Mean Food Chain Length Research Articles

Colonization–competition dynamics of basal species shape food web complexity in island metacommunities

Exploring how food web complexity emerges and evolves in island ecosystems remains a major challenge in ecology. Food webs assembled from multiple islands are commonly recognized as highly complex trophic networks that are dynamic in both space and time. In the context of global climate change, it remains unclear whether food web complexity will decrease in a monotonic fashion when undergoing habitat destruction (e.g., the inundation of islands due to sea-level rise). Here, we develop a simple yet comprehensive patch-dynamic framework for complex food web metacommunities subject to the competition-colonization tradeoff between basal species. We found that oscillations in food web topological complexity (characterized by species diversity, mean food chain length and the degree of omnivory) emerge along the habitat destruction gradient. This outcome is robust to changing parameters or relaxing the assumption of a strict competitive hierarchy. Having oscillations in food web complexity indicates that small habitat changes could have disproportionate negative effects on species diversity, thus the success of conservation actions should be evaluated not only on changes in biodiversity, but also on system robustness to habitat alteration. Overall, this study provides a parsimonious mechanistic explanation for the emergence of food web complexity in island ecosystems, further enriching our understanding of metacommunity assembly.

Spatial Heterogeneity of Food Webs in A River-Lake Ecotone under Flow Regulation – A Case Study in Northern China

The river–lake ecotone supports diverse aquatic life, but its food web structure and topology are poorly understood. Baiyangdian Lake, northern China’s largest shallow lake, depends on external environmental flows, of which the Fu River provides the most stable water supply. Here, we used stable isotopes and topological analysis to explore the food web structure along a spatial gradient using data from field surveys from 2018 to 2019. Carbon and nitrogen stable isotopes and the food web structure were associated with environmental factors for four ecosystem types (river, river mouth, lake mouth, lake). Detritus, phytoplankton, and zooplankton δ13C values became more depleted along the gradient from the river to the lake, whereas δ13C of submerged macrophytes was enriched in the ecotones compared to the river and lake. Higher δ15N values for basal resources and zooplankton occurred at the lake mouth and river mouth. The top consumers were omnivorous fish: Hemiculter leucisculus (trophic level [TL] = 3.85 ± 0.89) in the river and Pseudorasbora parva (TL = 4.54 ± 0.58) in the river mouth. Carnivorous Erythroculter dabryi occupied the highest TL (3.61 ± 0.36 and 4.46 ± 0.36, respectively) in the lake mouth and lake. These results together led to a change from a detritus-based to phytoplankton-based food web along the gradient from the river to the lake. The species richness, number of trophic links, link density, and mean food chain length all are greatest in the lake, followed by the lake mouth, and the lowest were in the river. Our results provide a holistic view of the ecotone ecosystem and its food web, suggesting that it supports a more diverse species assemblage and more complex food web structure than the adjacent river ecosystem, rather than the adjacent lake ecosystem. Therefore, management should emphasize the combined effects of altered hydrological regimes and poor water quality on the ecotone food webs to manage the river and lake more sustainably.

Human settlements in headwater catchments are associated with generalist stream food webs

While a variety of anthropogenic impacts on lotic biodiversity have been documented, food-web responses to catchment development are poorly understood. We selected 27 stream food webs of comparable quality and conducted an analysis to assess the effect of catchment development on food-web structure. We quantified population densities, built-up area, and proximity to urban centres, and calculated a Settlement Index for each catchment using Principal Components Analysis. We also calculated the percentage of agricultural land cover in each catchment. Next, we assessed the correlations between the structural properties of food-webs (species richness, connectance, mean food-chain length, linkage density, trophic generality, and trophic vulnerability) and the Settlement Index as well as agricultural land cover. We found that linkage density, trophic vulnerability (number of consumers per species), and trophic generality (number of resources per species) were higher in streams with greater Settlement Index, indicating a reduction in specialisation. However, no clear trends were observed for species richness, connectance, and mean food-chain length. Agricultural land cover was also not related to food-web structure. We propose that the reduction in specialisation may be driven by species turnover and feeding plasticity, as biotic invasion or species impoverishment was not evident.

Implications of scaled δ15N fractionation for community predator-prey body mass ratio estimates in size-structured food webs.

