Cascading effects from predator removal depend on resource availability in a benthic food web

We tested joint effects of predator loss and increased resource availability on the grazers’ trophic level and the propagation of trophic interactions in a benthic food web by excluding larger predatory fish from cages and manipulating nutrients in the coastal zone of the Baltic Sea. The combination of nutrient enrichment and excluding larger predators induced an increase in medium-sized predatory fish (three-spined stickleback). The meso-predator fish in turn did not change the total abundance of the invertebrate herbivores, but did cause a substantial shift in their community composition towards the dominance of gastropods by reducing amphipods by 40–60%, while gastropods were left unchanged. The shift in grazer composition generated a 23 times higher producer biomass, but only under nutrient enrichment. Our results show that top-predator declines can substantially shift the species composition at the grazers’ level, but that cascading effects on producers by a trophic cascade strongly depend on resource availability.     

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.     

A description of a highly diverse soft-bottom community in Kingston Harbour,Jamaica

A highly diverse soft-bottom community in Kingston Harbour, Jamaica is described to provide additional information on the structure of tropical benthic communities. It is shown that the community has a level of diversity unique to stable tropical environments, that dominance by a few species is greatly reduced,and that there are no true parallels to this community in similar environments. The community described seems to fit Sanders' stability-time hypothesis. The density and standing crop of the community are low, and possible explanations for this are discussed. Continuous year-round breeding, short life-spans, and fast turnover rates are suggested as the most important factors regulating biomass in the community described.     

Effects of an unusual natural temperature increase on a baltic soft-bottom community

The effects of a sudden, extraordinary temperature rise upon a soft-bottom community in the northern Baltic Sea were studied continuously for more than a month, during the summer of 1968. The mean bundance of the total macrofauna on the muddy bottom amounted to ca 6×10³ individuals · m^-2, on the sandy bottom to 2×10⁴ individuals· m^-2. The meiofauna, only sampled in the mud, averaged 1×10⁶ individuals · m^-2, with nematodes, oligochaetes and ostracods as dominating groups. During this period, the redoxpotential-discontinuity (RPD)-layer successively moved upwards in the muddy bottom, being relatively stable in the sand. The mud surface showed a succession of microbial communities. The initially oxidized, yellow surface film, with patches of diatoms and red Spirula, changed to a more grayish layer, with purple areas here interpreted as Chromatium and white spots of Beggiatoa. The latter dominated during the last part of the period, covering vast areas of the bottom like a gigantic cobweb. The endofauna was forced upwards by the emerging RPD-layer, the more sensitive components dying off, facultative anaerobes and nematodes flourishing. The described temperature increase of about 10 C° simulates the effects of a 3000 MW nuclear plant.     

Biomagnification of trace elements in a benthic food web: the case study of Deer Island (Northern Yellow Sea)

In this study, we investigated the concentrations of ten trace elements (Cu, Zn, Mn, Se, Ni, Cd, Cr, Pb, Hg, and As) and their trophodynamics in a benthic food chain of Deer Island, Northern Yellow Sea. The concentrations of Cu, Zn, Mn, Se, Ni, Cd, Cr, Pb, Hg, and As in the food chain ranged from 3.2 to 23.2, from 71 to 227, from 7.4 to 45.6, from 0.44 to 5.80, from 0.73 to 7.60, from 0.14 to 1.65, from 0.68 to 6.70, from 0.08 to 1.86, from 0.08 to 1.18, and from 0.24 to 3.92 mg kg^?1 dry weight, respectively. Among these trace elements, the linear regression between the log-transformed concentrations of Hg and Cd and δ¹⁵N values showed statistically significant increase (p<0.05) with the slopes of 0.134 and 0.144, indicating biomagnification of Hg and Cd occurred in the benthic food chain of Deer Island. While the linear regression for other eight trace elements (Cu, Zn, Mn, Se, Ni, Cr, Pb and As) were characterised by extensive scatter with non-significant correlation coefficients (R ²=0.002–0.235) and slopes (p=0.079–0.875), indicating there were not biomagnified or biodiluted of these trace elements.     

