Mercury bioaccumulation in estuarine food webs

We tested for unintended mercury contamination problems associated with estuarine floodplain restoration projects of the Louisiana coastal zone, USA. Barataria Bay and Breton Sound are two neighboring deltaic estuaries that were isolated by levees from the Mississippi River about 100 years ago. These estuaries recently have been reconnected to the nutrient-rich Mississippi River, starting major river diversion (input) flows in 1991 for Breton Sound and in 2004 for Barataria Bay. We collected > 2100 fish over five years from 20 stations in these estuaries to test two hypotheses about Hg bioaccumulation: (H1) Background Hg bioaccumulation in fish would be highest in low-salinity upper reaches of estuaries, and (H2) recent river inputs to these upper estuarine areas would increase Hg bioaccumulation in fish food webs. For H1, we surveyed fish Hg concentrations at several stations along a salinity gradient in Barataria Bay in 2003-2004, a time when this estuary lacked strong river inputs. Results showed that average Hg concentrations in fish communities were lowest (150 ng/g dry mass) in higher salinity areas and -2.4x higher (350 ng/g) in low-salinity oligohaline and freshwater upper reaches of the estuary. For H2, we tested for enhanced Hg bioaccumulation following diversion onset in both estuaries. Fish communities from Breton Sound that had long-term (> 10 years) diversion inputs had -1.7x higher average Hg contents of 610 ng/g Hg vs. 350 ng/g background values. Shorter-term diversion inputs over 2-3 years in upper Barataria Bay did not result in strong Hg enrichments or stable C isotope increases seen in Breton Sound, even though N and S stable-isotope values indicated strong river inputs in both estuaries. It may be that epiphyte communities on abundant submerged aquatic vegetation (SAV) are important hotspots for Hg cycling in these estuaries, and observed lesser development of these epiphyte communities in upper Barataria Bay during the first years of diversion inputs may account for the lessened Hg bioaccumulation in fish. A management consideration from this study is that river restoration projects may unintentionally fertilize SAV and epiphyte-based food webs, leading to higher Hg bioaccumulation in river-impacted floodplains and their food webs.     

Cyclic energy pathways in ecological food webs

Standard ecology textbooks typically maintain that nutrients cycle, but energy flows in unidirectional chains. However, here we use a new metric that allows for the identification and quantification of cyclic energy pathways. Some of these important pathways occur due to the contribution of dead organic matter to detrital pools and those organisms that feed on them, reintroducing some of that energy back into the food web. Recognition of these cyclic energy pathways profoundly impacts many aspects of ecology such as trophic levels, control, and the importance of indirect effects. Network analysis, specifically the maximum eigenvalue of the connectance matrix, is used to identify both the presence and strength of these structural cycles.     

Consumer-resource body-size relationships in natural food webs

It has been suggested that differences in body size between consumer and resource species may have important implications for interaction strengths, population dynamics, and eventually food web structure, function, and evolution. Still, the general distribution of consumer-'resource body-size ratios in real ecosystems, and whether they vary systematically among habitats or broad taxonomic groups, is poorly understood. Using a unique global database on consumer and resource body sizes, we show that the mean body-size ratios of aquatic herbivorous and detritivorous consumers are several orders of magnitude larger than those of carnivorous predators. Carnivorous predator-prey body-size ratios vary across different habitats and predator and prey types (invertebrates, ectotherm, and endotherm vertebrates). Predator-prey body-size ratios are on average significantly higher (1) in freshwater habitats than in marine or terrestrial habitats, (2) for vertebrate than for invertebrate predators, and (3) for invertebrate than for ectotherm vertebrate prey. If recent studies that relate body-size ratios to interaction strengths are general, our results suggest that mean consumer-resource interaction strengths may vary systematically across different habitat categories and consumer types.     

Topological structure and interaction strengths in model food webs

We report the results of carrying out a large number of simulations on a coevolutionary model of multispecies communities. A wide range of parameter values were investigated which allowed a rather complete picture of the change in behaviour of the model as these parameters were varied to be built up. Our main interest was in the nature of the community food webs constructed via the simulations. We identify the range of parameter values which give rise to realistic food webs and give arguments which allow some of the structure which is found to be understood in an intuitive way. Since the webs are evolved according to the rules of the model, the strengths of the predator–prey links are not determined a priori, and emerge from the process of constructing the web. We measure the distribution of these link strengths, and find that there are a large number of weak links, in agreement with recent suggestions. We also review some of the data on food webs available in the literature, and make some tentative comparisons with our results. The difficulties of making such comparisons and the possible future developments of the model are also briefly discussed.     

