The structure of a plant-pollinator food web

The pollination biology literature is dominated by examples of specialization between plants and their pollinators. However, a recent review shows that it is generalization that prevails in the field, with most plants having a number of pollinators and most pollinators visiting a number of plants. Consequently, the vast majority of plant–pollinator interactions are embedded in a complex web of plant–pollinator interactions. These plant-pollinator webs can be studied in the manner of conventional food webs and the aim of this paper is to illustrate how contemporary methods of web construction and analysis can be applied to plant-pollinator communities.     

Temperature alters food web body-size structure

The increased temperature associated with climate change may have important effects on body size and predator–prey interactions. The consequences of these effects for food web structure are unclear because the relationships between temperature and aspects of food web structure such as predator–prey body-size relationships are unknown. Here, we use the largest reported dataset for marine predator–prey interactions to assess how temperature affects predator–prey body-size relationships among different habitats ranging from the tropics to the poles. We found that prey size selection depends on predator body size, temperature and the interaction between the two. Our results indicate that (i) predator–prey body-size ratios decrease with predator size at below-average temperatures and increase with predator size at above-average temperatures, and (ii) that the effect of temperature on predator–prey body-size structure will be stronger at small and large body sizes and relatively weak at intermediate sizes. This systematic interaction may help to simplify forecasting the potentially complex consequences of warming on interaction strengths and food web stability.     

The effects of stochastic population dynamics on food web structure

We develop a stochastic, individual-based model for food web simulation which in the large-population limit reduces to the well-studied Webworld model, which has been used to successfully construct model food webs with several realistic features. We demonstrate that an almost exact match is found between the population dynamics in fixed food webs, and that the demographic fluctuations have systematic effects when the new model is used to construct food webs due to the presence of species with small populations.     

土壤食物网:结构、能流及稳定性

在国外4个土壤生态学项目的基础上,结合其他学者的研究成果,从连通网、能流网及功能网3个层次分别阐述了土壤食物网的结构、能流与稳定性。在连通网中,主要描述土壤食物网的功能群、营养位及格局;在能流网中,主要描述了面向过程食物网模型及其应用,主要涉及到土壤中的碳流与氮流;在功能网中,以作用强度为中心,描述了土壤食物网的稳定性问题。     

Integrating food web diversity, structure and stability

Given the unprecedented rate of species extinctions facing the planet, understanding the causes and consequences of species diversity in ecosystems is of paramount importance. Ecologists have investigated both the influence of environmental variables on species diversity and the influence of species diversity on ecosystem function and stability. These investigations have largely been carried out without taking into account the overarching stabilizing structures of food webs that arise from evolutionary and successional processes and that are maintained through species interactions. Here, we argue that the same large-scale structures that have been purported to convey stability to food webs can also help to understand both the distribution of species diversity in nature and the relationship between species diversity and food web stability. Specifically, the allocation of species diversity to slow energy channels within food webs results in the skewed distribution of interactions strengths that has been shown to confer stability to complex food webs. We end by discussing the processes that might generate and maintain the structured, stable and diverse food webs observed in nature.     

Species traits as drivers of food web structure

The use of functional traits to describe community structure is a promising approach to reveal generalities across organisms and ecosystems. Plant ecologists have demonstrated the importance of traits in explaining community structure, competitive interactions as well as ecosystem functioning. The application of trait‐based methods to more complex communities such as food webs is however more challenging owing to the diversity of animal characteristics and of interactions. The objective of this study was to determine how functional structure is related to food web structure. We consider that food web structure is the result of 1) the match between consumer and resource traits, which determine the occurence of a trophic interaction between them, and 2) the distribution of functional traits in the community. We implemented a statistical approach to assess whether or not 35 466 pairwise interactions between soil organisms are constrained by trait‐matching and then used a Procrustes analysis to investigate correlations between functional indices and network properties across 48 sites. We found that the occurrence of trophic interactions is well predicted by matching the traits of the resource with those of the consumer. Taxonomy and body mass of both species were the most important traits for the determination of an interaction. As a consequence, functional evenness and the variance of certain traits in the community were correlated to trophic complementarity between species, while trait identity, more than diversity, was related to network topology. The analysis was however limited by trait data availability, and a coarse resolution of certain taxonomic groups in our dataset. These limitations explain the importance of taxonomy, as well as the complexity of the statistical model needed. Our results outline the important implications of trait composition on ecological networks, opening promising avenues of research into the relationship between functional diversity and ecosystem functioning in multi‐trophic systems.     

