Species richness and trophic diversity increase decomposition in a co-evolved food web

Ecological communities show great variation in species richness, composition and food web structure across similar and diverse ecosystems. Knowledge of how this biodiversity relates to ecosystem functioning is important for understanding the maintenance of diversity and the potential effects of species losses and gains on ecosystems. While research often focuses on how variation in species richness influences ecosystem processes, assessing species richness in a food web context can provide further insight into the relationship between diversity and ecosystem functioning and elucidate potential mechanisms underpinning this relationship. Here, we assessed how species richness and trophic diversity affect decomposition rates in a complete aquatic food web: the five trophic level web that occurs within water-filled leaves of the northern pitcher plant, Sarracenia purpurea. We identified a trophic cascade in which top-predators--larvae of the pitcher-plant mosquito--indirectly increased bacterial decomposition by preying on bactivorous protozoa. Our data also revealed a facultative relationship in which larvae of the pitcher-plant midge increased bacterial decomposition by shredding detritus. These important interactions occur only in food webs with high trophic diversity, which in turn only occur in food webs with high species richness. We show that species richness and trophic diversity underlie strong linkages between food web structure and dynamics that influence ecosystem functioning. The importance of trophic diversity and species interactions in determining how biodiversity relates to ecosystem functioning suggests that simply focusing on species richness does not give a complete picture as to how ecosystems may change with the loss or gain of species.     

Autochthonous or allochthonous resources determine the characteristic population dynamics of ecosystem engineers and their impacts

Ecosystem engineering, or the modification of physical environments by organisms, can influence trophic interactions and thus food web dynamics. Although existing theory exclusively considers engineers using autochthonous resources, many empirical studies show that they often depend on allochthonous resources. By developing a simple mathematical model involving an ecosystem engineer that modifies the physical environment through its activities, its resource, and physical environment modified by the engineer, we compare the effects of autochthonous and allochthonous resources on the dynamics and stability of community with ecosystem engineers. To represent a variety of real situations, we consider engineers that alter either resource productivity, engineer feeding rate on the resource, or engineer mortality, and incorporate time-lagged responses of the physical environment. Our model shows that the effects of ecosystem engineering on community dynamics depend greatly on resource types. When the engineer consumes autochthonous resources, the community can exhibit oscillatory dynamics if the engineered environment affects engineer’s feeding rate or mortality. These cyclic behaviors are, however, stabilized by a slowly responding physical environment. When allochthonous resources are supplied as donor-controlled, on the other hand, the engineer population is unlikely to oscillate but instead can undergo unbounded growth if the engineered environment affects resource productivity or engineer mortality. This finding suggests that ecosystem engineers utilizing allochthonous resources may be more likely to reach high abundance and cause strong impacts on ecosystems. Our results highlight that community-based, compounding effects of trophic and physical biotic interactions of ecosystem engineers depend crucially on whether the engineers utilize autochthonous or allochthonous resources.     

Biodiversity, ecosystem functioning and food webs in fresh waters: assembling the jigsaw puzzle

1. Dramatic advances have been made recently in the study of biodiversity–ecosystem functioning (B-EF) relations and food web ecology. These fields are now starting to converge, and this fusion has the potential to improve our understanding of how environmental stressors modulate ecosystem processes and the supply of 'goods and services'.
2. Food web structure and dynamics can exert particularly strong influences on B-EF relations in fresh waters, as consumer–resource interactions (e.g. trophic cascades) are often more important than horizontal interactions within trophic levels. For instance, many freshwater food webs are size structured, with large organisms tending to occupy the higher trophic levels and often exerting powerful effects on ecosystem processes. However, because they are also vulnerable to perturbations, non-random losses of these large taxa can alter both food web structure and ecosystem functioning profoundly.
3. Recently, the focus of food web research has shifted away from exploring patterns, towards developing an understanding of processes (e.g. quantifying fluxes of individuals, biomass, energy, nutrients) and how the two interact. Many of the best-characterized food webs are from fresh waters, and these ecosystems are now being used to address some of the shortcomings of earlier B-EF studies. I have identified several key gaps in our current knowledge and highlighted potentially fruitful avenues of future B-EF and food web research.
4. A major challenge for this newly emerging research is to place it within a unified theoretical framework. The application of metabolic theory and ecological stoichiometry may help to achieve this goal by considering biological systems within the constraints imposed upon them by physical and chemical laws.     

