Plant species identity and diversity effects on different trophic levels of nematodes in the soil food web

Previous studies on biodiversity and soil food web composition have mentioned plant species identity, as well as plant species diversity as the main factors affecting the abundance and diversity of soil organisms. However, most studies have been carried out under limitations of time, space, or appropriate controls. In order to further examine the relation between plant species diversity and the soil food web, we conducted a three-year semi-field experiment in which eight plant species (4 forb and 4 grass species) were grown in monocultures and mixtures of two, four and eight plant species. In addition there were communities with 16 plant species. We analyzed the abundance and identity of the nematodes in soil and roots, including feeding groups from various trophic levels (primary and secondary consumers, carnivores, and omnivores) in the soil food web.
Plant species diversity and plant identity affected the diversity of nematodes. The effect of plant diversity was attributed to the complementarity in resource quality of the component plant species rather than to an increase in total resource quantity. The nematode diversity varied more between the different plant species than between different levels of plant species diversity, so that plant identity is more important than plant diversity. Nevertheless the nematode diversity in plant mixtures was higher than in any of the plant monocultures, due to the reduced dominance of the most abundant nematode taxa in the mixed plant communities. Plant species identity affected the abundances of the lower trophic consumer levels more than the higher trophic levels of nematodes. Plant species diversity and plant biomass did not affect nematode abundance. Our results, therefore, support the hypothesis that resource quality is more important than resource quantity for the diversity of soil food web components and that plant species identity is more important than plant diversity per se.     

Linking aboveground and belowground communities: the indirect influence of aphid species identity and diversity on a three trophic level soil food web

There has been a growing recent interest in how foliar herbivory may indirectly affect the belowground sub-system, but little is known about the belowground consequences of the identity, species composition or diversity of foliar herbivores. We performed an experiment, utilising model grassland communities containing three plant species, in which treatments consisted of addition of each of eight aphid species in single and in two- four- and eight-species combinations, as well as an aphid-free treatment. While aphid species treatments did not affect total plant biomass or productivity, aphid species identity had important effects on the relative abundance of the three plant species. This in turn affected the abundances of each of three groups of secondary consumers in the soil food web (bacterial- and fungal-feeding nematodes, and enchytraeids) but not primary consumers (microbes, herbivorous nematodes) or tertiary consumers (predatory nematodes). The fact that some trophic levels responded to treatments while others did not is consistent with trophic dynamic theory. Aphid species treatments also affected the community composition within each of the herbivorous, microbe-feeding and top predatory nematode groups, as well as diversity within the first two of these groups. However, aphid species diversity per se had few effects. There were specific instances in which specific aboveground and belowground response variables in two aphid species combinations differed significantly from those in both of the corresponding single aphid species treatments (apparently as a consequence of resource use complementarity between coexisting aphid species), but no instance in which increasing aphid diversity beyond two species had any effect. Our results provide evidence that the identity of aboveground consumers can have effects that propagate through multiple trophic levels in soil food webs in terms of consumer abundance, and composition and diversity within trophic levels.     

Compensation masks trophic cascades in complex food webs

Ecological networks, or food webs, describe the feeding relationships between interacting species within an ecosystem. Understanding how the complexity of these networks influences their response to changing top-down control is a central challenge in ecology. Here, we provide a model-based investigation of trophic cascades — an oft-studied ecological phenomenon that occurs when changes in the biomass of top predators indirectly effect changes in the biomass of primary producers — in complex food webs that are representative of the structure of real ecosystems. Our results reveal that strong cascades occur primarily in low richness and weakly connected food webs, a result in agreement with some prior predictions. The primary mechanism underlying weak or absent cascades was a strong compensatory response; in most webs, predators induced large population level cascades that were masked by changes in the opposite direction by other species in the same trophic guild. Thus, the search for a general theory of trophic cascades in food webs should focus on uncovering features of real ecosystems that promote biomass compensation within functional guilds or trophic levels.     

