Parasites Affect Food Web Structure Primarily through Increased Diversity and Complexity

Comparative research on food web structure has revealed generalities in trophic organization, produced simple models, and allowed assessment of robustness to species loss. These studies have mostly focused on free-living species. Recent research has suggested that inclusion of parasites alters structure. We assess whether such changes in network structure result from unique roles and traits of parasites or from changes to diversity and complexity. We analyzed seven highly resolved food webs that include metazoan parasite data. Our analyses show that adding parasites usually increases link density and connectance (simple measures of complexity), particularly when including concomitant links (links from predators to parasites of their prey). However, we clarify prior claims that parasites “dominate” food web links. Although parasites can be involved in a majority of links, in most cases classic predation links outnumber classic parasitism links. Regarding network structure, observed changes in degree distributions, 14 commonly studied metrics, and link probabilities are consistent with scale-dependent changes in structure associated with changes in diversity and complexity. Parasite and free-living species thus have similar effects on these aspects of structure. However, two changes point to unique roles of parasites. First, adding parasites and concomitant links strongly alters the frequency of most motifs of interactions among three taxa, reflecting parasites'' roles as resources for predators of their hosts, driven by trophic intimacy with their hosts. Second, compared to free-living consumers, many parasites'' feeding niches appear broader and less contiguous, which may reflect complex life cycles and small body sizes. This study provides new insights about generic versus unique impacts of parasites on food web structure, extends the generality of food web theory, gives a more rigorous framework for assessing the impact of any species on trophic organization, identifies limitations of current food web models, and provides direction for future structural and dynamical models.     

Lake size and fish diversity determine resource use and trophic position of a top predator in high‐latitude lakes

Prey preference of top predators and energy flow across habitat boundaries are of fundamental importance for structure and function of aquatic and terrestrial ecosystems, as they may have strong effects on production, species diversity, and food‐web stability. In lakes, littoral and pelagic food‐web compartments are typically coupled and controlled by generalist fish top predators. However, the extent and determinants of such coupling remains a topical area of ecological research and is largely unknown in oligotrophic high‐latitude lakes. We analyzed food‐web structure and resource use by a generalist top predator, the Arctic charr Salvelinus alpinus (L.), in 17 oligotrophic subarctic lakes covering a marked gradient in size (0.5–1084 km²) and fish species richness (2–13 species). We expected top predators to shift from littoral to pelagic energy sources with increasing lake size, as the availability of pelagic prey resources and the competition for littoral prey are both likely to be higher in large lakes with multispecies fish communities. We also expected top predators to occupy a higher trophic position in lakes with greater fish species richness due to potential substitution of intermediate consumers (prey fish) and increased piscivory by top predators. Based on stable carbon and nitrogen isotope analyses, the mean reliance of Arctic charr on littoral energy sources showed a significant negative relationship with lake surface area, whereas the mean trophic position of Arctic charr, reflecting the lake food‐chain length, increased with fish species richness. These results were supported by stomach contents data demonstrating a shift of Arctic charr from an invertebrate‐dominated diet to piscivory on pelagic fish. Our study highlights that, because they determine the main energy source (littoral vs. pelagic) and the trophic position of generalist top predators, ecosystem size and fish diversity are particularly important factors influencing function and structure of food webs in high‐latitude lakes.     

Food‐web structure and mercury dynamics in a large subarctic lake following multiple species introductions

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Trophic interactions and community structure in the upwelling system off Central Chile (33-39°S)