Nitrogen stable isotope ratios (δ(15) N) may be used to estimate community-level relationships between trophic level (TL) and body size in size-structured food webs and hence the mean predator to prey body mass ratio (PPMR). In turn, PPMR is used to estimate mean food chain length, trophic transfer efficiency and rates of change in abundance with body mass (usually reported as slopes of size spectra) and to calibrate and validate food web models. When estimating TL, researchers had assumed that fractionation of δ(15) N (Δδ(15) N) did not change with TL. However, a recent meta-analysis indicated that this assumption was not as well supported by data as the assumption that Δδ(15) N scales negatively with the δ(15) N of prey. We collated existing fish community δ(15) N-body size data for the Northeast Atlantic and tropical Western Arabian Sea with new data from the Northeast Pacific. These data were used to estimate TL-body mass relationships and PPMR under constant and scaled Δδ(15) N assumptions, and to assess how the scaled Δδ(15) N assumption affects our understanding of the structure of these food webs. Adoption of the scaled Δδ(15) N approach markedly reduces the previously reported differences in TL at body mass among fish communities from different regions. With scaled Δδ(15) N, TL-body mass relationships became more positive and PPMR fell. Results implied that realized prey size in these size-structured fish communities are less variable than previously assumed and food chains potentially longer. The adoption of generic PPMR estimates for calibration and validation of size-based fish community models is better supported than hitherto assumed, but predicted slopes of community size spectra are more sensitive to a given change or error in realized PPMR when PPMR is small.

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.

Multiple anthropogenic stressors and the structural properties of food webs

Coastal environments are among the most productive on the planet, providing a wide range of ecosystem services. Development and exploitation mean that they are faced with stresses from a number of anthropogenic sources. Such stresses are typically studied in isolation, but multiple stressors can combine in unexpected ways to alter the structure of ecological systems. Here, we experimentally explore the impacts of inorganic nutrients and organic matter on a range of food web properties. We find that these two stressors combine additively to produce significant increases in connectance and mean food chain length. Such increases are typically associated with enhanced robustness to secondary extinctions and productivity, respectively. Despite these apparent beneficial effects, we find a simplification of web structure in terms of taxon richness and diversity, and altered proportions of basal and top species. These effects are driven by a reduction in community assembly and lower consistency in a range of system properties as a result of the multiple stressors. Consequently, impacted food webs are likely to be more vulnerable to human- or climate-induced perturbations in the long-term.

Algivory in food webs of three temperate Andean rivers

AbstractFood web analyses have been fundamental in understanding community organization and ecosystem functioning. To date, a number of studies demonstrate that stream food webs depend to a large extent on allochthonous detritus, but there are more recent studies that show a high degree of autochthony. Our food‐web study was carried out in three Andean rivers (Coilaco, Guampoe and Trancura) within the catchment area of Toltén River in southern Chile. Based on the analyses of 4251 invertebrate gut contents, we found that these Andean stream food webs are dominated by herbivores (range: 50–73% of all species) supported by a species‐rich algal (basal) component, and characterized by a low proportion of omnivores (range: 8–27% of all species) and predatory species (range: 10–24%). Significant differences in the number of feeding links of the herbivores Meridialis diguillina and Antarctoperla michaelseni and the omnivore Smicridea chilensis were found between seasons. The spring herbivore Aubertoperla sp. showed significant differences between rivers. S. chilensis fed on 50 different prey items as compared with the herbivores whose maximum number of links ranged between 37 and 40. Web sizes ranged between 93 and 131 species and the proportion of top species was distinctly lower than those of basal (up to 0.651 in Coilaco River) and intermediate species. Direct connectance (links per species2) values were low and similar among rivers (range: 0.051–0.074), whereas mean food chain length ranged between 2.23 and 2.90. The distributions of web property values from the Andean rivers differed from those previously published. In contrast to previous predictions, mean food chain length in these Andean streams displayed a scale‐invariant pattern across different web sizes, but it was significantly related to the proportion of intermediate species.

