Direct and indirect effects of ants on a forest-floor food web

Interactions among predators that prey on each other and are potential competitors for shared prey (intraguild [IG] predators) are widespread in terrestrial ecosystems and have the potential to strongly influence the dynamics of terrestrial food webs. Ants and spiders are abundant and ubiquitous terrestrial IG predators, yet the strength and consequences of interactions between them are largely unknown. In the leaf-litter food web of a deciduous forest in Kentucky (USA), we tested the direct and indirect effects of ants on spiders and a category of shared prey (Collembola) by experimentally subsidizing ants in open plots in two field experiments. In the first experiment, ant activity was increased, and the density of ants in the litter was doubled, by placing carbohydrate and protein baits in the center of each plot. Gnaphosa spiders were almost twice as abundant and Schizocosa spiders were half as abundant in baited plots relative to controls. There were more tomocerid Collembola in baited plots, suggesting possible indirect effects on Collembola caused by ant-spider interactions. The second experiment, in which screening of two mesh sizes selectively excluded large and small worker ants from a sugar bait, revealed that the large ants, primarily Camponotus, could alone induce similar effects on spiders. Gnaphosa biomass density was almost twice as high in the plots where large ants were more active, whereas Schizocosa biomass density was reduced by half in these plots. Although tomocerid densities did not differ between treatments, tomocerid numbers were negatively correlated with the activity of Formica, another large ant species. Path analysis failed to support the hypothesis that the ant Camponotus indirectly affected tomocerid Collembola through effects on densities of spiders. However, path analysis also revealed other indirect effects of Camponotus affecting tomocerids. These results illustrate the complexity of interactions between and within two major IG predator groups with disparate predatory behaviors, complexities that will have consequences for functioning of the forest-floor food web.     

Effects of Drainage‐Basin Geomorphology on Insectivorous Bird Abundance in Temperate Forests

Abstract: Interfaces between terrestrial and stream ecosystems often enhance species diversity and population abundance of ecological communities beyond levels that would be expected separately from both the ecosystems. Nevertheless, no study has examined how stream configuration within a watershed influences the population of terrestrial predators at the drainage‐basin scale. We examined the habitat and abundance relationships of forest insectivorous birds in eight drainage basins in a cool temperate forest of Japan during spring and summer. Each basin has different drainage‐basin geomorphology, such as the density and frequency of stream channels. In spring, when terrestrial arthropod prey biomass is limited, insectivorous birds aggregated in habitats closer to streams, where emerging aquatic prey was abundant. Nevertheless, birds ceased to aggregate around streams in summer because terrestrial prey became plentiful. Watershed‐scale analyses showed that drainage basins with longer stream channels per unit area sustained higher densities of insectivorous birds. Moreover, such effects of streams on birds continued from spring through summer, even though birds dispersed out of riparian areas in the summer. Although our data are from only a single year, our findings imply that physical modifications of stream channels may reduce populations of forest birds; thus, they emphasize the importance of landscape‐based management approaches that consider both stream and forest ecosystems for watershed biodiversity conservation.     

Allochthonous subsidy of periodical cicadas affects the dynamics and stability of pond communities

Periodical cicadas emerge from below ground every 13 or 17 years in North American forests, with individual broods representing the synchronous movement of trillions of individuals across geographic regions. Due to predator satiation, most individuals escape predation, die, and become deposited as detritus. Some of this emergent biomass falls into woodland aquatic habitats (small streams and woodland ponds) and serves as a high-quality allochthonous detritus pulse in early summer. We present results of a two-part study in which we (1) quantified deposition of Brood X periodical cicada detritus into woodland ponds and low-order streams in southwestern Ohio, and (2) conducted an outdoor mesocosm experiment in which we examined the effects of deposition of different amounts of cicada detritus on food webs characteristic of forest ponds. In the mesocosm experiment, we manipulated the amount of cicada detritus input to examine if food web dynamics and stability varied with the magnitude of this allochthonous resource subsidy, as predicted by numerous theoretical models. Deposition data indicate that, during years of periodical cicada emergence, cicada carcasses can represent a sizable pulse of allochthonous detritus to forest aquatic ecosystems. In the mesocosm experiment, cicada carcass deposition rapidly affected food webs, leading to substantial increases in nutrients and organism biomass, with the magnitude of increase dependent upon the amount of cicada detritus. Deposition of cicada detritus impacted the stability of organism functional groups and populations by affecting the temporal variability and biomass minima. However, contrary to theory, stability measures were not consistently related to the size of the allochthonous pulse (i.e., the amount of cicada detritus). Our study underscores the need for theory to further explore consequences of pulsed allochthonous subsidies for food web stability.     

