Combined stable isotope and gut contents analysis of food webs in plant-dominated, shallow lakes

Summary 1. To determine feeding links between primary producers, invertebrates and fish, stable isotope analyses and gut content analyses of fish were conducted on the components of four shallow, eutrophic to hypertrophic, plant-dominated lakes.
2. Although separation of basal resources was possible, the diets of both fish and invertebrates were broad, comprising food from different compartments (planktonic, epiphytic/benthic), as well as from different trophic levels.
3. Mixing models were used to determine the extent to which periphyton production supported higher trophic levels. Only one species of invertebrate relied upon periphyton production exclusively.
4. Fish density affected the diets of invertebrates. The response was different for planktonic and epiphytic/benthic invertebrates. The proportion of periphyton production in the diets of zooplankton appeared to increase with fish density, whilst it decreased for other invertebrates.
5. As all zooplankton samples were collected in the open water at dusk, these results are further evidence for the diurnal horizontal migration of zooplankton. Although not conclusive, they are consistent with a behavioural response by invertebrates and zooplankton in the presence of fish.     

Unravelling complexities in benthic food webs using a dual stable isotope (hydrogen and carbon) approach

1. Stable carbon isotope studies have been an essential component of research regarding the contribution of methane (CH₄)-derived carbon to freshwater food webs and results have suggested that benthic macroinvertebrates in billabongs, streams and lakes may be partially, and in some instances, significantly 'fuelled' by methanotrophic biomass. However, the singular use of carbon isotopes can lead to ambiguous interpretations concerning the origin of carbon, especially in systems where phototrophs are likely to be using carbon respired sources and hence show more ¹³C-depleted values.
2. These uncertainties can be further resolved by the inclusion of additional isotopic data. Stable hydrogen isotopes are being increasingly used in food web studies with a marked advantage that sources may be isotopically distinct by one or two orders of magnitude greater than stable carbon or nitrogen, the isotopes most commonly used to delineate trophic interactions. By using hydrogen as a second biogeochemical tracer we provide further supportive evidence for the assimilation of methanotrophic microbial biomass by chironomid larvae.
3. Moreover, the hydrogen and carbon isotope values we found in chironomid tissues appear to reflect the original substrate used during methanogenesis; either acetate fermentation or carbonate reduction. Use of the former tends to result in relatively heavy carbon and light hydrogen isotope values due to kinetic isotope effects, whereas use of the latter results in relatively lighter carbon and heavier hydrogen isotope values.
4. We provide preliminary evidence to suggest that hydrogen and carbon isotope values in macroinvertebrates may be used to distinguish between CH₄ formation pathways and help to explain inter-depth and inter-specific differences between co-existing chironomid species found in the same lake.     

Arthropod food webs in organic and conventional wheat farming systems of an agricultural long‐term experiment: a stable isotope approach

• 1 Agricultural intensification not only alters the structure of arthropod communities, but also may affect biotic interactions by altering the availability of basal resources. We analyzed variations in stable isotope ratios (¹⁵N/¹⁴N and ¹³C/¹²C) of fertilizers, plants, prey and generalist predators in organic and conventional farming systems in a long‐term agricultural experiment [DOK trial (bioDynamic, bioOrganic, Konventionell)]. Two basal resources with pronounced differences in carbon isotope signatures, wheat litter (C₃ plant) and maize litter (C₄ plant), were used to uncover differences in food web properties between the two farming systems (conventional versus organic).
• 2 Predators incorporated significantly higher proportions of carbon from wheat sources in organically managed fields, suggesting that they were more closely linked to wheat‐consuming prey in this system. The δ¹⁵N values of three predaceous species were more than 2‰ greater in summer than in spring.
• 3 The results obtained suggest that generalist predators consumed higher proportions of herbivore prey in the organic system and that starvation and intraguild predation rates increased in some predator species with time.
• 4 Because the effects of farming system and sampling date on predators were species‐specific, conserving a diverse natural enemy community including species with different phenologies and sensitivities to management practices may, in the long term, be a good strategy for maintaining high pest suppression throughout the growing season.

