Food Web Variability Research Articles

Diversity, food web structure and the temporal stability of total plant and animal biomasses

Food‐web complexity and species richness are predicted to have tremendous effects on ecosystem functioning stability. Yet, our understanding of the relation between diversity and the stability of ecosystem functioning is still mainly limited to single trophic level communities and highly simplified food webs. To start filling this knowledge gap, we model allometric food webs and use structural equation modelling (SEM) to investigate the relations between diversity, food web structure and the temporal variability of total plant and animal biomasses and their components, i.e. synchrony and mean population variability. We find that stability responds differently depending on the metric used. When looking at total biomass variability, higher total biomasses and biomass stored at higher trophic level have the strongest stabilising effects, while species richness has lower impact than previously found. We confirm that synchrony among species is important for the stability of the total plant biomass. However, synchrony only weakly explains variations in the stability of total animal biomass among food webs. Species richness and food web variables often have opposite impacts on synchrony and mean population variability, leading to more ambiguous results for the plant community as synchrony matters only for plant stability. Our approach thus provides new insight on the complexity of the stability of ecosystem functioning in complex food webs.

RNA Sequencing of Lake Charr Epidermal Mucus to Assess Molecular Effects of Diluted Bitumen Exposure in a Boreal Lake

Transport of diluted bitumen (dilbit) from Canada’s oil sands region poses risk for leaks and spills of petroleum-derived contaminants into the environment. Exposure of fish to dilbit is known to cause cardiotoxicity, developmental deformities, and impairment in swim performance. However, previous studies have examined the toxicity of dilbit in laboratory settings which does not account for environmental and biological food-web variables that may alter exposure and/or toxicity of dilbit. Moreover, most methods of assessing organism health following oil exposure require lethal sampling. This work is a part of a larger set of experiments where dilbit spills were simulated within enclosures on a lake; the present study assesses the impacts of residual levels of dilbit that may have entered the surrounding lake environment from the enclosures following model spill cleanup. In order to understand the impacts of residual dilbit in an ecosystem setting without use of lethal sampling, epidermal mucus was collected and sequenced from lake charr (Salvelinus namaycush) exposed to residual dilbit in a boreal lake. While concentrations reached a maximum of 2.29 μg/L total polycyclic aromatic compounds (ΣPAC) within surface waters, surface water ΣPAC concentrations generally remained below 1 μg/L. Results of RNA sequencing were compared to sequencing data from mucus collected prior to dilbit additions. Differential gene expression and pathway analyses indicated dysregulation of genes associated with intermediary and energy metabolism as well as a trend in upregulation of cyp1a3 in epidermal mucus following dilbit exposure. Thus, results of the present study suggest that lake charr undergo consistent biological responses after exposure to residual levels of dilbit following a model spill, and that mRNA-based analysis of mucus may be a viable method for non-lethal oil exposure assessment. Overall, the results provide insight on the response of wild fish to very dilute dilbit exposures after a model spill cleanup.

Spatio-temporal patterns in stable isotope composition of a benthic intertidal food web reveal limited influence from salt marsh vegetation and green tide

Assessing fluxes of matter and energy in food webs within and across benthic habitats is important to understand the ecological functioning in bays and estuaries, where the productivity is favoured by a wide diversity of primary producers. The temporal variability (March vs September 2019) in the carbon and nitrogen stable isotope composition of primary food sources and benthic invertebrates consumers was investigated in a large intertidal area (Western English-Channel, France). The study area is influenced by megatidal conditions and characterised by salt marshes in the sheltered part, and seasonal Ulva spp. blooms. The spatio-temporal variability in the structure of the benthic food web was analysed at the scales of both the whole bay and the different assemblages, which constitute the mosaic of habitats. Inferences on potential sources fuelling the food web were supported by spatio-temporal patterns based on covariations and stable isotope trajectory analysis. Results highlighted that phytoplankton, microphytobenthos and SOM were, most likely, the main food sources. The trophic connectivity between salt marsh and benthic habitats within the bay was limited to some macrofauna species inhabiting muddy creeks within the salt marsh. Unexpectedly, the influence of Ulva spp. blooms appeared also limited. Spatial patterns illustrates the constancy of the spatial variability in the benthic pelagic coupling, with a higher influence of microphytobenthos in the upper shore compared to low shore assemblages. This first attempt to characterize intertidal benthic food web constitutes a relevant baseline for the conservation of the bay of Saint-Brieuc where a national Nature Reserve has been created in 1998 for the conservation of overwintering birds. The spatial and temporal patterns of the benthic food web observed in this study (1) confirm the importance to consider food web variability at spatial and temporal scales from sampling designs to data analysis, and (2) demonstrate the ability of the stable isotope trajectory analysis framework to highlight food web dynamics.

