Food Web Stability Research Articles

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.

THE FEAR OF BEING EATEN REDUCES ENERGY TRANSFER IN A SIMPLE FOOD CHAIN

Food chain length is an important property of ecosystems, but the mechanisms maintaining it remain elusive. Classical views suggest that energetic inefficiencies (the "energy-flow hypothesis") limit food chain length, but others have argued that better explanations reside in more complex scenarios that consider the stability of food webs or the combined effects of productivity and ecosystem size. We argue that abandonment of the energy-flow hypothesis is premature. For a simple tritrophic rocky intertidal food chain, we show that the efficiency of energy transfer is strongly influenced by predation risk and consumer density. Effects tied to predation risk were particularly strong, explaining 32% of the variation in growth efficiency (compared to 15% for density effects) and reducing it by 44-76%. Hence, the stress (fear of being eaten) that predators impose on prey may be instrumental in limiting energy transfer up the food chain and thus contribute to the shortening of food chains.

Hydrology and nutrient effects on food-web structure in ten lake superior coastal wetlands

ABSTRACT We examined the effects of hydrology and nutrients on the food webs of ten coastal wetlands on Lake Superior, using published stable isotope food web data for three wetlands and original data from seven additional systems in order to span regional hydrologic and nutrient enrichment gradients. We used a dual-source isotope mixing model to estimate the proportion of carbon in fish that originated from planktonic versus periphytic invertebrates, and we related carbon source to 1) nutrient enrichment, 2) hydraulic residence time, and 3) an index of nutrient loading that incorporates residence time and nutrient concentrations. There was no relationship between nutrient enrichment and the proportion of planktonic versus periphytic C in fish. Proportion of planktonic C in fish increased significantly with hydraulic residence time (F = 5.68, R2 = 0.42, p = 0.044). Riverine wetlands generally had lowest proportions of planktonic C in fish, dendritic wetlands were intermediate, and lagoon wetlands ha...

Allometric scaling enhances stability in complex food webs

Classic local stability theory predicts that complex ecological networks are unstable and are unlikely to persist despite empiricists' abundant documentation of such complexity in nature. This contradiction has puzzled biologists for decades. While some have explored how stability may be achieved in small modules of a few interacting species, rigorous demonstrations of how large complex and ecologically realistic networks dynamically persist remain scarce and inadequately understood. Here, we help fill this void by combining structural models of complex food webs with nonlinear bioenergetic models of population dynamics parameterized by biological rates that are allometrically scaled to populations' average body masses. Increasing predator-prey body mass ratios increase population persistence up to a saturation level that is reached by invertebrate and ectotherm vertebrate predators when being 10 or 100 times larger than their prey respectively. These values are corroborated by empirical predator-prey body mass ratios from a global data base. Moreover, negative effects of diversity (i.e. species richness) on stability (i.e. population persistence) become neutral or positive relationships at these empirical ratios. These results demonstrate that the predator-prey body mass ratios found in nature may be key to enabling persistence of populations in complex food webs and stabilizing the diversity of natural ecosystems.

Parasites dominate food web links.

Parasitism is the most common animal lifestyle, yet food webs rarely include parasites. The few earlier studies have indicated that including parasites leads to obvious increases in species richness, number of links, and food chain length. A less obvious result was that adding parasites slightly reduced connectance, a key metric considered to affect food web stability. However, reported reductions in connectance after the addition of parasites resulted from an inappropriate calculation. Two alternative corrective approaches applied to four published studies yield an opposite result: parasites increase connectance, sometimes dramatically. In addition, we find that parasites can greatly affect other food web statistics, such as nestedness (asymmetry of interactions), chain length, and linkage density. Furthermore, whereas most food webs find that top trophic levels are least vulnerable to natural enemies, the inclusion of parasites revealed that mid-trophic levels, not low trophic levels, suffered the highest vulnerability to natural enemies. These results show that food webs are very incomplete without parasites. Most notably, recognition of parasite links may have important consequences for ecosystem stability because they can increase connectance and nestedness.