Legacy effects of drought on plant growth and the soil food web

Soils deliver important ecosystem services, such as nutrient provision for plants and the storage of carbon (C) and nitrogen (N), which are greatly impacted by drought. Both plants and soil biota affect soil C and N availability, which might in turn affect their response to drought, offering the potential to feed back on each other’s performance. In a greenhouse experiment, we compared legacy effects of repeated drought on plant growth and the soil food web in two contrasting land-use systems: extensively managed grassland, rich in C and with a fungal-based food web, and intensively managed wheat lower in C and with a bacterial-based food web. Moreover, we assessed the effect of plant presence on the recovery of the soil food web after drought. Drought legacy effects increased plant growth in both systems, and a plant strongly reduced N leaching. Fungi, bacteria, and their predators were more resilient after drought in the grassland soil than in the wheat soil. The presence of a plant strongly affected the composition of the soil food web, and alleviated the effects of drought for most trophic groups, regardless of the system. This effect was stronger for the bottom trophic levels, whose resilience was positively correlated to soil available C. Our results show that plant belowground inputs have the potential to affect the recovery of belowground communities after drought, with implications for the functions they perform, such as C and N cycling.     

Trophic cascades and trophic trickles in pelagic food webs

Prey-dependent models, with the predation rate (per predator) a function of prey numbers alone, predict the existence of a trophic cascade. In a trophic cascade, the addition of a top predator to a two-level food chain to make a three-level food chain will lead to increases in the population size of the primary producers, and the addition of nutrients to three-level chains will lead to increases in the population numbers at only the first and third trophic levels. In contrast, ratio-dependent models, with the predation rate (per predator) dependent on the ratio of predator numbers to prey, predict that additions of top predators will not increase the population sizes of the primary producers, and that the addition of nutrients to a three-level food chain will lead to increases in population numbers at all trophic levels. Surprisingly, recent meta-analyses show that freshwater pelagic food web patterns match neither prey-dependent models (in pelagic webs, ''prey'' are phytoplankton, and ''predators'' are zooplankton), nor ratio-dependent models. In this paper we use a modification of the prey-dependent model, incorporating strong interference within the zooplankton trophic level, that does yield patterns matching those found in nature. This zooplankton interference model corresponds to a more reticulate food web than in the linear, prey-dependent model, which lacks zooplankton interference. We thus reconcile data with a new model, and make the testable prediction that the strength of trophic cascades will depend on the degree of heterogeneity in the zooplankton level of the food chain.     

Bottom-up trophic cascades and material transfer in terrestrial food webs

In contrast to top-down trophic cascades, few reviews have appeared of bottom-up trophic cascades. We review the recent development of research on bottom-up cascades in terrestrial food webs, focusing on tritrophic systems consisting of plants, herbivorous insects, and natural enemies, and attempt to integrate bottom-up cascade and material transfer among trophic levels. Bottom-up cascades are frequently reported in various tritrophic systems, and are important to determine community structure, population dynamics, and individual performance of higher trophic levels. In addition, we highlight several features of bottom-up cascades. Accumulation or dilution of plant nutritional and defensive materials by herbivorous insects provides a mechanistic base for several bottom-up cascades. Such a stoichiometric approach has the potential to improve our understanding of bottom-up cascading effects in terrestrial food webs. We suggest a future direction for research by integration of bottom-up cascades and material transfer among trophic levels.     

Omnivory does not prevent trophic cascades in pelagic food webs

1. Strong trophic cascades have been well documented in pelagic food webs of temperate lakes. In contrast, the limited available evidence suggests that strong cascades are less typical in tropical lakes.
2. To measure the effects of omnivorous tilapia on planktonic communities and water transparency of a small man-made tropical lake, we performed a 5-week in situ enclosure experiment with five densities of fish randomly allocated to 20 enclosures. Zooplankton and Phytoplankton biomasses as well as water transparency were measured weekly.
3. Results show that omnivorous tilapia significantly decreased the abundance of large Cladocerans, increased the abundance of small algae (greatest axial linear dimension <50  μ m) and decreased water transparency as predicted by trophic cascade theory.
4. Therefore, omnivory was not a sufficient factor to prevent a trophic cascade in this pelagic community, although the cascade effect was weaker than reported from many north temperate, nutrient-rich lakes.     