Resolving the predator first paradox: Arthropod predator food webs in pioneer sites of glacier forelands.

Primary succession on bare ground surrounded by intact ecosystems is, during its first stages, characterized by predator‐dominated arthropod communities. However, little is known on what prey sustains these predators at the start of succession and which factors drive the structure of these food webs. As prey availability can be extremely patchy and episodic in pioneer stages, trophic networks might be highly variable. Moreover, the importance of allochthonous versus autochthonous food sources for these pioneer predators is mostly unknown. To answer these questions, the gut content of 1,832 arthropod predators, including four species of carabid beetles, two lycosid and several linyphiid spider species caught in early and late pioneer stages of three glacier forelands, was screened molecularly to track intraguild and extraguild trophic interactions among all major prey groups occurring in these systems. Two‐thirds of the 2,310 identified food detections were collembolans and intraguild prey, while one‐third were allochthonous flying insects. Predator identity and not successional stage or valley had by far the strongest impact on the trophic interaction patterns. Still, the variability of prey spectra increased significantly from early to late pioneer stage, as did the niche width of the predators. As such the structure of pioneer arthropod food webs in recently deglaciated Alpine habitats seems to be driven foremost by predator identity while site and early successional effects contribute to a lesser extent to food web variability. Our findings also suggest that in these pioneer sites, predatory arthropods depend less on allochthonous aeolian prey but are mainly sustained by prey of local production.

Using a qualitative model to explore the impacts of ecosystem and anthropogenic drivers upon declining marine survival in Pacific salmon

SUMMARYCoho salmon (Oncorhynchus kisutch), Chinook salmon (Oncorhynchus tshawytscha) and steelhead (Oncorhynchus mykiss) in Puget Sound and the Strait of Georgia have exhibited declines in marine survival over the last 40 years. While the cause of these declines is unknown, multiple factors, acting cumulatively or synergistically, have likely contributed. To evaluate the potential contribution of a broad suite of drivers on salmon survival, we used qualitative network modelling (QNM). QNM is a conceptually based tool that uses networks with specified relationships between the variables. In a simulation framework, linkages are weighted and then the models are subjected to user-specified perturbations. Our network had 33 variables, including: environmental and oceanographic drivers (e.g., temperature and precipitation), primary production variables, food web components from zooplankton to predators and anthropogenic impacts (e.g., habitat loss and hatcheries). We included salmon traits (survival, abundance, residence time, fitness and size) as response variables. We invoked perturbations to each node and to suites of drivers and evaluated the responses of these variables. The model showed that anthropogenic impacts resulted in the strongest negative responses in salmon survival and abundance. Additionally, feedbacks through the food web were strong, beginning with primary production, suggesting that several food web variables may be important in mediating effects on salmon survival within the system. With this model, we were able to compare the relative influence of multiple drivers on salmon survival.

