Trophic relationships of juvenile blue crabs (Callinectes sapidus) in estuarine habitats

Salt marshes and shallow-water macroalgal beds are known to provide nursery habitat for many species of fish and invertebrates. The role of these habitats as refuge from predation is well established, but the degree to which indigenous primary production within the nursery provides food for growth and development of estuarine species remains unresolved. In this study, we tested the hypothesis that juvenile blue crabs depend on indigenous primary production, directly or indirectly, during their entire stay within the nursery. To test this hypothesis, we conducted isotopic studies and stomach content analyses of juveniles from habitats near the mouth of Delaware Bay and from an adjacent lagoonal estuary (ca. 39.5° N, 75.1° W). Primary producers, marsh detritus, various life-history stages of blue crabs and potential prey species were sampled in the main estuary and in an adjacent marsh during the summer and early fall of two consecutive years. Newly settled juveniles (<15 mm carapace width) from the marsh were about 1.8‰ lighter in carbon (−17.2‰) relative to larger juveniles from the marsh (15–30 mm carapace width) and appeared to have retained a carbon isotopic signature indicative of the phytoplankton-based food web associated with larval stages. However, the signature of juveniles changed as a function of size. Large juveniles and crabs >60 mm were enriched in δ¹³C (−14.7 ± 0.1‰) compared to small crabs, suggesting a gradual shift in diet from a planktonic to a detritus-based food web with increasing size. As with crabs from Delaware Bay, the δ¹³C signature of juvenile crabs sampled from macroalgal beds in the lagoonal estuary (Rehoboth Bay) changed as a function of size. Also, δ¹³C ratios of crabs varied among the various species of macroalgae. The δ¹⁵N composition of primary producers in the marsh and main estuary also was reflected in the δ¹⁵N values of crabs and other benthic consumers in the respective habitats. Results of stomach-content analysis in this study were consistent with isotope data. Observed changes in prey preferences were related to changes in size of juvenile crabs and also differed among habitats. Gut content analyses of the three size classes of juveniles in macroalgal beds from Rehoboth Bay indicated that the crabs depend heavily on various amphipod species that occur on the seaweeds. These amphipods graze directly on the macroalgae and are among the most abundant invertebrates in the macroalgal beds. This implies a direct trophic relationship between the juvenile crabs and the macroalgae. In summary, our study provides strong evidence that the value of nursery areas such as salt marshes and macroalgal beds goes beyond that of providing refuge from predation, and that species using these nurseries (e.g. juvenile blue crabs) are ultimately dependent on primary production originating in benthic plants indigenous to the nursery.     

Can stable isotope (δ<Superscript>13</Superscript>C and δ<Superscript>15</Superscript>N) measurements of little auk (<Emphasis Type="Italic">Alle alle</Emphasis>) adults and chicks be used to track changes in high-Arctic marine foodwebs?

The little auk (Alle alle), a small and abundant planktivorous seabird that breeds in the high Arctic, has the potential to be used as a monitor of the composition and abundance of lower trophic-level zooplankton. We investigated age- and sex-related sources of variation in diet and stable isotope (δ¹³C and δ¹⁵N) values of little auks breeding in Spitsbergen during the summer of 2002 to evaluate this possibility. Stable isotope profiles of both adult and chick blood changed over the breeding season, with blood δ¹⁵N values increasing and δ¹³C values decreasing. This could represent a switch to higher trophic-level prey derived from more pelagic sources. However, while chick blood δ¹³C values followed those values in their meals, this was not the case for blood δ¹⁵N values, suggesting additional physiological mechanisms influencing blood δ¹⁵N values in growing chicks. Chicks had consistently lower δ¹⁵N values than their parents, which may indicate they were being fed on lower trophic-level prey items or may alternatively reflect complexities in chick blood δ¹⁵N values through the growth period. These results have several important implications for use of stable isotope analysis as a tool to detect changes in seabird diet and availability of lower trophic-level prey in high-Arctic marine environments. Until physiological aspects of stable isotope discrimination are well understood, we caution against using chicks of this seabird as any form of isotopic monitor.     