Structural asymmetry and the stability of diverse food webs

Untangling the influence of human activities on food-web stability and persistence is complex given the large numbers of species and overwhelming number of interactions within ecosystems. Although biodiversity has been associated with stability, the actual structures and processes that confer stability to diverse food webs remain largely unknown. Here we show that real food webs are structured such that top predators act as couplers of distinct energy channels that differ in both productivity and turnover rate. Our theoretical analysis shows that coupled fast and slow channels convey both local and non-local stability to food webs. Alarmingly, the same human actions that have been implicated in the loss of biodiversity also directly erode the very structures and processes that we show to confer stability on food webs.

Estimating trophic link density from quantitative but incomplete diet data

The trophic link density and the stability of food webs are thought to be related, but the nature of this relation is controversial. This article introduces a method for estimating the link density from diet tables which do not cover the complete food web and do not resolve all diet items to species level. A simple formula for the error of this estimate is derived. Link density is determined as a function of a threshold diet fraction below which diet items are ignored (“diet partitioning function”). Furthermore, analytic relationships between this threshold-dependent link density and the generality distribution of food webs are established. A preliminary application of the method to field data suggests that empirical results relating link density to diversity might need to be revisited.

Nutrient-limited food webs with up to three trophic levels: Feasibility, stability, assembly rules, and effects of nutrient enrichment

Community structure is controlled, among multiple factors, by competition and predation. Using the R* rule and graphical analysis, we analyse here the feasibility, stability and assembly rules of resource-based food webs with up to three trophic levels. In particular, we show that (1) the stability of a food web with two plants and two generalist herbivores does not require that plants’ resource exploitation abilities trade-off with resistance to the two herbivores, and (2) food webs with two plants and either one generalist herbivore and a carnivore or two generalist herbivores and two generalist carnivores are not feasible because of cascade competition between top consumers. The relative strength of species interactions and the relative impacts of plants and herbivores on factors which control their growth also play a critical role. We discuss how community structure constrains assembly rules and yields cascades of extinctions in food webs.

Omnivory and the stability of food webs

The ecological concept of omnivory, feeding at more than a single trophic level, is formulated as an intermediate stage between any two of three classical three-dimensional species interaction systems—tritrophic chain, competition, and polyphagy. It is shown that omnivory may be either stabilizing or destabilizing, depending, in part, on the conditions of the parent systems from which it derives. It is further conjectured that the tritrophic to competition gradient cannot be entirely stable, that there must be an instability at some level of intermediate omnivory.

Factors affecting the expression of inducible defences in Euplotes: genotype, predator density and experience

Summary Inducible defences alter the strength of interaction in food webs. Their effectiveness depends both on the maximum level of induction and the speed at which induction happens. Maximum level and speed of induction should therefore evolve in concert. We examined the effect of genotype, number of predators and previous exposure to predators on speed and maximum level of induction in a morphological defence expressed by eight clones in three species of the ciliate Euplotes. Both speed and maximum level of induction, and the reaction to predator density varied among genotypes. These results show that there is genetic variance for all aspects of this inducible defence and the potential for complex evolutionary change under selection. Higher predator densities led to higher maximum levels of defence and, for one measure of induction speed, more rapid induction. Previous exposure to predators had no detectable effect on either speed or maximum level of induction. Our results demonstrate that Euplotes can precisely and rapidly adjust their morphological defence to the magnitude of predation risk. The speed of induction and the maximum level of defence varied among genotypes and this will lead to variation in defence level and vulnerability under natural conditions. Variation in prey vulnerability is a key factor promoting stability in food webs.

Temporal stability of aquatic food webs: partitioning the effects of species diversity, species composition and enrichment

AbstractTheory predicts that species diversity can enhance stability of community‐level biomass while simultaneously decreasing population‐level stability. Enrichment can theoretically destabilize communities but effects may become weaker with increasing diversity because of the inclusion of consumer‐resistant prey. Few experiments using direct manipulations of species diversity have tested these predictions. We used laboratory‐based aquatic food webs to examine the effects of species composition, diversity and enrichment on temporal variability of population‐ and community‐level biomass. We found weak effects of enrichment on population‐ and community‐level stability. However, diversity enhanced community‐level stability while species composition had no influence. In contrast, composition effects outweighed diversity effects when stability was measured at the population level. We found no negative effects of diversity on population‐level stability, in opposition to theory. Our results indicate that diversity can enhance stability in multitrophic systems, but effects vary with the scale of biological organization at which stability is measured.

