Algal δ13C Research Articles

Reconstructing midge consumer-resource dynamics using carbon stable isotope signatures of archived specimens.

Population cycles can be caused by consumer-resource interactions. Confirming the role of consumer-resource interactions, however, can be challenging due to an absence of data for the resource candidate. For example, interactions between midge larvae and benthic algae likely govern the high-amplitude population fluctuations of Tanytarsus gracilentus in Lake Mývatn, Iceland, but there are no records of benthic resources concurrent with adult midge population counts. Here, we investigate consumer population dynamics using the carbon stable isotope signatures of archived T. gracilentus specimens collected from 1977 to 2015, under the assumption that midge δ13 C values reflect those of resources they consumed as larvae. We used the time series for population abundance and δ13 C to estimate interactions between midges and resources while accounting for measurement error and possible preservation effects on isotope values. Results were consistent with consumer-resource interactions: high δ13 C values preceded peaks in the midge population, and δ13 C values tended to decline after midges reached high abundance. One interpretation of this dynamic coupling is that midge isotope signatures reflect temporal variation in benthic algal δ13 C values, which we expected to mirror primary production. Following from this explanation, high benthic production (enriched δ13 C values) would contribute to increased midge abundance, and high midge abundance would result in declining benthic production (depleted δ13 C values). An additional and related explanation is that midges deplete benthic algal abundance once they reach peak densities, causing midges to increase their relative reliance on other resources including detritus and associated microorganisms. Such a shift in resource use would be consistent with the subsequent decline in midge δ13 C values. Our study adds evidence that midge-resource interactions drive T. gracilentus fluctuations and demonstrates a novel application of stable isotope time-series data to understand consumer population dynamics.

Stable carbon and nitrogen isotopes explain methylmercury concentrations in stream food webs of Lake George, New York (USA).

Mercury has been studied extensively in lakes due to health risks associated with the consumption of contaminated fish, while stream ecosystems have received less attention. To better understand mercury bioavailability in the lower food web of streams, we collected macroinvertebrates (predators and detritivore) along with autochthonous (epilithic algae) and allochthonous (leaf litter) basal resources in eight streams entering Lake George. Samples were analyzed for methylmercury (MeHg), total mercury, and carbon and nitrogen isotopes (δ13C & δ15N) to determine how mercury concentrations in basal resources, biomagnification rates, and environmental factors (watershed characteristics and water chemistry) effected MeHg concentrations in predatory macroinvertebrates. While biomagnification rates, calculated as trophic magnification slope, explained between 68% and 98% of MeHg variability within a stream food web, the range was small (0.310-0.387) resulting in the biotic components following a consistent pattern of increasing MeHg among streams. The stream order was negatively related to basin slope for all biotic components and explained 70% of MeHg variability in predatory macroinvertebrates. Methylmercury concentrations were significantly and negatively related to δ13C in predators, epilithic algae, and leaf litter. We believe the biofilms on leaf litter utilized bacterial-respired carbon dioxide decreasing δ13C (<-28‰) and increasing MeHg while epilithic algal δ13C increased due to enhanced primary production resulting in biodilution of MeHg. Methylmercury in basal resources responded to δ13C similarly but through different processes. Our findings show shallow slopes elevate MeHg in basal resources and explain most of the predator MeHg variation among streams with little influence of biomagnification rates.

Stable carbon and nitrogen isotopic characterization and tracing nutrient sources of Ulva blooms around Jeju coastal areas

