Food web structure in Mediterranean streams: exploring stabilizing forces in these ecosystems

We constructed the food webs of six Mediterranean streams in order to determine ecological generalities derived from analysis of their structure and to explore stabilizing forces within these ecosystems. Fish, macroinvertebrates, primary producers and detritus are the components of the studied food webs. Analysis focused on a suite of food web properties that describe species’ trophic habits, linkage complexity and food chains. A great structural similarity was found in analyzed food webs; we therefore suggest average values for the structural properties of Mediterranean stream food webs. Percentage of omnivorous species was positively correlated with connectance, and there was a predominance of intermediate trophic level species that had established simple links with detritus. In short, our results suggest that omnivory and the weak interactions of detritivores have a stabilizing role in these food webs.     

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.     

Estimating morphologically determined connectance and structure for food webs of freshwater invertebrates

1. Connectance is a parameter of central importance in determining food-web structure, but the processes determining its value remain unclear. In evaluating possible explanations it is useful to know what patterns, and values, of connectance occur in food webs assembled at random from a set of species in a regional species pool; i.e. where the number of links is determined by the morphological features of the species present, not by the immediate effects of energetics or stability on the particular web. 2. This study examines, by means of laboratory experiments, the occurrence of potential feeding interactions among a set of freshwater invertebrate species randomly selected from different freshwater sites in a geographical region. The results from pairwise feeding trials are used to construct two ‘theoretical’ food webs, in which the patterns and values of connectance are examined. 3. Analyses of these webs indicate that their structure is consistent with the observed values in previously documented ‘real’ webs. Directed connectance values of 0.12–0.16 (or less) suggest that the assembled webs are no more connected than many freshwater webs from natural systems. The number of links per species increases curvilinearly with the number of species, during web assembly, consistent with recent hypotheses. 4. These results also indicate that quantifying, and understanding the determinants of, trophic generalism or specialism does have implications for understanding how connectance is constrained in real webs. Freshwater invertebrates seem to be relatively generalist, and freshwater food webs perhaps correspondingly highly connected. Such arguments have implications for interpreting other aspects of food-web structure in these systems, and for parameterizing models that are based on connectance.     

Global synthesis suggests that food web connectance correlates to invasion resistance

Biological invasions are a key component of global change, and understanding the drivers of global invasion patterns will aid in assessing and mitigating the impact of invasive species. While invasive species are most often studied in the context of one or two trophic levels, in reality species invade communities comprised of complex food webs. The complexity and integrity of the native food web may be a more important determinant of invasion success than the strength of interactions between a small subset of species within a larger food web. Previous efforts to understand the relationship between food web properties and species invasions have been primarily theoretical and have yielded mixed results. Here, we present a synthesis of empirical information on food web connectance and species invasion success gathered from different sources (estimates of food web connectance from the primary literature and estimates of invasion success from the Global Invasive Species Database as well as the primary literature). Our results suggest that higher‐connectance food webs tend to host fewer invaders and exert stronger biotic resistance compared to low‐connectance webs. We argue that while these correlations cannot be used to infer a causal link between food web connectance and habitat invasibility, the promising findings beg for further empirical research that deliberately tests for relationships between food web connectance and invasion.     

Food web structure in a tropical stream ecosystem

Abstract This study investigated the structure and properties of a tropical stream food web in a small spatial scale, characterizing its planktonic, epiphytic and benthic compartments. The study was carried out in the Potreirinho Creek, a second‐order stream located in the south‐east of Brazil. Some attributes of the three subwebs and of the conglomerate food web, composed by the trophic links of the three compartments plus the fish species, were determined. Among compartments, the food webs showed considerable variation in structure. The epiphytic food web was consistently more complex than the planktonic and benthic webs. The values of number of species, number of links and maximum food chain length were significantly higher in the epiphytic compartment than in the other two. Otherwise, the connectance was significantly lower in epiphyton. The significant differences of most food web parameters were determined by the increase in the number of trophic species, represented mainly by basal and intermediate species. High species richness, detritus‐based system and high degree of omnivory characterized the stream food web studied. The aquatic macrophytes probably provide a substratum more stable and structurally complex than the sediment. We suggest that the greater species richness and trophic complexity in the epiphytic subweb might be due to the higher degree of habitat complexity supported by macrophyte substrate. Despite differences observed in the structure of the three subwebs, they are highly connected by trophic interactions, mainly by fishes. The high degree of fish omnivory associated with their movements at different spatial scales suggests that these animals have a significant role in the food web dynamic of Potreirinho Creek. This interface between macrophytes and the interconnections resultant from fish foraging, diluted the compartmentalization of the Potreirinho food web.     