Increased primary production shifts the structure and composition of a terrestrial arthropod community

Numerous studies have examined relationships between primary production and biodiversity at higher trophic levels. However, altered production in plant communities is often tightly linked with concomitant shifts in diversity and composition, and most studies have not disentangled the direct effects of production on consumers. Furthermore, when studies do examine the effects of plant production on animals in terrestrial systems, they are primarily confined to a subset of taxonomic or functional groups instead of investigating the responses of the entire community. Using natural monocultures of the salt marsh cordgrass Spartina alterniflora, we were able to examine the impacts of increased plant production, independent of changes in plant composition and/or diversity, on the trophic structure, composition, and diversity of the entire arthropod community. If arthropod species richness increased with greater plant production, we predicted that it would be driven by: (1) an increase in the number of rare species, and/or (2) an increase in arthropod abundance. Our results largely supported our predictions: species richness of herbivores, detritivores, predators, and parasitoids increased monotonically with increasing levels of plant production, and the diversity of rare species also increased with plant production. However, rare species that accounted for this difference were predators, parasitoids, and detritivores, not herbivores. Herbivore species richness could be simply explained by the relationship between abundance and diversity. Using nonmetric multidimensional scaling (NMDS) and analysis of similarity (ANOSIM), we also found significant changes in arthropod species composition with increasing levels of production. Our findings have important implications in the intertidal salt marsh, where human activities have increased nitrogen runoff into the marsh, and demonstrate that such nitrogen inputs cascade to affect community structure, diversity, and abundance in higher trophic levels.     

Above- and belowground responses of Arctic tundra ecosystems to altered soil nutrients and mammalian herbivory

Theory and observation indicate that changes in the rate of primary production can alter the balance between the bottom-up influences of plants and resources and the top-down regulation of herbivores and predators on ecosystem structure and function. The exploitation ecosystem hypothesis (EEH) posited that as aboveground net primary productivity (ANPP) increases, the additional biomass should support higher trophic levels. We developed an extension of EEH to include the impacts of increases in ANPP on belowground consumers in a similar manner as aboveground, but indirectly through changes in the allocation of photosynthate to roots. We tested our predictions for plants aboveground and for phytophagous nematodes and their predators belowground in two common arctic tundra plant communities subjected to 11 years of increased soil nutrient availability and/or exclusion of mammalian herbivores. The less productive dry heath (DH) community met the predictions of EEH aboveground, with the greatest ANPP and plant biomass in the fertilized plots protected from herbivory. A palatable grass increased in fertilized plots while dwarf evergreen shrubs and lichens declined. Belowground, phytophagous nematodes also responded as predicted, achieving greater biomass in the higher ANPP plots, whereas predator biomass tended to be lower in those same plots (although not significantly). In the higher productivity moist acidic tussock (MAT) community, aboveground responses were quite different. Herbivores stimulated ANPP and biomass in both ambient and enriched soil nutrient plots; maximum ANPP occurred in fertilized plots exposed to herbivory. Fertilized plots became dominated by dwarf birch (a deciduous shrub) and cloudberry (a perennial forb); under ambient conditions these two species coexist with sedges, evergreen dwarf shrubs, and Sphagnum mosses. Phytophagous nematodes did not respond significantly to changes in ANPP, although predator biomass was greatest in control plots. The contrasting results of these two arctic tundra plant communities suggest that the predictions of EEH may hold for very low ANPP communities, but that other factors, including competition and shifts in vegetation composition toward less palatable species, may confound predicted responses to changes in productivity in higher ANPP communities such as the MAT studied here.     