The present investigation was aimed to characterize the Ulva blooms and to identify the probable sources for Ulva blooms along the Jeju Island coast for pertinent control measures. Algal isotope signatures (δ13C, δ15N and δ18O) and tissue nitrogen and carbon were analyzed to map nutrient sources around the Jeju coastal areas. The algal δ13C values were ranged from −20.52 to −4.39‰, while δ15N and δ18O values ranged from 4.26 to 8.29‰ and 12.80–17.34‰, respectively. Moreover, site-specific significant differences were observed in algal stable isotope (δ13C, δ15N and δ18O) values. The bi-plot (δ15N vs δ18O) diagram indicated four dominant nitrogen sources along the Jeju coast, with 1) soil organic nitrogen mixed with livestock wastes (spring water samples and E), the 2) synthetic fertilizer input (A3 and B2), 3) sewage discharge (D1, D2 and I3) and 4) aquaculture waste (fish farm samples, A4, A5, B1, G and I2). Present findings revealed the different potential nitrogen sources for localized increase of algal growth along the Jeju coast. Finally, the present findings could be used as baseline data for efficient nutrient management to remediate Ulva blooms along Jeju coastal environment.

Isotopic evidence of subtle nutrient enrichment in mangrove habitats of Golfo Dulce, Costa Rica

AbstractMangroves are of great ecological and socio‐economic importance, yet they are under threat from urban development on the southern Pacific coast of Costa Rica. To test for possible nutrient‐related impacts, we compared water‐column nutrient concentrations, C and N stable isotope values and other environmental variables between mangroves with known sewage loading (three “nutrient loaded” locations) and those without such loading (three “reference” locations). Instantaneous nutrient concentrations were low at all locations, Secchi depth was greater at reference locations, and chlorophyll concentrations were higher at nutrient loaded mangroves. Suspended matter did not vary between reference and nutrient loaded mangroves, and nor did bivalve and algal δ13C and δ15N values. Enrichment of δ15N and δ13C of red mangrove leaves at the nutrient loaded locations is attributed to pulsed inputs of materials that were not detected in the instantaneous nutrient data. We provide evidence of isotopic enrichment at nutrient loaded locations from mangrove material and recommend that adequate waste water treatment be carried out on all anthropogenic discharges into this vulnerable marine system.

Long-lived coralline alga records multidecadal variability in Labrador Sea carbon isotopes

While the recent decline in the δ13C composition of oceanic dissolved inorganic carbon (DIC) can be attributed to increasing anthropogenic CO2 emissions (13C Suess effect), the causes of natural variability in the δ13C of oceanic DIC (δ13CDIC) are far less understood. Unfortunately, instrumental oceanic DIC measurements are not available prior to the 1970s, prohibiting the observation and study of long-term variability in oceanic carbon isotope dynamics. Thus, in order to identify the main driving forces of changes in oceanic δ13CDIC, multicentury carbon isotope time series that extend from the present into the preindustrial period are required. Such time series may be extracted from the carbonate skeletons of long-lived marine organisms, which have been shown to be robust recorders of fluctuations and trends in oceanic δ13CDIC. In this study, we use an annually-banded coralline alga live-collected from the Labrador shelf to generate a 266-year time series of δ13CDIC changes in the Labrador Sea. Our results indicate that from the 1960s onwards, the rate of δ13CDIC decline in the Labrador Sea slightly exceeds the rate of δ13C decline in the atmosphere, providing support for the enhanced CO2 uptake ability of the Labrador Sea. In addition, the detrended algal δ13C time series displays multidecadal variability with typical Atlantic Multidecadal Oscillation (AMO) frequencies. We show that prior to the late 1980s, algal δ13C compositions significantly correlate with regional sea ice cover (SIC) variability, post-1850 instrumental, reconstructed AMO indices, and solar variability. We speculate that these low-frequency oscillations in δ13C reflect changes in marine primary productivity modulated by a mechanism involving solar changes, the AMO and SIC variability. Our algal carbon isotope time series suggests that while the anthropogenic Suess effect has influenced Labrador Sea δ13CDIC since the 1960s, its influence may have been obscured by the effects of natural climatic variability up until the late 1980s.