Benthic-pelagic trophic interactions within the fish assemblage in the western Bering Sea shelf area according to stomach content analysis and ratios of C and N stable isotopes

The composition, abundance, diet and trophic status of zooplankton, bottom invertebrates, fish and nekton were analyzed based on the data collected by the staff of the TINRO-Center during complex bottom trawl catches on the Bering Sea shelf in the fall of 2004. The stomach contents of mass fish species were analyzed and the nitrogen and carbon isotopic composition of 36 mass species of plankton, benthos, nekton and nektobenthos, which together make up the basis of pelagic and bottom communities, was determined. It was found that zooplankton noticeably differ from benthic invertebrates in carbon isotopic composition: δ¹³C values in zooplankton varied from −20.3‰ to −17.9‰; in benthos—from −17.5‰to −13.0‰; and in fish—from −19.2‰ (juvenile walleye pollock) to −15.3‰ (saffron cod). The levels of ¹³C isotope in the tissues of fish depended mostly on the share of pelagic or benthic animals in their diet. δ¹⁵N values in the studied species ranged from 8.6‰ (in sea urchins) to 17.2‰ (in large Pacific cods), which corresponds to a trophic level of 2.8. Obviously the δ¹⁵N values reflect the degree of predation and generally show the ratio of primary, secondary and tertiary consumers in a fish’s diet. Trophic interactions manifest a high degree of interdependence between benthic and pelagic communities (even without taking into account such lower components of the food web as phytoplankton, bacteria, and protozoa) occurring in most nektonic species that depend on both bottom and pelagic food.     

Trophic polymorphism in bluegill sunfish (<Emphasis Type="Italic">Lepomis macrochirus</Emphasis>) introduced into Lake Biwa: evidence from stable isotope analysis

Trophic polymorphism was recently reported in introduced bluegill (Lepomis macrochirus) in Lake Biwa, Japan, where three morphs are specialized in benthic invertebrates (benthivorous type), submerged aquatic plants (herbivorous type), and zooplankton (planktivorous type). We evaluated the long-term effects of food resource utilization by these trophic morphs using stable isotope ratios, δ¹⁵N and δ¹³C. A significant difference in δ¹⁵N was found between the benthivorous and planktivorous types. The planktivorous type had the higher δ¹⁵N value, which corresponded with the value expected from its prey, zooplankton. The lower δ¹⁵N value of the benthivorous type would be derived from the lower δ¹⁵N values of benthic prey organisms compared to zooplankton. These results support previous findings that the benthivorous and planktivorous types have different food resource utilization. In contrast, the δ¹⁵N and δ¹³C values of the herbivorous type were distinctly different from the expected values, indicating that this type was unlikely to utilize aquatic plants substantially, contradicting the results of the dietary analysis.     

Evaluating the utility of stable isotope analyses of archived freshwater sample materials

We evaluated the potential utility of stable isotope analysis of tissues commonly archived by aquatic biologists. Previous studies with chemically preserved samples have shown contradictory results, which present an obstacle for the use of archived sample materials. We tested the effects of ethanol and formalin preservation on zooplankton and of ethanol on benthic macroinvertebrate δ¹³C and δ¹⁵N values. We found that neither formalin nor ethanol had a significant effect on δ¹³C and δ¹⁵N values of preserved zooplankton. Nor did ethanol significantly affect δ¹³C or δ¹⁵N values of macroinvertebrates. However, ethanol preservation slightly, but significantly decreased C:N ratios of both zooplankton and macroinvertebrates, probably reflecting some extraction of lipids. Overall, the effects of preservatives on δ¹³C and δ¹⁵N values that we observed were minor. We also compared δ¹³C and δ¹⁵N values analysed from roach scales and perch operculum bones with those analysed from muscle tissue. Decalcification of scales and operculum bones only slightly improved our comparison to muscle tissue δ¹³C and δ¹⁵N values. Decalcified scales had slightly higher δ¹³C and lower δ¹⁵N values. Similarly, decalcified operculum bones showed slightly increased δ¹³C and decreased δ¹⁵N values to those for fish muscle. Our results confirm that scales and operculum bones can provide a suitable proxy for fish muscle in isotope studies with minor correction. We conclude that various archived sample materials can indeed be used with confidence for historical reconstructions of freshwater food webs by stable isotope analysis. Handling editor: K. Martens     