Nutrient subsidies to belowground microbes impact aboveground food web interactions

Historically, terrestrial food web theory has been compartmentalized into interactions among aboveground or belowground communities. In this study we took a more synthetic approach to understanding food web interactions by simultaneously examining four trophic levels and investigating how nutrient (nitrogen and carbon) and detrital subsidies impact the ability of the belowground microbial community to alter the abundance of aboveground arthropods (herbivores and predators) associated with the intertidal cord grass Spartina alterniflora. We manipulated carbon, nitrogen, and detrital resources in a field experiment and measured decomposition rate, soil nitrogen pools, plant biomass and quality, herbivore density, and arthropod predator abundance. Because carbon subsidies impact plant growth only indirectly (microbial pathways), whereas nitrogen additions both directly (plant uptake) and indirectly (microbial pathways) impact plant primary productivity, we were able to assess the effect of both belowground soil microbes and nutrient availability on aboveground herbivores and their predators. Herbivore density in the field was suppressed by carbon supplements. Carbon addition altered soil microbial dynamics (net potential ammonification, litter decomposition rate, DON [dissolved organic N] concentration), which limited inorganic soil nitrogen availability and reduced plant size as well as predator abundance. Nitrogen addition enhanced herbivore density by increasing plant size and quality directly by increasing inorganic soil nitrogen pools, and indirectly by enhancing microbial nitrification. Detritus adversely affected aboveground herbivores mainly by promoting predator aggregation. To date, the effects of carbon and nitrogen subsidies on salt marshes have been examined as isolated effects on either the aboveground or the belowground community. Our results emphasize the importance of directly addressing the soil microbial community as a factor that influences aboveground food web structure by affecting plant size and aboveground plant nitrogen.     

Using stable isotopes to reveal shifts in prey consumption by generalist predators.

The effectiveness of generalist predators in biological control may be diminished if increased availability of alternative prey causes individual predators to decrease their consumption of crop pests. Farming practices that enhance densities of microbidetritivores in the detrital food web can lead to increased densities of generalist predators that feed on pest species. The ability to predict the net biocontrol impact of increased predator densities depends upon knowing the extent to which individual predators may shift to detrital prey and feed less on crop pests when prey of the detritus-based food web are more abundant. We addressed this question by comparing ratios of stable isotopes of carbon (delta13C) and nitrogen (delta15N) in generalist ground predators and two types of prey (crop pests and microbidetritivores) in replicated 8 x 8 m cucurbit gardens subjected to one of two treatments: a detrital subsidy or no addition of detritus (control). Small sheet-web spiders (Linyphiidae) and small wolf spiders (Lycosidae) had delta13C values similar to those of Collembola in both the detrital and control treatments, indicating that small spiders belong primarily to the detrital food web. In control plots the larger generalist predators had delta13C values similar to those of the major insect pests, consistent with their known effectiveness as biocontrol agents. Adding detritus may have caused delta13C of one species of large wolf spider to shift toward that of the microbi-detritivores, although evidence is equivocal. In contrast, another large wolf spider displayed no shift in delta13C in the detrital treatment. Thus, stable isotopes revealed which generalist predators will likely continue to feed on pest species in the presence of greater densities of alternative prey.     