The stable isotopic composition of &amp;lt;i&amp;gt;Daphnia&amp;lt;/i&amp;gt; ephippia reflects changes in δ&amp;lt;sup&amp;gt;13&amp;lt;/sup&amp;gt;C and δ&amp;lt;sup&amp;gt;18&amp;lt;/sup&amp;gt;O values of food and water

Abstract. The stable isotopic composition of fossil resting eggs (ephippia) of Daphnia spp. is being used to reconstruct past environmental conditions in lake ecosystems. However, the underlying assumption that the stable isotopic composition of the ephippia reflects the stable isotopic composition of the parent Daphnia, of their diet and of the environmental water have yet to be confirmed in a controlled experimental setting. We performed experiments with Daphnia pulicaria cultures, which included a control treatment conducted at 12 °C in filtered lake water and with a diet of fresh algae and three treatments in which we manipulated the stable carbon isotopic composition (δ13C value) of the algae, stable oxygen isotopic composition (δ18O value) of the water and the water temperature, respectively. The stable nitrogen isotopic composition (δ15N value) of the algae was similar for all treatments. At 12 °C, differences in algal δ13C values and in δ18O values of water were reflected in those of Daphnia. The differences between ephippia and Daphnia stable isotope ratios were similar in the different treatments (δ13C: +0.2 ± 0.4 ‰ (standard deviation); δ15N: −1.6 ± 0.4 ‰; δ18O: −0.9 ± 0.4 ‰), indicating that changes in dietary δ13C values and in δ18O values of water are passed on to these fossilizing structures. A higher water temperature (20 °C) resulted in lower δ13C values in Daphnia and ephippia than in the other treatments with the same food source and in a minor change in the difference between δ13C values of ephippia and Daphnia (to −1.3 ± 0.3 ‰). This may have been due to microbial processes or increased algal respiration rates in the experimental containers, which may not affect Daphnia in natural environments. There was no significant difference in the offset between δ18O and δ15N values of ephippia and Daphnia between the 12 and 20 °C treatments, but the δ18O values of Daphnia and ephippia were on average 1.2 ‰ lower at 20 °C than at 12 °C. We conclude that the stable isotopic composition of Daphnia ephippia provides information on that of the parent Daphnia and of the food and water they were exposed to, with small offsets between Daphnia and ephippia relative to variations in Daphnia stable isotopic composition reported from downcore studies. However, our experiments also indicate that temperature may have a minor influence on the δ13C, δ15N and δ18O values of Daphnia body tissue and ephippia. This aspect deserves attention in further controlled experiments.

Characterising sediments of a tropical sediment-starved shelf using cluster analysis of physical and geochemical variables

Environmental context Australia's tropical marine estate is a biodiversity hotspot that is threatened by human activities. Analysis and interpretation of large physical and geochemistry data sets provides important information on processes occurring at the seafloor in this poorly known area. These processes help us to understand how the seafloor functions to support biodiversity in the region. Abstract Baseline information on habitats is required to manage Australia's northern tropical marine estate. This study aims to develop an improved understanding of seafloor environments of the Timor Sea. Clustering methods were applied to a large data set comprising physical and geochemical variables that describe organic matter (OM) reactivity, quantity and source, and geochemical processes. Arthropoda (infauna) were used to assess different groupings. Clusters based on physical and geochemical data discriminated arthropods better than geomorphic features. Major variations among clusters included grain size and a cross-shelf transition from authigenic-Mn–As enrichments (inner shelf) to authigenic-P enrichment (outer shelf). Groups comprising raised features had the highest reactive OM concentrations (e.g. low chlorin indices and C:N ratios, and high reaction rate coefficients) and benthic algal δ13C signatures. Surface area-normalised OM concentrations higher than continental shelf norms were observed in association with: (i) low δ15N, inferring Trichodesmium input; and (ii) pockmarks, which impart bottom–up controls on seabed chemistry and cause inconsistencies between bulk and pigment OM pools. Low Shannon–Wiener diversity occurred in association with low redox and porewater pH and published evidence for high energy. Highest β-diversity was observed at euphotic depths. Geochemical data and clustering methods used here provide insight into ecosystem processes that likely influence biodiversity patterns in the region.