Sympatric variability of isotopic baselines influences modeling of fish trophic patterns

Stable isotope signatures of freshwater snails and mussels have been established as a convenient baseline measurement at the primary consumer level for food-web coupling studies. We measured δ¹⁵N and δ¹³C of primary consumers, including mussels (Anodonta woodiana, Cristaria plicata, and Unio douglasiae), snails (Bellamya aeruginosa and Hippeutis sp.), and zooplankton from the same habitat within a shallow eutrophic lake. Primary consumers showed positive relationship between δ¹⁵N and δ¹³C, indicating a linkage between planktonic and benthic habitats in this system. The variation in isotope ratios was higher in short-lived primary consumers (zooplankton) compared with the long-lived primary consumers (mussels and snails), suggesting limited availability of short-lived primary consumers as isotopic baselines in aquatic food-web assessment. Significant differences in isotope ratios were also found among three species of mussels, and when using these mussels separately as pelagic baselines to calculate trophic position and contribution of planktonic and benthic sources of fishes, bias and even misestimates were observed. This finding suggests that caution is needed when multiple primary consumers coexist in the same habitat, and it is important to assess potential effects of different baselines used.     

Effect of light and zooplankton on skeletal δ13C values in the eastern Pacific corals Pavona clavus and Pavona gigantea

 Skeletal δ¹³C levels in symbiotic reef corals are believed to be predominantly influenced by metabolic fractionation. Therefore, environmental variables influencing coral metabolism should also affect skeletal δ¹³C levels. To test this hypothesis, we measured the effects of light (which drives photosynthesis) and relative zooplankton levels (heterotrophy) on skeletal δ¹³C values in the corals Pavona clavus and P. gigantea at two depths (1 m and 7 m). For both species, decreases in light or increases in zooplankton resulted in significant decreases in skeletal δ¹³C levels. A significant decrease in δ¹³C values with depth was observed in Pavona gigantea only. Thus, light and zooplankton directly affect coral skeletal δ¹³C values, supporting the hypothesis that metabolic fractionation significantly contributes to skeletal δ¹³C levels. Simultaneous decreases in both light and zooplankton resulted in decreases in skeletal δ¹³C values, reflecting decreases in light. In Pavona clavus, intra-annual variation in skeletal δ¹³C values over one year correlated with seasonal changes in irradiance. Further study is needed to resolve how skeletal δ¹³C values vary at intermediate levels of irradiance and zooplankton, and in other coral species. Accepted: 14 July 1998     

Contributions of Different Organic Carbon Sources to <Emphasis Type="Italic">Daphnia</Emphasis> in the Pelagic Foodweb of a Small Polyhumic Lake: Results from Mesocosm DI<Superscript>13</Superscript>C-Additions

Abstract Freshwater ecosystems derive organic carbon from both allochthonous and autochthonous sources. We studied the relative contributions of different carbon sources to zooplankton in a small, polyhumic, steeply stratified lake, using six replicate surface-to-sediment enclosures established during summer and autumn 2004. We added ¹³C-enriched bicarbonate to the epilimnion of half the enclosures for three weeks during each season and monitored carbon stable isotope ratios of DIC, DOC, POC and Daphnia, along with physical, chemical and biological variables. During summer, ¹³C-enriched DIC (δ¹³C up to 44 ± 7.2‰) was soon taken up by phytoplankton (δ¹³C up to −5.1 ± 13.6‰) and was transmitted to Daphnia (δ¹³C up to −1.7 ± 7.2‰), demonstrating consumption of phytoplankton. In contrast, during autumn, ¹³C-enriched DIC (δ¹³C up to 56.3 ± 9.8‰) was not transmitted to Daphnia, whose δ¹³C became progressively lower (δ¹³C down to −45.6 ± 3.3‰) concomitant with decreasing methane concentration. Outputs from a model suggested phytoplankton contributed 64–84% of Daphnia diet during summer, whereas a calculated pelagic carbon mass balance indicated only 30–40% could have come from phytoplankton. Although autumn primary production was negligible, zooplankton biomass persisted at the summer level. The model suggested methanotrophic bacteria contributed 64–87% of Daphnia diet during autumn, although the calculated carbon mass balance indicated a contribution of 37–112%. Thus methanotrophic bacteria could supply virtually all the carbon requirement of Daphnia during autumn in this lake. The strongly ¹³C-depleted Daphnia values, together with the outputs from the models and the calculated carbon mass balance showed that methanotrophic bacteria can be a greater carbon source for Daphnia in lakes than previously suspected.     