Multiple Patterns of Food Web Dynamics Revealed by a Minimal Non-deterministic Model

Understanding and predicting patterns arising from the dynamics of marine food webs is central to trophic and community ecology and numerical models of food webs constitute a primary tool to simulate these dynamics. Food web simulation models are often highly complex while at the same time often too constrained to reproduce the level of variability observed in real systems. The recently developed non-deterministic network dynamics (NDND) modelling framework has been suggested as a simulation alternative, which can generate multiple patterns of food web variability despite great structural simplicity. Two important aspects of the NDND modelling framework remain unexplored: first the derivation of model input parameters from empirical or theoretical studies and second the evaluation of the model simulations against observations. We provide a methodology for the derivation of model parameters based on empirical observations, the metabolic theory of ecology and life-history theory and apply it to the specific case of the Barents Sea food web. We then evaluate the ability of the NDND simulations to reproduce a wide range of patterns of food web dynamics against observations collected in the Barents Sea during 28 years. Patterns emerging from the simulations include trends and cycles in biomass, trophic levels and transfer efficiency, density-dependent growth, top-down vs bottom-up oscillations, ecosystem level stability and synchrony and trophic functional responses. The ability of the NDND to generate so many patterns observed empirically in the Barents Sea is remarkable given that it is based only on random trophic interactions operating within few constraints set by ecological rules. Our results show that investigations of food web dynamics in marine ecosystems, including the definition of reference states and responses to climate and exploitation pressures, may be achieved with models that are structurally simple and based on few well-established assumptions.

Microscale variations of food web functioning within a rocky shore invertebrate community

The assessment of relevant spatial scales at which ecological processes occur is of special importance for a thorough understanding of ecosystem functioning. In coastal ecosystems, the variability of trophic interactions has been studied at different spatial scales, but never at scales from centimetres to metres. In the present study, we investigated the link between habitat structure and small-scale variability of food web functioning on intertidal boulder field ecosystems. Two microhabitats, boulder-top and boulder-bottom, were considered, and the trophic ecology of invertebrate consumers was studied using stable isotope tracers. We found for two of the main suspension feeders of northern Atlantic rocky shores (the sponges Halichondria panicea and Hymeniacidon sanguinea) consistent 15N enrichment for individuals sampled under boulders, suggesting that these consumers relied on different trophic resource according to the microhabitat inhabited, at a centimetre scale. The high δ15N signatures found underneath boulders suggested higher use of highly decomposed organic matter in this microhabitat. The isotopic difference between the two microhabitats decreased in higher trophic level consumers, which likely foraged at a spatial scale including both microhabitats. Finally, our results reveal that in highly heterogeneous habitats such as boulder fields, trophic interactions are likely to vary strongly in space, which should be considered in future researches. The link between habitat physical structure and food web variability might also contribute to the high biological diversity characterizing heterogeneous ecosystems.

Temporal variations in organic carbon utilization by consumers in a lowland river

ABSTRACTUnderstanding temporal trophic interactions in riverine food webs is essential for predicting river ecosystem function and improving management of these ecosystems. Temporal changes in energy flows through riverine food webs are readily assumed but are rarely tested. Temporal variability in food webs from two reaches of a lowland river (Ovens River, south‐eastern Australia) with differing levels of floodplain connectivity were examined over 12 months. We investigated how seasonal changes, flow variability and floodplain connectivity influence (i) stable isotope signatures of basal organic carbon sources (terrestrial sources: trees and grasses; aquatic sources: seston, biofilm and filamentous algae) and consumers (macroinvertebrates and fish) and (ii) the relative proportions of organic carbon sources contributing to consumer biomass using mixing models. We hypothesized (i) that during high flows, increased floodplain connectivity would increase the lateral exchange of terrestrial carbon subsidies to main channel consumers and (ii) that during low flow periods, main channel consumers would derive the majority of their carbon from aquatic benthic sources. Results indicated that isotope signatures for basal sources and for most of the consumers varied temporally and spatially. Mixing models indicated that increased floodplain connectivity did not increase terrestrial subsidies to consumers during high flows. Seston was the primary source during high flows whilst terrestrial vegetation increased in importance during low flows. Filamentous algae was also important during low flows for some consumers. These findings indicate that it is essential to include temporal variability in order to understand energy flows in lowland rivers, thus allowing for the dynamic nature of these ecosystems. Copyright © 2010 John Wiley & Sons, Ltd.

