Differences in nitrogen and carbon stable isotopes between planktonic and benthic microalgae

We compiled published data on the nitrogen and carbon stable isotope ratios of phytoplankton and benthic microalgae from lentic systems and explored the primary factors determining the isotope values among systems. Also, we investigated seasonal changes in nitrogen stable isotope ratios of phytoplankton and benthic microalgae in the strongly acidic lake, Lake Katanuma, which has only two dominant species, Pinnularia acidojaponica as a benthic diatom and Chlamydomonas acidophila, a planktonic green alga. From the published dataset, it may be concluded that δ¹³C of benthic diatoms were more enriched than those of phytoplankton at the same sites, although the nitrogen isotope of phytoplankton and benthic microalgae were similar. This differences in δ¹³C between benthic microalgae and phytoplankton could be explained by the boundary layer effect. On the other hand, nitrogen isotope values of both benthic microalgae and phytoplankton were primarily controlled by the same environmental factor, and boundary layer effects are not the primary factor determining the nitrogen isotope values of microalgae. Also, we showed temporal dynamics in nitrogen isotopes of benthic and planktonic microalgae species in Lake Katanuma, and the trends of nitrogen isotopes are similar between benthic and planktonic microalgae, as concluded from the published dataset.     

Movement of carbon among estuarine habitats and its assimilation by invertebrates

We measured the extent of movement of carbon and its assimilation by invertebrates among estuarine habitats by analysing carbon stable isotopes of invertebrates collected along transects crossing the boundary of two habitats. The habitats were dominated by autotrophs with distinct isotope values: (1) mudflats containing benthic microalgae (mean −22.6, SE 0.6‰) and (2) seagrass and its associated epiphytic algae (similar values, pooled mean −9.8, 0.5‰). Three species of invertebrates were analysed: a palaemonid shrimp, Macrobrachium intermedium, and two polychaete worms, Nephtys australiensis and Australonereis ehlersi. All species had a similar narrow range of isotope values (−9 to −14‰), and showed no statistically significant relationship between position along transect and isotope values. Animals were relying on carbon from seagrass meadows whether they were in seagrass or on mudflats hundreds of metres away. Particulate organic matter collected from superficial sediments along the transects had similar values to animals (mean −11.1, SE 1.3‰) and also showed no significant relationship with position. The isotope values of these relatively immobile invertebrates and the particulate detritus suggest that carbon moves from subtidal seagrass meadows to mudflats as particulate matter and is assimilated by invertebrates. This assimilation might be direct in the case of the detritivorous worm, A. ehlersi, but must be via invertebrate prey in the case of the carnivorous worm, N. australiensis and the scavenging shrimp, M. intermedium. The extent of movement of carbon among habitats, especially towards shallower habitats, is surprising since in theory, carbon is more likely to move offshore in situations such as the current study where habitats are in relatively open, unprotected waters.     

Incorporation of Exotic Spartina alterniflora into Diet of Deposit-Feeding Snails in the Yangtze River Estuary Salt Marsh: Stable Isotope and Fatty Acid Analyses

The response of deposit-feeding animals to plant invasions is still unclear, because their food sources are often difficult to identify. We examined the effect of the exotic plant species, Spartina alterniflora, on the food source composition of two dominant snail species, Assiminea latericea and Cerithidea largillierti, in the Yangtze River estuary salt marsh using a combination of stable isotope and fatty acid analyses. We collected the snails and their potential food materials (sediment organic matter, particulate organic matter, and plant material) in S. alterniflora and native plant Phragmites australis marshes and then determined the composition of food sources of snails based on fatty acid markers and stable isotope composition. Our results indicated that A. latericea and C. largillierti are deposit feeders grazing on sedimentary particles originating from diatoms, bacteria, and vascular plants. Invasive S. alterniflora did not result in a change in the relative contribution of microalgae, bacteria, and vascular plants to the food source of the snails. Spartina alterniflora was confirmed to be assimilated by both snail species. The higher assimilation of S. alterniflora by A. latericea compared with C. largillierti is probably related to the greater ability of A. latericea to assimilate plant materials from detritus, as evidenced by fatty acid composition. Overall, S. alterniflora can be incorporated into the food web of the estuarine salt marsh by the dominant snail species with generalist-feeding habits.     