Relevance of evolutionary history for food web structure

Explaining the structure of ecosystems is one of the great challenges of ecology. Simple models for food web structure aim at disentangling the complexity of ecological interaction networks and detect the main forces that are responsible for their shape. Trophic interactions are influenced by species traits, which in turn are largely determined by evolutionary history. Closely related species are more likely to share similar traits, such as body size, feeding mode and habitat preference than distant ones. Here, we present a theoretical framework for analysing whether evolutionary history--represented by taxonomic classification--provides valuable information on food web structure. In doing so, we measure which taxonomic ranks better explain species interactions. Our analysis is based on partitioning of the species into taxonomic units. For each partition, we compute the likelihood that a probabilistic model for food web structure reproduces the data using this information. We find that taxonomic partitions produce significantly higher likelihoods than expected at random. Marginal likelihoods (Bayes factors) are used to perform model selection among taxonomic ranks. We show that food webs are best explained by the coarser taxonomic ranks (kingdom to class). Our methods provide a way to explicitly include evolutionary history in models for food web structure.     

Genetic variation, predator-prey interactions and food web structure

Food webs are networks of species that feed on each other. The role that within-population phenotypic and genetic variation plays in food web structure is largely unknown. Here, I show via simulation how variation in two key traits, growth rates and phenology, by influencing the variability of body sizes present through time, can potentially affect several structural parameters in the direction of enhancing food web persistence: increased connectance, decreased interaction strengths, increased variation among interaction strengths and increased degree of omnivory. I discuss other relevant traits whose variation could affect the structure of food webs, such as morphological and additional life-history traits, as well as animal personalities. Furthermore, trait variation could also contribute to the stability of food web modules through metacommunity dynamics. I propose future research to help establish a link between within-population variation and food web structure. If appropriately established, such a link could have important consequences for biological conservation, as it would imply that preserving (functional) genetic variation within populations could ensure the preservation of entire communities.     

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.     

Simultaneously estimating food web connectance and structure with uncertainty

1. Food web models explain and predict the trophic interactions in a food web, and they can infer missing interactions among the organisms. The allometric diet breadth model (ADBM) is a food web model based on the foraging theory. In the ADBM, the foraging parameters are allometrically scaled to body sizes of predators and prey. In Petchey et al. (Proceedings of the National Academy of Sciences, 2008; 105: 4191), the parameterization of the ADBM had two limitations: (a) the model parameters were point estimates and (b) food web connectance was not estimated.
2. The novelty of our current approach is: (a) We consider multiple predictions from the ADBM by parameterizing it with approximate Bayesian computation, to estimate parameter distributions and not point estimates. (b) Connectance emerges from the parameterization, by measuring model fit using the true skill statistic, which takes into account prediction of both the presences and absences of links.
3. We fit the ADBM using approximate Bayesian computation to 12 observed food webs from a wide variety of ecosystems. Estimated connectance was consistently greater than previously found. In some of the food webs, considerable variation in estimated parameter distributions occurred and resulted in considerable variation (i.e., uncertainty) in predicted food web structure.
4. These results lend weight to the possibility that the observed food web data is missing some trophic links that do actually occur. It also seems likely that the ADBM likely predicts some links that do not exist. The latter could be addressed by accounting in the ADBM for additional traits other than body size. Further work could also address the significance of uncertainty in parameter estimates for predicted food web responses to environmental change.

     

The human complement component C8B gene: structure and phylogenetic relationship

The eighth component of human complement (C8) is a serum protein that consists of three chains (, and ), encoded by three separate genes, viz., C8A, C8B, and C8G. In serum, the -subunit is non-covalently bound to the disulfide-linked - subunit. Using a full-length C8 cDNA probe, we isolated several clones from human genomic DNA libraries. Four clones covering the complete cDNA sequence were characterized by TaqI restriction mapping and were shotgun subcloned into M13. C8-cDNA-positive clones were partially sequenced to characterize the 12 exons of the gene with sizes from 69 to 347 bp. All intron-exon junctions followed the GT-AG rule. By using polymerase chain reaction (PCR) primers located in the adjacent intron sequences, all 12 exons of the C8B gene could be amplified from genomic DNA. All fragments showed the expected sizes. The sizes of eight introns could be determined by using primer pairs that amplified two exons and the enclosed intron, and by restriction mapping. These analyses and the insert sizes of the genomic clones indicate that the C8B gene has a total size of approximately 40 kb. The polymorphic TaqI site of the C8B gene localized in intron 11 could be demonstrated by direct restriction fragment analysis of a PCR fragment containing exons 11 and 12, and the enclosed intron 11. Homology comparison of the C8B gene with C8A and C9 on the basis of the exon structure confirmed the ancestral relationship known from the protein level.     

