Experimental and Natural Warming Elevates Mercury Concentrations in Estuarine Fish

Marine food webs are the most important link between the global contaminant, methylmercury (MeHg), and human exposure through consumption of seafood. Warming temperatures may increase human exposure to MeHg, a potent neurotoxin, by increasing MeHg production as well as bioaccumulation and trophic transfer through marine food webs. Studies of the effects of temperature on MeHg bioaccumulation are rare and no study has specifically related temperature to MeHg fate by linking laboratory experiments with natural field manipulations in coastal ecosystems. We performed laboratory and field experiments on MeHg accumulation under varying temperature regimes using the killifish, Fundulus heteroclitus. Temperature treatments were established in salt pools on a coastal salt marsh using a natural temperature gradient where killifish fed on natural food sources. Temperatures were manipulated across a wider range in laboratory experiments with killifish exposed to MeHg enriched food. In both laboratory microcosms and field mesocosms, MeHg concentrations in killifish significantly increased at elevated temperatures. Moreover, in field experiments, other ancillary variables (salinity, MeHg in sediment, etc.) did not relate to MeHg bioaccumulation. Modeling of laboratory experimental results suggested increases in metabolic rate as a driving factor. The elevated temperatures we tested are consistent with predicted trends in climate warming, and indicate that in the absence of confounding factors, warmer sea surface temperatures could result in greater in bioaccumulation of MeHg in fish, and consequently, increased human exposure.     

Rapidly increasing methyl mercury in endangered ivory gull (Pagophila eburnea) feathers over a 130 year record

Mercury (Hg) is increasing in marine food webs, especially at high latitudes. The bioaccumulation and biomagnification of methyl mercury (MeHg) has serious effects on wildlife, and is most evident in apex predators. The MeHg body burden in birds is the balance of ingestion and excretion, and MeHg in feathers is an effective indicator of overall MeHg burden. Ivory gulls (Pagophila eburnea), which consume ice-associated prey and scavenge marine mammal carcasses, have the highest egg Hg concentrations of any Arctic bird, and the species has declined by more than 80% since the 1980s in Canada. We used feathers from museum specimens from the Canadian Arctic and western Greenland to assess whether exposure to MeHg by ivory gulls increased from 1877 to 2007. Based on constant feather stable-isotope (δ¹³C, δ¹⁵N) values, there was no significant change in ivory gulls'' diet over this period, but feather MeHg concentrations increased 45× (from 0.09 to 4.11 µg g⁻¹ in adults). This dramatic change in the absence of a dietary shift is clear evidence of the impact of anthropogenic Hg on this high-latitude threatened species. Bioavailable Hg is expected to increase in the Arctic, raising concern for continued population declines in high-latitude species that are far from sources of environmental contaminants.     

The transfer of cadmium, mercury, methylmercury, and zinc in an intertidal rocky shore food chain

We examined the transfer of cadmium (Cd), inorganic mercury [Hg(II)], methylmercury (MeHg), and zinc (Zn) in an intertidal rocky shore food chain, namely from marine phytoplankton to suspension-feeding rock oysters (Saccostrea cucullata) and finally to predatory whelks Thais clavigera. The uptake of metals from the dissolved phase was also concurrently quantified in the oysters and the whelks. Metal uptake by the oysters was not directly proportional, whereas metal uptake by the whelks was directly proportional to metal concentration in the water. The order of uptake was MeHg>Hg(II)>Zn>Cd, and was much higher in the oysters than in the whelks. The relative uptake of Zn and Cd was comparable between oysters and whelks, whereas MeHg and Hg(II) showed disproportionally higher uptake in oysters than in whelks as compared to Zn and Cd. The assimilation efficiencies (AEs) were in the order of MeHg>Zn>Cd=Hg(II) in oysters, whereas the AEs were highest for MeHg and comparable for Zn, Cd, and Hg(II) in the whelks. Pre-exposure of the oysters to different dissolved concentrations of Cd significantly elevated the AEs of Cd and Hg(II) but not of Zn, in association with the induction of metallothioneins in the oysters. The whelks significantly assimilated Cd and Zn from various prey (barnacles, oysters, mussels, and snails) with contrasting strageties of metal sequestration and storage. There was no significant relationship between the metal AE and the metal partitioning in the soluble fraction (including metallothionein-like proteins, heat stable protein, and organelles). The insoluble fraction of metals was also available for metal assimilation. Our calculations show that the dietary uptake of metals can be dominant in the overall bioaccumulation in the oysters and whelks, and the trophic transfer factor was >1 for all metals. Thus, the four metals have a high potential of being biomagnified in the intertidal rocky shore food chain. MeHg possessed the highest and Hg(II) and Cd the lowest potential of trophic transfer among the four metals considered.     