Stable carbon isotope variability associated with taxonomic composition of lotic benthic algae

SummaryWe examined the relationship between the δ13C and taxonomic composition of benthic algae collected from a riffle (fast current habitat) of a non‐shaded mountain stream, which is a tributary of the Kiso River, Japan. The benthic algal δ13C ranged from −20.6 to −14.2‰ and tended to be 13C‐depleted with increasing relative abundance of upright filamentous cyanobacteria and 13C‐enriched with increasing relative abundance of prostrate filamentous cyanobacteria. Using isotopic mass balance equations, the relative abundance of the dominant taxa, upright and prostrate filamentous cyanobacteria, small diatoms and others, explained 74% of δ13C variability. This study shows a case where the difference in taxonomic composition is a possible source of the isotopic variability of benthic algae, which is a mixture of taxa with distinct isotopic signatures.

Relationship between δ13C of chironomid remains and methane flux in Swedish lakes

Summary1. Methanogenic carbon can be incorporated by methane‐oxidising bacteria, leading to a 13C‐depleted stable carbon isotopic composition (δ13C) of chironomids that feed on these microorganisms. This has been shown for the chironomid tribe Chironomini, but very little information is available about the δ13C of other abundant chironomid groups and the relationship between chironomid δ13C and methane production in lakes.2. Methane flux was measured at the water surface of seven lakes in Sweden. Furthermore, fluxes from the sediments to the water column were measured in transects in two of the lakes. Methane fluxes were then compared with δ13C of chitinous chironomid remains isolated from the lake surface sediments. Several different chironomid groups were examined (Chironomini, Orthocladiinae, Tanypodinae and Tanytarsini).3. Remains of Orthocladiinae in the seven study lakes had the highest δ13C values (−31.3 to −27.0‰), most likely reflecting δ13C of algae and other plant‐derived organic matter. Remains of Chironomini and Tanypodinae had lower δ13C values (−33.2 to −27.6‰ and −33.6 to −28.0‰, respectively). A significant negative correlation was observed between methane fluxes at the lake surface and δ13C of Chironomini (r = −0.90, P = 0.006). Methane release from the sediments was also negatively correlated with δ13C of Chironomini (r = −0.67, P = 0.025) in the transect samples obtained from two of the lakes. The remains of other chironomid taxa were only weakly or not correlated with methane fluxes measured in our study lakes (P &gt; 0.05).4. Selective incorporation of methane‐derived carbon can explain the observed correlations between methane fluxes and δ13C values of Chironomini. Remains of this group might therefore have the potential to provide information about past changes in methane availability in lakes using sediment records. However, differences in productivity, algal δ13C composition and the importance of allochthonous organic matter input between the studied lakes may also have influenced Chironomini δ13C. More detailed studies with a higher number of analysed samples and detailed measurement of δ13C of different ecosystem components (e.g. methane, dissolved inorganic carbon) will be necessary to further resolve the relative contribution of different carbon sources to δ13C of chironomid remains.

Temporal variability of benthic algal δ13C signatures influences assessments of carbon flows in stream food webs

Stream food web function is often assessed using carbon stable isotope assessments of the relative contribution of autochthonous and allochthonous sources of organic matter to consumer diets. As a result, variability in source signatures can strongly influence the assessment of carbon flows. To examine the implications of temporal source variability on food web interpretations, benthic algal δ13C signatures were measured over 8 weeks in five streams in subtropical Queensland, Australia. All food webs were largely driven by benthic algal carbon; however, substantial week-to-week variation in benthic algal δ13C signatures modified the calculated contributions of algae to consumer diets, with differences in autochthonous contributions of up to 11% between weeks. In addition, variable algal signatures led to many occasions in which the δ13C signatures of some consumers was beyond the range of available sources, meaning the mixing model analyses did not have a valid solution. Together, these findings suggest that temporal variability in algal δ13C signatures can strongly influence the interpretation of carbon flows in stream food webs. Future food web studies should assess the temporal variability of sources prior to sampling consumers, in order to characterise end member signatures and their relevance to consumers at the time of collection.