Structural changes in lake functioning induced from nutrient loading and climate variability

Climate variability is increasingly recognized as an important regulatory factor, capable of influencing the structural properties of aquatic ecosystems. Lakes appear to be particularly sensitive to the ecological impacts of climate variability, and several long time series have shown a close coupling between climate, lake thermal properties and individual organism physiology, population abundance, community structure, and food web dynamics. Thus, understanding the complex interplay among meteorological forcing, hydrological variability, and ecosystem functioning is essential for improving the credibility of model-based water resources/fisheries management. Our objective herein is to examine the relative importance of the ecological mechanisms underlying plankton seasonal variability in Lake Washington, Washington State (USA), over a 35-year period (1964–1998). Our analysis is founded upon an intermediate complexity plankton model that is used to reproduce the limiting nutrient (phosphate)–phytoplankton–zooplankton–detritus (particulate phosphorus) dynamics in the lake. Model parameterization is based on a Bayesian calibration scheme that offers insights into the degree of information the data contain about model inputs and allows obtaining predictions along with uncertainty bounds for modeled output variables. The model accurately reproduces the key seasonal planktonic patterns in Lake Washington and provides realistic estimates of predictive uncertainty for water quality variables of environmental management interest. A principal component analysis of the annual estimates of the underlying ecological processes highlighted the significant role of the phosphorus recycling stemming from the zooplankton excretion on the planktonic food web variability. We also identified a moderately significant signature of the local climatic conditions (air temperature) on phytoplankton growth ( r = 0.41), herbivorous grazing ( r = 0.38), and detritus mineralization ( r = 0.39). Our study seeks linkages with the conceptual food web model proposed by Hampton et al. [Hampton, S.E., Scheuerell, M.D., Schindler, D.E., 2006b. Coalescence in the Lake Washington story: interaction strengths in a planktonic food web. Limnol. Oceanogr. 51, 2042–2051.] to emphasize the “bottom-up” control of the Lake Washington plankton phenology. The posterior predictive distributions of the plankton model are also used to assess the exceedance frequency and confidence of compliance with total phosphorus (15 μg L −1) and chlorophyll a (4 μg L −1) threshold levels during the summer-stratified period in Lake Washington. Finally, we conclude by underscoring the importance of explicitly acknowledging the uncertainty in ecological forecasts to the management of freshwater ecosystems under a changing global environment.

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.

Soil fauna–microbe interactions: towards a conceptual framework for research

We explore the potential for applying broad ecological theories to interactions between soil animals and micro-organisms to generate a predictive framework within which more hypothesis led research can be undertaken. The paper stems from discussions during a workshop at the XIVth International Symposium on Soil Zoology. The possible linkage between biodiversity and ecosystem functions forms a good example of how soil zoology research can be productively stimulated by addressing a broader ecological concept but also how the concept can be tested below ground at fundamentally different scales to those commonly used above ground. Other areas of theory rapidly developing above ground, which are yet to be fully tested below ground, include: spatial variability in food webs; indirect interactions mediated through changes in plant secondary chemistry; signalling, including tritrophic interactions; optimal foraging theory, including depletion theory when patches differ in quality as well as quantity; adaptive plasticity in life history traits in relation to temporal variability in resources; trade-offs and facultative non-symbiotic and symbiotic mutualism. We identify modelling of effects of climate change on the soil compartment of the global carbon cycle as an area in which understanding of soil fauna–microbe interaction may outstrip current ecological theory and as a major challenge facing soil biologists in the future.

Testing predictions of the lake food web theory on pelagic communities of Australian reservoirs