Stable isotope analysis of the origins of zooplankton carbon in lakes of differing trophic state

Carbon stable isotope analysis was carried out on zooplankton from 24 United Kingdom lakes to examine the hypothesis that zooplankton dependence on allochthonous sources of organic carbon declines with increasing lake trophy. Stable isotope analysis was also carried out on particulate and dissolved organic matter (POM and DOM) and, in 11 of the lakes, of phytoplankton isolates. In 21 of the 24 lakes, the zooplankton were depleted in ¹³C relative to bulk POM, consistent with previous reports. δ¹³C for POM showed relatively little variation between lakes compared to high variation in values for DOM and phytoplankton. δ¹³C values for phytoplankton and POM converged with increasing lake trophy, consistent with the expected greater contribution of autochthonous production to the total organic matter pool in eutrophic lakes. The difference between δ¹³C for zooplankton and that for POM was also greatest in oligotrophic lakes and reduced in mesotrophic lakes, in accordance with the hypothesis that increasing lake trophic state leads to greater dependence of zooplankton on phytoplankton production. However, the difference increased again in hypertrophic lakes, where higher δ¹³C values for POM may have been due to greater inputs of ¹³C-enriched organic matter from the littoral zone. The very wide variation in phytoplankton δ¹³C between lakes of all trophic categories made it difficult to detect robust patterns in the variation in δ¹³C for zooplankton. Received: 2 November 1998 / Accepted: 3 December 1999     

Spatial and temporal trends in stable carbon and oxygen isotope ratios of juvenile winter flounder otoliths

Isotopic ratios of fish otoliths have been used in numerous studies as natural tags or markers to aid in the study of connectivity among fish populations. We investigated the use of spatial and temporal changes in the stable carbon and oxygen isotope ratios of otoliths to differentiate juvenile habitats of winter flounder (Pseudopleuronectes americanus). Young-of-the-year (YOY) juvenile winter flounder were collected annually over a three-year period from 18 stations along the coast of Rhode Island, USA. Sagittal otoliths were removed from fish and analyzed for stable carbon (¹³C/¹²C or δ¹³C) and oxygen (¹⁸O/¹⁶O or δ¹⁸O) isotope ratios using continuous flow isotope ratio mass spectrometry. Differences in isotope ratios were observed among stations and along salinity gradients in the Narragansett Bay estuary and an estuarine river system (Narrow River). Overall, the isotope ratio patterns observed among stations were consistent over the three sampling years; however, differences were noted in isotope ratios and the magnitude of the isotope ratio gradients among years. Significant positive correlations were noted between salinity and δ¹³C for two of the three years. For each of the three years sampled there was a highly significant positive correlation (2002, r = 0.93, P < 0.01; 2003, r = 0.85, P < 0.01; 2004, r = 0.97, P < 0.01) between δ¹⁸O and the salinity of the collection site. Also, there was a significant negative correlation between the number of months of above average river flow and δ¹⁸O for the three sampling years (r = 0.99, P < 0.05). These findings suggest that yearly changes in the volume of freshwater inputs to these estuarine habitats may be related to the differences observed in otolith δ¹⁸O isotope ratios. Because of these year-to-year differences, sampling of each cohort may be necessary in order to use this isotopic technique for winter flounder connectivity studies.     