Using trophic hierarchy to understand food web structure

Link arrangement in food webs is determined by the species' feeding habits. This work investigates whether food web topology is organized in a gradient of trophic positions from producers to consumers. To this end, we analyzed 26 food webs for which the consumption rate of each species was specified. We computed the trophic positions and the link densities of all species in the food webs. Link density measures how much each species contributes to the distribution of energy in the system. It is expressed as the number of links species establish with other nodes, weighted by their magnitude. We computed these two metrics using various formulations developed in the ecological network analysis framework. Results show a positive correlation between trophic position and link density across all the systems, regardless the specific formulas used to measure the two quantities. We performed the same analysis on the corresponding binary matrices (i.e. removing information about rates). In addition, we investigated the relation between trophic position and link density in: a) simulated binary webs with same connectance as the original ones; b) weighted webs with constant topology but randomized link strengths and c) weighted webs with constant connectance where both topology and link strengths are randomized. The correlation between the two indices attenuates, vanishes or becomes negative in the case of binary food webs and simulated data (weighted and unweighted).
According to our analysis, link density in food webs decreases with trophic position so that it is greatly reduced toward the top of the trophic hierarchy. This outcome, that seems to challenge previous conclusions based on null models, strongly depends on link quantification. Including interaction strengths may improve substantially our understanding of food web organization, and possibly contradict results based on the analysis of binary webs.     

Evaluating the effects of trophic complexity on a keystone predator by disassembling a partial intraguild predation food web

1. Many taxa can be found in food webs that differ in trophic complexity, but it is unclear how trophic complexity affects the performance of particular taxa. In pond food webs, larvae of the salamander Ambystoma opacum occupy the intermediate predator trophic position in a partial intraguild predation (IGP) food web and can function as keystone predators. Larval A. opacum are also found in simpler food webs lacking either top predators or shared prey. 2. We conducted an experiment where a partial IGP food web was simplified, and we measured the growth and survival of larval A. opacum in each set of food webs. Partial IGP food webs that had either a low abundance or high abundance of total prey were also simplified by independently removing top predators and/or shared prey. 3. Removing top predators always increased A. opacum survival, but removal of shared prey had no effect on A. opacum survival, regardless of total prey abundance. 4. Surprisingly, food web simplification had no effect on the growth of A. opacum when present in food webs with a low abundance of prey but had important effects on A. opacum growth in food webs with a high abundance of prey. Simplifying a partial IGP food web with a high abundance of prey reduced A. opacum growth when either top predators or shared prey were removed from the food web and the loss of top predators and shared prey influenced A. opacum growth in a non-additive fashion. 5. The non-additive response in A. opacum growth appears to be the result of supplemental prey availability augmenting the beneficial effects of top predators. Top predators had a beneficial effect on A. opacum populations by reducing the abundance of A. opacum present and thereby reducing the intensity of intraspecific competition. 6. Our study indicates that the effects of food web simplification on the performance of A. opacum are complex and depend on both how a partial IGP food web is simplified and how abundant prey are in the food web. These findings are important because they demonstrate how trophic complexity can create variation in the performance of intermediate predators that play important roles in temporary pond food webs.     

Intraguild indirect effects through trophic cascades between stream-dwelling fishes