Several predictions of the theory developed for pelagic food webs of the Northern Hemisphere were tested on water bodies of Eastern Australia. Eleven reservoirs, representing trophic and latitudinal gradients were sampled for nutrients, phytoplankton, zooplankton and pelagic fish. Two models of regression analysis, which analysed possible interactions between trophic levels were based on different sets of data. In one, each reservoir was represented by only one pair of observations – annual mean or single observation (“regional model”). In the other, seasonal means of four frequently sampled reservoirs similar in productivity were used (“temporal model”). Significant variation in total phytoplankton biovolume (TPB) was predicted by total phosphorus concentration (TP), total nitrogen concentration (TN), mean crustacean length and acoustic biomass of planktivorous fish in both models. This suggested that nutrient limitation, zooplankton grazing and positive effects of fish were probably important in controlling the biomass of primary producers at both regional and temporal scales. In the regional model, the biomass of fish was also negatively correlated with Daphnia biomass and mean crustacean length, suggesting that the trophic cascade hypothesis may be applicable to Eastern Australia for the considered range of reservoir productivities.The biovolume of cyanobacteria was not correlated to any variables tested in the regional model. In contrast, nutrient and food web variables had significant effects on cyanobacterial biovolume in the temporal model. This suggested that factors governing seasonal succession were probably more important for cyanobacteria than variation in reservoir productivity or location.Contrary to previous views, no negative relationship between total biomass of zooplankton and TPB was found in both models, suggesting that the community structure of zooplankton rather than its total biomass mediates top‐down effects. Many predictions of the food web theory remained robust in spite of substantial differences in animal taxonomy and physical environment of Australian ecosystems.

Spatial and temporal variability of food web structure during the spring bloom in the Skagerrak

During the intense spring diatom bloom, the spatial (km) and temporal (h) variability of the pelagic food web structure was investigated in the Skagerrak. Three surface water masses could be identified in the study area based on their temperature, salinity and biogeochemical characteris- tics. When the measured biological parameters were separated into water masses instead of stations, food web variability decreased by up to 41%. However, specific egg production rate by Acartia clausi and Oithona similis did not correlate with water masses and was probably influenced by other fac- tors, e.g. small-scale turbulence. Bacterial production and Calanus-grazing followed a diurnal cycle with increased activity during the night. The grazing impact by the copepod community was assessed by the egg production and the gut fluorescence method. Despite the Calanus population, the cope- pod community only grazed <3% of daily primary production. The protozooplankton, however, ingested 2 to 4 times more of primary production than the copepods. Overall, the zooplankton com- munity only grazed 17% of daily primary production, and consequently the majority of the spring bloom will leave the euphotic zone ungrazed as a potential food source for the benthos.

Temporal food web variability in an upper Missouri River backwater: energy origination points and transfer mechanisms

Abstract – Energy transfer mechanisms often cannot be identified with either an isotope or a food habits analysis; however, a combined assessment provided collaborative support for depicting backwater food webs. Lateral dimensions in riverine function substantially influence backwater ecology; however, backwater communities are highly dynamic and infrequently investigated due to logistical difficulties. We created seasonal food webs for an upper Missouri River backwater using a combined food habits and stable isotope analysis. Temporal transition of energy origins was apparent. In the spring, carbon primarily originated from sediments, detritus and periphyton. During mid‐summer, carbon was derived from a variety of sources, but included large proportions of terrestrial plants and particulate organic matter. By early fall, the backwater was uncoupled from the river, and autochthonous phytoplankton appeared to be the dominant carbon source. During all sample periods, Chironomidae was an important trophic link conveying energy from the primary producers to the higher consumers. Additionally, zooplankton were an important energy transfer taxa to the abundant age‐0 fish community in the fall, and Corixidae were important in the spring.

Food web structure variability in the surface layer, at a fixed station influenced by the North Western Mediterranean Current

The structure of the planktonic community and the influence of mesozooplankton migration on the microbial food web were investigated during six diel studies from June 92 to June 93 in the surface waters of a station in the North-Western Mediterranean Sea. Each diel study consisted of sampling at 5 and 40 m every 3 h over 24 h. Most of the times diel cycles did not show any convincing diel patterns in any of the variables studied. Clear zooplankton migration was evident in only two diel studies. The ratio of heterotrophic/autotrophic biomasses varied from 0.68 to 3.0, with a strong dominance of the heterotrophic biomass under oligotrophic conditions. Differences in food web structure were probably related to the influences of coastal water and the North-Western Mediterranean Current. Thus we found that the planktonic food web variability relatable to hydrodynamic variability, to be greater than diel variability. However, very large differences in food web structure among dates were evident. For example proportion of Chl a found in the <10 µm fraction varied from 18 to 96%.