The significance of partial migration for food web and ecosystem dynamics

Migration is ubiquitous and can strongly shape food webs and ecosystems. Less familiar, however, is that the majority of life cycle, seasonal and diel migrations in nature are partial migrations: only a fraction of the population migrates while the other individuals remain in their resident ecosystem. Here, we demonstrate different impacts of partial migration rendering it fundamental to our understanding of the significance of migration for food web and ecosystem dynamics. First, partial migration affects the spatiotemporal distribution of individuals and the food web and ecosystem-level processes they drive differently than expected under full migration. Second, whether an individual migrates or not is regularly correlated with morphological, physiological, and/or behavioural traits that shape its food-web and ecosystem-level impacts. Third, food web and ecosystem dynamics can drive the fraction of the population migrating, enabling the potential for feedbacks between the causes and consequences of migration within and across ecosystems. These impacts, individually and in combination, can yield unintuitive effects of migration and drive the dynamics, diversity and functions of ecosystems. By presenting the first full integration of partial migration and trophic (meta-)community and (meta-)ecosystem ecology, we provide a roadmap for studying how migration affects and is affected by ecosystem dynamics in a changing world.     

Unravelling the complex structure of forest soil food webs: higher omnivory and more trophic levels

Food web topologies depict the community structure as distributions of feeding interactions across populations. Although the soil ecosystem provides important functions for aboveground ecosystems, data on complex soil food webs is notoriously scarce, most likely due to the difficulty of sampling and characterizing the system. To fill this gap we assembled the complex food webs of 48 forest soil communities. The food webs comprise 89 to 168 taxa and 729 to 3344 feeding interactions. The feeding links were established by combining several molecular methods (stable isotope, fatty acid and molecular gut content analyses) with feeding trials and literature data. First, we addressed whether soil food webs (n = 48) differ significantly from those of other ecosystem types (aquatic and terrestrial aboveground, n = 77) by comparing 22 food web parameters. We found that our soil food webs are characterized by many omnivorous and cannibalistic species, more trophic chains and intraguild‐predation motifs than other food webs and high average and maximum trophic levels. Despite this, we also found that soil food webs have a similar connectance as other ecosystems, but interestingly a higher link density and clustering coefficient. These differences in network structure to other ecosystem types may be a result of ecosystem specific constraints on hunting and feeding characteristics of the species that emerge as network parameters at the food‐web level. In a second analysis of land‐use effects, we found significant but only small differences of soil food web structure between different beech and coniferous forest types, which may be explained by generally strong selection effects of the soil that are independent of human land use. Overall, our study has unravelled some systematic structures of soil food‐webs, which extends our mechanistic understanding how environmental characteristics of the soil ecosystem determine patterns at the community level.     

Dynamics of compartmented and reticulate food webs in relation to energetic flows

Using simple food webs, we address how the interactions of food web structure and energetic flows influence dynamics. We examine the effect of food web topologies with equivalent energetics (i.e., trophic interactions are equivalent at each trophic level), following which we vary energetic flows to include weak and strong interactions or nonequivalent energetics. In contrast to some work (Pimm 1979), we find that compartmented webs are more stable than reticulate webs. However, we find that nonequivalent energetics can stabilize previously unstable reticulate structures. It is not only weak flows that can be stabilizing but also the arrangement of the flows that emphasizes stabilizing mechanisms. We find that the main stabilizing mechanism is asynchrony, where structures and energetic arrangements that decrease synchrony such as internal segregation or competition will stabilize dynamics. Since compartments allow prey dynamics to behave somewhat independently, compartmentation readily promotes stability. In addition, these results can be scaled from simple food webs to more complex webs with many interacting subsystems so that linking weak subsystems to strong ones can stabilize dynamics. We show that food web dynamics are determined not only by topology but also the arrangement of weak and strong energetic flows.     