Changes in trophic linkages to shortfin eels (<Emphasis Type="Italic">Anguilla australis</Emphasis>) since the collapse of submerged macrophytes in Lake Ellesmere,New Zealand

Lake Ellesmere (Te Waihora) is a nationally important coastal brackish lake in New Zealand, however degradation in water quality and loss of submerged macrophytes over past decades have raised concerns in regards to the declining status of the lake’s commercial and customary fisheries, predominantly targeted at shortfin eels (Anguilla australis). We investigated foodweb dynamics and trophic linkages to shortfin eels in Lake Ellesmere using a combination of abundance assessments, dietary studies, and stable isotope analyses. Data from our study are compared with historical data sets on benthic invertebrate community composition and shortfin eel diets to trace changes in the trophic linkages to top predators that have occurred since the late 1960s. Stable isotope analyses indicate that the foodweb is predominantly driven by epipelic and phytoplankton derived carbon sources, although it was difficult to discriminate between these two carbon pools because of wind-driven resuspension of lake sediments. Comparison of our survey results with historical data sets indicates a clear shift in benthic biota from being dominated by phytofaunal species such as Potamopyrgus antipodarum (comprising 90% of total invertebrate biomass) during the 1960s, to now being almost entirely comprised of subterranean species such as Chironomus zealandicus and oligochaetes (together comprising 82% of total invertebrate biomass). This shift in benthic communities has resulted in significant changes in the size-specific diet of juvenile shortfin eels (<400 mm) from those reported for Lake Ellesmere during the mid 1970s, with Chironomus larvae now comprising 65% of the diets of juvenile eels, whereas historically P. antipodarum was the dominant food item (>30% of total biomass). This shift towards foraging on smaller sediment-dwelling species could have implications for juvenile eel bioenergetics, and may help explain why juvenile shortfin growth rates have significantly decreased in past decades. Juvenile shortfins now appear to switch to foraging on preyfish (mainly common bullies, Gobiomorphus cotidianus) at a smaller size (≈400 mm) than historically recorded (>500 mm). Dietary and stable isotope signatures indicated that small shortfins (100–299 mm) have considerable overlap in trophic position (δ¹³C = −20.4‰, δ¹⁵N = 13.6‰) with common bullies (δ¹³C = −20.5‰, δ¹⁵N = 13.7‰), the dominant fish in Lake Ellesmere (92% of total abundance CPUE), potentially indicating that these two species may directly compete for food resources. These findings again highlighted the importance of C. zealandicus in sustaining the fish populations of the lake. Handling editor: S. Declerck     

Mechanism for the small-scale movement of carbon among estuarine habitats: organic matter transfer not crab movement

In theory, carbon is highly mobile in aquatic systems. Recent evidence from carbon stable isotopes of crabs (Parasesarma erythrodactyla and Australoplax tridentata), however, shows that in subtropical Australian waters, measurable carbon movement between adjacent mangrove and saltmarsh habitats is limited to no more than a few metres. We tested whether the pattern in crab δ¹³C values across mangrove and saltmarsh habitats was explained by crab movement, or the movement of particulate organic matter. We estimated crab movement in a mark–recapture program using an array of pitfall traps on 13 transects (a total of 65 traps) covering an area of 600 m² across the interface of these two habitats. Over a 19-day period, the majority of crabs (91% for P. erythrodactyla, 93% for A. tridentata) moved <2 m from the place of initial capture. Crab movement cannot, therefore, explain the patterns in δ¹³C values of crabs. δ¹³C values of detritus collected at 2-m intervals across the same habitat interface fitted a sigmoidal curve of a similar form to that fitting the δ¹³C values of crabs. δ¹³C values of detritus were 2–4‰ more depleted in saltmarsh (−18.5±0.6‰), and 4–7‰ more depleted in mangroves (−25.9±0.1‰) than δ¹³C values of crabs recorded previously in each habitat. Assimilation by crabs of very small detrital fragments or microphytobenthos, more enriched in ¹³C, may explain the disparity in δ¹³C values. Nevertheless, the pattern in δ¹³C values of detritus suggests that crabs obtain their carbon from up to several metres away, but without themselves foraging more then a metre or so from their burrow. Such detailed measurements of carbon movement in estuaries provide a spatially explicit understanding of the functioning of food webs in saltmarsh and mangrove habitats.     

