and material cycling in streams, the choice of taxa in contaminant monitoring studies, and in understanding the fate of Hg in stream food webs.     

Temperature-associated population diversity in salmon confers benefits to mobile consumers

Habitat heterogeneity can generate intraspecific diversity through local adaptation of populations. While it is becoming increasingly clear that population diversity can increase stability in species abundance, less is known about how population diversity can benefit consumers that can integrate across population diversity in their prey. Here we demonstrate cascading effects of thermal heterogeneity on trout-salmon interactions in streams where rainbow trout rely heavily on the seasonal availability of anadromous salmon eggs. Water temperature in an Alaskan stream varied spatially from 5 degrees C to 17.5 degrees C, and spawning sockeye salmon showed population differentiation associated with this thermal heterogeneity. Individuals that spawned early in cool regions of the 5 km long stream were genetically differentiated from those spawning in warmer regions later in the season. Sockeye salmon spawning generates a pulsed resource subsidy that supports the majority of seasonal growth in stream-dwelling rainbow trout. The spatial and temporal structuring of sockeye salmon spawn timing in our focal stream extended the duration of the pulsed subsidy compared to a thermally homogeneous stream with a single population of salmon. Further, rainbow trout adopted movement strategies that exploited the multiple pulses of egg subsidies in the thermally heterogeneous stream. Fish that moved to track the resource pulse grew at rates about 2.5 times higher than those that remained stationary or trout in the reference stream with a single seasonal pulse of eggs. Our results demonstrate that habitat heterogeneity can have important effects on the population diversity of dominant species, and in turn, influence their value to species that prey upon them. Therefore, habitat homogenization may have farther-reaching ecological effects than previously considered.     

Quantity and quality: unifying food web and ecosystem perspectives on the role of resource subsidies in freshwaters

Although the study of resource subsidies has emerged as a key topic in both ecosystem and food web ecology, the dialogue over their role has been limited by separate approaches that emphasize either subsidy quantity or quality. Considering quantity and quality together may provide a simple, but previously unexplored, framework for identifying the mechanisms that govern the importance of subsidies for recipient food webs and ecosystems. Using a literature review of > 90 studies of open-water metabolism in lakes and streams, we show that high-flux, low-quality subsidies can drive freshwater ecosystem dynamics. Because most of these ecosystems are net heterotrophic, allochthonous inputs must subsidize respiration. Second, using a literature review of subsidy quality and use, we demonstrate that animals select for high-quality food resources in proportions greater than would be predicted based on food quantity, and regardless of allochthonous or autochthonous origin. This finding suggests that low-flux, high-quality subsidies may be selected for by animals, and in turn may disproportionately affect food web and ecosystem processes (e.g., animal production, trophic energy or organic matter flow, trophic cascades). We then synthesize and review approaches that evaluate the role of subsidies and explicitly merge ecosystem and food web perspectives by placing food web measurements in the context of ecosystem budgets, by comparing trophic and ecosystem production and fluxes, and by constructing flow food webs. These tools can and should be used to address future questions about subsidies, such as the relative importance of subsidies to different trophic levels and how subsidies may maintain or disrupt ecosystem stability and food web interactions.     

Omnivory and grazer functional composition moderate cascading trophic effects in experimental <Emphasis Type="Italic">Fucus vesiculosus</Emphasis> habitats

We tested the relative strength of direct versus indirect effects of an aquatic omnivore depending on the functional composition of grazers by manipulating the presence of gastropod and amphipod grazers and omnivorous shrimp in outdoor mesocosms. By selectively preying upon amphipods and reducing their abundance by 70–80%, omnivorous shrimp favoured the dominance of gastropods. While gastropods were the main microalgal grazers, amphipods controlled macroalgal biomass in the experiment. However, strong predation on the amphipod by the shrimp had no significant indirect effects on macroalgal biomass, indicating that when amphipod abundances declined, complementary feeding by the omnivore on macroalgae may have suppressed a trophic cascade. Accordingly, in the absence of amphipods, the shrimp grazed significantly on green algae and thereby suppressed the diversity of the macroalgal community. Our experiment demonstrates direct consumer effects by an omnivore on both the grazer and producer trophic levels in an aquatic food web, regulated by prey availability.     

Stable C and N isotope analysis elucidated the importance of zooplankton in a tropical seagrass bed of Santiago Island,Northwestern Philippines

The role of zooplankton in a tropical seagrass ecosystem was investigated in milkfish farms pollution-impacted and -unimpacted seagrass beds in Santiago Island coral reefs, Northwestern Philippines. The aim was to compare between the two sites: (1) abiotic factors and zooplankton community parameters, and (2) the trophic structure using C and N stable isotopes. Low water (98–119?mV) and sediment (–121 to ?138?mV) Oxidation Reduction Potential values indicated a reducing environment in the impacted site. Zooplankton in the impacted site showed the typical community response to eutrophication (low diversity, but high total abundance due to the dominance of the cyclopoid copepod Oithona oculata), generally few elevated δ¹⁵N values, but a significant shift towards depleted ¹³C due to the organic enrichment of fish-farm feeds. Apart from suggesting a highly complex food web with POM and zooplankton as main food sources in the unimpacted site, the Bayesian mixing model simulation generated reduced complexity in feeding interactions between basal sources, zooplankton, and fish including adults of a key fish species, Siganus fuscescens, in the impacted sites. In this study, C and N stable isotope analysis has clarified the importance of zooplankton as fish prey in a seagrass bed food web.     