Chemical information transfer in freshwater plankton

The structure of aquatic ecosystems is determined by complex interactions among individual organisms at different trophic levels. Although our basic understanding of how top-down and bottom-up processes interact to determine food-web dynamics has advanced, we still lack insights into how complex interactions and feedbacks affect the dynamics and structure of food webs. It is now becoming increasingly clear that, in addition to energy transfer from one trophic level to the other, there is exchange of information between these levels facilitated by the release of infochemicals by the organisms. There is evidence from recent studies that the exchange of chemical information in freshwater ecosystems is likely to play a decisive role in shaping structure and functioning of these systems. Chemical communication among freshwater organisms mediates many aspects of both predation and interspecific competition, which play key roles in determining community structure and ecosystem functioning. For example, consumer-induced defences in phytoplankton and zooplankton include modifications in the characteristics relating to life history, behaviour, morphology and biochemistry. These inducible defences affect trophic interactions by altering predator feeding rates through changes in attack rate or handling time, or both. Also host-specific fungal parasitism in phytoplankton is probably controlled by infochemicals. The motile fungi recognise their host by host-secreted compounds. Until now models describing the functioning of ecosystems mainly considered flows of biomass and energy. Integration of new knowledge about the role of chemical communication in these models may be one of the aims of ecological informatics. In this chapter I discuss how infochemicals may affect the dynamics and structure of planktonic food webs.     

Predicting food‐web structure with metacommunity models

Synthesis Metacommunity theory aims to elucidate the relative influence of local and regional‐scale processes in generating diversity patterns across the landscape. Metacommunity research has focused largely on assemblages of competing organisms within a single trophic level. Here, we test the ability of metacommunity models to predict the network structure of the aquatic food web found in the leaves of the northern pitcher plant Sarracenia purpurea. The species‐sorting and patch‐dynamics models most accurately reproduced nine food web properties, suggesting that local‐scale interactions play an important role in structuring Sarracenia food webs. Our approach can be applied to any well‐resolved food web for which data are available from multiple locations. The metacommunity framework explores the relative influence of local and regional‐scale processes in generating diversity patterns across the landscape. Metacommunity models and empirical studies have focused mostly on assemblages of competing organisms within a single trophic level. Studies of multi‐trophic metacommunities are predominantly restricted to simplified trophic motifs and rarely consider entire food webs. We tested the ability of the patch‐dynamics, species‐sorting, mass‐effects, and neutral metacommunity models, as well as three hybrid models, to reproduce empirical patterns of food web structure and composition in the complex aquatic food web found in the northern pitcher plant Sarracenia purpurea. We used empirical data to determine regional species pools and estimate dispersal probabilities, simulated local food‐web dynamics, dispersed species from regional pools into local food webs at rates based on the assumptions of each metacommunity model, and tested their relative fits to empirical data on food‐web structure. The species‐sorting and patch‐dynamics models most accurately reproduced nine food web properties, suggesting that local‐scale interactions were important in structuring Sarracenia food webs. However, differences in dispersal abilities were also important in models that accurately reproduced empirical food web properties. Although the models were tested using pitcher‐plant food webs, the approach we have developed can be applied to any well‐resolved food web for which data are available from multiple locations.     

