Relationships between DOC concentration and epilithon stable isotopes in boreal lakes

1. Littoral biota in boreal lakes are known to assimilate epilithon. Being able to predict the stable isotopic composition of these alga will help to identify those systems in which δ¹³C and δ¹⁵N analysis can be used in foodweb investigations of allochthony and biomagnification.
2. In a survey of 15 boreal lakes, the concentration of dissolved organic carbon (DOC) explained 76% of the variation in epilithon δ¹³C, and 86% of the variation in epilithon δ¹⁵N.
3. Because both δ¹³C and δ¹⁵N values were depressed and similar to terrestrial values in humic (high DOC) lakes, it will be more difficult to successfully employ stable isotopic techniques for estimating allochthony in such systems. Lower δ¹⁵N values in humic lakes also indicate that trophic positions estimated by stable isotopes are not directly comparable to those of similar biota inhabiting clearwater lakes, unless autochthonous baseline corrections are made.     

Tracing energy flow in stream food webs using stable isotopes of hydrogen

1. Use of the natural ratios of carbon and nitrogen stable isotopes as tracers of trophic interactions has some clear advantages over alternative methods for food web analyses, yet is limited to situations where organic materials of interest have adequate isotopic separation between potential sources. This constrains the use of natural abundance stable isotope approaches to a subset of ecosystems with biogeochemical conditions favourable to source separation. 2. Recent studies suggest that stable hydrogen isotopes (δD) could provide a robust tracer to distinguish contributions of aquatic and terrestrial production in food webs, but variation in δD of consumers and their organic food sources are poorly known. To explore the utility of the stable hydrogen isotope approach, we examined variation in δD in stream food webs in a forested catchment where variation in δ¹³C has been described previously. 3. Although algal δD varied by taxa and, to a small degree, between sites, we found consistent and clear separation (by an average of 67‰) from terrestrial carbon sources. Environmental conditions known to affect algal δ¹³C, such as water velocity and stream productivity did not greatly influence algal δD, and there was no evidence of seasonal variation. In contrast, algal δ¹³C was strongly affected by environmental factors both within and across sites, was seasonally variable at all sites, and partially overlapped with terrestrial δ¹³C in all streams with catchment areas larger than 10 km². 4. While knowledge of isotopic exchange with water and trophic fractionation of δD for aquatic consumers is limited, consistent source separation in streams suggests that δD may provide a complementary food web tracer to δ¹³C in aquatic food webs. Lack of significant seasonal or spatial variation in δD is a distinct advantage over δ¹³C for applications in many aquatic ecosystems.     

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.     

Trophic fractionation of carbon and nitrogen stable isotopes in Chironomus riparius reared on food of aquatic and terrestrial origin

1. Trophic fractionation was studied in short‐term laboratory feeding experiments with larvae of the deposit‐feeding midge Chironomus riparius. Larvae were fed food of terrestrial (oats, peat) and aquatic origin (Spirulina, Tetraphyll^®). 2. By analysing both whole larvae and isolated gut contents we were able to distinguish between the isotopic signature of recently ingested food and that of assimilated carbon and nitrogen in body tissue. Additionally we studied the effects of microbial conditioning, i.e. the colonisation and growth on food particles of microbes, on the isotopic signal of food resources. 3. Nitrogen fractionation for the different food types ranged from 0.67‰ to 2.68‰ between consumer and diet and showed that isotopic fractionation can be much lower than the value of 3.4‰ that is commonly assumed. 4. Microbial degradation of food particles resulted in an approximate doubling of the δ¹⁵N in 8 days, from 6.24 ± 0.05‰ to 11.36 ± 0.56‰. Values for δ¹³C increased only marginally, from ?20.66 ± 0.11‰ to ?20.34 ± 0.12‰. These results show that microbial conditioning of food may affect dietary isotope signatures (in particular N) and, unless accounted for, could introduce an error in measures of trophic fractionation. Microbial conditioning could well account for some of the variation in fractionation reported in the literature.     