A simulation analysis of continental shelf food webs

Energy flow through continental shelf food webs was examined using a simulation model. The model structure expands the two traditional marine food chains of phytoplankton-zooplankton-pelagic fish and benthos-demersal fish into a complex web which includes detritus, dissolved organic matter (DOM), bacteria, protozoa, and mucus net feeders. Simulation of energy flux for different shelf systems using the expanded web revealed that heterotrophic microorganisms and their predators account for a significant component of the energy flux in the continental shelf ecosystem. Contrary to previous models, where all phytoplankton were considered to be grazed by zooplankton, our simulation results indicate that only slightly more than 50% of the annual net primary production is grazed. A substantial quantity of the phytoplankton production directly becomes detritus. Bacteria mineralize detritus and DOM produced by phytoplankton and other components of the food web, converting these to biomass with high efficiency. Consequently, the model predicts that planktonic bacterial production is equivalent to zooplankton production. Exclusion of the bacteria requires the assumption that all DOM is either exported from the system or consumed by another component of the food web. Neither of these assumptions can be supported by present knowledge of the dynamics of DOM in the sea. Model simulations were also employed to test the hypothesis that production exceeds consumption on continental shelves, resulting in exports of 50% of the annual primary production. Simulations of shelves with high rates of primary production resulted in a particulate export of 27% and realistic estimates of secondary production. Results of other simulations suggest that shelves with lower primary production cannot export production and still maintain the macrobenthos and their predators. General properties about continental shelves can also be inferred from the model. From simulations of shelves of differing primary production, nanoplankton are predicted to account for a greater proportion of the primary production in nutrient limited systems. Benthic production appears to be related to both the quantity of primary production and the sinking rates of the phytoplankton. The model indicates that zooplankton fecal inputs to the shelf benthos are only a small portion of the total detrital flux, leading to the prediction that fecal pellets are of little significance in determining benthic production. Finally, the model generates production efficiencies that are highly variable depending on the type of system and kind of populations involved. We argue that the assumed ecological efficiency of 10% should be abandoned for continental shelves and other ecosystems.     

Response of complex food webs to realistic extinction sequences

Although an ecosystem's response to biodiversity loss depends on the order in which species are lost, the extinction sequences generally used to explore such responses in food webs have been ecologically unrealistic. We investigate how several extinction orders affect the minimum number of secondary extinctions expected within pelagic food webs from 34 temperate freshwater lakes. An ecologically plausible extinction order is derived from the geographically nested pattern of species composition among the lakes and is corroborated by species' pH tolerances. Simulations suggest that lake communities are remarkably robust to this realistic extinction order and highly sensitive to the reverse sequence of species loss. This sensitivity is not well explained by the known sensitivity of networks to the loss of highly connected species but appears to be better explained by our observation that trophic specialists preferentially consume widely distributed species at low risk of extinction. Our results highlight an important aspect of community organization that may help to maintain biodiversity amidst changing environments.     

Trophic structure of coastal Antarctic food webs associated with changes in sea ice and food supply

Predicting the dynamics of ecosystems requires an understanding of how trophic interactions respond to environmental change. In Antarctic marine ecosystems, food web dynamics are inextricably linked to sea ice conditions that affect the nature and magnitude of primary food sources available to higher trophic levels. Recent attention on the changing sea ice conditions in polar seas highlights the need to better understand how marine food webs respond to changes in such broad-scale environmental drivers. This study investigated the importance of sea ice and advected primary food sources to the structure of benthic food webs in coastal Antarctica. We compared the isotopic composition of several seafloor taxa (including primary producers and invertebrates with a variety of feeding modes) that are widely distributed in the Antarctic. We assessed shifts in the trophic role of numerically dominant benthic omnivores at five coastal Ross Sea locations. These locations vary in primary productivity and food availability, due to their different levels of sea ice cover, and proximity to polynyas and advected primary production. The delta15N signatures and isotope mixing model results for the bivalves Laternula elliptica and Adamussium colbecki and the urchin Sterechinus neumeyeri indicate a shift from consumption of a higher proportion of detritus at locations with more permanent sea ice in the south to more freshly produced algal material associated with proximity to ice-free water in the north and east. The detrital pathways utilized by many benthic species may act to dampen the impacts of large seasonal fluctuations in the availability of primary production. The limiting relationship between sea ice distribution and in situ primary productivity emphasizes the role of connectivity and spatial subsidies of organic matter in fueling the food web. Our results begin to provide a basis for predicting how benthic ecosystems will respond to changes in sea ice persistence and extent along environmental gradients in the high Antarctic.     

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.     