Food‐web structure and ecosystem function in the Laurentian Great Lakes—Toward a conceptual model

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Trophic disruption: a meta‐analysis of how habitat fragmentation affects resource consumption in terrestrial arthropod systems

Habitat fragmentation is a complex process that affects ecological systems in diverse ways, altering everything from population persistence to ecosystem function. Despite widespread recognition that habitat fragmentation can influence food web interactions, consensus on the factors underlying variation in the impacts of fragmentation across systems remains elusive. In this study, we conduct a systematic review and meta‐analysis to quantify the effects of habitat fragmentation and spatial habitat structure on resource consumption in terrestrial arthropod food webs. Across 419 studies, we found a negative overall effect of fragmentation on resource consumption. Variation in effect size was extensive but predictable. Specifically, resource consumption was reduced on small, isolated habitat fragments, higher at patch edges, and neutral with respect to landscape‐scale spatial variables. In general, resource consumption increased in fragmented settings for habitat generalist consumers but decreased for specialist consumers. Our study demonstrates widespread disruption of trophic interactions in fragmented habitats and describes variation among studies that is largely predictable based on the ecological traits of the interacting species. We highlight future prospects for understanding how changes in spatial habitat structure may influence trophic modules and food webs.     

A framework for understanding physical ecosystem engineering by organisms

While well‐recognized as an important kind of ecological interaction, physical ecosystem engineering by organisms is diverse with varied consequences, presenting challenges for developing and using general understanding. There is also still some uncertainty as to what it is, and some skepticism that the diversity of engineering and its effects is amenable to conceptual integration and general understanding. What then, are the key cause/effect relationships and what underlies them? Here we develop, enrich and extend our extant understanding of physical ecosystem engineering into an integrated framework that exposes the essential cause/effect relationships, their underpinnings, and the interconnections that need to be understood to explain or predict engineering effects. The framework has four cause/effect relationships linking four components: 1. An engineer causes structural change; 2. Structural change causes abiotic change; 3. Structural and abiotic change cause biotic change; 4. Structural, abiotic and biotic change can feedback to the engineer. The first two relationships describe an ecosystem engineering process and abiotic dynamics, while the second two describe biotic consequence for other species and the engineer. The four relationships can be parameterized and linked using time‐indexed equations that describe engineered system dynamics. After describing the relationships we discuss the utility of the framework; how it might be enriched; and briefly how it can be used to identify intersections of ecosystem engineering with fields outside ecology.     

Biodiversity maintenance in food webs with regulatory environmental feedbacks

Although the food web is one of the most fundamental and oldest concepts in ecology, elucidating the strategies and structures by which natural communities of species persist remains a challenge to empirical and theoretical ecologists. We show that simple regulatory feedbacks between autotrophs and their environment when embedded within complex and realistic food-web models enhance biodiversity. The food webs are generated through the niche-model algorithm and coupled with predator-prey dynamics, with and without environmental feedbacks at the autotroph level. With high probability and especially at lower, more realistic connectance levels, regulatory environmental feedbacks result in fewer species extinctions, that is, in increased species persistence. These same feedback couplings, however, also sensitize food webs to environmental stresses leading to abrupt collapses in biodiversity with increased forcing. Feedback interactions between species and their material environments anchor food-web persistence, adding another dimension to biodiversity conservation. We suggest that the regulatory features of two natural systems, deep-sea tubeworms with their microbial consortia and a soil ecosystem manifesting adaptive homeostatic changes, can be embedded within niche-model food-web dynamics.     

Complexity and structural properties of food webs in the Barents Sea

A food web topology describes the diversity of species and their trophic interactions, i.e. who eats whom, and structural analysis of food web topologies can provide insight into ecosystem structure and function. It appears simple, at first sight, to list all species and their trophic interactions. However, the very large number of species at low trophic levels and the impossibility to monitor all trophic interactions in the ocean makes it impossible to construct complete food web topologies. In practice, food web topologies are simplified by aggregating species into groups termed trophospecies. It is not clear though, how much simplified versions of food webs retain the structural properties of more detailed networks. Using the most comprehensive Barents Sea food web to date, we investigate the performance of methods to construct simplified food webs using three approaches: taxonomic, structural and regular clustering. We then evaluate how topological properties vary with the level of network simplification. Results show that alteration of food web structural properties due to aggregation are highly sensitive to the methodology used for grouping species and trophic links. In the specific case of the Barents Sea, we show that it is possible to preserve key structural properties of the original complex food web in simplified versions when using taxonomic or structural clustering combined with intermediate 25% linkage for trophic aggregation.     

