Examining the potential effects of species aggregation on the network structure of food webs

One of the key measures that have been used to describe the topological properties of complex networks is the “degree distribution”, which is a measure that describes the frequency distribution of number of links per node. Food webs are complex ecological networks that describe the trophic relationships among species in a community, and the topological properties of empirical food webs, including degree distributions, have been examined previously. Previously, the “niche model” has been shown to accurately predict degree distributions of empirical food webs, however, the niche model-generated food webs were referenced against empirical food webs that had their species grouped together based on their taxonomic and/or trophic relationships (aggregated food webs). Here, we explore the effects of species aggregation on the ability of the niche model to predict the total- (sum of prey and predator links per node), in- (number of predator links per node), and out- (number of prey links per node) degree distributions of empirical food webs by examining two food webs that can be aggregated at different levels of resolution. The results showed that (1) the cumulative total- and out-degree distributions were consistent with the niche model predictions when the species were aggregated, (2) when the species were disaggregated (i.e., higher resolution), there were mixed conclusions with regards to the niche model's ability to predict total- and out-degree distributions, (3) the model's ability to predict the in-degree distributions of the two food webs was generally inadequate. Although it has been argued that universal functional form based on the niche model could describe the degree distribution patterns of empirical food webs, we believe there are some limitations to the model's ability to accurately predict the structural properties of food webs.     

Connectance and parasite diet breadth in flea-mammal webs

The number of links in webs of species interactions, which lies at the heart of the biodiversity-stability debate, has given rise to controversy during the last 20 yr. Studies exploring these web properties have mainly focused on symmetric webs where each species can potentially feed on any other species; asymmetric webs such as host-parasite webs, where one set of species feed on another set of species, have been overlooked. However, food webs are incomplete without parasites and the study of parasite-host sub-web properties deserves attention. Here, using a large database involving 33 regional interaction webs between mammals and their flea parasites, we found a negative relationship between species richness and host-parasite connectance. We suggest that some phylogenetic constraints on flea diet may explain our observed patterns because we found that parasite diet breadth, measured as host taxonomic diversity, was invariant along our host richness gradient. We found that the slope of the logarithmic relationship between the number of realized links and species richness is lower than slope values reported for food webs. We suggest that connectance may not respond to increasing species richness as rapidly in host-parasite webs as in predator-prey food webs due to stronger coevolutionary requirements.     

Constraints and variation in food web link-species space

Predicting food web structure in future climates is a pressing goal of ecology. These predictions may be impossible without a solid understanding of the factors that structure current food webs. The most fundamental aspect of food web structure—the relationship between the number of links and species—is still poorly understood. Some species interactions may be physically or physiologically ‘forbidden''—like consumption by non-consumer species—with possible consequences for food web structure. We show that accounting for these ‘forbidden interactions'' constrains the feasible link-species space, in tight agreement with empirical data. Rather than following one particular scaling relationship, food webs are distributed throughout this space according to shared biotic and abiotic features. Our study provides new insights into the long-standing question of which factors determine this fundamental aspect of food web structure.     

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.     

Systematic variation in food web body-size structure linked to external subsidies

The relationship between body mass (M) and size class abundance (N) depicts patterns of community structure and energy flow through food webs. While the general assumption is that M and N scale linearly (on log–log axes), nonlinearity is regularly observed in natural systems, and is theorized to be driven by nonlinear scaling of trophic level (TL) with M resulting in the rapid transfer of energy to consumers in certain size classes. We tested this hypothesis with data from 31 stream food webs. We predicted that allochthonous subsidies higher in the web results in nonlinear M–TL relationships and systematic abundance peaks in macroinvertebrate and fish size classes (latter containing salmonids), that exploit terrestrial plant material and terrestrial invertebrates, respectively. Indeed, both M–N and M–TL significantly deviated from linear relationships and the observed curvature in M–TL scaling was inversely related to that observed in M–N relationships. Systemic peaks in M–N, and troughs in M–TL occurred in size classes dominated by generalist invertebrates, and brown trout. Our study reveals how allochthonous resources entering high in the web systematically shape community size structure and demonstrates the relevance of a generalized metabolic scaling model for understanding patterns of energy transfer in energetically ‘open’ food webs.     