Food web complexity and allometric scaling relationships in stream mesocosms: implications for experimentation

1. Mesocosms are used extensively by ecologists to gain a mechanistic understanding of ecosystems based on the often untested assumption that these systems can replicate the key attributes of natural assemblages. 2. Previous investigations of stream mesocosm utility have explored community composition, but here for the first time, we extend the approach to consider the replicability and realism of food webs in four outdoor channels (4 m(2)). 3. The four food webs were similarly complex, consisting of diverse assemblages (61-71 taxa) with dense feeding interactions (directed connectance 0.09-0.11). Mesocosm food web structural attributes were within the range reported for 82 well-characterized food webs from natural streams and rivers. When compared with 112 additional food webs from standing freshwater, marine, estuarine and terrestrial environments, stream food webs (including mesocosms) had similar characteristic path lengths, but typically lower mean food chain length and exponents for the species-link relationship. 4. Body size (M) abundance (N) allometric scaling coefficients for trivariate taxonomic mesocosm food webs (-0.53 to -0.49) and individual size distributions (-0.60 to -0.58) were consistent and similar to those from natural systems, suggesting that patterns of energy flux between mesocosm consumers and resources were realistic approximations. 5. These results suggest that stream mesocosms of this scale can support replicate food webs with a degree of biocomplexity that is comparable to 'natural' streams. The findings highlight the potential value of mesocosms as model systems for performing experimental manipulations to test ecological theories, at spatiotemporal scales of relevance to natural ecosystems.

Primary Consumer Body Size and Food-Chain Length in Terrestrial Communities

Using 21 community food webs, we tested Elton's two hypotheses regarding the main factors limiting food-chain length in terrestrial communities, namely, energy (energy limitation hypothesis—ELH) and body size (size limitation hypothesis—SLH). As predators tend to be larger than their prey, food-chains are size-structured: animal size increases with trophic position. We found a negative correlation between the size of the primary consumer and the length of the chain. Food-chains based on small primary consumers are longer than those based on large primary consumers, and size rather than energetic efficiency is the main contributing factor. We found no correlation between habitat productivity and mean food-chain length. All these findings support the SLH over the ELH. Our results suggest that, as in aquatic communities, a single factor—a predator/prey size-ratio greater than 1—governs the structure of terrestrial communities.

Global patterns of aquatic food chain length

Food chain length is a fundamental ecosystem property, and plays a central role in determining ecosystem functioning. Recent advances in the field of stable isotope ecology allow the estimation of food chain length (FCL) from stable nitrogen isotope (δ15N) data. We conducted a global literature synthesis and estimated FCL for 219 lake, stream, and marine ecosystems. Streams had shorter food chains (∼3.5 trophic levels) than marine and lake ecosystems (∼4.0 trophic levels). In marine systems, inclusion of marine mammals increased FCL by 2/3 of a trophic level. For each ecosystem type, estimates of FCL were normally distributed and spanned two full trophic levels. Comparison with published connectance food webs revealed similar mean FCL values, though stable isotope‐derived FCL estimates were less variable. At the global scale, FCL showed weak or no relationships with ecosystem size, mean annual air temperature, or latitude. Our study highlights the utility of stable isotopes for quantifying among‐system food web variability, and the application of this approach for assessing global‐scale patterns of food chain length.

Global patterns of aquatic food chain length

Food chain length is a fundamental ecosystem property, and plays a central role in determining ecosystem functioning. Recent advances in the field of stable isotope ecology allow the estimation of food chain length (FCL) from stable nitrogen isotope (δ15N) data. We conducted a global literature synthesis and estimated FCL for 219 lake, stream, and marine ecosystems. Streams had shorter food chains (∼3.5 trophic levels) than marine and lake ecosystems (∼4.0 trophic levels). In marine systems, inclusion of marine mammals increased FCL by 2/3 of a trophic level. For each ecosystem type, estimates of FCL were normally distributed and spanned two full trophic levels. Comparison with published connectance food webs revealed similar mean FCL values, though stable isotope-derived FCL estimates were less variable. At the global scale, FCL showed weak or no relationships with ecosystem size, mean annual air temperature, or latitude. Our study highlights the utility of stable isotopes for quantifying among-system food web variability, and the application of this approach for assessing global-scale patterns of food chain length.

Food web structure and function in two arctic streams with contrasting disturbance regimes