Trophic interactions and community structure in the upwelling system off Central Chile (USCCh) (33-39°S) are analyzed using biological and ecological data concerning the main trophic groups and the Ecopath with Ecosim software version 5.0 (EwE). The model encompasses the fisheries, cetaceans, sea lion, marine birds, cephalopods, large-sized pelagic fish (sword fish), medium-sized pelagic fish (horse mackerel, hoki), small-sized pelagic fish (anchovy, common sardine), demersal fish (e.g. Chilean hake, black conger-eel), benthic invertebrates (red squat lobster, yellow squat lobster) and other groups such as zooplankton, phytoplankton and detritus. Input data was gathered from published and unpublished reports and our own estimates. Trophic interactions, system indicators and food web attributes are calculated using network analysis routines included in EwE. Results indicate that trophic groups are aligned around four trophic levels (TL) with phytoplankton and detritus at the TL=1, while large-sized pelagic fish and cetaceans are top predators (TL>4.0). The fishery is located at an intermediate to low trophic level (TL=2.97), removing about 15% of the calculated system primary production. The pelagic realm dominates the system, with medium-sized pelagic fish as the main fish component in biomass, while small-sized pelagic fish dominate total landings. Chilean hake is by far the main demersal fish component in both, biomass and yield. Predators consume the greater part of the production of the most important fishery resources, particularly juvenile stages of Chilean hake. Consequently, mortality by predation is an important component of total mortality. However, fishery also removes a large fraction of common sardine, anchovy, horse mackerel, and Chilean hake. The analysis of direct and indirect trophic impacts reveals that Chilean hake is a highly cannibalistic species. Chilean hake is also an important predator on anchovy, common sardine, benthic invertebrates, and demersal fish. The fisheries heavily impact on Chilean hake, common sardine, anchovy, and horse mackerel. Total system biomass (B=476 t km⁻² year⁻¹) and throughput (T=89454 t km⁻² year⁻¹) estimated in the USCCh model are in accordance with models of comparable systems. Considering system attributes derived from network analysis, the USCCh can be characterized as an immature system, with short trophic chains and low trophic transfer efficiency. Finally, we suggest that trophic interactions should be considered in stock assessment and management programs in USCCh. In addition, future research programs should be carried out in order to understand the ecosystem effects of fishing and trophic control in this highly productive food web.     

Altered trophic pathway and parasitism in a native predator (Lepomis gibbosus) feeding on introduced prey (Dreissena polymorpha)

Populations of invasive species tend to have fewer parasites in their introduced ranges than in their native ranges and are also thought to have fewer parasites than native prey. This ‘release’ from parasites has unstudied implications for native predators feeding on exotic prey. In particular, shifts from native to exotic prey should reduce levels of trophically transmitted parasites. We tested this hypothesis in native populations of pumpkinseed sunfish (Lepomis gibbosus) in Lake Opinicon, where fish stomach contents were studied intensively in the 1970s, prior to the appearance of exotic zebra mussels (Dreissena polymorpha) in the mid-1990s. Zebra mussels were common in stomachs of present-day pumpkinseeds, and stable isotopes of carbon and nitrogen confirmed their importance in long-term diets. Because historical parasite data were not available in Lake Opinicon, we also surveyed stomach contents and parasites in pumpkinseed in both Lake Opinicon and an ecologically similar, neighboring lake where zebra mussels were absent. Stomach contents of pumpkinseed in the companion lake did not differ from those of pre-invasion fish from Lake Opinicon. The companion lake, therefore, served as a surrogate “pre-invasion” reference to assess effects of zebra mussel consumption on parasites in pumpkinseed. Trophically transmitted parasites were less species-rich and abundant in Lake Opinicon, where fish fed on zebra mussels, although factors other than zebra mussel consumption may contribute to these differences. Predation on zebra mussels has clearly contributed to a novel trophic coupling between littoral and pelagic food webs in Lake Opinicon.     

Complex life cycles in a pond food web: effects of life stage structure and parasites on network properties,trophic positions and the fit of a probabilistic niche model

Most food webs use taxonomic or trophic species as building blocks, thereby collapsing variability in feeding linkages that occurs during the growth and development of individuals. This issue is particularly relevant to integrating parasites into food webs because parasites often undergo extreme ontogenetic niche shifts. Here, we used three versions of a freshwater pond food web with varying levels of node resolution (from taxonomic species to life stages) to examine how complex life cycles and parasites alter web properties, the perceived trophic position of organisms, and the fit of a probabilistic niche model. Consistent with prior studies, parasites increased most measures of web complexity in the taxonomic species web; however, when nodes were disaggregated into life stages, the effects of parasites on several network properties (e.g., connectance and nestedness) were reversed, due in part to the lower trophic generality of parasite life stages relative to free-living life stages. Disaggregation also reduced the trophic level of organisms with either complex or direct life cycles and was particularly useful when including predation on parasites, which can inflate trophic positions when life stages are collapsed. Contrary to predictions, disaggregation decreased network intervality and did not enhance the fit of a probabilistic niche model to the food webs with parasites. Although the most useful level of biological organization in food webs will vary with the questions of interest, our results suggest that disaggregating species-level nodes may refine our perception of how parasites and other complex life cycle organisms influence ecological networks.     