1. To examine the strength and role of indirect effects through trophic cascades by omnivorous fish on algal biomass in streams, we introduced one of four fish species (ayu Plecoglossus altivelis altivelis, pike gudgeon Pseudogobio esocinus esocinus, Japanese dace Tribolodon hakonensis and pale chub Zacco platypus) in experimental pools. We also investigated the indirect effects of gudgeon, dace and chub on the growth of ayu. 2. We chose the four fish species across a continuum of omnivory. Ayu fed mainly on benthic algae, and gudgeon predominantly on invertebrates. Dace and chub fed on both algae and invertebrates, but dace preyed on invertebrates more than chub. 3. The presence of gudgeon, dace and chub reduced the number of algal-feeding invertebrates and increased the algal biomass through trophic cascades. Consequently, ayu growth rate over the experimental period in pools with one of the three fish species was 25.9-44.1 times greater than the growth rate in pools with only ayu. 4. The positive indirect effect on ayu growth was large for gudgeon and dace and small for chub, whereas the addition of ayu reduced ayu growth considerably due to intraspecific competition. 5. The relative intensity of indirect effects on ayu growth through trophic cascades was predictable from the food overlap between ayu and the other fishes. However, the similar strength of indirect effects by gudgeon and dace that fed differently on algae and invertebrates suggests that feeding behaviour, prey preference and trait-mediated indirect interactions were also important in the prediction.     

Predicting the effects of temperature on food web connectance

Few models concern how environmental variables such as temperature affect community structure. Here, we develop a model of how temperature affects food web connectance, a powerful driver of population dynamics and community structure. We use the Arrhenius equation to add temperature dependence of foraging traits to an existing model of food web structure. The model predicts potentially large temperature effects on connectance. Temperature-sensitive food webs exhibit slopes of up to 0.01 units of connectance per 1°C change in temperature. This corresponds to changes in diet breadth of one resource item per 2°C (assuming a food web containing 50 species). Less sensitive food webs exhibit slopes down to 0.0005, which corresponds to about one resource item per 40°C. Relative sizes of the activation energies of attack rate and handling time determine whether warming increases or decreases connectance. Differences in temperature sensitivity are explained by differences between empirical food webs in the body size distributions of organisms. We conclude that models of temperature effects on community structure and dynamics urgently require considerable development, and also more and better empirical data to parameterize and test them.     

Subsidy from the detrital food web, but not microhabitat complexity, affects the role of generalist predators in an aboveground herbivore food web

The activity and density of generalist predators, such as carabid beetles, rove beetles and spiders, may increase in response to: (1) increased availability of prey from the belowground subsystem and/or (2) enhanced complexity of aboveground vegetation. Organic farming practices support decomposer populations and enhance habitat complexity due to an increased weed density. A response by generalist predators to such below‐ or aboveground changes could affect predation rates on herbivores in the aboveground food web. We tested this hypothesis in a replicated field experiment conducted in a winter wheat field, where increased predator activity could lead to improved control of herbivorous pests. In a crossed design, we increased and lowered densities of decomposer prey, and manipulated vegetation complexity using artificial plants in order to examine the effect of structural complexity in isolation from effects of plant‐attracted additional prey. Isotomid Collembola exhibited lowest activity‐densities (AD) in plots treated with soil insecticide and had gradually increasing AD in untreated plots and plots receiving detrital subsidies. Carabid beetles and cursorial spiders did not respond to increased availability of isotomid prey, and they unexpectedly displayed higher AD in the structurally less‐complex plots. Aphid density mirrored the positive response of isotomids to detrital subsidies, suggesting that aphids benefited from reduced predation due to predators switching to abundant prey in the decomposer subsystem. The absence of a numerical response by surface‐active predators apparently strengthened this indirect effect of isotomids on aphids. Our results suggest that indirect predator‐mediated prey‐prey interactions can reduce beneficial effects of detrital subsidies on pest suppression. We further demonstrated that generalist predators may not per se benefit from structural complexity. Both results document the challenges associated with management practices that support generalist predators, as these measures may not necessarily improve herbivore suppression.     