Potential trophic biomasses and trace-substance concentrations in unstructured marine food webs

It is proposed that unstructured food webs may more closely resemble real marine food webs than does the conventional, structured model. An unstructured food-web model leads to a set of very simple expressions for the potential partition of matter in the food web in steady state, including the potential fluxes of material and biomasses of trophic types and the concentration of trace substances in the members and materials of such a food web. The approach may explain some anomalies of relative predator-prey biomasses and of trace-element distribution, and may be of further use for analyzing and predicting (a) the tropho-dynamic parameters of marine systems, (b) the trophic positions, and the steadystate fluxes and biomasses of marine organisms, (c) the distribution of trace materials in marine biota; and for relating findings among these areas. Other matters, such as limitations of food conversion, indicated by concentration factors of trace substances, the possibilities of non-causal association of anomalously high levels of trace substances (including pollutants) with diseased or otherwise abnormal marine creatures, and an inverse relationship of early concentrations of newly introduced trace substances and their eventual concentrations in various organisms, are also developed in this approach.     

Sources of primary production supporting food webs in an arid coastal embayment

Penaeid shrimp can be useful ecological indicators of linkages between shallow tropical coastal habitats, acting as integrators of carbon and nitrogen sources due to their generalist feeding habits and their mobility between habitats and with tidal cycles. In the current study, the contribution of mangrove, seagrass and microbial mat to the nutrition of two penaeid shrimp species, Penaeus semisulcatus and Metapeneus ensis, in a shallow arid embayment in the Arabian Gulf was assessed through a combination of analysis of stomach contents and dual carbon and nitrogen stable isotope signatures. Shrimp tissue stable isotope signatures identified seagrass as a major source of carbon and nitrogen for both species, contributing 21–38?% (1–99?‰). Microbial mat was also detected as a significant nutritional source for early-stage Metapenaeus affinis postlarvae (1–27?%). However, mangroves were not identified as a significant source, with the range of results including the possibility of a zero contribution. Moreover, the greatest possible contribution of mangroves as source of carbon was less than for the other primary producers. This may be due the high salinity and wide temperature range limiting mangrove productivity as well as the low export of dissolved and particulate organic material out of the mangroves due to low rainfall.     

Measuring terrestrial subsidies to aquatic food webs using stable isotopes of hydrogen

Understanding river food webs requires distinguishing energy derived from primary production in the river itself (autochthonous) from that produced externally (allochthonous), yet there are no universally applicable and reliable techniques for doing so. We compared the natural abundance stable isotope ratios of hydrogen (deltaD) of allochthonous and autochthonous energy sources in four different aquatic ecosystems. We found that autochthonous organic matter is uniformly far more depleted in deuterium (lower deltaD values) than allochthonous: an average difference of approximately 100% per hundred. We also found that organisms at higher trophic levels, including both aquatic invertebrates and fish, have deltaD values intermediate between aquatic algae and terrestrial plants. The consistent differences between leaves and algae in deltaD among these four watersheds, along with the intermediate values in higher trophic levels, indicate that natural abundance hydrogen isotope signatures are a powerful tool for partitioning energy flow in aquatic ecosystems.     

Trophic levels and trophic tangles: the prevalence of omnivory in real food webs

The concept of trophic levels is one of the oldest in ecology and informs our understanding of energy flow and top-down control within food webs, but it has been criticized for ignoring omnivory. We tested whether trophic levels were apparent in 58 real food webs in four habitat types by examining patterns of trophic position. A large proportion of taxa (64.4%) occupied integer trophic positions, suggesting that discrete trophic levels do exist. Importantly however, the majority of those trophic positions were aggregated around integer values of 0 and 1, representing plants and herbivores. For the majority of the real food webs considered here, secondary consumers were no more likely to occupy an integer trophic position than in randomized food webs. This means that, above the herbivore trophic level, food webs are better characterized as a tangled web of omnivores. Omnivory was most common in marine systems, rarest in streams, and intermediate in lakes and terrestrial food webs. Trophic-level-based concepts such as trophic cascades may apply to systems with short food chains, but they become less valid as food chains lengthen.     