Vertebrate diets derived from trophically transmitted fish parasites in the Bothnian Bay

Parasites that are transmitted through predator–prey interactions may be used as indicators of trophic relationships between organisms. Yet, they are rarely used as such in the construction of topological (predator–prey) food webs. We constructed food webs of vertebrate trophic interactions using observed diet alone, trophically transmitted parasites alone, and the combination of the two based on data from 31 species of fish from the Bothnian Bay, Finland. The fish food web contained 530 links derived from observed diet, 724 links inferred from parasitism, and 1,058 links calculated from a combination of both stomach contents and parasites. This sub-web constructed from stomach contents had a mean of 17.1 links per fish species, while that using parasites had 23.4 links per fish. Combining the two diet indicators yielded 34.1 links per fish species, illustrating the complementarity of the two methods. Mean number of prey species per fish species was 12.5 using observed diet items, 15.8 using parasites, and 24.5 using both measures together. Mean number of predators per fish species was 7.4 using observed diet, 11.7 using parasites and 15.0 using both. A positive correlation was found between the mean number of parasites and the number of prey taxa in the diet among the fishes. Omnivorous fish had the highest diversity of both parasite species and prey items, while benthophagous fish had among the lowest. Mean total abundance and mean total prevalence of parasites correlated positively with fish size, with piscivores being the largest with the highest abundance and prevalence, while planktivores and benthivores had the lowest. Trophically transmitted parasites may be used to help construct vertebrate sub-webs and derive information about food web processes. Parasites alone provided equivalent if not more information than observed diet. However, resolution is improved by using parasites and observed diet together.     

Trophic Structure of One Deep-sea Benthic Fish Community in the Eastern Canadian Arctic: Application of Food, Parasites and Multivariate Analysis

Synopsis As new arctic marine fisheries develop there is need for a comprehensive ecosystem approach to long-term management. This approach recognizes the importance of community interactions such as food web structure and trophic patterns. We determined whether hierarchical clustering (guild formation) is an effective method of trophic evaluation in deep-sea Artic fish communities using stomach content and parasite data with size class, and evaluated the application of endohelminth communities (parasite species transmitted in the food) as indicators of trophic status. Cluster analysis using food group abundance with size class of fish revealed the presence of 11 guilds within the community, however the same analysis using parasite data showed little correlation between food and parasites. Redundancy analysis (RDA) within the 11 guilds also revealed no significant correlations between food group and parasite abundance suggesting that this type of ordination is not suited for environments containing mainly generalist feeders. RDA of individual taxa without a priori guild designation found that taxa in benthic deep-sea communities are defined by their ability to exploit prey species in more than one habitat zone. Benthic fish species were significantly correlated with benthic food groups and parasites that utilize benthic intermediate hosts whereas benthopelagic–pelagic species fed on a higher diversity of prey species and were infected by a larger number of non-host specific parasites. Eigenanalysis and Monte Carlo results showed that parasites and food groups are highly correlated, indicating that parasite community analysis is an effective tool for predicting feeding strategies in Arctic marine environments. It also suggests that in most cases endoparasite infections alone could be used for trophic evaluation in the absence of stomach content data.     

东太平洋中部中上层鲨鱼群落营养生态位分化

鲨鱼在大洋生态系统中占据着重要的生态地位,其作为顶级捕食者,通过下行效应直接影响生态系统的稳定.稳定同位素技术是目前研究摄食生态学强有力的手段之一,可利用碳氮稳定同位素在食物网中的特性分别指示鲨鱼的食物来源和营养级.本研究选取8种130尾采集自东太平洋中部的中上层鲨鱼,应用稳定同位素绘制其种群生态位图谱,比较不同种群间的生态地位及资源分配方式上的差异.结果表明:不同鲨鱼种群碳、氮稳定同位素比值存在显著差异;8种鲨鱼在东太平洋生态系统中的营养级为4.3~5.4,大青鲨、尖吻鲭鲨与其他6种鲨鱼存在摄食隔离,表现出独特的营养生态地位.这些结果充分证明大洋性中上层鲨鱼并非生态系统的冗余种,其营养生态位的独特性不会被其他捕食者简单地替代和弥补.     