Bioaccumulation and Trophic Transfer of Mercury and Selenium in African Sub-Tropical Fluvial Reservoirs Food Webs (Burkina Faso)

The bioaccumulation and biomagnification of mercury (Hg) and selenium (Se) were investigated in sub-tropical freshwater food webs from Burkina Faso, West Africa, a region where very few ecosystem studies on contaminants have been performed. During the 2010 rainy season, samples of water, sediment, fish, zooplankton, and mollusks were collected from three water reservoirs and analysed for total Hg (THg), methylmercury (MeHg), and total Se (TSe). Ratios of δ¹³C and δ¹⁵N were measured to determine food web structures and patterns of contaminant accumulation and transfer to fish. Food chain lengths (FCLs) were calculated using mean δ¹⁵N of all primary consumer taxa collected as the site-specific baseline. We report relatively low concentrations of THg and TSe in most fish. We also found in all studied reservoirs short food chain lengths, ranging from 3.3 to 3.7, with most fish relying on a mixture of pelagic and littoral sources for their diet. Mercury was biomagnified in fish food webs with an enrichment factor ranging from 2.9 to 6.5 for THg and from 2.9 to 6.6 for MeHg. However, there was no evidence of selenium biomagnification in these food webs. An inverse relationship was observed between adjusted δ¹⁵N and log-transformed Se:Hg ratios, indicating that Se has a lesser protective effect in top predators, which are also the most contaminated animals with respect to MeHg. Trophic position, carbon source, and fish total length were the factors best explaining Hg concentration in fish. In a broader comparison of our study sites with literature data for other African lakes, the THg biomagnification rate was positively correlated with FCL. We conclude that these reservoir systems from tropical Western Africa have low Hg biomagnification associated with short food chains. This finding may partly explain low concentrations of Hg commonly reported in fish from this area.     

Climate change–contaminant interactions in marine food webs: Toward a conceptual framework

Climate change is reshaping the way in which contaminants move through the global environment, in large part by changing the chemistry of the oceans and affecting the physiology, health, and feeding ecology of marine biota. Climate change‐associated impacts on structure and function of marine food webs, with consequent changes in contaminant transport, fate, and effects, are likely to have significant repercussions to those human populations that rely on fisheries resources for food, recreation, or culture. Published studies on climate change–contaminant interactions with a focus on food web bioaccumulation were systematically reviewed to explore how climate change and ocean acidification may impact contaminant levels in marine food webs. We propose here a conceptual framework to illustrate the impacts of climate change on contaminant accumulation in marine food webs, as well as the downstream consequences for ecosystem goods and services. The potential impacts on social and economic security for coastal communities that depend on fisheries for food are discussed. Climate change–contaminant interactions may alter the bioaccumulation of two priority contaminant classes: the fat‐soluble persistent organic pollutants (POPs), such as polychlorinated biphenyls (PCBs), as well as the protein‐binding methylmercury (MeHg). These interactions include phenomena deemed to be either climate change dominant (i.e., climate change leads to an increase in contaminant exposure) or contaminant dominant (i.e., contamination leads to an increase in climate change susceptibility). We illustrate the pathways of climate change–contaminant interactions using case studies in the Northeastern Pacific Ocean. The important role of ecological and food web modeling to inform decision‐making in managing ecological and human health risks of chemical pollutants contamination under climate change is also highlighted. Finally, we identify the need to develop integrated policies that manage the ecological and socioeconomic risk of greenhouse gases and marine pollutants.     