Food webs robustness to biodiversity loss: the roles of connectance, expansibility and degree distribution

We analyse the robustness of food webs against species loss by considering the influence of several structural factors of the networks, such as connectance, degree distribution and expansibility. The last concept refers to the absence of structural bottlenecks in the food web, whose removal separate the network into large isolate clusters. In theory networks with identical connectance can display different expansibility characteristics. Using the spectral scaling method we studied 17 food networks and classified them as good expansion (GE) and not-GE networks. The combination of GE properties and degree distribution of species permitted the classification of food webs into six different classes. These classes characterize the differences in robustness of food webs to species loss. While the webs having uniform degree distributions and displaying GE properties are the most robust to species loss, the presence of bottlenecks and skewed distribution of the number of links per species make food webs very vulnerable to primary removal of species.     

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.     

Food webs: a plea for parasites

Parasites have the capacity to regulate host populations and may be important determinants of community structure, yet they are usually neglected in studies of food webs. Parasites can provide much of the information on host biology, such as diet and migration, that is necessary to construct accurate webs. Because many parasites have complex life cycles that involve several different hosts, and often depend on trophic interactions for transmission, parasites provide complementary views of web structure and dynamics. Incorporation of parasites in food webs can substantially after baste web properties, Including connectance, chain length and proportions of top and basal species, and can allow the testing of specific hypotheses related to food-web dynamics.     

Forecasting fine‐scale changes in the food‐web structure of coastal marine communities under climate change

Climate change is inducing deep modifications in local communities worldwide as a consequence of individualistic species range shifts. Understanding how complex interaction networks will be reorganized under climate change represents a major challenge in the fields of ecology and biogeography. However, forecasting the potential effects of climate change on local communities, and more particularly on food‐web structure, requires the consideration of highly structuring processes, such as trophic interactions. A major breakthrough is therefore expected by combining predictive models integrating habitat selection processes, the physiological limits of marine species and their trophic interactions. In this study, we forecasted the potential impacts of climate change on the local food‐web structure of the highly threatened Gulf of Gabes ecosystem located in the south of the Mediterranean Sea. We coupled the climatic envelope and habitat models to an allometric niche food web model, hence taking into account the different processes acting at regional (climate) and local scales (habitat selection and trophic interactions). Our projections under the A2 climate change scenario showed that future food webs would be composed of smaller species with fewer links, resulting in a decrease of connectance, generality, vulnerability and mean trophic level of communities and an increase of the average path length, which may have large consequences on ecosystem functioning. The unified framework presented here, by connecting food‐web ecology, biogeography and seascape ecology, allows the exploration of spatial aspects of interspecific interactions under climate change and improves our current understanding of climate change impacts on local marine food webs.     

Resource tracking in marine parasites: going with the flow?

Understanding how diversity interacts with energy supply is of broad ecological interest. Most studies to date have investigated patterns within trophic levels, reflecting a lack of food webs which include information on energy flow. We added parasites to a published marine energy‐flow food web, to explore whether parasite diversity is correlated with energy flow to host taxa. Parasite diversity was high with 36 parasite taxa affecting 40 of the 51 animal taxa. Adding parasites increased the number of trophic links per species, trophic link strength, connectance, and food chain lengths. There was evidence of an asymptotic relationship between energy flowing through a food chain and parasite diversity, although there were clear outliers. High parasite diversity was associated with host taxa which were highly connected within the food web. This suggests that energy flow through a taxon may favour parasite diversity, up to a maximal value. The evolutionary and energetic basis for that limitation is of key interest in understanding the basis for parasite diversity in natural food webs and thus their role in food web dynamics.     

Indirect Energy Flows in Niche Model Food Webs: Effects of Size and Connectance

Indirect interactions between species have long been of interest to ecologists. One such interaction type takes place when energy or materials flow via one or more intermediate species between two species with a direct predator-prey relationship. Previous work has shown that, although each such flow is small, their great number makes them important in ecosystems. A new network analysis method, dynamic environ approximation, was used to quantify the fraction of energy flowing from prey to predator over paths of length greater than 1 (flow indirectness or FI) in a commonly studied food web model. Web structure was created using the niche model and dynamics followed the Yodzis-Innes model. The effect of food web size (10 to 40 species) and connectance (0.1 to 0.48) on FI was examined. For each of 250 model realizations run for each pair of size and connectance values, the FI of every predator-prey interaction in the model was computed and then averaged over the whole network. A classification and regression tree (CART) analysis was then used to find the best predictors of FI. The mean FI of the model food webs is 0.092, with a standard deviation of 0.0279. It tends to increase with system size but peaks at intermediate connectance levels. Of 27 potential predictor variables, only five (mean path length, dominant eigenvalue of the adjacency matrix, connectance, mean trophic level and fraction of species belonging to intermediate trophic levels) were selected by the CART algorithm as best accounting for variation in the data; mean path length and the dominant eigenvalue of the adjacency matrix were dominant.     