Parasites reduce food web robustness because they are sensitive to secondary extinction as illustrated by an invasive estuarine snail

A robust food web is one in which few secondary extinctions occur after removing species. We investigated how parasites affected the robustness of the Carpinteria Salt Marsh food web by conducting random species removals and a hypothetical, but plausible, species invasion. Parasites were much more likely than free-living species to suffer secondary extinctions following the removal of a free-living species from the food web. For this reason, the food web was less robust with the inclusion of parasites. Removal of the horn snail, Cerithidea californica, resulted in a disproportionate number of secondary parasite extinctions. The exotic Japanese mud snail, Batillaria attramentaria, is the ecological analogue of the native California horn snail and can completely replace it following invasion. Owing to the similarities between the two snail species, the invasion had no effect on predator–prey interactions. However, because the native snail is host for 17 host-specific parasites, and the invader is host to only one, comparison of a food web that includes parasites showed significant effects of invasion on the native community. The hypothetical invasion also significantly reduced the connectance of the web because the loss of 17 native trematode species eliminated many links.     

Piscivore enhancement effects on food webs depend on planktivore body size and species composition in replicated whole lake experiments

Recent syntheses of trophic cascade and biomanipulation research have suggested that the effects of piscivores on planktivorous fish populations are reduced, when planktivores are capable of outgrowing predator gape limitation and in systems with complex food web interactions. These hypotheses, however, have not been tested in long-term, whole-lake, experiments where processes such as fish recruitment and compensatory food web responses may be important. We conducted a replicated whole-lake experiment to test for the effects of supplemental piscivore introductions on food webs of eutrophic lakes dominated by deep-bodied planktivores. Responses to piscivore enhancement were compared between lakes differing in food web structure due to the presence of omnivorous gizzard shad (Dorosoma cepedianum). A significant decrease in the relative abundance of juvenile planktivorous fish, and an increase in total benthic macroinvertebrate density was observed in lakes containing mainly bluegills (Lepomus machrochirus). In contrast, lakes containing gizzard shad exhibited no significant responses to piscivore manipulation. Our results support the hypothesis that food webs in lakes dominated by deep-bodied planktivorous fish species respond weakly to piscivore enhancement. In addition, our findings support the hypothesis that cascading trophic interactions are weaker in lake ecosystems with more complex food web interactions such as those containing gizzard shad.     

Influences of fish on food web structure and function in mountain lakes

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Host centrality in food web networks determines parasite diversity

Background

Parasites significantly alter topological metrics describing food web structure, yet few studies have explored the relationship between food web topology and parasite diversity.

Methods/Principal Findings

This study uses quantitative metrics describing network structure to investigate the relationship between the topology of the host food web and parasite diversity. Food webs were constructed for four restored brackish marshes that vary in species diversity, time post restoration and levels of parasitism. Our results show that the topology of the food web in each brackish marsh is highly nested, with clusters of generalists forming a distinct modular structure. The most consistent predictors of parasite diversity within a host were: trophic generality, and eigenvector centrality. These metrics indicate that parasites preferentially colonise host species that are highly connected, and within modules of tightly interacting species in the food web network.

Conclusions/Significance

These results suggest that highly connected free-living species within the food web may represent stable trophic relationships that allow for the persistence of complex parasite life cycles. Our data demonstrate that the structure of host food webs can have a significant effect on the establishment of parasites, and on the potential for evolution of complex parasite life cycles.     