Mercury bioaccumulation in estuarine food webs

We tested for unintended mercury contamination problems associated with estuarine floodplain restoration projects of the Louisiana coastal zone, USA. Barataria Bay and Breton Sound are two neighboring deltaic estuaries that were isolated by levees from the Mississippi River about 100 years ago. These estuaries recently have been reconnected to the nutrient-rich Mississippi River, starting major river diversion (input) flows in 1991 for Breton Sound and in 2004 for Barataria Bay. We collected > 2100 fish over five years from 20 stations in these estuaries to test two hypotheses about Hg bioaccumulation: (H1) Background Hg bioaccumulation in fish would be highest in low-salinity upper reaches of estuaries, and (H2) recent river inputs to these upper estuarine areas would increase Hg bioaccumulation in fish food webs. For H1, we surveyed fish Hg concentrations at several stations along a salinity gradient in Barataria Bay in 2003-2004, a time when this estuary lacked strong river inputs. Results showed that average Hg concentrations in fish communities were lowest (150 ng/g dry mass) in higher salinity areas and -2.4x higher (350 ng/g) in low-salinity oligohaline and freshwater upper reaches of the estuary. For H2, we tested for enhanced Hg bioaccumulation following diversion onset in both estuaries. Fish communities from Breton Sound that had long-term (> 10 years) diversion inputs had -1.7x higher average Hg contents of 610 ng/g Hg vs. 350 ng/g background values. Shorter-term diversion inputs over 2-3 years in upper Barataria Bay did not result in strong Hg enrichments or stable C isotope increases seen in Breton Sound, even though N and S stable-isotope values indicated strong river inputs in both estuaries. It may be that epiphyte communities on abundant submerged aquatic vegetation (SAV) are important hotspots for Hg cycling in these estuaries, and observed lesser development of these epiphyte communities in upper Barataria Bay during the first years of diversion inputs may account for the lessened Hg bioaccumulation in fish. A management consideration from this study is that river restoration projects may unintentionally fertilize SAV and epiphyte-based food webs, leading to higher Hg bioaccumulation in river-impacted floodplains and their food webs.     

Introduced trout sever trophic connections in watersheds: consequences for a declining amphibian

Trophic linkages between terrestrial and aquatic ecosystems are increasingly recognized as important yet poorly known features of food webs. Here we describe research to understand the dynamics of lake food webs in relation to a native riparian amphibian and its interaction with introduced trout. The mountain yellow-legged frog Rana muscosa is endemic to alpine watersheds of the Sierra Nevada Mountains and the Transverse Ranges of California, but it has declined to a small fraction of its historical distribution and abundance. Although remaining frogs and introduced trout feed in different habitats of alpine lakes, our stable-isotope analyses clearly show that the same resource base of benthic invertebrates sustains their growth. During one period, insect emergence from naturally fishless lakes was nearly 20-fold higher compared to adjacent lakes with trout, showing that fish reduce availability of aquatic prey to amphibious and terrestrial consumers. Although trout cannot prey on adult frogs due to gape limitation, foraging post-metamorphic frogs are 10 times more abundant in the absence of trout, suggesting an important role for competition for prey by trout in highly unproductive alpine watersheds. Most Sierran lakes contain fish, and those that do not are usually small isolated ponds; in our study, these two lake types supported the lowest densities of post-metamorphic frogs, and these frogs were less reliant on local, benthic sources of productivity. Since Rana muscosa was formerly the most abundant vertebrate in the Sierra Nevada, the reduction in energy flow from lake benthos to this consumer due to fish introductions may have had negative consequences for its numerous terrestrial predators, many of which have also declined. We suggest that disruptions of trophic connections between aquatic and terrestrial food webs are an important but poorly understood consequence of fish introduction to many thousands of montane lakes and streams worldwide and may contribute to declines of native consumers in riparian habitats.     