Invasions and extinctions reshape coastal marine food webs

The biodiversity of ecosystems worldwide is changing because of species loss due to human-caused extinctions and species gain through intentional and accidental introductions. Here we show that the combined effect of these two processes is altering the trophic structure of food webs in coastal marine systems. This is because most extinctions ( approximately 70%) occur at high trophic levels (top predators and other carnivores), while most invasions are by species from lower trophic levels (70% macroplanktivores, deposit feeders, and detritivores). These opposing changes thus alter the shape of marine food webs from a trophic pyramid capped by a diverse array of predators and consumers to a shorter, squatter configuration dominated by filter feeders and scavengers. The consequences of the simultaneous loss of diversity at top trophic levels and gain at lower trophic levels is largely unknown. However, current research suggests that a better understanding of how such simultaneous changes in diversity can impact ecosystem function will be required to manage coastal ecosystems and forecast future changes.     

Biodiversity, ecosystem functioning and food webs in fresh waters: assembling the jigsaw puzzle

1. Dramatic advances have been made recently in the study of biodiversity–ecosystem functioning (B-EF) relations and food web ecology. These fields are now starting to converge, and this fusion has the potential to improve our understanding of how environmental stressors modulate ecosystem processes and the supply of 'goods and services'.
2. Food web structure and dynamics can exert particularly strong influences on B-EF relations in fresh waters, as consumer–resource interactions (e.g. trophic cascades) are often more important than horizontal interactions within trophic levels. For instance, many freshwater food webs are size structured, with large organisms tending to occupy the higher trophic levels and often exerting powerful effects on ecosystem processes. However, because they are also vulnerable to perturbations, non-random losses of these large taxa can alter both food web structure and ecosystem functioning profoundly.
3. Recently, the focus of food web research has shifted away from exploring patterns, towards developing an understanding of processes (e.g. quantifying fluxes of individuals, biomass, energy, nutrients) and how the two interact. Many of the best-characterized food webs are from fresh waters, and these ecosystems are now being used to address some of the shortcomings of earlier B-EF studies. I have identified several key gaps in our current knowledge and highlighted potentially fruitful avenues of future B-EF and food web research.
4. A major challenge for this newly emerging research is to place it within a unified theoretical framework. The application of metabolic theory and ecological stoichiometry may help to achieve this goal by considering biological systems within the constraints imposed upon them by physical and chemical laws.     

Major food web properties of two sandy beaches with contrasting morphodynamics, and effects on the stability

We determined major structural properties influencing the food webs of two sandy beaches with contrasting morphodynamics in the Atlantic coast of Uruguay: reflective (narrow and steep) and dissipative beaches (wide and flat). Furthermore, we evaluated how these characteristics could influence the stability of the local food webs. To this end, we examined the correlation of several food web properties with different ecosystem types (including freshwater habitats, estuary, marine, and terrestrial environments) using a principal components analysis. Sandy beach food web components included detritus, phytoplankton, zooplankton, benthic invertebrates, fishes, and seabirds. Our results revealed that the dissipative beach presented higher trophic levels, a higher number of trophic species, more links per species, as well as a higher proportion of intermediate trophic species, but lower connectance and proportion of omnivorous species than the reflective beach. The variation in the food web properties was explained by two principal components. Sandy beach food webs contribute mainly to one dimension of the principal components analysis that was determined by the number of trophic species, links per species, the trophic similarity, and the characteristic path length. We suggest that species and link characteristics, such as predominance of scavengers and detritivorous, the relatively high connectance and the short path length are drivers in the food web structure and may play a role in the community dynamic.     