Feeding by larvae of intertidal invertebrates: assessing their position in pelagic food webs

One of the leading determinants of the structure and dynamics of marine populations is the rate of arrival of new individuals to local sites. While physical transport processes play major roles in delivering larvae to the shore, these processes become most important after larvae have survived the perils of life in the plankton, where they usually suffer great mortality. The lack of information regarding larval feeding makes it difficult to assess the effects of food supply on larval survival, or the role larvae may play in nearshore food webs. Here, we examine the spectrum of food sizes and food types consumed by the larvae of two intertidal barnacle species and of the predatory gastropod Concholepas concholepas. We conducted replicated experiments in which larvae were exposed to the food size spectrum (phytoplankton, microprotozoan and autotrophic picoplankton) found in nearshore waters in central Chile. Results show that barnacle nauplii and gastropod veligers are omnivorous grazers, incorporating significant fractions of heterotrophs in their diets. In accordance with their feeding mechanisms and body size, barnacle nauplii were able to feed on autotrophic picoplankton (<5 microm) and did not consume the largest phytoplankton cells, which made the bulk of phytoplankton biomass in spring-summer blooms. Balanoid nauplii exhibited higher ingestion rates than the smaller-bodied chthamaloid larvae. Newly hatched C. concholepas larvae also consumed picoplankton cells, while competent larvae of this species ingested mostly the largest phytoplankton cells and heterotrophic protozoans. Results suggest that persistent changes in the structure of pelagic food webs can have important effects on the species-specific food availability for invertebrate larvae, which can result in large-scale differences in recruitment rates of a given species, and in the relative recruitment success of the different species that make up benthic communities.     

Estimating nonlinear interaction strengths: an observation-based method for species-rich food webs

Efforts to estimate the strength of species interactions in species-rich, reticulate food webs have been hampered by the multitude of direct and indirect interactions such systems exhibit and have been limited by an assumption that pairwise interactions display linear functional forms. Here we present a new method for directly measuring, on a per capita basis, the nonlinear strength of trophic species interactions within such food webs. This is an observation-based method, requiring three pieces of information: (1) species abundances, (2) predator and prey-specific handling times, and (3) data from predator-specific feeding surveys in which the number of individuals observed feeding on each of the predator's prey species has been tallied. The method offers a straightforward way to assess the completeness of one's sampling effort in accurately estimating interaction strengths through the construction of predator-specific prey accumulation curves. The method should be applicable to a variety of systems in which empirical estimates of direct interaction strengths have thus far remained elusive.     

Chemically mediated competition between microbes and animals: microbes as consumers in food webs

Microbes are known to affect ecosystems and communities as decomposers, pathogens, and mutualists. However, they also may function as classic consumers and competitors with animals if they chemically deter larger consumers from using rich food-falls such as carrion, fruits, and seeds that can represent critical windfalls to both microbes and animals. Microbes often use chemicals (i.e., antibiotics) to compete against other microbes. Thus using chemicals against larger competitors might be expected and could redirect significant energy subsidies from upper trophic levels to the detrital pathway. When we baited traps in a coastal marine ecosystem with fresh vs. microbe-laden fish carrion, fresh carrion attracted 2.6 times as many animals per trap as microbe-laden carrion. This resulted from fresh carrion being found more frequently and from attracting more animals when found. Microbe-laden carrion was four times more likely to be uncolonized by large consumers than was fresh carrion. In the lab, the most common animal found in our traps (the stone crab Menippe mercenaria) ate fresh carrion 2.4 times more frequently than microbe-laden carrion. Bacteria-removal experiments and feeding bioassays using organic extracts of microbe-laden carrion showed that bacteria produced noxious chemicals that deterred animal consumers. Thus bacteria compete with large animal scavengers by rendering carcasses chemically repugnant. Because food-fall resources such as carrion are major food subsidies in many ecosystems, chemically mediated competition between microbes and animals could be an important, common, but underappreciated interaction within many communities.     

Historical metal concentrations in lacustrine food webs revealed using fossil ephippia from Daphnia.

Metal contamination of freshwater ecosystems is increasingly prevalent due to anthropogenic activities such as metal smelting and fossil fuel combustion. While toxicological studies focus on aqueous metal concentrations that result in lethal or sublethal responses, currently the only method for reconstructing a lake's metal contamination history is through an examination of the sedimentary deposits. In this paper, we suggest that cladoceran diapausing eggs (ephippia), which are abundant in nature and accumulate maternally derived metals, can be used to measure historical variations in biologically relevant metals that derive from the water column (water, diet). Linear regressions of total metal content against ephippia density or mass were strong (R2 > 0.80, P < 0.04) and revealed that metals were incorporated into ephippia with little contamination from the sediment matrix. Comparison of metal concentrations in ephippia and bulk sediments from three lakes demonstrated that some metals associated with urban sources (Cd, Cr, Mo) were preferentially concentrated in ephippia, whereas concentrations of other metals indicating landscape erosion (Al, Ca, Fe, Mn) exhibited greater concentrations in bulk sediments than in diapausing eggs. Because historical changes in metals within fossils and bulk sediments were uncorrelated in most instances, past variation in the metal content of ephippia provided a unique history of food web exposure to metals in the water column.     