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

Summary1. 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.

Comparison of methods to determine algal δ13C in freshwater

To accurately assess the flux of mass and energy to higher trophic levels in a food web using stable isotopes, the isotopic signature of basal sources is required. When studying aquatic food webs, it is difficult to obtain a signature for algae because of challenges associated with isolating small organisms from a bulk sample. In this study, we compared freshwater algal δ13C values obtained using five approaches from the literature. Results indicated that the signatures derived from a primary consumer such as Daphnia sp., from particulate organic carbon with a correction for algal biomass, and from isolated algal samples were comparable. By contrast, algal δ13C values based on the signature of carbon dioxide and algal carbon fractionation were significantly lower than those of the other approaches. The inconsistent values produced by this method were likely due to problems in determining fractionation values based on current models and were potentially related to bicarbonate uptake by algae.

Stable isotope analysis indicates microalgae as the predominant food source of fauna in a coastal forest stream, south‐east Brazil

Abstract: Stable isotope studies of food webs in floodplains, large rivers, mangroves, and seagrasses have shown that, although a large proportion of the biomass may come from higher plants, microalgae provide a disproportionate amount of carbon assimilated by metazoan consumers. Evidence is building that this may also be the case for streams, especially those in the tropics. At the level of individual consumer species we also see that the apparent diet may not be reflected in the carbon assimilated. Tropical streams commonly have omnivore‐detritivore species that potentially show this phenomenon. We tested these concepts in four moderately shaded sites in a stream in well‐preserved Atlantic rainforest at Ilha Grande, Rio de Janeiro. We sampled aquatic insects, shrimps and fish as well as potential terrestrial and aquatic primary food sources. Carbon stocks from terrestrial sources predominated over carbon of algal origin (&gt;99% of total). The primary sources of carbon showed distinctly different isotopic signatures: terrestrial sources had δ13C values close to −30‰, microalgae were −20‰ and macroalgae were −25‰. All fauna had δ13C values consistent with a carbon source derived from microalgae. Baetid mayflies and atyid shrimps exert a strong grazing pressure on periphyton and organic sediments but appear to assimilate predominantly microalgae. The palaemonid shrimp Macrobrachium olfersi also ingests large amounts of detritus of terrestrial origin, but apparently assimilates animal prey with algal δ13C signatures. These results support the growing view that tropical stream food chains are primarily algal based.

SPATIAL SCALES OF CARBON FLOW IN A RIVER FOOD WEB

Spatial extents of food webs that support stream and river consumers are largely unknown, but such information is essential for basic understanding and management of lotic ecosystems. We used predictable variation in algal δ13C with water velocity, and measurements of consumer δ13C and δ15N to examine carbon flow and trophic structure in food webs of the South Fork Eel River in Northern California. Analyses of δ13C showed that the most abundant macroinvertebrate groups (collector-gatherers and scrapers) relied on algae from local sources within their riffle or shallow pool habitats. In contrast, filter-feeding invertebrates in riffles relied in part on algal production derived from upstream shallow pools. Riffle invertebrate predators also relied in part on consumers of pool-derived algal carbon. One abundant taxon drifting from shallow pools and riffles (baetid mayflies) relied on algal production derived from the habitats from which they dispersed. The trophic linkage from pool algae to riffle invertebrate predators was thus mediated through either predation on pool herbivores dispersing into riffles, or on filter feeders. Algal production in shallow pool habitats dominated the resource base of vertebrate predators in all habitats at the end of the summer. We could not distinguish between the trophic roles of riffle algae and terrestrial detritus, but both carbon sources appeared to play minor roles for vertebrate consumers. In shallow pools, small vertebrates, including three-spined stickleback (Gasterosteus aculeatus), roach (Hesperoleucas symmetricus), and rough-skinned newts (Taricha granulosa), relied on invertebrate prey derived from local pool habitats. During the most productive summer period, growth of all size classes of steelhead and resident rainbow trout (Oncorhynchus mykiss) in all habitats (shallow pools, riffles, and deep unproductive pools) was largely derived from algal production in shallow pools. Preliminary data suggest that the strong role of shallow pool algae in riffle steelhead growth during summer periods was due to drift of pool invertebrates to riffles, rather than movement of riffle trout. Data for δ15N showed that resident rainbow trout (25–33 cm standard length) in deep pools preyed upon small size classes of juvenile steelhead that were most often found in riffles or shallow pools. While many invertebrate consumers relied primarily on algal production derived from local habitats, our study shows that growth of top predators in the river is strongly linked to food webs in adjacent habitats. These results suggest a key role for emigration of aquatic prey in determining carbon flow to top predators.