Intraguild predation,invertebrate predators,and trophic cascades in lake food webs

The top-down and bottom-up properties of model food webs that include intraguild predation and self-limiting factors such as cannibalism are investigated. Intraguild predation can dampen or even reverse the top-down effects predicted by food chain theory. The degree of self-limitation among the intraguild prey is a key factor in determining the direction and strength of the top-down response. Intraguild predation and self-limiting factors can also substantially alter the bottom-up effects of enrichment. These results can help explain the disparate results of trophic cascade experiments in lakes, where cascades are usually seen when large Daphnia are the primary herbivores, but not when smaller-bodied herbivores are dominant. Top-down manipulations should cascade at least modestly to phytoplankton in those lakes whose food web can be reasonably approximated by a chain (typically, those where Daphnia is the dominant herbivore), as predicted by food chain theory. On the other hand, smaller-bodied zooplankton are often preyed upon heavily by invertebrate predators as well as by planktivorous fish, thereby introducing elements of intraguild predation into these food webs. In this case, conventional food chain theory is likely to give incorrect predictions. Very large cascade effects may be due primarily to regime shifts between intraguild predation-dominated food webs and those that more resemble food chains, rather than due to the simple food chain cascade usually considered.     

Allochtonous input and trophic level heterogeneity: impact on an aquatic food web

The impact of food web complexity in open and closed food webs was explored experimentally under controlled laboratory conditions. We used an aquatic model community consisting of two forms of primary producers, phytoplankton ( Scenedesmus obtusiuscusculus ) and periphyton ( Nitzschia perminuta ) and two types of consumers, Daphnia pulex feeding on phytoplankton and Chydorous sphaericus feeding on both periphyton and phytoplankton. Three different food webs all having the phytoplankton and periphyton, but having either one of the consumers or both were set up. These food webs were studied in an open and closed treatment. In the open treatment, phytoplankton was continuously flowing through the aquaria whereas in the closed system all the phytoplankton was delivered at the start of the experiment.
D. pulex had a positive effect on the density of C. sphaericus in both the open and closed treatments. In the open treatment C. sphaericus increased to very high numbers and was able to depress phytoplankton and thereby negatively affect D. pulex . Our study shows that the explicit handling of the population dynamics of both grazers allowed us to show how a compensatory increase in one primary producer due to increased grazing on another primary producer creates a negative feedback between consumers: C. sphaericus increased and negatively affected D. pulex .     

The response of a three trophic level soil food web to the identity and diversity of plant species and functional groups

Despite considerable recent interest in how biodiversity may influence ecosystem properties, the issue of how plant diversity and composition may affect multiple trophic levels in soil food webs remains essentially unexplored. We conducted a glasshouse experiment in which three plant species of each of three functional groups (grasses, N‐fixing legumes and forbs) were grown in monoculture and in mixtures of three species (with the three species being in the same or different functional groups) and all nine species. Plant species identity had important effects on the biomasses or population densities of belowground primary consumers (microbial biomass, herbivorous nematodes) and two groups of secondary consumers (microbe‐feeding nematodes and enchytraeids); the third consumer trophic level (predatory nematodes) was marginally not significantly affected at P=0.05. Plant species also influenced the relative importance of the bacterial‐based and fungal‐based energy channels for both the primary and secondary consumer trophic levels. Within‐group diversity of only the soil microflora and herbivorous nematodes (both representing the basal consumer trophic level) were affected by plant species identity. However, community composition within all trophic groupings considered (herbivorous nematodes, microbes, microbe‐feeding nematodes, predatory nematodes) was strongly influenced by what plant species were present. Despite the strong responses of the soil biota to plant species identity, there were few effects of plant species or functional group richness on any of the belowground response variables measured. Further, net primary productivity (NPP) was unaffected by plant diversity. Since some belowground response variables were correlated with NPP across treatments, it is suggested that belowground responses to plant diversity might become more apparent in situations when NPP itself responds to plant diversity. Our results point to plant species identity as having important multitrophic effects on soil food webs, both at the whole trophic group and within‐group levels of resolution, and suggest that differences in plant traits across species may be important in driving the decomposer subsystem.     