Interplay between the paradox of enrichment and nutrient cycling in food webs

Nutrient cycling is fundamental to ecosystem functioning. Despite recent major advances in the understanding of complex food web dynamics, food web models have so far generally ignored nutrient cycling. However, nutrient cycling is expected to strongly impact food web stability and functioning. To make up for this gap, we built an allometric and size structured food web model including nutrient cycling. By releasing mineral nutrients, recycling increases the availability of limiting resources for primary producers and links each trophic level to the bottom of food webs. We found that nutrient cycling can provide a significant part of the total nutrient supply of the food web, leading to a strong enrichment effect that promotes species persistence in nutrient poor ecosystems but leads to a paradox of enrichment at high nutrient inputs. The presence of recycling loops linking each trophic level to the basal resources weakly affects species biomass temporal variability in the food web. Recycling loops tend to slightly dampen the destabilising effect of nutrient enrichment on consumer temporal variability while they have opposite effects for primary producers. By considering nutrient cycling, this new model improves our understanding of the response of food webs to nutrient availability and opens perspectives to better link studies on food web dynamics and ecosystem functioning.     

Effects of extinction on food web structures on an evolutionary time scale

Extinction affected food web structure in paleoecosystems. Recent theoretical studies that examined the effects of extinction intensity on food web structure on ecological time scales have considered extinction to involve episodic events, with pre-extinction food webs becoming established without dynamics. However, in terms of the paleontological time scale, food web structures are generated from feedback with repeated extinctions, because extinction frequency is affected by food web structure, and food web structure itself is a product of previous extinctions. We constructed a simulation model of changes in tri-trophic-level food webs to examine how continual extinction events affect food webs on an evolutionary time scale. We showed that under high extinction intensity (1) species diversity, especially that of consumer species, decreased; (2) the total population density at each trophic level decreased, while the densities of individual species increased; and (3) the trophic link density of the food web increased. In contrast to previous models, our results were based on an assumption of long-term food web development and are able to explain overall trends posited by empirical investigations based on fossil records.     

Integrating Ecosystem Engineering and Food Web Ecology: Testing the Effect of Biogenic Reefs on the Food Web of a Soft-Bottom Intertidal Area

The potential of ecosystem engineers to modify the structure and dynamics of food webs has recently been hypothesised from a conceptual point of view. Empirical data on the integration of ecosystem engineers and food webs is however largely lacking. This paper investigates the hypothesised link based on a field sampling approach of intertidal biogenic aggregations created by the ecosystem engineer Lanice conchilega (Polychaeta, Terebellidae). The aggregations are known to have a considerable impact on the physical and biogeochemical characteristics of their environment and subsequently on the abundance and biomass of primary food sources and the macrofaunal (i.e. the macro-, hyper- and epibenthos) community. Therefore, we hypothesise that L. conchilega aggregations affect the structure, stability and isotopic niche of the consumer assemblage of a soft-bottom intertidal food web. Primary food sources and the bentho-pelagic consumer assemblage of a L. conchilega aggregation and a control area were sampled on two soft-bottom intertidal areas along the French coast and analysed for their stable isotopes. Despite the structural impacts of the ecosystem engineer on the associated macrofaunal community, the presence of L. conchilega aggregations only has a minor effect on the food web structure of soft-bottom intertidal areas. The isotopic niche width of the consumer communities of the L. conchilega aggregations and control areas are highly similar, implying that consumer taxa do not shift their diet when feeding in a L. conchilega aggregation. Besides, species packing and hence trophic redundancy were not affected, pointing to an unaltered stability of the food web in the presence of L. conchilega.     