The probabilistic niche model reveals substantial variation in the niche structure of empirical food webs

The structure of food webs, complex networks of interspecies feeding interactions, plays a crucial role in ecosystem resilience and function, and understanding food web structure remains a central problem in ecology. Previous studies have shown that key features of empirical food webs can be reproduced by low-dimensional "niche" models. Here we examine the form and variability of food web niche structure by fitting a probabilistic niche model to 37 empirical food webs, a much larger number of food webs than used in previous studies. The model relaxes previous assumptions about parameter distributions and hierarchy and returns parameter estimates for each species in each web. The model significantly outperforms previous niche model variants and also performs well for several webs where a body-size-based niche model performs poorly, implying that traits other than body size are important in structuring these webs' niche space. Parameter estimates frequently violate previous models' assumptions: in 19 of 37 webs, parameter values are not significantly hierarchical, 32 of 37 webs have nonuniform niche value distributions, and 15 of 37 webs lack a correlation between niche width and niche position. Extending the model to a two-dimensional niche space yields networks with a mixture of one- and two-dimensional niches and provides a significantly better fit for webs with a large number of species and links. These results confirm that food webs are strongly niche-structured but reveal substantial variation in the form of the niche structuring, a result with fundamental implications for ecosystem resilience and function.     

Regulation of intertidal food webs by avian predators on New England rocky shores

Although there is a large body of research on food webs in rocky intertidal communities, most of the emphasis has been on the marine benthic components. Effects of avian predation on highly mobile predators such as crabs, remains practically unstudied in rocky shore ecosystems. The crab, Cancer borealis, is an important component of the diet of gulls (Larus marinus, L. argentatus) at the Isles of Shoals, Maine, USA. C. borealis prey include the predatory gastropod Nucella lapillus L., the herbivore Littorina littorea, and mussels Mytilus edulis L. We hypothesized that gulls reduce abundance of C. borealis in the low intertidal and shallow subtidal, thereby allowing C. borealis prey to persist in high numbers. A study of crab tidal migration showed that C. borealis density nearly doubled at high tide compared to low tide; thus, crabs from a large subtidal source population migrate into the intertidal zone during high tides and either emigrate or are removed by gulls during low tides. Results from a small-scale (1 m2) predator caging experiment in the low intertidal zone indicated that enclosed crabs significantly reduced L. littorea abundance when protected from gull predation. In a much larger-scale gull exclusion experiment, densities of C. borealis increased significantly during low and high tides in exclosures relative to the controls. C. borealis density was inversely correlated with changes in the abundance of two mesopredators Carcinus maenas and Nucella lapillus, and with the space-occupier M. edulis. There was a similar negative correlation between abundance of C. borealis and the change in abundance of the herbivore L. littorea, but the trend was not significant. Mortality of tethered L. littorea was associated with C. borealis density across sites. However, preferred algae did not change in response to L. littorea density during the experiment. Thus, we found suggestive, but not conclusive, evidence for a three-level cascade involving gulls, crabs, and L. littorea. Our studies strongly suggest that gulls, as apex predators, generate three-level trophic cascades in rocky intertidal food webs by preventing the highly mobile subtidal predator, C. borealis, from establishing substantial populations in the low-mid intertidal zone thereby indirectly enhancing densities of two key mesopredators (N. lapillus, Carcinus) and blue mussels (M. edulis).     

Native fish diversity alters the effects of an invasive species on food webs

Aquatic communities have been altered by invasive species, with impacts on native biodiversity and ecosystem function. At the same time, native biodiversity may mitigate the effects of an invader. Common carp (Cyprinus carpio) is a ubiquitous, invasive fish species that strongly influences community and ecosystem processes. We used common carp to test whether the potential effects of an invasive species are altered across a range of species diversity in native communities. In mesocosms, treatments of zero, one, three, and six native fish species were used to represent the nested subset patterns observed in fish communities of lakes in Illinois, USA. The effect of the invader was tested across fish richness treatments by adding common carp to the native community and substituting native biomass with common carp. Native species and intraspecific effects reduced invader growth. The invader reduced native fish growth; however, the negative effect was minimized with increasing native richness. The zooplankton grazer community was modified by a top-down effect from the invader that increased the amount of phytoplankton. Neither the invader nor richness treatments influenced total phosphorus or community metabolism. Overall, the invader reduced resources for native species; and the effect scaled with how the invader was incorporated into the community. Higher native diversity mitigated the impact of the invader, confirming the need to consider biodiversity when predicting the impacts of invasive species.