     

Structure of tropical river food webs revealed by stable isotope ratios

Fish assemblages in tropical river food webs are characterized by high taxonomic diversity, diverse foraging modes, omnivory, and an abundance of detritivores. Feeding links are complex and modified by hydrologic seasonality and system productivity. These properties make it difficult to generalize about feeding relationships and to identify dominant linkages of energy flow. We analyzed the stable carbon and nitrogen isotope ratios of 276 fishes and other food web components living in four Venezuelan rivers that differed in basal food resources to determine 1) whether fish trophic guilds integrated food resources in a predictable fashion, thereby providing similar trophic resolution as individual species, 2) whether food chain length differed with system productivity, and 3) how omnivory and detritivory influenced trophic structure within these food webs. Fishes were grouped into four trophic guilds (herbivores, detritivores/algivores, omnivores, piscivores) based on literature reports and external morphological characteristics. Results of discriminant function analyses showed that isotope data were effective at reclassifying individual fish into their pre-identified trophic category. Nutrient-poor, black-water rivers showed greater compartmentalization in isotope values than more productive rivers, leading to greater reclassification success. In three out of four food webs, omnivores were more often misclassified than other trophic groups, reflecting the diverse food sources they assimilated. When fish δ¹⁵N values were used to estimate species position in the trophic hierarchy, top piscivores in nutrient-poor rivers had higher trophic positions than those in more productive rivers. This was in contrast to our expectation that productive systems would promote longer food chains. Although isotope ratios could not resolve species-level feeding pathways, they did reveal how top consumers integrate isotopic variability occurring lower in the food web. Top piscivores, regardless of species, had carbon and nitrogen profiles less variable than other trophic groups.     

Food webs: a plea for parasites

Parasites have the capacity to regulate host populations and may be important determinants of community structure, yet they are usually neglected in studies of food webs. Parasites can provide much of the information on host biology, such as diet and migration, that is necessary to construct accurate webs. Because many parasites have complex life cycles that involve several different hosts, and often depend on trophic interactions for transmission, parasites provide complementary views of web structure and dynamics. Incorporation of parasites in food webs can substantially after baste web properties, Including connectance, chain length and proportions of top and basal species, and can allow the testing of specific hypotheses related to food-web dynamics.     

Linking aboveground and belowground food webs through carbon and nitrogen stable isotope analyses

Carbon and nitrogen stable isotope ratios (δ¹³C and δ¹⁵N) have been used for more than two decades in analyses of food web structure. The utility of isotope ratio measurements is based on the observation that consumer δ¹³C values are similar (<1‰ difference) to those of their diet, while consumer δ¹⁵N values are about 3‰ higher than those of their diet. The technique has been applied most often to aquatic and aboveground terrestrial food webs. However, few isotope studies have examined terrestrial food web structure that includes both above- and belowground (detrital) components. Here, we review factors that may influence isotopic signatures of terrestrial consumers in above- and belowground systems. In particular, we emphasize variations in δ¹³C and δ¹⁵N in belowground systems, e.g., enrichment of ¹³C and ¹⁵N in soil organic matter (likely related to soil microbial metabolism). These enrichments should be associated with the high ¹³C (~3‰) enrichment in belowground consumers relative to litter and soil organic matter and with the large variation in δ¹⁵N (~6‰) of the consumers. Because such enrichment and variation are much greater than the trophic enrichment generally used to estimate consumer trophic positions, and because many general predators are considered dependent on energy and material flows from belowground, the isotopic variation in belowground systems should be taken into account in δ¹³C and δ¹⁵N analyses of terrestrial food webs. Meanwhile, by measuring the δ¹³C of key predators, the linkage between above- and belowground systems could be estimated based on observed differences in δ¹³C of primary producers, detritivores and predators. Furthermore, radiocarbon (¹⁴C) measurements will allow the direct estimation of the dependence of predators on the belowground systems.     

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.

Trophic ecology of the Lake Superior wave zone: a stable isotope approach

Stable carbon and nitrogen isotope ratio analyses were used to characterize the primary energy sources and trophic positions of 16 common Lake Superior wave zone invertebrate species. Isotope data from six tributary species that were taxonomically and ecologically matched with common wave zone species revealed broad energetic separation between these similarly structured benthic food webs. Previously published stable isotope data for Lake Superior wetland and pelagic food webs were used to assess the relative importance of inter-habitat energy flow within the Lake Superior ecosystem. The results of these comparisons indicate that the Lake Superior wave zone is energetically distinct from its tributaries, wetlands, and to a lesser extent from its vast pelagic realm. This information and approach should prove useful in future studies on the bioenergetics of inter-zonal migrants and other species that forage in multiple habitats within the lake and also in revealing energetic connections among terrestrial, riverine, littoral, and pelagic food webs in the coastal ecosystems of Lake Superior.     