Food web modularity and biodiversity promote species persistence in polluted environments

Pollution represents a major threat to biodiversity. A wide class of pollutants tends to accumulate within organisms and propagate within communities via trophic interactions. Thus the final effects of accumulable pollutants may be determined by the structure of food webs and not only by the susceptibility of their constituent species. Species within real food webs are typically arranged into modules, which have been proposed to be determinants of network stability. In this study we evaluate the effect of network modularity and species richness on long‐term species persistence in communities perturbed by pollutant stress. We built model food webs with different levels of modularity and used a bioenergetic model to project the dynamics of species. Further, we modeled the dynamics of bioaccumulated and environmental pollutants. We found that modularity promoted the stability of food webs subjected to pollutant stress. We also found that richer food webs were more robust at all modularity levels. Nevertheless, modularity did not promote stability of communities facing a perturbation that shared most features with the pollutant perturbation, but does not spread through trophic interactions. The positive effect of both modularity and species richness on species persistence was cancelled and even reversed when the structure of food web departed from a realistic body size distribution or a hierarchical feeding structure. Our results support the idea that modularity implies important dynamic consequences for communities facing pollution, highlighting a main role of network structure on ecosystem stability.     

Topological keystone species: measures of positional importance in food webs

The local extinction or large fluctuation in abundance of a species may seriously affect other species in the community. The effects spread through the community by direct and indirect interactions. The network perspective on ecology can help map the pathways of these effects, for food webs, the pathways of indirect trophic interactions. Indirect interactions typically decay in intensity as they spread. Therefore, there is a conceptual maximum range in topological space beyond which interactions have no effects, even though all species remain connected. Neither the local characteristics of species, nor the global characteristics of entire webs, suitably quantify this range. We therefore apply intermediate scale indices that reflect the limitations imposed by effect damping in networks. We present a complex analysis of the topological positional importance of species in the Chesapeake Bay web. This web is a carbon-flow network that represents trophic interactions. We present several different indices reflecting different properties and discuss which questions the different indices best answer. We look for the best indices for identifying the key players in ecosystem functioning. Our study contributes to the quantification of relative species importance and provides an exact and a priori determination of a class of candidate keystone species that can inform applied and conservation ecology as well as theoretical concerns.     

How habitat-modifying organisms structure the food web of two coastal ecosystems

The diversity and structure of ecosystems has been found to depend both on trophic interactions in food webs and on other species interactions such as habitat modification and mutualism that form non-trophic interaction networks. However, quantification of the dependencies between these two main interaction networks has remained elusive. In this study, we assessed how habitat-modifying organisms affect basic food web properties by conducting in-depth empirical investigations of two ecosystems: North American temperate fringing marshes and West African tropical seagrass meadows. Results reveal that habitat-modifying species, through non-trophic facilitation rather than their trophic role, enhance species richness across multiple trophic levels, increase the number of interactions per species (link density), but decrease the realized fraction of all possible links within the food web (connectance). Compared to the trophic role of the most highly connected species, we found this non-trophic effects to be more important for species richness and of more or similar importance for link density and connectance. Our findings demonstrate that food webs can be fundamentally shaped by interactions outside the trophic network, yet intrinsic to the species participating in it. Better integration of non-trophic interactions in food web analyses may therefore strongly contribute to their explanatory and predictive capacity.     

Connecting ecology to daily life: how students and teachers relate food webs to the food they eat

This study used sociocultural learning theory to better understand how middle and high school environmental science and biology students and pre- and in-service science teachers connect the daily life activity of eating to the food web model learned in school. We sought to understand how student and teacher perceptions of the environment and their experiences influenced their responses to interview questions regarding this topic. Findings, based on transcribed interviews with 54 study participants, indicate that three quarters of teachers and students were unable to connect the food they eat with ecosystem food webs. Even so, many respondents particularly those from elite public schools, did not demonstrate common food web misconceptions identified by other researchers, instead showing a sophisticated understanding of food web interactions. These findings indicate that even though participants were proficient in their school science understanding of food web interactions, they did not readily think about how their everyday out of school activities, like eating, relate to those interactions. This may be representative of a more general disconnect between formal ecology instruction and daily life activities. We provide several recommendations for how this disconnect can be remedied in our classrooms.