A modified niche model for generating food webs with stage‐structured consumers: The stabilizing effects of life‐history stages on complex food webs

1. Almost all organisms grow in size during their lifetime and switch diets, trophic positions, and interacting partners as they grow. Such ontogenetic development introduces life‐history stages and flows of biomass between the stages through growth and reproduction. However, current research on complex food webs rarely considers life‐history stages. The few previously proposed methods do not take full advantage of the existing food web structural models that can produce realistic food web topologies.
2. We extended the niche model developed by Williams and Martinez (Nature, 2000, 404, 180–183) to generate food webs that included trophic species with a life‐history stage structure. Our method aggregated trophic species based on niche overlap to form a life‐history structured population; therefore, it largely preserved the topological structure of food webs generated by the niche model. We applied the theory of allometric predator–prey body mass ratio and parameterized an allometric bioenergetic model augmented with biomass flow between stages via growth and reproduction to study the effects of a stage structure on the stability of food webs.
3. When life‐history stages were linked via growth and reproduction, more food webs persisted, and persisting food webs tended to retain more trophic species. Topological differences between persisting linked and unlinked food webs were small to modest. The slopes of biomass spectra were lower, and weak interaction links were more prevalent in the linked food webs than the unlinked ones, suggesting that a life‐history stage structure promotes characteristics that can enhance stability of complex food webs.
4. Our results suggest a positive relationship between the complexity and stability of complex food webs. A life‐history stage structure in food webs may play important roles in dynamics of and diversity in food webs.

     

Species richness, species range size and ecological specialisation among African primates: geographical patterns and conservation implications

The geographical distribution of species richness and species range size of African anthropoid primates (catarrhines) is investigated and related to patterns of habitat and dietary niche breadth. Catarrhine species richness is concentrated in the equatorial regions of central and west Africa; areas that are also characterised by low average species range sizes and increased ecological specificity. Species richness declines with increasing latitude north and south of the equator, while average species range size, habitat and dietary breadth increase. Relationships between species richness, species range size and niche breadth remain once latitudinal and longitudinal effects have been removed. Among areas of lowest species richness, however, there is increased variation in terms of average species range size and niche breadth, and two trends are identified. While most such areas are occupied by a few wide-ranging generalists, others are occupied by range-restricted specialist species. That conservation efforts increasingly focus on regions of high species richness may be appropriate if these regions are also characterised by species that are more restricted in both their range size and their ecological versatility, although special consideration may be required for some areas of low species richness.     

Macroecological Patterns of Climatic Niche Breadth Variation in Lacertid Lizards

Measuring climatic niche position and breadth may help to determine where species can occur over space and time. Using GIS-based and phylogenetic comparative methods, we investigated global patterns of variation in climatic niche breadth in lacertid lizards to test the following three hypotheses about climatic niche widths. First, does a species' temperature or precipitation niche breadth relate to its temperature or precipitation niche position(the mean value of annual mean temperature or annual precipitation across sampled localities in the range of each species)? Second, are there trade-offs between a species' temperature niche breadth and precipitation niche breadth? Third, does a species' temperature or precipitation niche breadth relate to altitude or latitude? We expect that:(1) species distributed in cold regions are specialized for low-temperature environments(i.e. narrow niche breadth center around low temperatures);(2) a negative relationship between species niche breadth on temperature and precipitation axes according to the tradeoff hypothesis(i.e. species that tolerate a broad range of precipitation regimes cannot also tolerate a broad range of temperatures);(3) precipitation niche breadth decreases with altitude or latitude, whereas temperature climatic niche breadth increases with altitude or latitude. Based on the analytical results we found that:(1) temperature niche breadth and position are negatively related, while precipitation niche breadth and position are positively related;(2) there is no trade-off between temperature and precipitation niche breadths; and (3) temperature niche breadth and latitude/altitude are positively related, but precipitation niche breadth and latitude/altitude are not significantly related. Our results show many similarities with previous studies on climatic niche widths reported for amphibians and lizards, which provide further evidence that such macroecological patterns of variation in climatic niche breadths may be widespread.     

The coexistence of fish species in streams: relationships between assemblage attributes and trophic and environmental variables

Diet overlap and niche breadth are well-known species traits from trophic ecology that can assist in explaining how species interact and coexist as well as the ecological mechanisms that influence biodiversity. In the present study, we analyzed the relationships between these trophic variables and indicators of resource availability with some attributes of fish assemblages (species richness, Shannon diversity index, evenness, density and individual body size). The physical and chemical characteristics of the biotopes (topography, water quality and conservation of slopes) were examined to identify possible patterns. Monthly sampling using electrofishing was conducted in 2003 along five streams located in the Cuiabá River watershed. The relationships between environmental variables and attributes of fish assemblages were evaluated using Spearman correlation. Species richness and abundance varied among streams, with higher values (54 and 82 species) found in low-gradient streams that drained small swampy areas discharging in Cuiabá River. Diet overlap showed significant and negative correlations with species richness, the diversity index, abundance, variation in body size and food availability and positive correlations with evenness and niche breadth. Environments that had greater food availability had a reduced variety of food items (smaller amplitude) and distinct values in terms of dietary overlap. Nevertheless, to explain resource partitioning in a fish assemblage (overlap), it is necessary to consider food availability and niche breadth (degree of trophic specialization) beyond the interaction of these variables with species richness and density. In conclusion, high diversity and abundance values were strongly associated with a high degree of trophic specialization (low amplitude of trophic niche) and a small degree of overlap in the diet.     