Trophic interactions among invertebrates in termitaria in the African savanna: a stable isotope approach

Abstract 1. Termites (Isoptera) in tropical savannas are known as ecosystem engineers, affecting the spatial and temporal distribution of water, carbon, cations, and nutrients through their mound structures. Their mounds, however, also offer habitation to diverse taxa and feeding guilds of other invertebrates; a keystone role that has not been properly quantified. 2. The aim of this study was to explore the ecosystem role of termitaria in determining invertebrate diversity and their potential trophic interactions. We used stable isotopes to distinguish termite‐feeding invertebrates from invertebrates merely living in termite mounds under field conditions. 3. The results suggest that inquiline spiders (Arachnida) do not feed on termites directly, but on other invertebrates within the termitaria that are termitophagous, elevating the spiders three trophic levels higher than the termites. 4. This study is the first to demonstrate food web interactions among inquiline invertebrates with a stable isotope approach. It provides evidence that termites play a keystone role in the system by providing habitat for various, trophically interacting invertebrates. These results illustrate a rather unexplored ecosystem property of savanna termites.     

A stable carbon and nitrogen isotope study of lake food webs in Canada's Boreal Plain

1.  Stable carbon and nitrogen isotope and fish stomach content analyses were used to investigate food webs in five relatively undisturbed lakes on the Boreal Plain of Canada. Stable isotope analysis was also used to determine the importance of external and internal carbon sources.
2.  Overlap in the carbon and nitrogen signatures of primary producers made it difficult to determine unambiguously the feeding habits of many invertebrates. However, isotope analysis suggested that external carbon inputs were detectable in the aquatic food chains of the one lake with a short water residence time («1 year). In the other four lakes, with water residence times ≥1 year, autochthonous carbon was the only detectable carbon source in the food webs.
3.  Food webs in these lakes spanned a range of four to five trophic levels. Both invertebrates and fish appeared to eat a variety of food, often feeding at more than one trophic level.
4.  With the exception of one lake (SPH20), top predators in these lakes, northern pike ( Esox lucius ) and fathead minnows ( Pimephales promelas ), occupied similar trophic positions despite large differences in body size and trophic morphology. In SPH20, where there were two additional fish species, pike occupied a higher trophic position. However, all the top predators in each lake appeared to be omnivores and generalists.
5.  The prevalence of omnivory and the apparent generalist feeding habits of fish in these lakes suggest that organisms are flexible in their feeding habits and that these food webs will be resilient to disturbance.     

Terrestrial detritus supports the food webs in lowland intermittent streams of south‐eastern Australia: a stable isotope study

• 1 Large amounts of terrestrial detritus enter many low‐order forested streams, and this organic material is often the major basal resource in the metazoan food webs of such systems. However, despite their apparently low biomass, algae are the dominant food of organisms in a number of aquatic communities which conventionally would have been presumed to be dependent on allochthonous detritus, particularly those in the tropics and also in lowland intermittent streams in arid Australia.
• 2 The dual stable isotope signatures (δ¹³C and δ¹⁵N) of potential primary food sources were compared with the isotopic signatures of common aquatic animals in lowland intermittent streams in south‐eastern Australia, in both spring and summer, to determine whether allochthonous detritus was an important nutritional resource in these systems. The isotopic signatures of the major potential allochthonous plant food sources (Eucalyptus, Phalaris and Juncus) overlapped, but were distinct from algae and the dominant macrophytes growing in the study reaches. The isotopic signatures of biofilm were more spatially and temporally variable than those of the other basal resources.
• 3 Despite allochthonous detritus having relatively high C : N ratios compared to other potential basal resources, results from isosource mixing model calculations demonstrated that this detritus, and the associated biofilm, were the major energy sources assimilated by macroinvertebrate primary consumers in both spring and summer. The importance of these energy sources was also reflected in animals higher in the food web, including predatory macroinvertebrates and fish. These resources were supplemented by autochthonous sources of higher nutritional value (i.e. filamentous algae and macrophytes, which had relatively low C : N ratios) when they became more prolific as the streams dried to disconnected pools in summer.
• 4 The results highlight the importance of allochthonous detritus (particularly from Eucalyptus) as a dependable energy source for benthic macroinvertebrates and fish in lowland intermittent streams of south‐eastern Australia. This contrasts with previous stable isotope studies conducted in lowland intermittent streams in arid Australia, which have reported that the fauna are primarily dependent on autochthonous algae.