Habitat subdivision causes changes in food web structure

Theory suggests that the response of communities to habitat subdivision depends on both species' characteristics and the extent to which species interact. For species with dynamics that are independent of other species, subdivision is expected to promote regional extinction as populations become small and isolated. By contrast, intermediate levels of subdivision can facilitate persistence of strongly interacting species. Consistent with this prediction, experimental subdivision lengthened persistence of some species, altering the extent of food web collapse through extinction. Extended persistence was associated with immigration rescuing a basal prey species from local extinction. As predicted by food web theory, habitat subdivision reduced population density of a top predator. Removal of this top predator from undivided microcosms increased the abundance of two other predator species, and these changes paralleled those produced by habitat subdivision. These results show that species interactions structured this community, and illustrate the need for investigations of other communities.     

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.     

Predation and food web structure along a habitat duration gradient

Food web statistics showed a complex relationship with measures of habitat variability in temporary ponds. Connectance was highest in short-duration, highly variable habitats, and lowest in habitats of intermediate duration and variability. The number of links and links/taxon increased with increasing duration. Much of the variation in the food web statistics could be explained by a strong linear relationship between number of taxa and number of links/taxon and a quadratic relationship of taxa number with the number of links. However, after accounting for this variation, there remained a relationship of duration with links and links/taxon. The relationship between the food web statistics and duration corresponded to experimental evaluations of predation in these habitats that showed an increasing importance of predation in long-duration habitats. The food web statistics, however, missed threshold effects in the relationship between predation and habitat duration. Differences in food web statistics before and after a regional drought could be explained by a decrease in taxa number after the drought. Connectance was the most robust statistic in relation to taxa number, but was also the least sensitive to changes in habitat characteristics. Received: 11 March 1996 / Accepted: 8 January 1997     

The predatory Chironomidae of an iron-rich stream: feeding ecology and food web structure

Abstract. 1. Three species of Tanypodinae (Chironomidae) were found in an acid and iron-rich stream in southern England. Maximum abundance was achieved in summer and they were sparse at other times. Individuals were aggregated on the stream bed and were overrepresented in accumulations of leaf litter.
2. The diets of all three species consisted of a mixture of prey (prominently detritivorous chironomid larvae) and detritus. More detritus and fewer prey were taken in winter than in summer.
3. When comparing large tanypod species with small and, intraspecifically, late instars with early, the proportion of guts containing prey increased with increasing body size.
4. Stonefly larvae were more prominent in the diet of Zavrelimyia barbatipes (Kieffer) in summer than in winter but for the other two species the reverse was true. A bigger proportion of Trissopelopia longimana (Staeger) guts contained prey in early summer than in August whereas more Macropelopia goetghebueri (Kieffer) guts contained prey in August. This was apparently a consequence of seasonal differences in the distribution of body size among the populations of these two species.
5. The stream contains two further common predators, Plectrocnemia conspersa (Curtis) and Sialis fuliginosa Pict. These are important predators of tanypod larvae but might also compete with them since they severely deplete populations of prey taken in common.
6. Analysis of the food-web in Broadstone Stream reveals remarkably high values of connectance (C and C_max) and of species richness times connectance (SC_max). Such characteristics are theoretically associated with fragile and dynamically unstable food webs, and may be found in 'constant' environments. There is also an apparently unusual prevalence of omnivory in the community.     