Food Web Topology in High Mountain Lakes

Although diversity and limnology of alpine lake systems are well studied, their food web structure and properties have rarely been addressed. Here, the topological food webs of three high mountain lakes in Central Spain were examined. We first addressed the pelagic networks of the lakes, and then we explored how food web topology changed when benthic biota was included to establish complete trophic networks. We conducted a literature search to compare our alpine lacustrine food webs and their structural metrics with those of 18 published lentic webs using a meta-analytic approach. The comparison revealed that the food webs in alpine lakes are relatively simple, in terms of structural network properties (linkage density and connectance), in comparison with lowland lakes, but no great differences were found among pelagic networks. The studied high mountain food webs were dominated by a high proportion of omnivores and species at intermediate trophic levels. Omnivores can exploit resources at multiple trophic levels, and this characteristic might reduce competition among interacting species. Accordingly, the trophic overlap, measured as trophic similarity, was very low in all three systems. Thus, these alpine networks are characterized by many omnivorous consumers with numerous prey species and few consumers with a single or few prey and with low competitive interactions among species. The present study emphasizes the ecological significance of omnivores in high mountain lakes as promoters of network stability and as central players in energy flow pathways via food partitioning and enabling energy mobility among trophic levels.     

Food web stability and weighted connectance: the complexity-stability debate revisited

How the complexity of food webs relates to stability has been a subject of many studies. Often, unweighted connectance is used to express complexity. Unweighted connectance is measured as the proportion of realized links in the network. Weighted connectance, on the other hand, takes link weights (fluxes or feeding rates) into account and captures the shape of the flux distribution. Here, we used weighted connectance to revisit the relation between complexity and stability. We used 15 real soil food webs and determined the feeding rates and the interaction strength matrices. We calculated both versions of connectance, and related these structural properties to food web stability. We also determined the skewness of both flux and interaction strength distributions with the Gini coefficient. We found no relation between unweighted connectance and food web stability, but weighted connectance was positively correlated with stability. This finding challenges the notion that complexity may constrain stability, and supports the ‘complexity begets stability’ notion. The positive correlation between weighted connectance and stability implies that the more evenly flux rates were distributed over links, the more stable the webs were. This was confirmed by the Gini coefficients of both fluxes and interaction strengths. However, the most even distributions of this dataset still were strongly skewed towards small fluxes or weak interaction strengths. Thus, incorporating these distribution with many weak links via weighted instead of unweighted food web measures can shed new light on classical theories.     

水生生态系统食物网复杂性与多样性的关系

探索食物网的复杂结构是生态学的中心问题之一。基于构建的黄河口海草床食物网并耦合实际食物网的数据集，整理了包含河口、湖泊、海洋和河流四种水生生态系统类型的48个实际食物网案例。以食物网的节点数反映食物网多样性，物种之间的营养链接数、链接密度和连通度来表示食物网的复杂性，采用营养缩尺模型描述水生生态系统食物网的复杂性特征与节点数的普适性规律。结果表明：所涉及的48个水生生态系统食物网的多样性和复杂性跨度较大，其中，节点数的分布范围为4-124，链接数为3-1830，链接密度为0.75-15.71，连通度为0.06-0.25。不同类型水生生态系统间的连通度存在显著性差异（P=0.01），节点数、链接数、链接密度不存在显著性差异。各类型生态系统的食物网链接数、链接密度均随节点数的增加而增加（R²=0.92，P<0.001和R²=0.82，P<0.001）。湖泊生态系统的连通度随节点数的变化不明显，围绕在0.20附近；而其他3种类型生态系统的食物网连通度随节点数的增加而降低（R²=0.06-0.41，P<0.001）。对全球尺度的水生食物网多样性和复杂性的定量化研究对于提升对食物网的复杂结构的科学认识，从系统尺度探究多样性和复杂性的关系提供数据支撑。     

The structure of food webs along river networks

Do changes in the species composition of riverine fish assemblages along river networks lead to predictable changes in food‐web structure? We assembled empirical “fish‐centered” river food webs for three rivers located along a latitudinal gradient in the South Saskatchewan River Basin (SSRB) that differ in land‐use impacts and geomorphology but flow through similar mountain, foothill, and prairie physiographic regions. We then calculated 17 food‐web properties to determine whether the nine river food webs differed according to physiographic region or river sub‐basin. There were no statistically significant differences in the 17 food‐web properties calculated among the rivers. In contrast, fish species richness, connectance, the proportion of herbivores, and the proportion of cannibals changed longitudinally along the river network. Our results suggest that regional changes in river geomorphology and physicochemistry play an important role in determining longitudinal variation in food‐web properties such as fish species richness and connectance. In contrast, the overall structure of river food webs may be relatively similar and insensitive to regional influences such as zoogeography. Further explorations of river and other food webs would greatly illuminate this suggestion.     