     

An assessment of sample processing methods for stable isotope analyses of marine food webs

Carbon and nitrogen stable isotope ratios are commonly used in the study of marine food webs. However, different sample processing methods can influence the measurement of these stable isotope ratios. The purpose of this study is to define an adequate methodology to be used in the construction of whole food webs. It is demonstrated that acidification of the samples results in a decrease in carbon stable isotope values for sedimentary organic matter, suspended particulate organic matter, plankton and invertebrates with carbonated structures. The response was variable for nitrogen isotope ratios. Based on our results we recommend sample acidification for carbon analysis in these compartments where effects of this treatment were observed. We observed a decrease in δ¹³C values after washing with distilled water, so we do not recommend washing with water after acidification. For nitrogen analysis, acidification should be avoided. The various dehydration treatments studied caused significant differences only in nitrogen isotope ratios.     

Food webs in Mediterranean rivers

River food webs are subject to two regimes of longitudinally varying ecological control: productivity and disturbance. Light-limited productivity increases as channels widen downstream. Time windows for growth, however, shrink as discharge increases, substrate particle size decreases, and the frequency of flood-driven bed mobilization increases downstream. Mediterranean rivers are periodically reset by hydrologic events with somewhat predictable timing. Typically, a rainy winter with high river discharge is followed by summer drought with little or no rainfall and slowly declining river flow. The magnitude and timing of winter floods and severity of subsequent summer drought can vary considerably from year to year, however. Episodic scouring floods or prolonged periods of drought are experienced as disturbances, stressors, or opportunities by river biota. The timing, duration, and intensity of these hydrologic controls affect performances of individuals, distribution and abundances of populations, and outcomes and consequences of species interactions. These interactions in turn determine how river food webs will assemble, develop, and reconfigure after disturbance. We discuss how spatial variation in solar radiation and spatial and temporal variations in disturbance affects river food webs under Mediterranean climate seasonality, focusing primarily on long-term observations in the Eel River of northwestern California, USA.     

Food webs in river networks

Food webs and river drainages are both hierarchical networks and complex adaptive systems. How does living within the second affect the first? Longitudinal gradients in productivity, disturbance regimes and habitat structure down rivers have long interested ecologists, but their effects on food web structure and dynamics are just beginning to be explored. Even less is known about how network structure per se influences river and riparian food webs and their members. We offer some preliminary observations and hypotheses about these interactions, emphasizing observations on upstream–downstream changes in food web structure and controls, and introducing some ideas and predictions about the unexplored question of food web responses to some of the network properties of river drainages.     

Untangling the complex issue of dissolved organic carbon uptake: a stable isotope approach

1. We estimated uptake of stream water dissolved organic carbon (DOC) through a whole-stream addition of a ¹³C-DOC tracer coupled with laboratory measurements of bioavailability of the tracer and stream water DOC.
2. The tracer, a leachate of ¹³C-labelled tree tissues, was added to the head waters of White Clay Creek, Pennsylvania, U.S.A., over a 2-h period and followed 1.27 km downstream to generate mass transfer coefficients for DOC lability classes within the tracer.
3. From the longitudinal ¹³C uptake curve, we resolved labile and semi-labile DOC classes within the ¹³C-DOC tracer comprising 82% and 18% of the tracer respectively.
4. Plug-flow laboratory bioreactors colonized and maintained with stream water were used to determine the concentration of stream water DOC fractions that had a similar lability to the labile and semi-labile classes within the tracer and we assumed that stream water DOC and tracer DOC with comparable lability fractions in the bioreactors behaved similarly in the stream, i.e. they had the same mass transfer coefficients.
5. A small fraction (8.6%) of the stream water DOC was labile, travelling 238 m downstream before being taken up. The remaining bioavailable stream water DOC was semi-labile and transported 4.5 km downstream before being taken up. These uptake lengths suggest that the labile DOC is an energy source within a stream reach, while the semi-labile DOC is exported out of the reach to larger rivers and the downstream estuary, where it may provide energy for marine microbial communities or simply be exported to the oceans.     