Predator diversity and identity drive interaction strength and trophic cascades in a food web

Declining predator diversity may drastically affect the biomass and productivity of herbivores and plants. Understanding how changes in predator diversity can propagate through food webs to alter ecosystem function is one of the most challenging ecological research topics today. We studied the effects of predator removal in a simple natural food web in the Sierra Nevada mountains of California (USA). By excluding the predators of the third trophic level of a food web in a full-factorial design, we monitored cascading effects of varying predator diversity and composition on the herbivorous beetle Chrysomela aeneicollis and the willow Salix orestera, which compose the first and second trophic levels of the food web. Decreasing predator diversity increased herbivore biomass and survivorship, and consequently increased the amount of plant biomass consumed via a trophic cascade. Despite this simple linear mean effect of diversity on the strength of the trophic cascade, we found additivity, compensation, and interference in the effects of multiple predators on herbivores and plants. Herbivore survivorship and predator-prey interaction strengths varied with predator diversity, predator identity, and the identity of coexisting predators. Additive effects of predators on herbivores and plants may have been driven by temporal niche separation, whereas compensatory effects and interference occurred among predators with a similar phenology. Together, these results suggest that while the general trends of diversity effects may appear linear and additive, other information about species identity was required to predict the effects of removing individual predators. In a community that is not temporally well-mixed, predator traits such as phenology may help predict impacts of species loss on other species. Information about predator natural history and food web structure may help explain variation in predator diversity effects on trophic cascades and ecosystem function.     

Grazer traits, competition, and carbon sources to a headwater-stream food web

We investigated the effect of grazing by a dominant invertebrate grazer (the caddisfly Glossosoma penitum) on the energy sources used by other consumers in a headwater-stream food web. Stable isotope studies in small, forested streams in northern California have shown that G. penitum larvae derive most of their carbon from algae, despite low algal standing crops. We hypothesized that the caddisfly competes with other primary consumers (including mayflies) for algal food and increases their reliance on terrestrial detritus. Because Glossosoma are abundant and defended from predators by stone cases, their consumption of algal energy may reduce its transfer up the food chain. We removed Glossosoma (natural densities >1000 caddisflies/m2) from five approximately 4 m2) stream sections during the summer of 2000 and measured responses of algae, invertebrate primary consumers, and invertebrate predators. The treatment reduced Glossosoma biomass by 80-90%. We observed a doubling in chlorophyll a per area in sections with reduced Glossosoma abundance and aggregative increases in the biomass of undefended primary consumers. Heptageniid mayfly larvae consumed more algae (as measured by stable carbon isotope ratios and gut content analysis) in caddisfly removal plots at the end of the 60-day experiment, although not after one month. We did not see isotopic evidence of increased algal carbon in invertebrate predators, however. Patterns of caddisfly and mayfly diets in the surrounding watershed suggested that mayfly diets are variable and include algae and detrital carbon in variable proportions, but scraping caddisflies consume primarily algae. Caddisfly and mayfly diets are more similar in larger, more productive streams where the mayflies assimilate more algae. Isotopic analysis, in combination with measurements of macroinvertebrate abundance and biomass in unmanipulated plots, suggested that a substantial portion of the invertebrate community (>50% of biomass) was supported, at least partially, by local algal carbon during midsummer. These data suggest that algae may be more important to community dynamics in headwater streams than their relatively low productivity would suggest. Through their high densities and relative invulnerability to predation, armored grazers may also affect community structure and flow of algal and detrital carbon in headwater streams.     

Suppression of soybean aphid by generalist predators results in a trophic cascade in soybeans.

Top-down regulation of herbivores in terrestrial ecosystems is pervasive and can lead to trophic cascades that release plants from herbivory. Due to their relatively simplified food webs, agroecosystems may be particularly prone to trophic cascades, a rationale that underlies biological control. However, theoretical and empirical studies show that, within multiple enemy assemblages, intraguild predation (IGP) may lead to a disruption of top-down control by predators. We conducted a factorial field study to test the separate and combined effects of predators and parasitoids in a system with asymmetric IGP. Specifically we combined ambient levels of generalist predators (mainly Coccinellidae) of the soybean aphid, Aphis glycines Matsumura, with controlled releases of the native parasitoid Lysiphlebus testaceipes (Cresson) and measured their impact on aphid population growth and soybean biomass and yield. We found that generalist predators provided strong, season-long aphid suppression, which resulted in a trophic cascade that doubled soybean biomass and yield. However, contrary to our expectations, L. testaceipes provided minor aphid suppression and only when predators were excluded, which resulted in nonadditive effects when both groups were combined. We found direct and indirect evidence of IGP, but because percentage parasitism did not differ between predator exclusion and ambient predator treatments, we concluded that IGP did not disrupt parasitism during this study. Our results support theoretical predictions that intraguild predators which also provide strong herbivore suppression do not disrupt top-down control of herbivores.     