Patterns of stable carbon isotope turnover in gag, <Emphasis Type="Italic">Mycteroperca microlepis</Emphasis>, an economically important marine piscivore determined with a non-lethal surgical biopsy procedure

To determine the feasibility of using stable isotopes to track diet shifts in wild gag, Mycteroperca microlepis, populations over seasonal timescales, we conducted a repeated measures diet-shift experiment on four adult gag held in the laboratory. Fish were initially fed a diet of Atlantic mackerel, Scomber scombrus, (mean δ¹³C = −21.3‰ ± 0.2, n = 20) for a period of 56 days and then shifted to a diet of pinfish, Lagodon rhomboids, (mean δ¹³C = −16.6‰ ± 0.6, n = 20) for the 256 day experiment. We developed a non-lethal surgical procedure to obtain biopsies of the muscle, liver, and gonad tissue monthly from the same four fish. We then determined the δ¹³C value of each tissue by isotope ratio mass spectrometry. For the gonad tissue we used the relationship between C/N and lipid content to correct for the influence of lipids on δ¹³C value. We observed a significant shift in the δ¹³C values of all of the tissues sampled in the study. Carbon turnover rates varied among the three tissues, but the shift in diet from mackerel to pinfish was clearly traceable through analysis of δ¹³C values. The turnover rates for muscle tissue were 0.005‰ day⁻¹, and for gonad tissue was 0.009‰ day⁻¹. Although it is generally thought that tissue turnover rates in ectotherms are driven primarily by growth, we found that metabolic rate can be a major factor driving tissue turnover in adult gag.     

Disentangling effects of growth and nutritional status on seabird stable isotope ratios

A growing number of studies suggest that an individual’s physiology affects its carbon and nitrogen stable isotope signatures, obscuring a signal often assumed to be only a reflection of diet and foraging location. We examined effects of growth and moderate food restriction on red blood cell (RBC) and feather δ¹⁵N and δ¹³C in rhinoceros auklet chicks (Cerorhinca monocerata), a piscivorous seabird. Chicks were reared in captivity and fed either control (75 g/day; n = 7) or ~40% restricted (40 g/day; n = 6) amounts of high quality forage fish. We quantified effects of growth on isotopic fractionation by comparing δ¹⁵N and δ¹³C in control chicks to those of captive, non-growing subadult auklets (n = 11) fed the same diet. To estimate natural levels of isotopic variation, we also collected blood from a random sample of free-living rhinoceros auklet adults and chicks in the Gulf of Alaska (n = 15 for each), as well as adult feather samples (n = 13). In the captive experiment, moderate food restriction caused significant depletion in δ¹⁵N of both RBCs and feathers in treatment chicks compared to control chicks. Growth also induced depletion in RBC δ¹⁵N, with chicks exhibiting lower δ¹⁵N when they were growing the fastest. As growth slowed, δ¹⁵N increased, resulting in an overall pattern of enrichment over the course of the nestling period. Combined effects of growth and restriction depleted δ¹⁵N in chick RBCs by 0.92‰. We propose that increased nitrogen-use efficiency is responsible for ¹⁵N depletion in both growing and food-restricted chicks. δ¹⁵N values in RBCs of free-ranging auklets fell within a range of only 1.03‰, while feather δ¹⁵N varied widely. Together, our captive and field results suggest that both growth and moderate food restriction can affect stable isotope ratios in an ecologically meaningful way in RBCs although not feathers due to greater natural variability in this tissue.     

Does avian malaria infection affect feather stable isotope signatures?