Habitat size influences food web structure in drying streams

Biodiversity in running waters is threatened by an increased severity and incidence of low‐flow extremes resulting from global climate change and a growing human demand for freshwater resources. Although it is unknown how and to what extent riverine communities will change in the face of these threats, considerable insight will be gained from efforts aimed at quantifying habitat size‐related controls on the trophic relationships among taxa in streams experiencing extreme flow loss. Here we report on a detailed space‐for‐time survey of replicate stream food webs sampled along the perennial‐ to‐drying continuum in each of fourteen different intermittent South Island, New Zealand streams. We quantified several structural attributes of food webs at fifty‐eight sites, including two taxonomically‐based metrics (web size, predator:prey ratio) and three stable isotope‐based metrics (food chain length [FCL], trophic area, δ¹³C range); we also quantified habitat size‐, disturbance‐, and resource‐related covariates at each site. Food web structure varied widely across sample sites within and across study streams and much of this variation was explained by habitat size. Consistent with our predictions, we found that food webs became smaller (ca 30 to ca 15 taxa, ca 20‐fold reduction in stable isotope‐based trophic area) and shorter (maximum trophic position [FCL] from 4.1 to 2.0, 25% reduction in predator:prey ratio) as we moved from the largest to smaller habitats. These results, and a comparison of our findings with those from a similar assessment conducted in perennial streams, suggest that there are perturbation thresholds which may trigger food web collapse when exceeded, and further imply that food webs may ultimately be ‘sized’ to minimum flows rather than average flow conditions. Our work provides a basis for making general predictions about how habitat contraction, and flow loss in particular, may affect communities and additionally provides insight on mechanisms warranting further attention.     

Effects of spatial scale of sampling on food web structure

This study asks whether the spatial scale of sampling alters structural properties of food webs and whether any differences are attributable to changes in species richness and connectance with scale. Understanding how different aspects of sampling effort affect ecological network structure is important for both fundamental ecological knowledge and the application of network analysis in conservation and management. Using a highly resolved food web for the marine intertidal ecosystem of the Sanak Archipelago in the Eastern Aleutian Islands, Alaska, we assess how commonly studied properties of network structure differ for 281 versions of the food web sampled at five levels of spatial scale representing six orders of magnitude in area spread across the archipelago. Species (S) and link (L) richness both increased by approximately one order of magnitude across the five spatial scales. Links per species (L/S) more than doubled, while connectance (C) decreased by approximately two‐thirds. Fourteen commonly studied properties of network structure varied systematically with spatial scale of sampling, some increasing and others decreasing. While ecological network properties varied systematically with sampling extent, analyses using the niche model and a power‐law scaling relationship indicate that for many properties, this apparent sensitivity is attributable to the increasing S and decreasing C of webs with increasing spatial scale. As long as effects of S and C are accounted for, areal sampling bias does not have a special impact on our understanding of many aspects of network structure. However, attention does need be paid to some properties such as the fraction of species in loops, which increases more than expected with greater spatial scales of sampling.     

基于稳定同位素的湿地食物源判定和食物网构建研究进展

湿地生物营养动力学是湿地生态系统结构和功能评价研究的基础.碳、氮稳定同位素作为识别营养关系的方法,已在湿地生态系统食物来源、组成和食物链传递研究中得到广泛运用.本文系统综述了稳定同位素食物贡献度计算模型和营养级确定的基本方法和理论;讨论了动物营养分馏值和基线的选择依据;概括了湿地生态系统典型食物源及其稳定同位素变化特征;总结了草食、杂食和肉食等不同营养级动物的食物来源.指出了稳定同位素在湿地食物源溯源和食物网研究中的不足;基于国内外研究现状和发展趋势及需求,展望了未来同位素技术在湿地食物网生态学研究中的运用前景和研究重点,提出需要加强稳定同位素营养分馏和基线的影响因素、样品处理和保存方式研究以及胃含物、分子标记物和多元素同位素结合分析.     

Stability of pitcher-plant microfaunal populations depends on food web structure

Enrichment (increasing K) destabilizes simple consumer–resource interactions, but increasing food web complexity in various ways can remove this paradox of enrichment. We varied resources and number of omnivorous predators (mosquitoes) and tested for effects on the stability (persistence and temporal variability) of microfaunal populations living in pitcher plants. Top-down (omnivorous) effects were destabilizing, decreasing the persistence time of a rotifer, Habrotrocha rosa, and perhaps a microflagellate, Bodo sp. Enrichment effects were more complex, in part due to effects of shredding midges on resource availability, and in part due to interactions with predation. The persistence of Bodo increased with resource availability (more bacteria due to shredding by midges; no paradox of enrichment). Increasing resources by adding ants decreased persistence of H. rosa when mosquitoes were rare (paradox of enrichment), but the effect was reversed in leaves with significant colonization by mosquitoes. Thus, in the microfaunal community of pitcher plants, omnivorous predation tends to be destabilizing, and also tends to remove the paradox of enrichment.