Microhabitat influence on chironomid community structure and stable isotope signatures in West Greenland lakes

Most functional feeding types are represented within the species rich group of aquatic chironomids. Thus, we hypothesized that different lake types and microhabitats within lakes would (1) host specific chironomid communities and (2) that the individual communities would show specific δ ¹³C stable isotope signatures reflecting the prevailing origin of food source. To test our hypotheses, five lakes in southwest Greenland were investigated at a high taxonomic resolution and with detailed information on δ ¹³C signature of the chironomids and of individual microhabitats (macrophytes, sediment, stones, and profundal). We found that there was a significant difference in δ ¹³C between the chironomid assemblages of freshwater lakes and oligosaline lakes, while assemblages of the littoral microhabitats did not differ significantly. The δ ¹³C of chironomids reflected the wide variety of habitat signals, particularly in the freshwater lakes. Our results indicate that many chironomid taxa are ubiquitous and are found in several microhabitats, suggesting that they can adjust their feeding strategy according to the habitat. The implication is that chironomid assemblage composition has only limited use as indicator of littoral microhabitats in the Arctic. On the other hand, the δ ¹³C signature of fossil chironomids might have a potential as indicator of microhabitats in freshwater lakes.     

A stable isotope study of linkages between stream and terrestrial food webs through spider predation

SUMMARY 1. Transfer of carbon from freshwater to terrestrial ecosystems can occur through predation on adult aquatic insects, but the significance of this trophic pathway to the energetics of riparian communities is poorly understood. We used stable isotopes of carbon and nitrogen to explore linkages between aquatic insect production and the nutrition of web‐building and free‐living spiders alongside two streams in the North Island of New Zealand. 2. δ¹³C values for riparian tree leaves (means for each site = ?32.2 and ?30.3‰) were distinct from those of lichens collected from stream channel rocks and instream algae, both of which were similar (?23.4 to ?22.4‰). δ¹⁵N values for leaves were similar at both sites (?3.4 and ?2.7‰), but algae were considerably more depleted in δ¹⁵N atonesite suggesting significant differences in instream nitrogen sources between the twostreams. 3. Isotope values for potential aquatic prey of spiders indicated that aquatic algal production was their primary carbon source at both sites. Terrestrial invertebrates collected and assumed to be potential prey reflected a range of carbon sources and represented several trophic levels. 4. At one site, δ¹³C values indicated a primarily algae‐aquatic insect pathway of carbon transfer to both web‐building and free‐living spider guilds. The other site appeared to have a primarily terrestrial carbon pathway for the free‐living spider guild, and a mixed aquatic‐terrestrial pathway for the web‐building guild. 5. Overall, web‐building spiders were estimated to obtain around 61% of their body carbon from aquatic production compared with 55% for free‐living spiders. Our findings suggest that consumption of prey derived from aquatic sources can provide significant nutrition for spiders living along some stream channels. This pathway may represent an important feedback mechanism contributing to the energetics of riparian communities at sites where aquatic insect production is high.     

Food choice of Antarctic soil arthropods clarified by stable isotope signatures

Antarctic soil ecosystems are amongst the most simplified on Earth and include only few soil arthropod species, generally believed to be opportunistic omnivorous feeders. Using stable isotopic analyses, we investigated the food choice of two common and widely distributed Antarctic soil arthropod species using natural abundances of ¹³C and ¹⁵N and an isotope labelling study. In the laboratory we fed the isotomid springtail Cryptopygus antarcticus six potential food sources (one algal species, two lichens and three mosses). Our results showed a clear preference for algae and lichens rather than mosses. These results were corroborated by field data comparing stable isotope signatures from the most dominant cryptogams and soil arthropods (C. antarcticus and the oribatid mite Alaskozetes antarcticus). Thus, for the first time in an Antarctic study, we present clear evidence that these soil arthropods show selectivity in their choice of food and have a preference for algae and lichens above mosses.