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.

Global change alters the stability of food webs

AbstractRecent research has generally shown that a small change in the number of species in a food web can have consequences both for community structure and ecosystem processes. However ‘change’ is not limited to just the number of species in a community, but might include an alteration to such properties as precipitation, nutrient cycling and temperature. How such changes might affect species interactions is important, not just through the presence or absence of interactions, but also because the patterning of interaction strengths among species is intimately associated with community stability. Interaction strengths encompass such properties as feeding rates and assimilation efficiencies, and encapsulate functionally important information with regard to ecosystem processes. Interaction strengths represent the pathways and transfer of energy through an ecosystem. We review the best empirical data available detailing the frequency distribution of interaction strengths in communities. We present the underlying (but consistent) pattern of species interactions and discuss the implications of this patterning. We then examine how such a basic pattern might be affected given various scenarios of ‘change’ and discuss the consequences for community stability and ecosystem functioning.

Establishment of trophic continuum in the food web of the Yellow Sea and East China Sea ecosystem: Insight from carbon and nitrogen stable isotopes

Stable carbon and nitrogen isotope ratios (delta (13)C and delta (15)N) are used to study the trophic structure of food web in the Yellow Sea and East China Sea ecosystem. The trophic continuum of pelagic food web from phytoplankton to top preyer was elementarily established, and a trophic structure diagram in the Yellow Sea and East China Sea was outlined in combination with carbon isotopic data of benthic organisms, which is basically consistent with and makes some improvements on the simplified Yellow Sea food web and the trophic structure diagram drawn based on the biomass of main resource population during 1985-1986. This result indicates that the stable isotope method is a potential useful means for further studying the complete marine food web trophic continuum from viruses to top predators and food web stability.

Stable isotope food web studies: a case for standardized sample treatment

The enrichment of the stable isotopes 13C and 15N across trophic levels is a commonly used tool in studies on organic matter flow and food webs. An accepted standard for pre-analysis sample preparation, however, is still missing. Thus, potential methodological bias in single studies may hamper comparability and scalability of data from different sources. Sample CaCO3 content introduces a positive bias in ?13C measurements and a negative bias in ?15N measurements. The acidification of samples to remove inorganic carbonate significantly reduces both ?13C and ?15N. As a standard procedure we recommend (i) to acidify samples with as little hydrochloric acid (HCl) as possible using the drop-by-drop technique and (ii) to restrain from rinsing after HCl application.

Stabilization of Large Generalized Lotka-Volterra Foodwebs By Evolutionary Feedback

Conventional ecological models show that complexity destabilizes foodwebs, suggesting that foodwebs should have neither large numbers of species nor a large number of interactions. However, in nature the opposite appears to be the case. Here we show that if the interactions between species are allowed to evolve within a generalized Lotka-Volterra model such stabilizing feedbacks and weak interactions emerge automatically. Moreover, we show that trophic levels also emerge spontaneously from the evolutionary approach, and the efficiency of the unperturbed ecosystem increases with time. The key to stability in large foodwebs appears to arise not from complexity per se but from evolution at the level of the ecosystem which favors stabilizing (negative) feedbacks.

Predator–prey body size, interaction strength and the stability of a real food web

SummaryWe examined the empirical relationship between predator–prey body size ratio and interaction strength in the Ythan Estuary food web.We have refined a previously published version of the food web and explored how size‐based predatory effects might affect food web dynamics. To do so, we used four predatory speciesCrangon crangon(Linnaeus),Carcinus maenas(Linnaeus),Pomatoschistus microps(Krøyer) andPlatichthys flesus(Linnaeus) and one common prey speciesCorophium volutator(Pallas) from the food web.All predators and prey were sorted into small, medium and large size classes and placed into mesocosms in all possible pairwise combinations of size and species identity to determine per capita effects of predators on prey (aij).Using Lotka–Volterra dynamics the empirical body size relationships obtained from these experiments and other relationships already available for the Ythan Estuary, we parameterized a food web model for this system. The local stability properties of the resulting food web models were then determined.We found that by choosing interaction strengths using an empirically defined scaling law, the resulting food web models are always dynamically stable, despite the residual uncertainties in the modelling approach. This contrasts with the statistical expectation that random webs with random parameters have a vanishingly improbable chance of stability.The patterning of predator and prey body sizes in real ecosystems affects the arrangement of interaction strengths, which in turn determines food web stability.