An inverse model for the analysis of the Venice lagoon food web

A steady-state model of the Venice lagoon food web was constructed, based on a comprehensive set of data, which were collected in the years 2001-2005. Energy flows were estimated by means of an inverse methodology of constrained optimization based on the Minimum Norm criterion, i.e. on the minimization of both the sum of squares of the residuals and of the sum of squares of energy flows. The solution was constrained by a set inequalities, which were derived from general eco-physiological knowledge and site specific data on energy flows. The trophic network was represented by thirty-two nodes, including single-species compartments for the species of high economical or ecological relevance. Mass balance equations were weighted, in order to obtain meaningful results in presence of large differences, up to 5 orders of magnitude, among biomasses. A perturbation technique was applied, with the purpose of reducing the risk of finding solutions heavily affected by the set of constraints and of obtaining a more robust representation of the energy flows. The main patterns of energy flow are consistent with those obtained in previous attempts at modelling the Venice lagoon food web. Micro- and macro-phytobenthos account for the largest fraction of the primary production. Energy is then transferred towards higher trophic levels by means of two main pathways: the recycling of dead biomass through the detritus compartment and the direct consumption by grazers. The first pathway is the most important and accounts for approximately two/thirds of the energy transferred to the second trophic level.     

Response of a soft-bottom harpacticoid community to stingray (Dasyatis sabina) disturbance

The recolonization by harpacticoid copepods of patches disturbed by stingrays [Dasyatis sabina (LeSeur)] feeding in a subtidal (2–3 m depth) site off the Florida gulf coast (Lat. 29°54.55N, Long. 84°31.45W) was followed. Samples from disturbed patches could no longer be distinguished from control samples in terms of numbers of harpacticoids 29 h after disturbance. No species became significantly disproportionately abundant during the recovery suggesting that small-scale natural disturbances are not an important organizing force in this community.     

Adaptations in a hierarchical food web of southeastern Lake Michigan

Two issues in ecological network theory are: (1) how to construct an ecological network model and (2) how do entire networks (as opposed to individual species) adapt to changing conditions? We present a novel method for constructing an ecological network model for the food web of southeastern Lake Michigan (USA) and we identify changes in key system properties that are large relative to their uncertainty as this ecological network adapts from one time point to a second time point in response to multiple perturbations. To construct our food web for southeastern Lake Michigan, we followed the list of seven recommendations outlined in Cohen et al. [Cohen, J.E., et al., 1993. Improving food webs. Ecology 74, 252–258] for improving food webs. We explored two inter-related extensions of hierarchical system theory with our food web; the first one was that subsystems react to perturbations independently in the short-term and the second one was that a system's properties change at a slower rate than its subsystems’ properties. We used Shannon's equations to provide quantitative versions of the basic food web properties: number of prey, number of predators, number of feeding links, and connectance (or density). We then compared these properties between the two time-periods by developing distributions of each property for each time period that took uncertainty about the property into account. We compared these distributions, and concluded that non-overlapping distributions indicated changes in these properties that were large relative to their uncertainty. Two subsystems were identified within our food web system structure (p < 0.001). One subsystem had more non-overlapping distributions in food web properties between Time 1 and Time 2 than the other subsystem. The overall system had all overlapping distributions in food web properties between Time 1 and Time 2. These results supported both extensions of hierarchical systems theory. Interestingly, the subsystem with more non-overlapping distributions in food web properties was the subsystem that contained primarily benthic taxa, contrary to expectations that the identified major perturbations (lower phosphorous inputs and invasive species) would more greatly affect the subsystem containing primarily pelagic taxa. Future food-web research should employ rigorous statistical analysis and incorporate uncertainty in food web properties for a better understanding of how ecological networks adapt.     

