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

Terrestrial carbon and nutrients can subsidize the detrital pool of freshwater ecosystems; yet, the importance of terrestrial subsidies to lake and pond food webs is uncertain and debated. Terrestrial detritus is expected to have the greatest impact on food webs when water bodies are small and shallow with low levels of incident light. Temporary forested ponds fit this description and are often assumed to have a leaf detritus-based food web, but this has not been quantified. In a whole-ecosystem experiment, we traced the flow of isotopically enriched leaf litter to primary producers and consumers in a small, forested pond. We found that terrestrial leaves provided nutrients to algae, offering an indirect pathway in which leaf litter can enter the food web. Terrestrial leaves were also consumed directly, and larval caddisfly ( Limnephilus sp.) shredders likely mobilized leaf nutrients to other consumers, a process overlooked in many previous small-scale experiments that did not incorporate shredders. Unexpectedly, most consumers relied heavily upon algal food pathways despite low light and net heterotrophic conditions. Overall, our study highlights the interconnectedness of algal and leaf litter pathways in small pond food webs, and emphasizes that algal pathways are prevalent and important even in small, shaded ponds with high loads of terrestrial leaf litter. [ABSTRACT FROM AUTHOR]

Stable isotope analysis has emerged as one of the primary means for examining the structure and dynamics of food webs, and numerous analytical approaches are now commonly used in the field. Techniques range from simple, qualitative inferences based on the isotopic niche, to Bayesian mixing models that can be used to characterize food-web structure at multiple hierarchical levels. We provide a comprehensive review of these techniques, and thus a single reference source to help identify the most useful approaches to apply to a given data set. We structure the review around four general questions: (1) what is the trophic position of an organism in a food web?; (2) which resource pools support consumers?; (3) what additional information does relative position of consumers in isotopic space reveal about food-web structure?; and (4) what is the degree of trophic variability at the intrapopulation level? For each general question, we detail different approaches that have been applied, discussing the strengths and weaknesses of each. We conclude with a set of suggestions that transcend individual analytical approaches, and provide guidance for future applications in the field. [ABSTRACT FROM AUTHOR]

Abstract: Stable isotope methods are powerful, frequently used tools which allow diet and trophic position reconstruction of organisms and the tracking of energy sources through ecosystems. The majority of ecosystems have multiple food sources which have distinct carbon and nitrogen isotopic signatures despite occupying a single trophic level. This difference in the starting isotopic composition of primary producers sets up an isotopic baseline that needs to be accounted for when calculating diet or trophic position using stable isotopic methods. This is particularly important when comparing animals from different regions or different times. Failure to do so can cause erroneous estimations of diet or trophic level, especially for organisms with mixed diets. The isotopic baseline is known to vary seasonally and in concert with a host of physical and chemical variables such as mean annual rainfall, soil maturity, and soil pH in terrestrial settings and lake size, depth, and distance from shore in aquatic settings. In the fossil record, the presence of shallowing upward suites of rock, or parasequences, will have a considerable impact on the isotopic baseline as basin size, depth and distance from shore change simultaneously with stratigraphic depth. For this reason, each stratigraphic level is likely to need an independent estimation of baseline even within a single outcrop. Very little is known about the scope of millennial or decadal variation in isotopic baseline. Without multi-year data on the nature of isotopic baseline variation, the impacts of time averaging on our ability to resolve trophic relationships in the fossil record will remain unclear. The use of a time averaged baseline will increase the amount of error surrounding diet and trophic position reconstructions. Where signal to noise ratios are low, due to low end member disparity (e.g., aquatic systems), or where the observed isotopic shift is small (≤1‰) the error introduced by time averaging may severely inhibit the scope of one''s interpretations and limit the types of questions one can reliably answer. In situations with strong signal strength, resulting from high amounts of end member disparity (e.g., terrestrial settings), this additional error maybe surmountable. Baseline variation that is adequately characterized can be dealt with by applying multiple end-member mixing models. [Copyright &y& Elsevier]