Trophic relationships of hydrothermal vent and non-vent communities in the upper sublittoral and upper bathyal zones off Kueishan Island,Taiwan: a combined morphological,gut content analysis and stable isotope approach

This study used morphological, gut content analysis and carbon- and nitrogen-stable isotope analysis to investigate the trophic structure of upper sublittoral (15–30 m deep) and upper bathyal (200–300 m deep) hydrothermal vents and the adjacent non-vent upper bathyal environment off Kueishan Island. The sublittoral vents host no chemosynthetic fauna, but green and red algae, epibiotic biofilm on crustacean surfaces, and zooplankton form the base of the trophic system. Suspension-feeding sea anemones and the generalist omnivorous vent crab Xenograpsus testudinatus occupy higher trophic levels. The upper bathyal hydrothermal vent is a chemoautotrophic-based system. The vent mussel Bathymodiolus taiwanensis forms a chemosynthetic component of this trophic system. Bacterial biofilm, surface plankton, and algae form the other dietary fractions of the upper bathyal fauna. The vent hermit crab Paragiopagurus ventilatus and the vent crab X. testudinatus are generalist omnivores. The vent-endemic tonguefish Symphurus multimaculatus occupies the top level of the trophic system. The adjacent non-vent upper bathyal region contains decapod crustaceans, which function as either predators or scavengers. The assemblages of X. testudinatus from sublittoral and upper bathyal vents exhibited distinct stable isotope values, suggesting that they feed on different food sources. The upper bathyal Xenograpsus assemblages displayed large variations in their stable isotope values and exhibited an ontogenetic shift in their δ¹³C and δ¹⁵N stable isotope signatures. Some individuals of Xenograpsus exhibited δ¹⁵N values close to those of non-vent species, suggesting that the highly mobile Xenograpsus may transfer energy between the upper bathyal hydrothermal vents and the adjacent non-vent upper bathyal environment.     

Can stable isotope ratios provide for community-wide measures of trophic structure?

Stable isotope ratios (typically of carbon and nitrogen) provide one representation of an organism's trophic niche and are widely used to examine aspects of food web structure. Yet stable isotopes have not been applied to quantitatively characterize community-wide aspects of trophic structure (i.e., at the level of an entire food web). We propose quantitative metrics that can be used to this end, drawing on similar approaches from ecomorphology research. For example, the convex hull area occupied by species in delta13C-delta15N niche space is a representation of the total extent of trophic diversity within a food web, whereas mean nearest neighbor distance among all species pairs is a measure of species packing within trophic niche space. To facilitate discussion of opportunities and limitations of the metrics, we provide empirical and conceptual examples drawn from Bahamian tidal creek food webs. These examples illustrate how this methodology can be used to quantify trophic diversity and trophic redundancy in food webs, as well as to link individual species to characteristics of the food web in which they are embedded. Building from extensive applications of stable isotope ratios by ecologists, the community-wide metrics may provide a new perspective on food web structure, function, and dynamics.     

Arthropod food web restoration following removal of an invasive wetland plant.

Restoration of habitats impacted by invasive plants is becoming an increasingly important tool in the management of native biodiversity, though most studies do not go beyond monitoring the abundance of particular taxonomic groups, such as the return of native vegetation. Yet, the reestablishment of trophic interactions among organisms in restored habitats is equally important if we are to monitor and understand how ecosystems recover. This study examined whether food web interactions among arthropods (as inferred by abundance of naturally occurring stable isotopes of C [delta13C] and N [delta15N]) were reestablished in the restoration of a coastal Spartina alterniflora salt marsh that had been invaded by Phragmites australis. From patterns of C and N stable isotopes we infer that trophic interactions among arthropods in the native salt marsh habitats are characterized by reliance on the dominant marsh plant Spartina as a basal resource. Herbivores such as delphacid planthoppers and mirid bugs have isotope signatures characteristic of Spartina, and predatory arthropods such as dolicopodid flies and spiders likewise have delta13C and delta15N signatures typical of Spartina-derived resources (approximately -13 per thousand and 10 per thousand, respectively). Stable isotope patterns also suggest that the invasion of Phragmites into salt marshes and displacement of Spartina significantly alter arthropod food web interactions. Arthropods in Phragmites-dominated sites have delta13C isotope values between -18 per thousand and -20 per thousand, suggesting reliance on detritus and/or benthic microalgae as basal resources and not on Phragmites, which has a delta13C approximately -26 per thousand. Since most Phragmites herbivores are either feeding internally or are rare transients from nearby Spartina, these resources do not provide significant prey resources for other arthropod consumers. Rather, predator isotope signatures in the invaded habitats indicate dependence on detritus/algae as basal resources instead of the dominant vegetation. The reestablishment of Spartina after removal of Phragmites, however, not only returned species assemblages typical of reference (uninvaded) Spartina, but stable isotope signatures suggest that the trophic interactions among the arthropods were also similar in reestablished habitats. Specifically, both herbivores and predators showed characteristic Spartina signatures, suggesting the return of the original grazer-based food web structure in the restored habitats.     