The trophic niche of sculpins Cottus spp. in forage fish assemblages of boreal lakes

We compared the trophic niches of freshwater sculpins Cottus spp. with those of other co-habiting forage fishes in two groups of boreal lakes with distinct habitats and fish communities. Near North Lakes (45° 00′ to 47° 30′ N) were deeper, stratified and contained lake trout Salvelinus namaycush as the apex piscivore, whereas Far North Lakes (51° 10′ to 52° 20′ N) were shallower, did not stratify and contained pike Esox lucius and walleye Sander vitreus as the apex piscivores. Trophic niches of sculpins and other forage fishes were compared based on niche metrics calculated from muscle stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope ratios. In Near North Lakes, sculpins were found almost exclusively in deep, offshore waters and their niche positions reflected a greater reliance on pelagic production (lower δ¹³C) and a higher trophic elevation (higher δ¹⁵N) compared with most other forage fishes. Furthermore, sculpins in Near North Lakes tended to have larger trophic niches (occupied greater area in δ¹³C– δ¹⁵N space), particularly in the food chain (δ¹⁵N) dimension, than other cohabiting forage fishes. In contrast, sculpins in Far North Lakes were commonly found in the nearshore and had trophic niche positions and sizes that were similar to those of the other cohabiting forage fishes. This study illustrates the flexibility in the realised trophic niches of sculpins in relation to habitat availability and fish community composition in boreal lakes.     

Interactions between invading benthivorous fish and native whitefish in subarctic lakes

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Piscivory and trophic position of Anguilla anguilla in two lakes: importance of macrozoobenthos density

The feeding habits of the European eel Anguilla anguilla (>300 mm total length, L _T) were compared in two lakes of different environmental state: Lake Großer Vätersee (LGV), Germany (clear water, mesotrophic and submerged macrophytes), and Lake Vallum (LV), Denmark (turbid, eutrophic and no submerged macrophytes). The density of macrozoobenthos was higher in LV (3500 individuals m⁻²) than in LGV (1500 individuals m⁻²). The abundance of small prey fishes (40–99 mm L _T) was highest in LV. In LV, A. anguilla fed on macrozoobenthos, in particular, chironomid larvae. In LGV, A. anguilla used fishes as the main food component. Stable isotope analyses confirmed the stomach contents dietary results. The estimated mean ± s.d . trophic positions of A. anguilla in LGV (3·7 ± 0·2) was one level higher than those of fish in LV (2·7 ± 0·2). Based on these results, it is concluded that piscivory among A. anguilla was generally controlled by the density of macrozoobenthos. Stable isotope analysis further indicated that A. anguilla may act as integrators between benthic and pelagic food webs when density of insect larvae is low.     

Terrestrial support of pelagic consumers: patterns and variability revealed by a multilake study

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Depth‐specific variation in carbon isotopes demonstrates resource partitioning among the littoral zoobenthos

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Origin and trophic importance of detritus-evidence from stable isotopes in the benthos of a small,temperate estuary

Ratios of ¹³C/¹²C and ¹⁵N/¹⁴N were measured in dissolved inorganic carbon (DIC), marginal vegetation, benthic macrodetritus (diameter > 1 mm) and selected invertebrate consumers in the Gamtoos estuary, South Africa to: (1) trace the provenance of benthic detrital deposits, and (2) determine the extent to which three abundant species of macroinvertebrates utilise this resource. DIC was strongly depleted in ¹³C with average ¹³C values (–9.5±0.5) being typical of limnetic waters. Benthic detrital particles (¹³C–24.1±0.3) originated mainly from marginal vegetation (¹³C–25.7±0.3), but their slightly elevated carbon ratio suggests additional input from ¹³C-rich sources-possibly C₄ plants cultivated on the floodplain. Populations of the fossorial ghost shrimp Callianassa kraussi, the bentho-pelagic amphipod Grandidierella lignorum and the epifaunal crab Hymenosoma robiculare together account for 96% of total benthic biomass in the upper regions of this estuary. Marked differences in trophic niches were evident among these three consumer species. Ghost shrimp (¹³C –32.5±0.3) foraged by filter-feeding on fine suspended particulate organic matter (¹³C–31.2±0.5). Amphipods (¹³C–28.0±0.6) utilised some benthic detritus but fed mainly on suspended material. Only the relatively rare crabs (¹³C–23.8±1.5) appeared to utilise benthic detrital particles to any significant extent. In the benthic consumer community of the upper Gamtoos estuary, suspension feeders make up 98% of biomass and thus clearly dominate over deposit feeders. This can be traced to the low contribution of higher plants (c. 13%) to overall carbon production, and detritus originating from macrophytes is consequently relatively unimportant in supporting invertebrate secondary production in this particular system.     