There are three hypothesized controls on food-chain length (FCL): energy supply (or “resource availability”), ecosystem size and disturbance (or “environmental variation”). In this article, the evidence for controls on FCL in freshwater ecosystems is evaluated. First, the various ways FCL can be measured are defined. Food-chain length typically is estimated as (1) connectance-based FCL—an average connectance between basal resources and top consumers, (2) functional FCL—by experimental determination of functionally significant effects of a top predator on lower trophic-level biomass patterns, and (3) realized FCL—an average connectance measure weighted by energy flow between basal consumers and the consumer occupying the maximum trophic position in the food web. Second, all evidence for relationships between the three hypothetical controls and FCL in freshwater ecosystems are evaluated. The review includes studies from streams, lakes, ponds, wetlands, phytotelmata, and experimental containers. Surprisingly, few studies of FCL in freshwaters that test the same suite of controls using the same methods are found. Equally compelling results arise from case studies based on functional, realized, and connectance-based measures of FCL. Third, 10 rules of thumb that could increase similarity of future studies, thereby facilitating synthesis across systems, are suggested. Fourth, it is discussed how FCL influences the concentration of contaminants in large-bodied animals (many of which are consumed by humans) as well as the efficacy of biocontrol applications in agriculture. Finally, there is a discussion of the potential relationships between global climate change, hydrology, and FCL in freshwaters. [ABSTRACT FROM AUTHOR]

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Streams experience frequent natural disturbance and are undergoing considerable anthropogenic disturbance due to dam construction and water diversion. Disturbance is known to impact community structure, but its effect on food chain length is still a matter of considerable debate. Theoretical models show that longer food chains are less resilient to disturbance, so food chain length is predicted to be shorter following a disturbance event. Here we experimentally test the effect of disturbance on food chain length in streams by diverting stream flow. We found that our experimental low-flow disturbance did not alter food chain length. We did see an effect on body-size structure in our food webs suggesting that food chain length may be an insensitive indicator of disturbance. We suggest that habitat heterogeneity and food web complexity buffer the effect of disturbance on food chain length. The theoretical predictions of disturbance on food chain length are only likely to be seen in homogeneous systems that closely approximate the linear food chains the models are based upon. [ABSTRACT FROM AUTHOR]

Ecologists have long struggled to explain variation in food-chain length among natural ecosystems. Food-chain length is predicted to be shorter in ecosystems subjected to greater disturbance because longer chains are theoretically less resilient to perturbation. Moreover, food-chain length is expected to be longer in larger ecosystems because increasing ecosystem size increases species richness and stabilizes predator-prey interactions, or increases total resource availability. Here we test the roles of disturbance and ecosystem size in-determining the food-chain length of terrestrial food webs on Bahamian islands. We found that disturbance affected the identity of top predators, but did not change food-chain length because alternative top predators occupied similar trophic positions. On the other hand, a 106- fold increase in ecosystem size elevated food-chain length by one trophic level. We suggest that the effect of disturbance on food-chain length is weak when alternate top predators are trophic omnivores and have similar trophic positions. This and previous work in lakes suggest that ecosystem size may be a strong determinant of food-chain length in both aquatic and terrestrial ecosystems [ABSTRACT FROM AUTHOR]

Understanding and explaining the causes of variation in food-chain length is a fundamental challenge for community ecology. The productive-space hypothesis, which suggests food-chain length is determined by the combination of local resource availability and ecosystem size, is central to this challenge. Two different approaches currently exist for testing the productive-space hypothesis: (1) the dual gradient approach that tests for significant relationships between food-chain length and separate gradients of ecosystem size (e.g., lake volume) and per-unit-size resource availability (e.g., g C m−1 year−2), and (2) the single gradient approach that tests for a significant relationship between food-chain length and the productive space (product of ecosystem size and per-unit-size resource availability). Here I evaluate the efficacy of the two approaches for testing the productive-space hypothesis. Using simulated data sets, I estimate the Type 1 and Type 2 error rates for single and dual gradient models in recovering a known relationship between food-chain length and ecosystem size, resource availability, or the combination of ecosystem size and resource ability, as specified by the productive-space hypothesis. The single gradient model provided high power (low Type 2 error rates) but had a very high Type 1 error rate, often erroneously supporting the productive-space hypothesis. The dual gradient model had a very low Type 1 error rate but suffered from low power to detect an effect of per-unit-size resource availability because the range of variation in resource availability is limited. Finally, I performed a retrospective power analysis for the Post et al. (Nature 405:1047–1049, ) data set, which tested and rejected the productive-space hypothesis using the dual gradient approach. I found that Post et al. (Nature 405:1047–1049, ) had sufficient power to reject the productive-space hypothesis in north temperate lakes; however, the productive-space hypothesis must be tested in other ecosystems before its generality can be fully addressed. [ABSTRACT FROM AUTHOR]