It is widely accepted that stable isotope ratios in inert tissues such as feather keratin reflect the dietary isotopic signature at the time of the tissue synthesis. However, some elements such as stable nitrogen isotopes can be affected by individual physiological state and nutritional stress. Using malaria infection experiment protocols, we estimated the possible effect of malaria parasite infections on feather carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope signatures in juvenile common crossbills Loxia curvirostra. The birds were experimentally infected with Plasmodium relictum (lineage SGS1) and P. ashfordi (GRW2), two widespread parasites of passerines. Experimental birds developed heavy parasitemia of both parasites and maintained high levels throughout the experiment (33 days). We found no significant difference between experimental and control birds in both δ¹³C and δ¹⁵N values of feathers re-grown. The study shows that even heavy primary infections of malaria parasites do not affect feather δ¹³C and δ¹⁵N isotopic signatures. The results of this experiment demonstrate that feather isotope values of wild-caught birds accurately reflect the dietary isotopic sources at the time of tissue synthesis even when the animal’s immune system might be challenged due to parasitic infection.     

A laboratory study on biochemical degradation and microbial utilization of organic matter comprising a marine diatom,land grass,and salt marsh plant in estuarine ecosystems

We studied the biochemical degradation of organic matter comprising marine diatom, land grass, and salt marsh plant in estuarine ecosystems in two laboratory microcosms consisting of estuarine sediments and coastal seawater. The materials were incubated separately and together under controlled oxic and anoxic conditions to test effects of co-metabolism and redox on overall degradation of organic matter. We followed variations of bulk parameters [total organic carbon (TOC), total nitrogen (TN), C/N ratio, δ¹³C_TOC, and δ¹⁵N_TN], fatty acid concentrations, and compound-specific δ¹³C values over 3 months. Coexistence of marine diatom (relatively labile) with land grass/salt marsh plant (relatively refractory) in the microcosms yielded a negative co-metabolism effect (retardation rather than acceleration) on the overall degradation of organic matter. The ratios of oxic to anoxic degradation rate constants (k _ox/k _an) of TOC and most fatty acids were in a range of 1.1–1.7, implying that redox conditions per se had a limited influence on degradation of fresh organic materials in estuarine ecosystems. Variations of two bacteria-specific fatty acids (iso- and anteiso-15:0) and their δ¹³C values indicated that bacterial metabolism could use organic carbon (OC) from any available material when only one single-source material was dominant in the ecosystems. However, bacteria probably utilized OC preferentially from labile marine diatom when multiple-source materials were almost equally present in the ecosystems.     

Preliminary assessment of Greenland halibut diet in Cumberland Sound using stable isotopes

We provide preliminary carbon (δ¹³C) and nitrogen (δ¹⁵N) stable isotope assessment of the Greenland halibut (Reinhardtius hippoglossoides) diet in Cumberland Sound, with focus on two possible prey sources: pelagic represented by capelin (Mallotus villosus) and epibenthic represented by shrimp (Lebbeus polaris). The δ¹³C for the Greenland halibut stock indicated a pelagic carbon source in Cumberland Sound while stable isotope mixing models, IsoSource and MixSIR, indicated a 99% dietary composition of capelin relative to the shrimp. The δ¹⁵N did not vary across Greenland halibut size ranges and placed them at a fourth trophic position relative to a primary herbivore. This study provides the starting point for more elaborate Cumberland Sound research on the local Greenland halibut feeding ecology by confirming pelagic feeding and establishing relative trophic position as well as identifying stable isotopes as a useful tool for the study of diet in cold water fish species.     

Establishing pathways of energy flow for insect predators using stable isotope ratios: field and laboratory evidence

 Quantifying pathways of energy transfer between plants, pests, and beneficial insects is a necessary step toward maintaining pest stable agroecosystems in the absence of chemical subsidies. A diet switching experiment utilizing a predatory ladybird beetle, Hippodamia variegata (Goeze), evaluated the use of naturally occurring stable C and N isotopes as an economically feasible and safe method for quantifying pathways of energy flow within agroecosystems. Stable isotope values of the ladybird beetle Coleomegilla maculata lengi (Timberlake) collected from an agroecosystem were used to estimate the relative amount of C and N derived from agricultural plants and incorporated by ladybird beetles based on mass balance equations. At the beginning of the diet-switching experiment δ¹³C and δ¹⁵N values of H. variegata (–12.0‰ and 6.3‰, respectively) differed by –0.2‰ and 2.9‰ from the aphids that were provided exclusively as their diet. These data are consistent with previous estimates of trophic level isotope effects. After switching the diet of H. variegata to an alternative food, isotope values of H. variegata gradually shifted toward expected values for individuals fed this diet (–22.9‰ and 8.8‰ for δ¹³C and δ¹⁵N values, respectively). Isotope values of another ladybird beetle, C. maculata, collected from the field indicated that in May, alfalfa and maize (pollen) obtained in the previous year contributed 32% and 68% of the C or N to the diets of these individuals and in August, 52%, 6%, and 42% of the C or N assimilated by these insects was derived from alfalfa, wheat, and maize, respectively. These data are consistent with expectations based on the relative abundance of C. maculata in various crops during the season. The field and laboratory data are a clear indication that isotope values are sensitive to dietary changes on a relatively short time scale (days) and provide a strong basis for the use stable C and N isotope to trace energy flow patterns of these beneficial organisms within agroecosystems. Received: 17 November 1995 / Accepted: 20 June 1996     