Species invasion shifts the importance of predator dependence

The strength of interference between foraging individuals can influence per capita consumption rates, with important consequences for predator and prey populations and system stability. Here we demonstrate how the replacement of a previously established invader, the predatory crab Carcinus maenas, by the recently invading predatory crab Hemigrapsus sanguineus shifts predation from a species that experiences strong predator interference (strong predator dependence) to one that experiences weak predator interference (weak predator dependence). We demonstrate using field experiments that differences in the strength of predator dependence persist for these species both when they forage on a single focal prey species only (the mussel Mytilus edulis) and when they forage more broadly across the entire prey community. This shift in predator dependence with species replacement may be altering the biomass across trophic levels, consistent with theoretical predictions, as we show that H. sanguineus populations are much larger than C. maenas populations throughout their invaded ranges. Our study highlights that predator dependence may differ among predator species and demonstrates that different predatory impacts of two conspicuous invasive predators may be explained at least in part by different strengths of predator dependence.     

Incorporating temporally dynamic baselines in isotopic mixing models

Stable isotopes (particularly C and N) are widely used to make inferences regarding food web structure and the phenology of consumer diet shifts, applications that require accurate isotopic characterization of trophic resources to avoid biased inferences of feeding relationships. For example, most isotope mixing models require that endmembers be adequately represented by a single probability distribution; yet, there is mounting evidence that the isotopic composition of aquatic organisms often used as mixing model endmembers can change over periods of weeks to months. A review of the literature indicated that the delta13C values of five aquatic primary consumer taxa, commonly used as proxies of carbon production sources (i.e., trophic baselines), express seasonally dynamic cycles characterized by an oscillation between summer maxima and winter minima. Based on these results, we built a dynamic baseline mixing model that allows a growing consumer to track temporal gradients in the isotopic baselines of a food web. Simulations showed that the ability of a consumer to maintain or approach isotopic equilibrium with its diet over a realistic growth season was strongly affected by both the rate of change of the isotopic baseline and equilibration rate of the consumer. In an empirical application, mixing models of varying complexity were used to estimate the relative contribution of benthic vs. pelagic carbon sources to nine species of juvenile fish in a fluvial lake of the St. Lawrence River system (Québec, Canada). Estimates of p (proportion of carbon derived from benthic sources) derived from a static mixing model indicated broad interspecific variation in trophic niche, ranging from complete benthivory to > 95% reliance on pelagic food webs. Output from the more realistic dynamic baseline mixing model increased estimated benthivory by an average of 36% among species. Taken together, our results demonstrate that failing to identify dynamic baselines when present, and (or) matching consumers with baseline taxa that possess substantially different equilibration rates can seriously bias interpretation of stable isotope data. Additionally, by providing a formalized framework that allows both resources and consumers to shift their isotopic value through time, our model demonstrates a feasible approach for incorporating temporally dynamic isotope conditions in trophic studies of higher consumers.     

Restoring piscivorous fish populations in the Laurentian Great Lakes causes seabird dietary change

Ecosystem change often affects the structure of aquatic communities thereby regulating how much and by what pathways energy and critical nutrients flow through food webs. The availability of energy and essential nutrients to top predators such as seabirds that rely on resources near the water's surface will be affected by changes in pelagic prey abundance. Here, we present results from analysis of a 25-year data set documenting dietary change in a predatory seabird from the Laurentian Great Lakes. We reveal significant declines in trophic position and alterations in energy and nutrient flow over time. Temporal changes in seabird diet tracked decreases in pelagic prey fish abundance. As pelagic prey abundance declined, birds consumed less aquatic prey and more terrestrial food. This pattern was consistent across all five large lake ecosystems. Declines in prey fish abundance may have primarily been the result of predation by stocked piscivorous fishes, but other lake-specific factors were likely also important. Natural resource management activities can have unintended consequences for nontarget ecosystem components. Reductions in pelagic prey abundance have reduced the capacity of the Great Lakes to support the energetic requirements of surface-feeding seabirds. In an environment characterized by increasingly limited pelagic fish resources, they are being offered a Hobsonian choice: switch to less nutritious terrestrial prey or go hungry.