Food-chain length is a central characteristic of ecological communities because of its strong influence on community structure and ecosystem function. While recent studies have started to better clarify the relationship between food-chain length and environmental gradients such as resource availability and ecosystem size, much less progress has been made in isolating the ultimate and proximate mechanisms that determine food-chain length. Progress has been slow, in part, because research has paid little attention to the proximate changes in food web structure that must link variation in food-chain length to the ultimate dynamic mechanism. Here we outline the structural mechanisms that determine variation in food-chain length. We explore the implications of these mechanisms for understanding how changes in food-web structure influence food-chain length using both an intraguild predation community model and data from natural ecosystems. The resulting framework provides the mechanisms for linking ultimate dynamic mechanisms to variation in food-chain length. It also suggests that simple linear food-chain models may make misleading predictions about patterns of variation in food-chain length because they are unable to incorporate important structural mechanisms that alter food-web dynamics and cause non-linear shifts in food-web structure. Intraguild predation models provide a more appropriate theoretical framework for understanding food-chain length in most natural ecosystems because they accommodate all of the proximate structural mechanisms identified here. [ABSTRACT FROM AUTHOR]

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 δ13C-δ15N 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. [ABSTRACT FROM AUTHOR]

Traditional approaches to the study of food webs emphasize the transfer of local primary productivity in the form of living plant organic matter across trophic levels. However, dead organic matter, or detritus, a common feature of most ecosystems plays a frequently overlooked role as a dynamic heterogeneous resource and habitat for many species. We develop an integrative framework for understanding the impact of detritus that emphasizes the ontogeny and heterogeneity of detritus and the various ways that explicit inclusion of detrital dynamics alters generalizations about the structure and functioning of food webs. Through its influences on food web composition and dynamics, detritus often increases system stability and persistence, having substantial effects on trophic structure and biodiversity. Inclusion of detrital heterogeneity in models of food web dynamics is an essential new direction for ecological research. [ABSTRACT FROM AUTHOR]

Food-chain length is a central characteristic of ecological communities that has attracted considerable attention for over 75 years because it strongly affects community structure, ecosystem processes and contaminant concentrations. Conventional wisdom holds that either resource availability or dynamical stability limit food-chain length; however, new studies and new techniques challenge the conventional wisdom and broaden the discourse on food-chain length. Recent results suggest that resource availability limits food-chain length only in systems with very low resource availability, and call into question the theoretical basis for dynamical stability as a determinant of food-chain length. Evidence currently points towards a complex and contingent framework of interacting constraints that includes the history of community organization, resource availability, the type of predator–prey interactions, disturbance and ecosystem size. Within this framework, the debate has shifted from a search for singular explanations to a search for when and where different constraints operate to determine food-chain length. [Copyright &y& Elsevier]

Develops methods and assumptions for generating isotopic baseline to estimate trophic position in the United States. Examination on predation ability of surface-grazing snails and filter-feeding mussels; Variation of aquatic food web; Importance of stable isotopes.

Tests three hypotheses that predict food-chain length to be determined by productivity alone, ecosystem size alone, or a combination of productivity and ecosystem size. Methods; Results; Conclusion that food-chain length increases with ecosystem size, but is not related to productivity.

The article presents the authors' views on the significance of stable isotope ratios on community-wide measures of trophic structure. The authors noted that applications of stable isotopes to disentangle food web complexity becomes increasingly problematic as the number of resource pools for a particular consumer increases. They suggested that food web ecologists should understand situations under which particular tools including stable isotope-based metrics will prove must useful.

Food chain length (FCL) is a fundamental component of food web structure. Studies in a variety of ecosystems suggest that FCL is determined by energy supply, environmental stability, and/or ecosystem size, but the nature of the relationship between environmental stability and FCL, and the mechanism linking ecosystem size to FCL, remain unclear. Here we show that FCL increases with drainage area and decreases with hydrologic variability and intermittency across 36 North American rivers. Our analysis further suggests that hydrologic variability is the mechanism underlying the correlation between ecosystem size and FCL in rivers. Ecosystem size lengthens river food chains by integrating and attenuating discharge variation through stream networks, thereby enhancing environmental stability in larger river systems. [ABSTRACT FROM AUTHOR]