Metabolism and foraging strategies of mid‐latitude mesozooplankton during cyanobacterial blooms as revealed by fatty acids,amino acids,and their stable carbon isotopes

Increasing sea surface temperatures (SST) and blooms of lipid‐poor, filamentous cyanobacteria can change mesozooplankton metabolism and foraging strategies in marine systems. Lipid shortage and imbalanced diet may challenge the build‐up of energy pools of lipids and proteins, and access to essential fatty acids (FAs) and amino acids (AAs) by copepods. The impact of cyanobacterial blooms on individual energy pools was assessed for key species temperate Temora longicornis and boreal Pseudo‐/Paracalanus spp. that dominated field mesozooplankton communities isolated by seasonal stratification in the central Baltic Sea during the hot and the cold summer. We looked at (a) total lipid and protein levels, (b) FA trophic markers and AA composition, and (c) compound‐specific stable carbon isotopes (δ¹³C) in bulk mesozooplankton and in a subset of parameters in particulate organic matter. Despite lipid‐poor cyanobacterial blooms, the key species were largely able to cover both energy pools, yet a tendency of lipid reduction was observed in surface animals. Omni‐ and carnivory feeding modes, FA trophic makers, and δ¹³C patterns in essential compounds emphasized that cyanobacterial FAs and AAs have been incorporated into mesozooplankton mainly via feeding on mixo‐ and heterotrophic (dino‐) flagellates and detrital complexes during summer. Foraging for essential highly unsaturated FAs from (dino‐) flagellates may have caused night migration of Pseudo‐/Paracalanus spp. from the deep subhalocline waters into the upper waters. Only in the hot summer (SST>19.0°C) was T. longicornis submerged in the colder subthermocline water (~4°C). Thus, the continuous warming trend and simultaneous feeding can eventually lead to competition on the preferred diet by key copepod species below the thermocline in stratified systems. A comparison of δ¹³C patterns of essential AAs in surface mesozooplankton across sub‐basins of low and high cyanobacterial biomasses revealed the potential of δ¹³C‐AA isoscapes for studies of commercial fish feeding trails across the Baltic Sea food webs.     

Differences in feeding behaviour among Chironomus species revealed by measurements of sulphur stable isotopes and cadmium in larvae

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River habitat homogenisation enhances trophic competition and promotes individual specialisation among young of the year fish

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Beyond trophic morphology: stable isotopes reveal ubiquitous versatility in marine turtle trophic ecology

The idea that interspecific variation in trophic morphology among closely related species effectively permits resource partitioning has driven research on ecological radiation since Darwin first described variation in beak morphology among Geospiza. Marine turtles comprise an ecological radiation in which interspecific differences in trophic morphology have similarly been implicated as a pathway to ecopartition the marine realm, in both extant and extinct species. Because marine turtles are charismatic flagship species of conservation concern, their trophic ecology has been studied intensively using stable isotope analyses to gain insights into habitat use and diet, principally to inform conservation management. This legion of studies provides an unparalleled opportunity to examine ecological partitioning across numerous hierarchical levels that heretofore has not been applied to any other ecological radiation. Our contribution aims to provide a quantitative analysis of interspecific variation and a comprehensive review of intraspecific variation in trophic ecology across different hierarchical levels marshalling insights about realised trophic ecology derived from stable isotopes. We reviewed 113 stable isotope studies, mostly involving single species, and conducted a meta‐analysis of data from adults to elucidate differences in trophic ecology among species. Our study reveals a more intricate hierarchy of ecopartitioning by marine turtles than previously recognised based on trophic morphology and dietary analyses. We found strong statistical support for interspecific partitioning, as well as a continuum of intraspecific trophic sub‐specialisation in most species across several hierarchical levels. This ubiquity of trophic specialisation across many hierarchical levels exposes a far more complex view of trophic ecology and resource‐axis exploitation than suggested by species diversity alone. Not only do species segregate along many widely understood axes such as body size, macrohabitat, and trophic morphology but the general pattern revealed by isotopic studies is one of microhabitat segregation and variation in foraging behaviour within species, within populations, and among individuals. These findings are highly relevant to conservation management because they imply ecological non‐exchangeability, which introduces a new dimension beyond that of genetic stocks which drives current conservation planning. Perhaps the most remarkable finding from our data synthesis is that four of six marine turtle species forage across several trophic levels. This pattern is unlike that seen in other large marine predators, which forage at a single trophic level according to stable isotopes. This finding affirms suggestions that marine turtles are robust sentinels of ocean health and likely stabilise marine food webs. This insight has broader significance for studies of marine food webs and trophic ecology of large marine predators. Beyond insights concerning marine turtle ecology and conservation, our findings also have broader implications for the study of ecological radiations. Particularly, the unrecognised complexity of ecopartitioning beyond that predicted by trophic morphology suggests that this dominant approach in adaptive radiation research likely underestimates the degree of resource overlap and that interspecific disparities in trophic morphology may often over‐predict the degree of realised ecopartitioning. Hence, our findings suggest that stable isotopes can profitably be applied to study other ecological radiations and may reveal trophic variation beyond that reflected by trophic morphology.     