Trophic ecology of reef sharks determined using stable isotopes and telemetry

Establishing the ecological role of predators within an ecosystem is central to understanding community dynamics and is useful in designing effective management and conservation strategies. We analysed differences in the trophic ecology of four species of reef sharks (Carcharhinus melanopterus, Carcharhinus amblyrhynchos, Triaenodon obesus and Negaprion acutidens) at Ningaloo Reef, Western Australia, by analysing tissue stable isotopes (δ¹⁵N and δ¹³C). We also monitored animals using acoustic telemetry to determine long-term residency patterns in a bay at the southern end of the reef, Coral Bay. Overall, mean δ¹³C was similar among species, ranging between −10.9 and −11.8‰, suggesting a food-web dependency on coastal producers. Classification and regression tree analysis identified an effect of species on δ¹⁵N that separated C. amblyrhynchos and C. melanopterus from N. acutidens and T. obesus. For C. amblyrhynchos and C. melanopterus, animals were also divided by size classes, with smaller sharks having lower average δ¹⁵N than larger animals; this suggests that δ¹⁵N increases with size for these two species. Juvenile C. melanopterus, juvenile N. acutidens and adult T. obesus had trophic levels of 3.7, for juvenile C. amblyrhynchos and adult C. melanopterus it was 4, and adult C. amblyrhynchos had a value of 4.3. Trophic-level estimates for C. melanopterus and C. amblyrhynchos corroborate previous conclusions based on diet studies. We found no evidence for a difference in isotopic composition between resident and non-resident sharks. The lack of variation in isotopic composition was consistent with high mean residency of these species recorded using acoustic telemetry, which was 79% (±0.09 SE) of days monitored for T. obesus, followed by N. acutidens (57 ± 19.55%), C. amblyrhynchos (54 ± 13%) and C. melanopterus (33 ± 8.28%). High δ¹³C composition in reef sharks and long-term residency behaviour suggest that coastal marine reserves might provide effective conservation refuges for some species.     

Size-dependent ontogenetic diet shifts to piscivory documented from stable isotope analyses in an introduced population of largemouth bass

Piscivorous largemouth bass (Micropterus salmoides) have been introduced in several regions outside of their native range in North America, resulting in significant disturbance to native fish communities. This species exhibits an ontogenetic diet shift from zooplanktivory to piscivory as juveniles. An early switch to piscivory allows 0+ bass to increase their growth rate prior to winter, resulting in reduced mortality. However, little is known about the dietary switch at the population level during the first year. We used carbon stable isotope analyses to examine the diets of age 0+ individuals in Lake Izunuma, Japan. The onset of the shift to piscivory occurred at a smaller size than in native or other non-native areas [>40 mm total length (TL)]. We found a positive correlation between TL and δ¹³C throughout summer and autumn. Small individuals had δ¹³C values that were similar to those of zooplankton, whereas large individuals had δ¹³C values that were similar to those of cyprinid prey species. This suggests that the smaller 0+ individuals remain zooplanktivorous until late autumn, whereas the larger individuals shift to piscivory as early as June, soon after the breeding season ends. Our results also suggest that a significant number of 0+ bass failed to switch to piscivory until the winter of their first year, despite the smaller size threshold for the onset of piscivory.