Environmental grain and polychaete species' diversity in a bathyal benthic community

Patterns of polychaete species' dispersion in the San Diego Trough, Southern California Continental Borderland, North Pacific Ocean, at a depth of 1230 m are analyzed. Samples consist of 4 replicate 0.25-m² box cores, each partitioned into twenty-five 0.01-m² subcores, and of 1 unpartitioned core. The sampler is biased but, when this bias is taken into account, few species show strong aggregation either between or within cores. If all species are considered together as replicates, intraspecific aggregation is detected between cores, but uniform dispersion dominates within cores. Patchiness or habitat partitioning on the assayed scales can thus account for little of the extremely high species' diversity observed. Apparently due to an environmental structure having a “grain” smaller than 0.01-m², the diversity of species assumed to be sedentary is higher than the diversity of species assumed to be mobile. One evidence of such structure is a negative covariance between paraonid abundance and density of “mudballs” constructed by a cirratulid. Consideration of these results suggests that either grain specialization or simultaneous, biogenic disequilibrium or both act on spatial scales smaller than 0.01-m² to maintain high polychaete species' diversity in the San Diego Trough.     

Quantitative approaches to the analysis of stable isotope food web data

Ecologists use stable isotopes (delta13C, delta15N) to better understand food webs and explore trophic interactions in ecosystems. Traditionally, delta13C vs. delta15N bi-plots have been used to describe food web structure for a single time period or ecosystem. Comparisons of food webs across time and space are increasing, but development of statistical approaches for testing hypotheses regarding food web change has lagged behind. Here we present statistical methodologies for quantitatively comparing stable isotope food web data. We demonstrate the utility of circular statistics and hypothesis tests for quantifying directional food web differences using two case studies: an arthropod salt marsh community across a habitat gradient and a freshwater fish community from Lake Tahoe, USA, over a 120-year time period. We calculated magnitude and mean angle of change (theta) for each species in food web space using mean delta13C and delta15N of each species as the x, y coordinates. In the coastal salt marsh, arthropod consumers exhibited a significant shift toward dependence on Spartina, progressing from a habitat invaded by Phragmites to a restored Spartina habitat. In Lake Tahoe, we found that all species from the freshwater fish community shifted in the same direction in food web space toward more pelagic-based production with the introduction of nonnative Mysis relicta and onset of cultural eutrophication. Using circular statistics to quantitatively analyze stable isotope food web data, we were able to gain significant insight into patterns and changes in food web structure that were not evident from qualitative comparisons. As more ecologists incorporate a food web perspective into ecosystem analysis, these statistical tools can provide a basis for quantifying directional food web differences from standard isotope data.     

Hydrology and grazing jointly control a large-river food web

Inputs of fresh water and grazing both can control aquatic food webs, but little is known about the relative strengths of and interactions between these controls. We use long-term data on the food web of the freshwater Hudson River estuary to investigate the importance of, and interactions between, inputs of fresh water and grazing by the invasive zebra mussel (Dreissena polymorpha). Both freshwater inputs and zebra mussel grazing have strong, pervasive effects on the Hudson River food web. High flow tended to reduce population size in most parts of the food web. High grazing also reduced populations in the planktonic food web, but increased populations in the littoral food web, probably as a result of increases in water clarity. The influences of flow and zebra mussel grazing were roughly equal (i.e., within a factor of 2) for many variables over the period of our study. Zebra mussel grazing made phytoplankton less sensitive to freshwater inputs, but water clarity and the littoral food web more sensitive to freshwater inputs, showing that interactions between these two controlling factors can be strong and varied.     