Summary1. We studied the effect of substratum movement on the communities of adjacent mountain and spring tributaries of the Ivishak River in arctic Alaska (69°1′N, 147°43′W). We expected the mountain stream to have significant bed movement during summer because of storm flows and the spring stream to have negligible bed movement because of constant discharge.2. We predicted that the mountain stream would be inhabited only by taxa able to cope with frequent bed movement. Therefore, we anticipated that the mountain stream would have lower macroinvertebrate species richness and biomass and a food web with fewer trophic levels and lower connectance than the spring stream.3. Substrata marked in situ indicated that 57–66% of the bed moved during summer in the mountain stream and 4–20% moved in the spring stream.4. Macroinvertebrate taxon richness was greater in the spring (25 taxa) than in the mountain stream (20 taxa). Mean macroinvertebrate biomass was also greater in the spring (4617 mg dry mass m−2) than in the mountain stream (635 mg dry mass m−2). Predators contributed 25% to this biomass in the spring stream, but only 7% in the mountain stream.5. Bryophyte biomass was >1000 times greater in the spring stream (88.4 g ash‐free dry mass m−2) than the mountain stream (0.08 g ash‐free dry mass m−2). We attributed this to differences in substratum stability between streams. The difference in extent of bryophyte cover between streams probably explains the high macroinvertebrate biomass in the spring stream.6. Mean food‐web connectance was similar between streams, ranging from 0.18 in the spring stream to 0.20 in the mountain stream. Mean food chain length was 3.04 in the spring stream and 1.83 in the mountain stream. Dolly Varden char (Salvelinus malma) was the top predator in the mountain stream and the American dipper (Cinclus mexicanus) was the top predator in the spring stream. The difference in mean food chain length between streams was due largely to the presence of C. mexicanus at the spring stream.7. Structural differences between the food webs of the spring and mountain streams were relatively minor. The difference in the proportion of macroinvertebrate biomass contributing to different trophic levels was major, however, indicating significant differences in the volume of material and energy flow between food‐web nodes (i.e. food web function).

Food web dynamics of irrigated rice fields at five elevations in Luzon, Philippines

AbstractThe above-water food webs of arthropod communities in irrigated rice fields on Luzon Island, Philippines, were studied over the growing season at five sites (Los Baños, Cabanatuan, Bayombong, Kiangan, Banaue) ranging in elevation from 22 m to 1524 m. Arthropod populations were vacuum-sampled at roughly weekly intervals from the date after seedlings were transplanted to flowering at each site. Site- and time-specific webs were constructed from a 687-taxa cumulative Philippines web and time-series of species present. Taxonomic composition, food web structure, and arthropod phenology were broadly similar across different sites. Arthropod abundance was inversely associated with altitude across the five sites, but numbers of taxa and links and six food web statistics showed no obvious increasing or decreasing trend with altitude. The rise of taxa, links and mean food chain length over the growing season at each site reflected an increase in plant size with age and, at some sites, an orderly accumulation of newly arriving herbivore, predator, parasitoid and omnivore species. At each site, herbivores built up faster than predators and parasitoids, and predators arrived faster than parasitoids; the difference between the latest and earliest sampling dates of first arrivals, averaged over the five sites, was 38, 63 and 73 days for herbivores, predators and parasitoids, respectively. Site-to-site consistencies in food web properties and first arrivals suggest that such patterns may be influenced more by crop age than by geography or altitude. Sampled predator, parasitoid and omnivore taxa potentially encountered only a subset of their lifetime prey and predator species at any particular time in the rice field. Prey lists cumulated over time may underestimate the temporal specificity of predation by potential biological control agents. Research opportunities linking rice food webs and integrated pest management with East Indies biogeography are proposed.

Spatial and Temporal Variation in the Structure of an Intermittent‐Stream Food Web

Food webs from the Lerderderg River, an intermittent stream in Victoria, Australia, were compiled with the aim of examining changes in food web structure in a highly variable habitat. Emphasis was placed on a high degree of taxonomic precision. Spatial and temporal variation in the food webs was assessed by partitioning the study area into three sites, located ≈1.5 km apart along the river. Sites differed in overall stream width and the length of the low streamflow period during summer. Three separate webs for each site were compiled for four different times of the year. Relatively little spatial variation in community structure was observed. In contrast, temporal variation was considerable, with species composition and the number of species in the community changing considerably over the year. The number of species increased dramatically as the period of constant streamflow lengthened. Detritivores dominated the community, both in terms of species and individual numbers. The proportion of predators in the community increased slightly by the end of the year, suggesting that recolonization of the community by predators lags behind that of detritivores. The increase in the number of predator species also resulted in an increase in the mean food chain length through the year. Patterns observed in the food webs tended to fall within the range of values reported from several previous studies, suggesting that underlying constraints may structure certain aspects of food webs. However, the constancy of certain food web statistics was attributable either to methodological decisions made during compilation, or to an inherent property of the statistic itself. The potential sensitivity of several food web statistics to the methodology used to compile a food web render between—web comparisons difficult due to the confounding effects of methodology. This suggests that comparisons between food webs should be restricted to webs derived from similar habitats using a comparable methodology.