STABLE-CARBON-ISOTOPE RATIOS OF RIVER BIOTA:IMPLICATIONS FOR ENERGY FLOW IN LOTIC FOOD WEBS

Stable-isotope ratios of carbon (13C/12C or δ13C) have been widely used to determine the energy base of stream food webs, but such use is controversial due to unexplained variability in algal δ13C. I used published δ13C data from temperate headwater streams through medium-sized rivers (0.2–4000 km2 watershed area) collected during summer baseflows and original data from streams in northern California to analyze energy pathways through river food webs. The analyses showed three important results. First, epilithic algal δ13C and watershed area are positively related, suggesting that effects of carbon limitation on algal carbon uptake result in 13C enrichment of algal δ13C in larger, more productive rivers. Second, epilithic algae and terrestrial detritus δ13C values are often distinct in small shaded streams but overlap in some larger unshaded streams and rivers. Measurements of δ13C values may be most useful in distinguishing algal and terrestrial energy sources in unproductive streams with supersaturated dissolved CO2 concentrations, and some productive rivers where CO2 concentrations are low relative to photosynthetic rates. Finally, consumer δ13C values are more strongly related to algal δ13C than terrestrial δ13C. The relative contribution of terrestrial and algal carbon sources often varied by functional feeding group within and between sites. However, with the exception of shredders and scrapers, which respectively relied on terrestrial and algal carbon sources, patterns of consumer δ13C clearly show a transition from terrestrial to algal carbon sources for many lotic food webs in streams with ≥10 km2 watershed area. The observed transition to algal carbon sources is likely related to increasing primary production rates as forest canopy cover declines in larger streams, although decreasing retention or quality of terrestrial carbon may also play a role. Improved analyses of algal δ13C and δ15N combined with quantitative study of organic matter dynamics and food web structure should allow the relative importance of these factors to be distinguished in future food web studies.

Critical examination of stable isotope analysis as a means for tracing carbon pathways in stream ecosystems

Existing data conflict as to the capability of stable isotope analysis (SIA) for distinguishing carbon pathways in stream ecosystems. I compiled 803 published measurements of δ13C for allochthonous litter and both lotic attached algae and consumers to search for general patterns that transcend individual studies. Half of the fishes and 30% of the invertebrates exhibited δ13C values outside the range that could be ascribed to allochthonous litter. The enormous variability in attached algal δ13C from −40 to −20‰ (mean ± SD = −29 ± 4) completely enveloped that observed for terrestrial leaf litter (mean = −28 ± 1‰). Therefore, for 50% of the fishes and 70% of the invertebrates it was impossible to discriminate between allochthonous and autochthonous carbon dependency. Faunal δ13C did not change in any quantitatively predictable manner with removal of riparian trees. Three conclusions are possible from this secondary analysis: (1) autotrophic pathways within forested headwaters are much more important to lotic foodwebs than would be suggested by their particulate inputs alone; (2) the great variability in attached algal δ13C may often preclude use of SIA for identifying carbon pathways in stream ecosystems; thus (3) the utility of carbon SIA in understanding anthropogenic alterations to the carbon budget of streams is presently minimal.