Terrestrial, benthic, and pelagic resource use in lakes: results from a three-isotope Bayesian mixing model

Fluxes of organic matter across habitat boundaries are common in food webs. These fluxes may strongly influence community dynamics, depending on the extent to which they are used by consumers. Yet understanding of basal resource use by consumers is limited, because describing trophic pathways in complex food webs is difficult. We quantified resource use for zooplankton, zoobenthos, and fishes in four low-productivity lakes, using a Bayesian mixing model and measurements of hydrogen, carbon, and nitrogen stable isotope ratios. Multiple sources of uncertainty were explicitly incorporated into the model. As a result, posterior estimates of resource use were often broad distributions; nevertheless, clear patterns were evident. Zooplankton relied on terrestrial and pelagic primary production, while zoobenthos and fishes relied on terrestrial and benthic primary production. Across all consumer groups terrestrial reliance tended to be higher, and benthic reliance lower, in lakes where light penetration was low due to inputs of terrestrial dissolved organic carbon. These results support and refine an emerging consensus that terrestrial and benthic support of lake food webs can be substantial, and they imply that changes in the relative availability of basal resources drive the strength of cross-habitat trophic connections.     

Asymmetry in community regulation: effects of predators and productivity

Eutrophication and predator additions and extinctions are occurring in ecosystems worldwide. Although theory predicts that both will strongly alter the distribution of biomass in whole communities, empirical evidence has not been consolidated to quantitatively determine whether these theoretical predictions are generally borne out in real ecosystems. Here we analyze data from two types of trophic cascade studies, predator removals in factorial combination with fertilization and observed productivity gradients, to assess the role of top-down and bottom-up forces in structuring multi-trophic communities and compare results from these analyses to those from an extensive database of trophic cascade studies. We find that herbivore biomass declines and plant biomass increases in the presence of predators, regardless of system productivity. In contrast, while plants are increased by fertilization, this effect does not significantly increase herbivores in either the presence or absence of predators. These patterns are consistent among marine, freshwater, and terrestrial ecosystems and are largely independent of study size and duration. Thus, top-down effects of predation are transferred through more trophic levels than are bottom-up effects of eutrophication, showing strong asymmetry in the direction of control of biomass distribution in communities.     

Nutrient enrichment reduces constraints on material flows in a detritus-based food web

Most aquatic and terrestrial ecosystems are experiencing increased nutrient availability, which is affecting their structure and function. By altering community composition and productivity of consumers, enrichment can indirectly cause changes in the pathways and magnitude of material flows in food webs. These changes, in turn, have major consequences for material storage and cycling in the ecosystem. Understanding mechanisms and predicting consequences of nutrient-induced changes in material flows requires a quantitative food web approach that combines information on consumer energetics and consumer-resource stoichiometry. We examined effects of a whole-system experimental nutrient enrichment on the trophic basis of production and the magnitude and pathways of carbon (C), nitrogen (N), and phosphorus (P) flows in a detritus-based stream food web. We compared the response of the treated stream to an adjacent reference stream throughout the study. Dietary composition and elemental flows varied considerably among invertebrate functional feeding groups. During nutrient enrichment, increased flows of leaf litter and amorphous detritus to shredders and gatherers accounted for most of the altered flows of C from basal resources to consumers. Nutrient enrichment had little effect on patterns of material flows but had large positive effects on the magnitude of C, N, and P flows to consumers (mean increase of 97% for all elements). Nutrient-specific food webs revealed similar flows of N and P to multiple functional groups despite an order of magnitude difference among groups in consumption of C. Secondary production was more strongly related to consumption of nutrients than C, and increased material flows were positively related to the degree of consumer-resource C:P and C:N imbalances. Nutrient enrichment resulted in an increased proportion of detrital C inputs consumed by primary consumers (from -15% to 35%) and a decreased proportion of invertebrate prey consumed by predators (from -80% to 55%). Our results demonstrate that nutrient enrichment of detritus-based systems may reduce stoichiometric constraints on material flows, increase the contribution of consumers to C, N, and P cycling, alter the proportion of C inputs metabolized by consumers, and potentially lead to reduced ecosystem-level storage of C.     