A role for brain size and cognition in food webs

Predators tend to be large and mobile, enabling them to forage in spatially distinct food web compartments (e.g. littoral and pelagic aquatic macrohabitats). This feature can stabilise ecosystems when predators are capable of rapid behavioural response to changing resource conditions in distinct habitat compartments. However, what provides this ability to respond behaviourally has not been quantified. We hypothesised that predators require increased cognitive abilities to occupy their position in a food web, which puts pressure to increase brain size. Consistent with food web theory, we found that fish relative brain size increased with increased ability to forage across macrohabitats and increased relative trophic positions in a lacustrine food web, indicating that larger brains may afford the cognitive capacity to exploit various habitats flexibly, thus contributing to the stability of whole food webs.     

Exploitation of asymmetric predator–prey interactions by trophically transmitted parasites

The directionality of asymmetric interactions between predators (definitive hosts) and prey (intermediate hosts) should impact trophic transmission in parasites. This study tests the prediction that trophically transmitted parasites are funneled towards asymmetric predator–prey interactions where intermediate hosts have few predators and definitive hosts feed upon many prey (‘downward asymmetry’). The distribution of trophically transmitted parasites was examined in four published food webs in relation to mismatch asymmetry of predator–prey interactions. We found that trophically transmitted parasites exploit downwardly asymmetric interactions in a nonrandom manner, and particular predator–prey pairs contain more trophically transmitted parasites than would be expected by random chance alone. These findings suggest that food web topology has great bearing on the ecology of trophically transmitted parasites, and that consideration of parasite life cycles in the context of food web organization can provide insights into the forces affecting the evolution of trophic transmission.     

Ecological role and historical trends of large pelagic predators in a subtropical marine ecosystem of the South Atlantic

Large pelagic predators occupy high positions in food webs and could control lower trophic level species by direct and indirect ecological interactions. In this study we aimed to test the hypotheses: (1) pelagic predators are keystone species, and their removals could trigger impacts on the food chain; (2) higher landings of pelagic predators could trigger fishing impacts with time leading to a drop in the mean trophic level of catches; and (3) recovery in the pelagic predators populations, especially for sharks, could be achieved with fishing effort reduction. We performed a food web approach using an Ecopath with Ecosim model to represent the Southeastern and Southern Brazil, a subtropical marine ecosystem, in 2001. We then calibrated the baseline model using catch and fishing effort time series from 2001 to 2012. Afterwards, we simulated the impact of fishing effort changes on species and assessed the ecological impacts on the pelagic community from 2012 to 2025. Results showed that the model was well fitted to landing data for the majority of groups. The pelagic predators species were classified as keystone species impacting mainly on pelagic community. The ecosystem was resilient and fisheries seem sustainable at that time. However, the temporal simulation, from 2001 to 2012, revealed declines in the biomass of three sharks, tuna and billfish groups. It was possible observe declines in the mean trophic level of the catch and in the mean total length of landings. Longline fisheries particularly affected the sharks, billfish and swordfish, while hammerhead sharks were mostly impacted by gillnet fishery. Model simulations showed that large sharks’ biomasses could be recovered or maintained only after strong fishing effort reduction.     

高营养级捕食者在浅水湖泊沿岸带与敞水区能量耦合的维持作用

研究利用保安湖沿岸带与敞水区两种生境中高营养级捕食者(达氏鲌与红鳍原鲌)的碳(δ13C)、氮(δ15N)稳定同位素数据,通过稳定同位素质量平衡混合模型、非度量多维尺度分析(NMDS)等方法,分析了两种生境中鲌类食物来源的差异.结果表明,两种生境中鲌类食物来源基本一致,且食物来源较为广泛,包括沿岸带饵料鱼类、敞水区饵料鱼...     

Trophic niche differences between coexisting omnivores silver carp and bighead carp in a pelagic food web