Some ecological implications of a neem (azadirachtin) insecticide disturbance to zooplankton communities in forest pond enclosures

A neem-based insecticide, Neemix ® 4.5, was applied to forest pond enclosures at concentrations of 10, 17, and 28 μg l −1 azadirachtin (the active ingredient). At these test concentrations, significant, concentration-dependent reductions in numbers of adult copepods were observed, but immature copepod and cladoceran populations were unaffected. There was no evidence of recovery of adult copepods within the sampling season (May to October). The ecological significance of this disturbance to the zooplankton community was examined by determining biomass as a measure of food availability for higher predators, plankton community respiration, dissolved oxygen (DO) concentrations, and conductivity as functional indicators of ecosystem stress, and zooplankton food web stability as a measure of effects on trophic structure. The selective removal or reduction of adult copepods was sufficient to measurably reduce total zooplankton biomass for several weeks mid-season. During the period of maximal impact (about 4–9 weeks after the applications), total plankton community respiration was significantly reduced, and this appeared to contribute to significant, concentration-dependent increases in dissolved oxygen and decreases in conductivity among treated enclosures. The reductions in adult copepods resulted in negative effects on zooplankton food web stability through eliminations of a trophic link and reduced interactions and connectance. Comparing the results here to those from a previous study with tebufenozide, which was selectively toxic to cladocerans and had little effect on food web stability, indicates that differential sensitivity among taxa can influence the ecological significance of pesticide effects on zooplankton communities.

Stabilization of chaotic and non-permanent food-web dynamics

Several decades of dynamical analyses of food-web networks (1-6) have led to important insights into the effects of complexity, omnivory and interaction strength on food-web stability (6-8). Several re- cent insights (7, 8) are based on nonlinear bioenergetic consumer-resource models (9) that display chaotic behavior in three species food chains (10, 11) which can be stabilized by omnivory (7) and weak interaction of a fourth species (8). We slightly relax feeding on low-density prey in these models by modifying stan- dard food-web interactions known as "type II" functional responses (12). This change drastically alters the dynamics of realistic systems containing up to ten species. Our modification stabilizes chaotic dynamics in three species systems and reduces or eliminates extinctions and non-persistent chaos (11) in ten species systems. This increased stability allows analysis of systems with greater biodiversity than in earlier work and suggests that dynamic stability is not as severe a constraint on the structure of large food webs as previously thought. The sensitivity of dynamical models to small changes in the predator-prey functional response well within the range of what is empirically observed suggests that functional response is a crucial aspect of species interactions that must be more precisely addressed in empirical studies.

Compartments revealed in food-web structure.

Compartments in food webs are subgroups of taxa in which many strong interactions occur within the subgroups and few weak interactions occur between the subgroups. Theoretically, compartments increase the stability in networks, such as food webs. Compartments have been difficult to detect in empirical food webs because of incompatible approaches or insufficient methodological rigour. Here we show that a method for detecting compartments from the social networking science identified significant compartments in three of five complex, empirical food webs. Detection of compartments was influenced by food web resolution, such as interactions with weights. Because the method identifies compartmental boundaries in which interactions are concentrated, it is compatible with the definition of compartments. The method is rigorous because it maximizes an explicit function, identifies the number of non-overlapping compartments, assigns membership to compartments, and tests the statistical significance of the results. A graphical presentation reveals systemic relationships and taxa-specific positions as structured by compartments. From this graphic, we explore two scenarios of disturbance to develop a hypothesis for testing how compartmentalized interactions increase stability in food webs.