Effects of trophic skewing of species richness on ecosystem functioning in a diverse marine community

Widespread overharvesting of top consumers of the world's ecosystems has "skewed" food webs, in terms of biomass and species richness, towards a generally greater domination at lower trophic levels. This skewing is exacerbated in locations where exotic species are predominantly low-trophic level consumers such as benthic macrophytes, detritivores, and filter feeders. However, in some systems where numerous exotic predators have been added, sometimes purposefully as in many freshwater systems, food webs are skewed in the opposite direction toward consumer dominance. Little is known about how such modifications to food web topology, e.g., changes in the ratio of predator to prey species richness, affect ecosystem functioning. We experimentally measured the effects of trophic skew on production in an estuarine food web by manipulating ratios of species richness across three trophic levels in experimental mesocosms. After 24 days, increasing macroalgal richness promoted both plant biomass and grazer abundance, although the positive effect on plant biomass disappeared in the presence of grazers. The strongest trophic cascade on the experimentally stocked macroalgae emerged in communities with a greater ratio of prey to predator richness (bottom-rich food webs), while stronger cascades on the accumulation of naturally colonizing algae (primarily microalgae with some early successional macroalgae that recruited and grew in the mesocosms) generally emerged in communities with greater predator to prey richness (the more top-rich food webs). These results suggest that trophic skewing of species richness and overall changes in food web topology can influence marine community structure and food web dynamics in complex ways, emphasizing the need for multitrophic approaches to understand the consequences of marine extinctions and invasions.     

Sources of organic matter and methylmercury in littoral macroinvertebrates: a stable isotope approach

The main objective of this study was to assess organic matter (OM) and methylmercury (MeHg) sources for freshwater littoral macroinvertebrate primary consumers. The carbon and nitrogen stable isotope ratios (δ¹³C, δ¹⁵N) of sources (epiphytes, macrophytes, suspended particulate matter _SPM) and of macroinvertebrate consumers were measured in a fluvial lake with extensive macrophyte beds (emergent and submerged). To determine the relative contribution of each OM source to macroinvertebrate diets we used the IsoSource model that examines all possible combinations of solutions for each source. Total and MeHg concentrations of consumers were also measured. Results show that epiphytes and macrophytes are dominant in the diet of macroinvertebrates, especially in early summer (July). In mid-summer (August), SPM constitutes a non-negligible OM source to the primary consumers. Hg concentrations were higher in epiphytes than in the other OM sources. The proportion of epiphytes in macroinvertebrate diet was positively correlated with the percentage of MeHg in their tissues. There was no relationship between SPM assimilation and Hg concentration in macroinvertebrate consumers. These results suggest that epiphytes and macrophytes constitute the main pathway of Hg bioaccumulation in littoral food webs.

Ocean acidification increases fatty acids levels of larval fish

Rising levels of anthropogenic carbon dioxide in the atmosphere are acidifying the oceans and producing diverse and important effects on marine ecosystems, including the production of fatty acids (FAs) by primary producers and their transfer through food webs. FAs, particularly essential FAs, are necessary for normal structure and function in animals and influence composition and trophic structure of marine food webs. To test the effect of ocean acidification (OA) on the FA composition of fish, we conducted a replicated experiment in which larvae of the marine fish red drum (Sciaenops ocellatus) were reared under a climate change scenario of elevated CO₂ levels (2100 µatm) and under current control levels (400 µatm). We found significantly higher whole-body levels of FAs, including nine of the 11 essential FAs, and altered relative proportions of FAs in the larvae reared under higher levels of CO₂. Consequences of this effect of OA could include alterations in performance and survival of fish larvae and transfer of FAs through food webs.     

Experimental Detritus Manipulations Unite Surface and Cave Stream Ecosystems Along a Common Energy Gradient