Population studies on benthic nematodes within a subtropical seagrass community

Examination of the benthic nematode fauna of the soft surface sediments of a turtle grass (Thalassia testudinum König) bed in Biscayne Bay (Miami, Florida) has revealed a high degree of homogeneity exemplified by the dominance of four species out of approximately 100 nematode taxa from the area. The dominant species, Metoncholaimus scissus, Theristus fistulatus, Spirinia parasitifera and Gomphionema typica, regularly comprised between 87 and 95% of the total number of nematodes present in samples collected during the winter and spring of 1966. T. fistulatus showed an abundance of 56% over the course of the study, i.e., 160 samples collected over a period of 14 months. Maximal peaks in population densities were noted and correlated with physiographic alterations in the environment. The M. scissus population declined concurrent with changes in the community; at the same time, with accumulation of sediment, the Terschellingia longicaudata population increased. Ratios of species, and especially shifts in the dominantforms present, with repeated collections, are extremely useful indicators of important biological and physical changes in a particular environment. Analysis of distributional data on dominant species in 64 samples from eight closely approximated positions showed that observed temporal and spatial variations were not significant statistically at the 5% level. It is concluded that erroneous observations can be made from ecological studies based on field data derived without proper replication or consideration of seasonal factors. The latter as well as intrinsic variability within the particular locality itself contribute to the basic faunistic composition of benthic communities.This work was supported by Grant 12482 from the National Institute of Health to the Institute of Marine Science, University of Miami, and is a contribution (No. 819) from the Institute of Marine Science and from the Nematology Section, Entomology Research Institute, Research Branch, Canada Department of Agriculture, Ottawa.     

Importance of predation by infaunal polychaetes in controlling the structure of a soft-bottom community in Maine,USA

Field manipulations in a Maine, USA intertidal mud flat showed that Nereis virens adults were an important factor in regulating the abundance of Corophium volutator, an amphipod which comprised 63% of the total number of individuals in the benthic community. Removal of N. virens adults resulted in an increase in C. volutator abundance, while addition of N. virens adults led to a decrease. There was a significant negative correlation between the abundance of N. virens adults and that of C. volutator. C. volutator was also negatively correlated with the density of all the other infaunal species combined. These results suggest that there is at least a 3-level interactive system (N. virens —C. volutator — other infaunal species) within the infauna in Maine; N. virens is a factor in controlling the abundance of C. volutator, and C. volutator may play a role in regulating the densities of other infaunal species. Models of generalized cropping in soft-bottom systems may be too simple if they ignore complex trophic relationships within the infauna.     

Temperature-dependent ranges of coexistence in a model of a two-prey-one-predator microbial food web

The objective of our study was to analyze the effects of temperature on the population dynamics of a three-species food web consisting of two prey bacteria (Pedobacter sp. and Acinetobacter johnsonii) and a protozoan predator (Tetrahymena pyriformis) as model organisms. We assessed the effects of temperature on the growth rates of all three species with the objective of developing a model with four differential equations based on the experimental data. The following hypotheses were tested at a theoretical level: Firstly, temperature changes can affect the dynamic behavior of a system by temperature-dependent parameters and interactions and secondly, food web response to temperature cannot be derived from the single species temperature response. The main outcome of the study is that temperature changes affect the parameter range where coexistence is possible within all three species. This has significant consequences on our ideas regarding the evaluation of effects of global warming.     

Nutrient enrichment reduces constraints on material flows in a detritus-based food web

Most aquatic and terrestrial ecosystems are experiencing increased nutrient availability, which is affecting their structure and function. By altering community composition and productivity of consumers, enrichment can indirectly cause changes in the pathways and magnitude of material flows in food webs. These changes, in turn, have major consequences for material storage and cycling in the ecosystem. Understanding mechanisms and predicting consequences of nutrient-induced changes in material flows requires a quantitative food web approach that combines information on consumer energetics and consumer-resource stoichiometry. We examined effects of a whole-system experimental nutrient enrichment on the trophic basis of production and the magnitude and pathways of carbon (C), nitrogen (N), and phosphorus (P) flows in a detritus-based stream food web. We compared the response of the treated stream to an adjacent reference stream throughout the study. Dietary composition and elemental flows varied considerably among invertebrate functional feeding groups. During nutrient enrichment, increased flows of leaf litter and amorphous detritus to shredders and gatherers accounted for most of the altered flows of C from basal resources to consumers. Nutrient enrichment had little effect on patterns of material flows but had large positive effects on the magnitude of C, N, and P flows to consumers (mean increase of 97% for all elements). Nutrient-specific food webs revealed similar flows of N and P to multiple functional groups despite an order of magnitude difference among groups in consumption of C. Secondary production was more strongly related to consumption of nutrients than C, and increased material flows were positively related to the degree of consumer-resource C:P and C:N imbalances. Nutrient enrichment resulted in an increased proportion of detrital C inputs consumed by primary consumers (from -15% to 35%) and a decreased proportion of invertebrate prey consumed by predators (from -80% to 55%). Our results demonstrate that nutrient enrichment of detritus-based systems may reduce stoichiometric constraints on material flows, increase the contribution of consumers to C, N, and P cycling, alter the proportion of C inputs metabolized by consumers, and potentially lead to reduced ecosystem-level storage of C.     