Molecular detection of trophic links in a complex insect host-parasitoid food web

Previously, host-parasitoid links have been unveiled almost exclusively by time-intensive rearing, while molecular methods were used only in simple agricultural host-parasitoid systems in the form of species-specific primers. Here, we present a general method for the molecular detection of these links applied to a complex caterpillar-parasitoid food web from tropical rainforest of Papua New Guinea. We DNA barcoded hosts, parasitoids and their tissue remnants and matched the sequences to our extensive library of local species. We were thus able to match 87% of host sequences and 36% of parasitoid sequences to species and infer subfamily or family in almost all cases. Our analysis affirmed 93 hitherto unknown trophic links between 37 host species from a wide range of Lepidoptera families and 46 parasitoid species from Hymenoptera and Diptera by identifying DNA sequences for both the host and the parasitoid involved in the interaction. Molecular detection proved especially useful in cases where distinguishing host species in caterpillar stage was difficult morphologically, or when the caterpillar died during rearing. We have even detected a case of extreme parasitoid specialization in a pair of Choreutis species that do not differ in caterpillar morphology and ecology. Using the molecular approach outlined here leads to better understanding of parasitoid host specificity, opens new possibilities for rapid surveys of food web structure and allows inference of species associations not already anticipated.     

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.     

Food web architecture and basal resources interact to determine biomass and stoichiometric cascades along a benthic food web

Understanding the effects of predators and resources on primary producers has been a major focus of interest in ecology. Within this context, the trophic cascade concept especially concerning the pelagic zone of lakes has been the focus of the majority of these studies. However, littoral food webs could be especially interesting because base trophic levels may be strongly regulated by consumers and prone to be light limited. In this study, the availability of nutrients and light and the presence of an omnivorous fish (Hyphessobrycon bifasciatus) were manipulated in enclosures placed in a humic coastal lagoon (Cabiúnas Lagoon, Macaé - RJ) to evaluate the individual and interactive effects of resource availability (nutrients and light) and food web configuration on the biomass and stoichiometry of periphyton and benthic grazers. Our findings suggest that light and nutrients interact to determine periphyton biomass and stoichiometry, which propagates to the consumer level. We observed a positive effect of the availability of nutrients on periphytic biomass and grazers' biomass, as well as a reduction of periphytic C∶N∶P ratios and an increase of grazers' N and P content. Low light availability constrained the propagation of nutrient effects on periphyton biomass and induced higher periphytic C∶N∶P ratios. The effects of fish presence strongly interacted with resource availability. In general, a positive effect of fish presence was observed for the total biomass of periphyton and grazer's biomass, especially with high resource availability, but the opposite was found for periphytic autotrophic biomass. Fish also had a significant effect on periphyton stoichiometry, but no effect was observed on grazers' stoichiometric ratios. In summary, we observed that the indirect effect of fish predation on periphyton biomass might be dependent on multiple resources and periphyton nutrient stoichiometric variation can affect consumers' stoichiometry.     

Temporal and spatial variability in soil food web structure

Heterogeneity is a prominent feature of most ecosystems. As a result of environmental heterogeneity the distribution of many soil organisms shows a temporal as well as horizontal and vertical spatial patterning. In spite of this, food webs are usually portrayed as static networks with highly aggregated trophic groups over broader scales of time and space. The variability in food web structure and its consequences have seldom been examined. Using data from a Scots pine forest soil in the Netherlands, we explored (1) the temporal and spatial variability of a detrital food web and its components, (2) the effect of taxonomic resolution on the perception of variability over time and across space, and (3) the importance of organic matter quality as an explanatory factor for variability in food web composition. Compositional variability, expressed using the Bray‐Curtis similarity index, was measured over 2.5 years using a stratified litterbag design with three organic horizons per litterbag set. Variability in community composition and organic matter degradation increased over time in the litter horizon only. Seasonal variation in community composition was larger than variation between samples from the same season in different years. Horizontal spatial variability in community composition and organic matter degradation was relatively low, with no increase in variability with increasing distance between samples. Vertically, communities and organic matter degradation was more different between the non‐adjacent litter and humus horizons than between adjacent layers. These findings imply that soil food webs, at least in temperate forest plantations, are more variable than is currently appreciated in experiments and model studies, and that organic matter turnover might be an important factor explaining variability in community composition. Species composition was more variable than functional group composition, which implies that aggregated food webs will seem less sensitive to local temporal and spatial changes than they in fact are.