Topological properties of food webs: from real data to community assembly models

We explore patterns of trophic connections between species in the largest and highest-quality empirical food webs to date, introducing a new topological property called the link distribution frequency (i.e. degree distribution), defined as the frequency of species S _L with L links. Non-trivial differences are shown in link distribution frequencies between species-rich and species-poor communities, which might have important consequences for the responses of ecosystems to disturbances. Coarse-grained topological properties observed, as species richness-connectance and number of links-species richness relationships, provide no support for the theory of links-species scaling law or constant connectance across empirical food webs investigated. We further explore these observations by means of simulated food webs resulting from multitrophic assembly models using different functional responses between species. Species richness-connectance and links-species richness relationships of empirical food webs are reproduced by our models, but degree distributions are not properly predicted, suggesting the need of new theoretical approximations to food web assembly. The best agreement between empirical and simulated webs occurs for low values of interaction strength between species, corroborating previous empirical and theoretical findings where weak interactions govern food web dynamics.     

Connectance determines invasion success via trophic interactions in model food webs

Human induced global change has greatly altered the structure and composition of food webs through the invasion of non‐native species and the extinction of native species. Much attention has been paid to the effects of species deletions on food web structure and stability. However, recent empirical evidence suggests that for most taxa local species richness has increased as successful invasions outpace extinctions at this scale. This pattern suggests that food webs, which represent feeding interactions at the local scale, may be increasing in species richness. Knowledge of how food web structure relates to invasive species establishment and the effect of successful invaders on subsequent food web structure remains an unknown but potentially important aspect of global change. Here we explore the effect of food web topology on invasion success in model food webs to develop hypotheses about how the distribution of biodiversity across trophic levels affects the success of invasion at each trophic level. Our results suggest a connectance (C) based framework for predicting invasion success in food webs due to the way that C constrains the number of species at each trophic level and thus the number of potential predators and prey for an invader at a given trophic level. We use the relationship between C and the proportion of species at each trophic level in 14 well studied food webs to make the following predictions; 1) the success of basal invaders will increase as C increases due to the decrease in herbivores in high C webs, 2) herbivore invasion success will decrease as C increases due to the decrease in the proportion of basal species and increase in intermediate species and omnivores in high C webs. 3) Top predator invasion success will increase as C increases due to the increase in intermediate prey species. However, it is not clear how the relative influence of trophic structure compares to empirically known predictors of invasion success such as invader traits, propagule pressure, and resource availability.     

The structure of assembled communities

A set of rules is formulated which expresses the random assembly of ecological communities by sequentially arriving species, subject to energetic constraints. It is shown that these “assembled communities” provide a reasonable model for 35 out of the 40 real food webs recently compiled by Briand (1981), on the basis of the statistics: species richness, proportion of herbivores, ratio of prey to predators, proportion of dietary specialists, number of trophic links, number of potential competitive links, connectance, and average maximal food chain length. However, the observed frequency of intervality among Briand's food webs deviates significantly from the value expected on the basis of random sampling from the mathematical universe of assembled webs. Finally, there are indications in this work that the process of community genesis may be fundamentally different in fluctuating and in constant environments.     

Geographical variation in food web structure in Nepenthes pitcher plants

Abstract. 1. Relative to Nepenthes species in West Malaysia near the evolutionary centre of the genus, outlying species of Nepenthes in the Seychelles, Sri Lanka and Madagascar have fewer species of both prey and predator living in them, fewer and smaller guilds of species, much apparently empty niche space, less complex food webs, and a greater connectance. The ratios of prey to predators, and of connectance (C₁) to the total number of trophic types present remain approximately constant.
2. Differences between the food webs appear to be related in a complex way to the size of the country and its degree of spatial and temporal isolation, the size of the local species pool capable of colonizing the pitchers, and the number of Nepenthes species present. However, the maximal length of food chains in the richest and most complex food webs is probably limited by energetic constraints or environmental predictability.
3. The data may illustrate how food webs change to become more complex, both by the addition of new guilds of species and the addition of species to existing guilds, while at the same time certain properties of the food web are kept approximately constant.