Whole-system nutrient enrichment increases secondary production in a detritus-based ecosystem

Although the effects of nutrient enrichment on consumer-resource dynamics are relatively well studied in ecosystems based on living plants, little is known about the manner in which enrichment influences the dynamics and productivity of consumers and resources in detritus-based ecosystems. Because nutrients can stimulate loss of carbon at the base of detrital food webs, effects on higher consumers may be fundamentally different than what is expected for living-plant-based food webs in which nutrients typically increase basal carbon. We experimentally enriched a detritus-based headwater stream for two years to examine the effects of nutrient-induced changes at the base of the food web on higher metazoan (predominantly invertebrate) consumers. Our paired-catchment design was aimed at quantifying organic matter and invertebrate dynamics in the enriched stream and an adjacent reference stream for two years prior to enrichment and two years during enrichment. Enrichment had a strong negative effect on standing crop of leaf litter, but no apparent effect on that of fine benthic organic matter. Despite large nutrient-induced reductions in the quantity of leaf litter, invertebrate secondary production during the enrichment was the highest ever reported for headwater streams at this Long Term Ecological Research site and was 1.2-3.3 times higher than predicted based on 15 years of data from these streams. Abundance, biomass, and secondary production of invertebrate consumers increased significantly in response to enrichment, and the response was greater among taxa with larval life spans < or = 1 yr than among those with larval life spans >1 yr. Production of invertebrate predators closely tracked the increased production of their prey. The response of invertebrates was largely habitat-specific with little effect of enrichment on food webs inhabiting bedrock outcrops. Our results demonstrate that positive nutrient-induced changes to food quality likely override negative changes to food quantity for consumers during the initial years of enrichment of detritus-based stream ecosystems. Longer-term enrichment may impact consumers through eventual reductions in the quantity of detritus.     

Disentangling trophic relationships in a High Arctic tundra ecosystem through food web modeling

Determining the manner in which food webs will respond to environmental changes is difficult because the relative importance of top-down vs. bottom-up forces in controlling ecosystems is still debated. This is especially true in the Arctic tundra where, despite relatively simple food webs, it is still unclear which forces dominate in this ecosystem. Our primary goal was to assess the extent to which a tundra food web was dominated by plant-herbivore or predator-prey interactions. Based on a 17-year (1993-2009) study of terrestrial wildlife on Bylot Island, Nunavut, Canada, we developed trophic mass balance models to address this question. Snow Geese were the dominant herbivores in this ecosystem, followed by two sympatric lemming species (brown and collared lemmings). Arctic foxes, weasels, and several species of birds of prey were the dominant predators. Results of our trophic models encompassing 19 functional groups showed that <10% of the annual primary production was consumed by herbivores in most years despite the presence of a large Snow Goose colony, but that 20-100% of the annual herbivore production was consumed by predators. The impact of herbivores on vegetation has also weakened over time, probably due to an increase in primary production. The impact of predators was highest on lemmings, intermediate on passerines, and lowest on geese and shorebirds, but it varied with lemming abundance. Predation of collared lemmings exceeded production in most years and may explain why this species remained at low density. In contrast, the predation rate on brown lemmings varied with prey density and may have contributed to the high-amplitude, periodic fluctuations in the abundance of this species. Our analysis provided little evidence that herbivores are limited by primary production on Bylot Island. In contrast, we measured strong predator-prey interactions, which supports the hypothesis that this food web is primarily controlled by top-down forces. The presence of allochthonous resources subsidizing top predators and the absence of large herbivores may partly explain the predominant role of predation in this low-productivity ecosystem.     