Subsidies of detritus from donor habitats are important energy sources for many ecosystems, but understanding their role in structuring recipient food webs requires comparative experimental studies along the full spectrum of detrital fluxes. Here we report results from an experimental addition of maize (Zea mays L.) litter to a detritus-poor cave stream ecosystem, which we then compare with analogous, past experiments using detritus-rich surface stream ecosystems that similarly have detritus-based food webs and extremely low in situ primary production. Bulk-tissue and compound-specific stable isotope analyses showed that maize litter carbon (C) was rapidly assimilated by microbes and transferred via successive trophic levels to the top of the cave stream food web (omnivorous crayfishes and predatory salamanders). All trophic levels increased in abundance and biomass, but only facultative cave taxa, that is those also found in surface streams, contributed to this numerical response. The lack of response by obligate cave species presumably occurred because evolutionary trade-offs associated with adaptations to low-C environments constrained their population-level responses during the one-year period of the litter addition. Comparison of the responses of the cave community with the analogous litter manipulation experiments in surface streams showed strong convergence in the functional relationship between invertebrate and detritus biomass (R ² = 0.72, P < 0.0001). Our results suggest that these seemingly disparate stream food webs lie along a single, common gradient of detritus supply, occupied at its extreme minimum by communities of obligate cave taxa adapted to low-energy environments.     

High‐resolution food webs based on nitrogen isotopic composition of amino acids

Food webs are known to have myriad trophic links between resource and consumer species. While herbivores have well‐understood trophic tendencies, the difficulties associated with characterizing the trophic positions of higher‐order consumers have remained a major problem in food web ecology. To better understand trophic linkages in food webs, analysis of the stable nitrogen isotopic composition of amino acids has been introduced as a potential means of providing accurate trophic position estimates. In the present study, we employ this method to estimate the trophic positions of 200 free‐roaming organisms, representing 39 species in coastal marine (a stony shore) and 38 species in terrestrial (a fruit farm) environments. Based on the trophic positions from the isotopic composition of amino acids, we are able to resolve the trophic structure of these complex food webs. Our approach reveals a high degree of trophic omnivory (i.e., noninteger trophic positions) among carnivorous species such as marine fish and terrestrial hornets.This information not only clarifies the trophic tendencies of species within their respective communities, but also suggests that trophic omnivory may be common in these webs.     

Structural Degradation in Mediterranean Sea Food Webs: Testing Ecological Hypotheses Using Stochastic and Mass-Balance Modelling

Human-mediated disturbances such as fishing, habitat modification, and pollution have resulted in significant shifts in species composition and abundance in marine ecosystems which translate into degradation of food-web structure. Here, we used a comparative ecological modelling approach and data from two food webs (North-Central Adriatic and South Catalan Sea) and two time periods (mid-late 1970s and 1990s) in the Mediterranean Sea to evaluate how changes in species composition and biomass have affected food-web properties and the extent of ecosystem degradation. We assembled species lists and ecological information for both regions and time periods into stochastic structural and mass-balance food-web models, and compared the outcomes of 22 food-web properties. Our results show strong similarities in structural food-web properties between the North-Central Adriatic and South Catalan Seas indicating similar ecosystem structure and levels of ecological degradation between regions and time periods. In contrast, a comparison with other published marine food webs (Caribbean, Benguela, and US continental shelf) suggested that Mediterranean webs are in an advanced state of ecological degradation. This was reflected by lower trophic height, linkage density, connectance, omnivory, species involved in looping, trophic chain length and fraction of biomass at higher trophic levels, as well as higher generality and fraction of biomass at lower trophic levels. An analysis of robustness to simulated species extinction revealed lower robustness to species removals in Mediterranean webs and corroborated their advanced state of degradation. Importantly, the two modelling approaches used delivered comparable results suggesting that they both capture fundamental information about how food webs are structured. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Trophic functioning of the Tiahura reef sector, Moorea Island, French Polynesia

 Energy-balanced steady-state models of the fringing and barrier reefs of Tiahura, Moorea Island, French Polynesia, are presented. A total of 43 and 46 trophic groups were identified on the two reef habitats respectively. The models’ outputs indicate that most of the substantial primary productivity is processed and recycled (59–69% of NPP) in the web through detritus based, microbially mediated food webs, with a substantial but secondary flux through grazer-based webs. This mechanism produces long pathways with low trophic efficiencies at the higher trophic levels. The trophic structure of both reef habitats efficiently conserves energy and materials within the reef ecosystem through two forms of internal recycling: a relatively large cycle produced through detritus and a microbial food web, and a relatively short one directly produced through predation. The models outputs suggest that bottom-up and top-down control are each ecologically important in both reef habitats. Accepted: 14 April 1997     