Restored oyster reefs and living shorelines can augment predator trophic dynamics

Coastal habitat loss has led to declines in many higher trophic level predators. These declines can be mitigated through habitat restoration, which putatively enhances predator populations and trophic dynamics by creating foraging opportunities in reestablished habitat, but this lacks empirical support. Here, we assess the prediction restored coastal habitat supports sportfish feeding relationships similar to natural habitat, at restored oyster reefs (Crassostrea virginica) and living shoreline habitats in Florida, U.S.A. Stable isotopes and gut contents were analyzed from young sportfish (i.e. predatory fish targeted by anglers) collected at control and restored sites for up to 3 years. The influence of habitat features, predator size, and prey availability on carbon and nitrogen isotope values were examined using a model species (mangrove/gray snapper [Lutjanus griseus]). In summary, sportfish species had distinct isotopic values, consuming similar prey but in different proportions between habitats. Prey abundance and richness, and reef height were influential predictors of L. griseus stable isotope values, with restored reefs contributing to their diet more than controls according to Bayesian mixing models. The trophic niche area of L. griseus and their predominant prey achieved trophic equivalence between restored and natural oyster reefs within 1 year following restoration. Stabilized living shorelines attained trophic equivalency to natural shorelines, but may take longer to accrue prey for some species. These findings generate fundamental ecological knowledge and actionable science, including targets, timelines, and indicator species, that natural resource managers and restoration practitioners can use to assess trophic structure and success of coastal habitat restoration.     

Patterns of macroinvertebrate abundance in inland saline wetlands: a trophic analysis

We used stable isotopes and gut-content analysis to compare trophic relations of macroinvertebrates between two types of saline wetlands in the Laramie Basin, Wyoming, USA. Amphipods (Hyallela azteca), chironomid larvae, and predatory insects (mainly Zygopteran larvae) occurred in both wetland types. However, in oligosaline wetlands (0.5–5‰ total dissolved solids) amphipods were dominant, whereas in mesosaline wetlands (5–18‰) amphipods were scarce and chironomid larvae and predatory insects were much more abundant. Salinity alone seemed inadequate to explain these differences, so we examined trophic interactions to address three main questions: (1) why are predatory insects more abundant in mesosaline wetlands, (2) why are chironomid larvae less abundant in oligosaline wetlands, and (3) why are amphipods uncommon in mesosaline wetlands? Zygopteran larvae ate mainly chironomid larvae and zooplankton, and did not eat amphipods; chironomid larvae were an average 93 dry mass of zygopteran intake. Guts of amphipods contained no animal parts and little vascular plant tissue, but rather mainly amorphous detritus. Chironomid larvae ate amorphous detritus as well as diatoms. There is potential for competition between amphipods and chironomids, but the nature of amorphous detritus in these wetlands needs further study. Because amphipods appear unavailable as prey for predatory insects, top-down impacts on chironomid larvae may be greater in oligosaline than in mesosaline wetlands. Our results suggest that (1) predatory zygopteran larvae and hemipterans are more abundant in mesosaline wetlands due to more abundant chironomid prey, (2) chironomids are less common in oligosaline wetlands because of both competition with amphipods and greater per capita consumption by predatory insects, and (3) amphipods are scarce in mesosaline wetlands because, lacking a resting stage and mode of direct dispersal, they do not cope well with extremes created by hydrologic instability in mesosaline wetlands. These mechanisms may apply to wetlands in other regions where similar patterns of invertebrate community structure have been reported.     