Evidence supporting the importance of terrestrial carbon in a large-river food web

Algal carbon has been increasingly recognized as the primary carbon source supporting large-river food webs; however, many of the studies that support this contention have focused on lotic main channels during low-flow periods. The flow variability and habitat-heterogeneity characteristic of these systems has the potential to significantly influence food web structure and must be integrated into models of large-river webs. We used stable-isotope analysis and IsoSource software to model terrestrial and algal sources of organic carbon supporting consumer taxa in the main channel and oxbow lakes of the Brazos River, Texas, USA, during a period of frequent hydrologic connectivity between these habitat types. Standardized sampling was conducted monthly to collect production sources and consumer species used in isotopic analysis. Predictability of hydrologic connections between habitat types was based on the previous 30 years of flow data. IsoSource mixing models identified terrestrial C3 macrophytes (riparian origin) as the primary carbon source supporting virtually all consumers in the main channel and most consumers in oxbow lakes. Small-bodied consumers (<100 mm) in oxbow lakes assimilated large fractions of algal carbon whereas this pattern was not apparent in the main channel. Estimates of detritivore trophic positions based on delta15N values indicated that terrestrial material was likely assimilated via invertebrates rather than directly from detritus. High flows in the river channel influenced algal standing stock, and differences in the importance of terrestrial and algal production sources among consumers in channel vs. oxbow habitats were associated with patterns of flooding. The importance of terrestrial material contradicts the findings of recent studies of large-river food webs that have emphasized the importance of algal carbon and indicates that there can be significant spatial, temporal, and taxonomic variation in carbon sources supporting consumers in large rivers.     

Direct and indirect effects of ants on a forest-floor food web

Interactions among predators that prey on each other and are potential competitors for shared prey (intraguild [IG] predators) are widespread in terrestrial ecosystems and have the potential to strongly influence the dynamics of terrestrial food webs. Ants and spiders are abundant and ubiquitous terrestrial IG predators, yet the strength and consequences of interactions between them are largely unknown. In the leaf-litter food web of a deciduous forest in Kentucky (USA), we tested the direct and indirect effects of ants on spiders and a category of shared prey (Collembola) by experimentally subsidizing ants in open plots in two field experiments. In the first experiment, ant activity was increased, and the density of ants in the litter was doubled, by placing carbohydrate and protein baits in the center of each plot. Gnaphosa spiders were almost twice as abundant and Schizocosa spiders were half as abundant in baited plots relative to controls. There were more tomocerid Collembola in baited plots, suggesting possible indirect effects on Collembola caused by ant-spider interactions. The second experiment, in which screening of two mesh sizes selectively excluded large and small worker ants from a sugar bait, revealed that the large ants, primarily Camponotus, could alone induce similar effects on spiders. Gnaphosa biomass density was almost twice as high in the plots where large ants were more active, whereas Schizocosa biomass density was reduced by half in these plots. Although tomocerid densities did not differ between treatments, tomocerid numbers were negatively correlated with the activity of Formica, another large ant species. Path analysis failed to support the hypothesis that the ant Camponotus indirectly affected tomocerid Collembola through effects on densities of spiders. However, path analysis also revealed other indirect effects of Camponotus affecting tomocerids. These results illustrate the complexity of interactions between and within two major IG predator groups with disparate predatory behaviors, complexities that will have consequences for functioning of the forest-floor food web.     