Understanding how omnivorous consumers are affected by their resources and how this is expressed through the food chain is a fundamental issue in ecology. We used stable isotope analysis of archived scales of two pelagic single-chain omnivorous fish species, bighead carp (Hypophthalmichthys nobilis) and silver carp (H. molitrix), to reconstruct historical trophic interactions patterns along a gradient of resources. We found that, although bighead carp and silver carp utilize the similar resources from the pelagic food chain, they can coexist and persist not only by regulating their trophic position and trophic dissimilarity, but also by regulating trophic niche width. Omnivorous fish often exhibit flexible foraging strategies, which is closely related to the availability of ecologic context. We found a positive relationship between trophic dissimilarity and zooplankton density, which may indicate that the competitive interactions induce strong top-down effects on zooplankton, and/or that high zooplankton availability release the between-population trophic interaction through bottom-up effect. The trophic niche width of bighead carp was positively related with zooplankton availability, probably reflecting that the niche of an omnivore at a higher trophic position is more sensitive to high quality resources. Our results indicate how different aspects of the trophic partitioning of coexisting omnivores may be regulated by different ecological contexts. These alternatives are not mutually exclusive and further theoretical work should include both these mechanisms to re-evaluate the effects of omnivory on food web properties.     

Defining and measuring trophic role similarity in food webs using regular equivalence

We present a graph theoretic model of analysing food web structure called regular equivalence. Regular equivalence is a method for partitioning the species in a food web into "isotrophic classes" that play the same structural roles, even if they are not directly consuming the same prey or if they do not share the same predators. We contrast regular equivalence models, in which two species are members of the same trophic group if they have trophic links to the same set of other trophic groups, with structural equivalence models, in which species are equivalent if they are connected to the exact same other species. Here, the regular equivalence approach is applied to two published food webs: (1) a topological web (Malaysian pitcher plant insect food web) and (2) a carbon-flow web (St. Marks, Florida seagrass ecosystem food web). Regular equivalence produced a more satisfactory set of classes than did the structural approach, grouping basal taxa with other basal taxa and not with top predators. Regular equivalence models provide a way to mathematically formalize trophic position, trophic group and trophic niche. These models are part of a family of models that includes structural models used extensively by ecologists now. Regular equivalence models uncover similarities in trophic roles at a higher level of organization than do the structural models. The approach outlined is useful for measuring the trophic roles of species in food web models, measuring similarity in trophic relations of two or more species, comparing food webs over time and across geographic regions, and aggregating taxa into trophic groups that reduce the complexity of ecosystem feeding relations without obscuring network relationships. In addition, we hope the approach will prove useful in predicting the outcome of predator-prey interactions in experimental studies.     

Niche segregation of coexisting Arctic charr (Salvelinus alpinus) and brown trout (Salmo trutta) constrains food web coupling in subarctic lakes

1. Generalist fish species are recognised as important couplers of benthic and pelagic food‐web compartments in lakes. However, interspecific niche segregation and individual specialisation may limit the potential for generalistic feeding behaviour. 2. We studied summer habitat use, stomach contents and stable isotopic compositions of the generalist feeder Arctic charr coexisting with its common resource competitor brown trout in five subarctic lakes in northern Norway to reveal population‐level and individual‐level niche plasticity. 3. Charr and trout showed partial niche segregation in all five lakes. Charr used all habitat types and a wide variety of invertebrate prey including zooplankton, whereas trout fed mainly on insects in the littoral zone. Hence, charr showed a higher potential to promote habitat and food‐web coupling compared to littoral‐dwelling trout. 4. The level of niche segregation between charr and trout and between pelagic‐caught and littoral‐caught charr depended on the prevailing patterns of interspecific and intraspecific resource competition. The two fish species had partially overlapping trophic niches in one lake where charr numerically dominated the fish community, whereas the most segregated niches occurred in lakes where trout were more abundant. 5. In general, pelagic‐caught charr had substantially narrower dietary and isotopic niches and relied less on littoral carbon sources compared to littoral‐caught conspecifics that included generalist as well as specialised benthivorous and planktivorous individuals. Despite the partially specialised planktivorous niche and thus reduced potential of pelagic‐dwelling charr to promote benthic–pelagic coupling, the isotopic compositions of both charr subpopulations suggested a significant reliance on both littoral and pelagic carbon sources in all five study lakes. 6. Our study demonstrates that both interspecific niche segregation between and individual trophic specialisation within generalist fish species can constrain food‐web coupling and alter energy mobilisation to top consumers in subarctic lakes. Nevertheless, pelagic and littoral habitats and food‐web compartments may still be highly integrated due to the potentially plastic foraging behaviour of top consumers.