Variable littoral‐pelagic coupling as a food‐web response to seasonal changes in pelagic primary production

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A comparison of the trophic ecology of the crayfishes (Orconectes nais (Faxon) and Orconectes neglectus (Faxon)) and the central stoneroller minnow ( Campostoma anomalum (Rafinesque)): omnivory in a tallgrass prairie stream

Omnivorous fish, such as the central stoneroller minnow (Campostoma anomalum(Rafinesque)), and crayfish often play important roles in the trophic dynamics of streams. The trophic role of these two omnivores has not been compared within a system even though they both consume algae, detritus and invertebrates and often co-occur in streams in the Midwestern United States. Natural abundance of ¹⁵N and ¹³C isotopes and a whole stream ¹⁵N-labeled ammonium chloride release were used to compare the trophic ecology of the central stoneroller minnow (Campostoma anomalum (Rafinesque)) and two species of crayfish (Orconectes neglectus (Faxon) and Orconectes nais (Faxon)) in a tallgrass prairie stream. The ¹⁵N and ¹³C values of Orconectes spp. were more similar to coarse benthic organic matter (CBOM) and filamentous green algae than to invertebrates, fine benthic organic matter (FBOM), and periphyton. Values for ¹⁵N and ¹³C in C. anomalum were more similar to grazer and collector invertebrates and filamentous green algae than to FBOM and periphyton. Results from a ¹⁵N tracer release also indicated a portion of algae and/or invertebrates were a component of nitrogen assimilated in Orconectes spp. and C. anomalum diets. Gut contents of C. anomalum were also analyzed. In contrast to stable isotope data, amorphous detritus was a significant component of C. anomalum guts, followed by diatoms and filamentous green algae. A significant percentage of invertebrate material was found in C. anomalum guts sampled in the spring. Experiments were conducted in artificial streams to determine if Orconectes spp. and C. anomalum could reduce epilithic algal biomass in small streams. Algal biomass on clay tile substrata was decreased relative to controls in artificial stream channels containing O. neglectus (3.4 fold, p=0.0002), C. anomalum (2.1 fold, p=0.0012), and both species combined (3.0 fold, p=0.0003). Results indicate that Orconectes spp. are functioning more as algal and detrital processors than as predators in Kings Creek. Isotope and gut content data show that C. anomalum includes invertebrates as well as algae and detritus in its diet. Both species have the potential to affect algal biomass and are important omnivores in the stream food web.     

The effects of preservation on fish tissue stable isotope signatures

The effects of formalin and ethanol preservation on the δ¹³C and δ¹⁵N isotope signatures of Arctic charr Salvelinus alpinus muscle tissue were examined. The lipid content of the tissue samples studied ranged from 3·6 to 6·1% and was not correlated with the magnitude of observed isotopic shifts in preserved samples. Ethanol and formalin significantly depleted and enriched, respectively, the δ¹³C isotope signatures of preserved tissues when compared to control samples. Ethanol did not significantly enrich δ¹⁵N signatures in comparison to controls, whereas formalin did. A meta-analysis of multiple species effects further demonstrated significant preservation effects in fish tissue. Statistical analysis of data obtained by correcting preserved tissue isotope signatures with literature, bootstrapped or meta-analysis derived correction factors demonstrated significant differences between corrected and control sample isotope signatures or failure to produce a unity slope when the data sets were regressed against one another. Species-specific, bootstrapped linear correction models resulted in no such errors. Results suggest that species-specific correction methods should be used for fishes because of the known wide variation in fish tissue lipid content and composition. Accordingly, the use of pilot studies will be required to develop correction factors that properly adjust for preservation effects when interpreting temporal patterns in historic analyses of food webs.