A mathematical model of competition and colonization in a community of marine benthic algae

A mathematical model has been constructed for the algal community on the rocky shores of a Norwegian fjord. We report here on the studies of competition and colonization along a vertical transect from the upper intertidal to the sublittoral habiats. Results on species abundance and distribution (patterns of zonation) and time to reach maturity have been compared to observations both in the fjord area and in other rocky shore areas.Competition coefficients for the algae were inferred from plant morphology and shown to be in agreement with observations of algal abundance and their zone-forming ability. Competition restricts the distribution of the species, especially at the lower elevations, but does not alter their relative position. However, increasing uniform competition prolongs the time in which zone-forming can occur, and it also decreases the overall biomass which an area can sustain. Colonization by a single species may create transient stages in community development of the same order of magnitude as algae longevity, and probably also alters the zonation pattern to some degree.The simulation results indicate that the large-scale algal distribution pattern in the Hardangerfjord area results from global stability of the rocky shore community.     

Disentangling trophic relationships in a High Arctic tundra ecosystem through food web modeling

Determining the manner in which food webs will respond to environmental changes is difficult because the relative importance of top-down vs. bottom-up forces in controlling ecosystems is still debated. This is especially true in the Arctic tundra where, despite relatively simple food webs, it is still unclear which forces dominate in this ecosystem. Our primary goal was to assess the extent to which a tundra food web was dominated by plant-herbivore or predator-prey interactions. Based on a 17-year (1993-2009) study of terrestrial wildlife on Bylot Island, Nunavut, Canada, we developed trophic mass balance models to address this question. Snow Geese were the dominant herbivores in this ecosystem, followed by two sympatric lemming species (brown and collared lemmings). Arctic foxes, weasels, and several species of birds of prey were the dominant predators. Results of our trophic models encompassing 19 functional groups showed that <10% of the annual primary production was consumed by herbivores in most years despite the presence of a large Snow Goose colony, but that 20-100% of the annual herbivore production was consumed by predators. The impact of herbivores on vegetation has also weakened over time, probably due to an increase in primary production. The impact of predators was highest on lemmings, intermediate on passerines, and lowest on geese and shorebirds, but it varied with lemming abundance. Predation of collared lemmings exceeded production in most years and may explain why this species remained at low density. In contrast, the predation rate on brown lemmings varied with prey density and may have contributed to the high-amplitude, periodic fluctuations in the abundance of this species. Our analysis provided little evidence that herbivores are limited by primary production on Bylot Island. In contrast, we measured strong predator-prey interactions, which supports the hypothesis that this food web is primarily controlled by top-down forces. The presence of allochthonous resources subsidizing top predators and the absence of large herbivores may partly explain the predominant role of predation in this low-productivity ecosystem.     

Latitudinal variation in top-down and bottom-up control of a salt marsh food web

The shrub Iva frutescens, which occupies the terrestrial border of U.S. Atlantic Coast salt marshes, supports a food web that varies strongly across latitude. We tested whether latitudinal variation in plant quality (higher at high latitudes), consumption by omnivores (a crab, present only at low latitudes), consumption by mesopredators (ladybugs, present at all latitudes), or the life history stage of an herbivorous beetle could explain continental-scale field patterns of herbivore density. In a mesocosm experiment, crabs exerted strong top-down control on herbivorous beetles, ladybugs exerted strong top-down control on aphids, and both predators benefited plants through trophic cascades. Latitude of plant origin had no effect on consumers. Herbivorous beetle density was greater if mesocosms were stocked with beetle adults rather than larvae, and aphid densities were reduced in the "adult beetle" treatment. Treatment combinations representing high and low latitudes produced patterns of herbivore density similar to those in the field. We conclude that latitudinal variation in plant quality is less important than latitudinal variation in top consumers and competition in mediating food web structure. Climate may also play a strong role in structuring high-latitude salt marshes by limiting the number of herbivore generations per growing season and causing high overwintering mortality.