Bottom-up trophic cascades and material transfer in terrestrial food webs

In contrast to top-down trophic cascades, few reviews have appeared of bottom-up trophic cascades. We review the recent development of research on bottom-up cascades in terrestrial food webs, focusing on tritrophic systems consisting of plants, herbivorous insects, and natural enemies, and attempt to integrate bottom-up cascade and material transfer among trophic levels. Bottom-up cascades are frequently reported in various tritrophic systems, and are important to determine community structure, population dynamics, and individual performance of higher trophic levels. In addition, we highlight several features of bottom-up cascades. Accumulation or dilution of plant nutritional and defensive materials by herbivorous insects provides a mechanistic base for several bottom-up cascades. Such a stoichiometric approach has the potential to improve our understanding of bottom-up cascading effects in terrestrial food webs. We suggest a future direction for research by integration of bottom-up cascades and material transfer among trophic levels.     

Resource partitioning between Pacific walruses and bearded seals in the Alaska Arctic and sub-Arctic

Climate-mediated changes in the phenology of Arctic sea ice and primary production may alter benthic food webs that sustain populations of Pacific walruses (Odobenus rosmarus divergens) and bearded seals (Erignathus barbatus). Interspecific resource competition could place an additional strain on ice-associated marine mammals already facing loss of sea ice habitat. Using fatty acid (FA) profiles, FA trophic markers, and FA stable carbon isotope analyses, we found that walruses and bearded seals partitioned food resources in 2009–2011. Interspecific differences in FA profiles were largely driven by variation in non-methylene FAs, which are markers of benthic invertebrate prey taxa, indicating varying consumption of specific benthic prey. We used Bayesian multi-source FA stable isotope mixing models to estimate the proportional contributions of particulate organic matter (POM) from sympagic (ice algal), pelagic, and benthic sources to these apex predators. Proportional contributions of FAs to walruses and bearded seals from benthic POM sources were high [44 (17–67)% and 62 (38–83)%, respectively] relative to other sources of POM. Walruses also obtained considerable contributions of FAs from pelagic POM sources [51 (32–73)%]. Comparison of δ¹³C values of algal FAs from walruses and bearded seals to those from benthic prey from different feeding groups from the Chukchi and Bering seas revealed that different trophic pathways sustained walruses and bearded seals. Our findings suggest that (1) resource partitioning may mitigate interspecific competition, and (2) climate change impacts on Arctic food webs may elicit species-specific responses in these high trophic level consumers.     

Stable isotope analysis of aquatic invertebrate communities in irrigated rice fields cultivated under different management regimes

In this study we have used stable isotope analysis to identify major food resources driving food webs in commercial rice agroecosystems and to examine the effects of agricultural management practices on the trophic structure of these food webs. Potential carbon sources and aquatic macroinvertebrate consumers were collected from large-scale rice farms in south-eastern Australia cultivated under three different crop management regimes conventional-aerial (agrochemicals applied, aerially sown), conventional-sod (agrochemicals applied, directly sown) and organic-sod (agrochemical-free, directly sown). Evidence from stable isotope analysis demonstrated the importance of food sources, such as biofilm and detritus, as the principal energy sources driving aquatic food webs in rice agroecosystems. Despite the greater diversity of potential food sources collected from the organic-sod regime across all sampling occasions, the range of food resources directly assimilated by macroinvertebrate consumers did not differ substantially across management regimes. Trophic complexity of aquatic food webs, as evidenced by the number of trophic levels identified using δ¹⁵N data, differed across management regimes at the early season sampling. Sites with low or no agrochemical applications contained more than two trophic levels, but at the site with the highest pesticide application no primary or secondary consumers were found. Our data demonstrates that the choice of agricultural management regime has a season-long influence on aquatic food webs in rice crops, and highlights the importance of conserving non-rice food resources that drive these trophic networks.