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.     

Seed availability and insect herbivory limit recruitment and adult density of native tall thistle

Understanding spatial and temporal variation in factors influencing plant regeneration is critical to predicting plant population growth. We experimentally evaluated seed limitation, insect herbivory, and their interaction in the regeneration and density of tall thistle (Cirsium altissimum) across a topographic ecosystem productivity gradient in tallgrass prairie over two years. On ridges and in valleys, we used a factorial experiment manipulating seed availability and insect herbivory to quantify effects of: seed input on seedling density, insect herbivory on juvenile density, and cumulative impacts of both seed input and herbivory on reproductive adult density. Seed addition increased seedling densities at three of five sites in 2006 and all five sites in 2007. Insect herbivory reduced seedling survival across all sites in both years, as well as rosette survival from the previous year's seedlings. In both years, insecticide treatment of seed addition plots led to greater adult tall thistle densities in the following year, reflecting the increase in juvenile thistle densities in the experimental year. Seedling survival was not density dependent. Our analytical projection model predicts a significant long-term increase in adult densities from seed input, with a greater increase under experimentally reduced insect herbivory. While plant community biomass and water stress varied significantly between ridges and valleys, the effects of seed addition and insect herbivory did not vary with gradient position. These results support conceptual models that predict seedling and adult densities of short-lived monocarpic perennial plants should be seed limited. Further, the experiment demonstrates that even at high juvenile plant densities, at which density dependence potentially could have overridden herbivore effects on plant survival, insect herbivory strongly affected juvenile thistle performance and adult densities of this native prairie species.     

Divergent composition but similar function of soil food webs of individual plants: plant species and community effects

Soils are extremely rich in biodiversity, and soil organisms play pivotal roles in supporting terrestrial life, but the role that individual plants and plant communities play in influencing the diversity and functioning of soil food webs remains highly debated. Plants, as primary producers and providers of resources to the soil food web, are of vital importance for the composition, structure, and functioning of soil communities. However, whether natural soil food webs that are completely open to immigration and emigration differ underneath individual plants remains unknown. In a biodiversity restoration experiment we first compared the soil nematode communities of 228 individual plants belonging to eight herbaceous species. We included grass, leguminous, and non-leguminous species. Each individual plant grew intermingled with other species, but all plant species had a different nematode community. Moreover, nematode communities were more similar when plant individuals were growing in the same as compared to different plant communities, and these effects were most apparent for the groups of bacterivorous, carnivorous, and omnivorous nematodes. Subsequently, we analyzed the composition, structure, and functioning of the complete soil food webs of 58 individual plants, belonging to two of the plant species, Lotus corniculatus (Fabaceae) and Plantago lanceolata (Plantaginaceae). We isolated and identified more than 150 taxa/groups of soil organisms. The soil community composition and structure of the entire food webs were influenced both by the species identity of the plant individual and the surrounding plant community. Unexpectedly, plant identity had the strongest effects on decomposing soil organisms, widely believed to be generalist feeders. In contrast, quantitative food web modeling showed that the composition of the plant community influenced nitrogen mineralization under individual plants, but that plant species identity did not affect nitrogen or carbon mineralization or food web stability. Hence, the composition and structure of entire soil food webs vary at the scale of individual plants and are strongly influenced by the species identity of the plant. However, the ecosystem functions these food webs provide are determined by the identity of the entire plant community.