Food Web Structure in a Harsh Glacier-Fed River

Glacier retreat is occurring across the world, and associated river ecosystems are expected to respond more rapidly than those in flowing waters in other regions. The river environment directly downstream of a glacier snout is characterised by extreme low water temperature and unstable channel sediments but these habitats may become rarer with widespread glacier retreat. In these extreme environments food web dynamics have been little studied, yet they could offer opportunities to test food web theories using highly resolved food webs owing to their low taxonomic richness. This study examined the interactions of macroinvertebrate and diatom taxa in the Ödenwinkelkees river, Austrian central Alps between 2006 and 2011. The webs were characterised by low taxon richness (13–22), highly connected individuals (directed connectance up to 0.19) and short mean food chain length (2.00–2.36). The dominant macroinvertebrates were members of the Chironomidae genus Diamesa and had an omnivorous diet rich in detritus and diatoms as well as other Chironomidae. Simuliidae (typically detritivorous filterers) had a diet rich in diatoms but also showed evidence of predation on Chironomidae larvae. Food webs showed strong species-averaged and individual size structuring but mass-abundance scaling coefficients were larger than those predicted by metabolic theory, perhaps due to a combination of spatial averaging effects of patchily distributed consumers and resources, and/or consumers deriving unquantified resources from microorganisms attached to the large amounts of ingested rock fragments. Comparison of food web structural metrics with those from 62 published river webs suggest these glacier-fed river food web properties were extreme but in line with general food web scaling predictions, a finding which could prove useful to forecast the effects of anticipated future glacier retreat on the structure of aquatic food webs.     

The latitudinal gradient in niche breadth: concepts and evidence

We examine Robert MacArthur's hypothesis that niche breadth is positively associated with latitude (the latitude-niche breadth hypothesis). This idea has been influential and long standing, yet no studies have evaluated its generality or the validity of its assumptions. We review the theoretical arguments suggesting a positive relationship between niche breadth and latitude. We also use available evidence to evaluate the assumptions and predictions of MacArthur's latitude-niche breadth hypothesis. We find that neither the assumptions nor the predictions of the hypothesis are supported by data. We propose an alternative hypothesis linking latitude with niche breadth. Unlike previous ideas, our conceptual framework does not require equilibrial assumptions and is based on recently uncovered patterns of species interactions.     

Temporal and spatial changes in benthic invertebrate trophic networks along a taxonomic richness gradient

Species interactions underlie most ecosystem functions and are important for understanding ecosystem changes. Representing one type of species interaction, trophic networks were constructed from biodiversity monitoring data and known trophic links to assess how ecosystems have changed over time. The Baltic Sea is subject to many anthropogenic pressures, and low species diversity makes it an ideal candidate for determining how pressures change food webs. In this study, we used benthic monitoring data for 20 years (1980–1989 and 2010–2019) from the Swedish coast of the Baltic Sea and Skagerrak to investigate changes in benthic invertebrate trophic interactions. We constructed food webs and calculated fundamental food web metrics evaluating network horizontal and vertical diversity, as well as stability that were compared over space and time. Our results show that the west coast of Sweden (Skagerrak) suffered a reduction in benthic invertebrate biodiversity by 32% between the 1980s and 2010s, and that the number of links, generality of predators, and vulnerability of prey have been significantly reduced. The other basins (Bothnian Sea, Baltic Proper, and Bornholm Basin) do not show any significant changes in species richness or consistent significant trends in any food web metrics investigated, demonstrating resilience at a lower species diversity. The decreased complexity of the Skagerrak food webs indicates vulnerability to further perturbations and pressures should be limited as much as possible to ensure continued ecosystem functions.     

Climate change impacts in multispecies systems: drought alters food web size structure in a field experiment

Experimental data from intergenerational field manipulations of entire food webs are scarce, yet such approaches are essential for gauging impacts of environmental change in natural systems. We imposed 2 years of intermittent drought on stream channels in a replicated field trial, to measure food web responses to simulated climate change. Drought triggered widespread losses of species and links, with larger taxa and those that were rare for their size, many of which were predatory, being especially vulnerable. Many network properties, including size–scaling relationships within food chains, changed in response to drought. Other properties, such as connectance, were unaffected. These findings highlight the need for detailed experimental data from different organizational levels, from pairwise links to the entire food web. The loss of not only large species, but also those that were rare for their size, provides a newly refined way to gauge likely impacts that may be applied more generally to other systems and/or impacts.     

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.     

Aquatic food‐web structure along a salinized dryland river

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Species richness and allometric scaling jointly determine biomass in model aquatic food webs

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Pyramids of species richness: the determinants and distribution of species diversity across trophic levels

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Variation in niche parameters along the diversity gradient of unicellular eukaryote assemblages

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