Contribution of potential organic matter sources to the macrobenthos nutrition in the Natori estuarine tidal flats,Japan

In this investigation, we used stable isotope and fatty acid biomarker analyses to estimate and compare the potential food sources that support macrobenthos (Nuttallia olivacea, Corbicula japonica, and Hediste sp.) in the Natori estuarine tidal flats of Japan. The δ¹³C and δ¹⁵N mean values for the sediment organic matter (SOM) were ?23.6‰ and 6.1‰, respectively, which were due to the contribution of terrestrial and/or aquatic vascular plant particulate organic matter (POM) from upper stream river or surrounding areas. Furthermore, from the results of the IsoSource mixing model, the contributions of estuarine POM to the diets of Hediste sp., C. japonica, and N. olivacea were 85.1%, 74.9%, and 48.9%, respectively. Moreover, essential fatty acids such as 20:5ω3, 18:2ω6 and 18:3ω3 highly contributed to the diets of macrobenthos from benthic diatoms, terrestrial and/or aquatic vascular plants. The contents of fatty acid markers of terrestrial OM (e.g., long chain fatty acids [LCFAs]) in the 3 species of macrobenthos were low in comparison to those of other food sources. Overall, the marine POM dietary contribution was minimal, while terrestrial OM, bacteria, and benthic diatoms constituted a significant portion of the macrobenthos diet, although the contribution varied among the 3 species of macrobenthos.     

Food habits of the farmer damselfish <Emphasis Type="Italic">Stegastes nigricans</Emphasis> inferred by stomach content,stable isotope,and fatty acid composition analyses

The territorial damselfish, Stegastes nigricans, maintains algal farms by excluding invading herbivores and weeding unpalatable algae from its territories. In Okinawa, Japan, S. nigricans farms are exclusively dominated by Polysiphonia sp., a highly digestible filamentous rhodophyte. This study was aimed at determining the diet of S. nigricans in Okinawa and its dependency on these almost-monoculture algal farms based on stomach content and chemical analyses. Stomach content analyses revealed that all available food items in the algal farms (i.e., algae, benthic animal inhabitants, trapped detritus) were contained in fish stomachs, but amorphous organic matter accounted for 68% of the contents. Therefore, carbon and nitrogen stable isotope ratios and fatty acid (FA) compositions were analyzed to trace items actually assimilated in their bodies. Stable isotope analyses showed that benthic animals were an important food source even for this farmer fish. Two essential fatty acids (EFAs), 20:4n6 and 20:5n3, which are produced only by rhodophytes among available food items, were rich in the muscle tissue of S. nigricans as well as in algal mats and detritus, suggesting that algal mats contribute EFAs to S. nigricans directly and indirectly through the food web. In conclusion, S. nigricans ingested algal mats, detritus, and benthic animals maintained within its farm. Algae and detritus were original sources of EFAs, and benthic animals, which were much more abundant in the farms than in outside territories, provided a nitrogen-rich dietary source for the fish.     

Water moss as a food item of the zoobenthos in the Yenisei River

Bryophytes are abundant in streams and are a habitat for many invertebrates, but their contribution to the diet of fluvial zoobenthos is still debated. To estimate the amount of bryophyte-derived organic matter assimilated by benthic invertebrates, we used a combination of fatty acid and stable isotope analyses during a four-year monthly study of a littoral site in the Yenisei River (Siberia, Russia). Acetylenic acids, which are highly specific biomarkers of the water moss Fontinalis antipyretica, were found in lipids of all dominant benthic animals: gammarids, ephemeropterans, chironomids and trichopterans. The dominant zoobenthic species, Eulimnogammarus viridis, had maximum levels of the biomarkers in its biomass during winter, and minimum levels in summer. The zoobenthos in the studied site regularly consume and assimilate bryophyte-derived organic matter as a minor supplemental food. This consumption increases in winter, when the main food source of the zoobenthos, epilithic biofilms, are probably scarce.     

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

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

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

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

Food web analysis of southern California coastal wetlands using multiple stable isotopes

Carbon, nitrogen, and sulfur stable isotopes were used to characterize the food webs (i.e., sources of carbon and trophic status of consumers) in Tijuana Estuary and San Dieguito Lagoon. Producer groups were most clearly differentiated by carbon, then by sulfur, and least clearly by nitrogen isotope measurements. Consumer ¹⁵N isotopic enrichment suggested that there are four trophic levels in the Tijuana Estuary food web and three in San Dieguito Lagoon. A significant difference in multiple isotope ratio distributions of fishes between wetlands suggested that the food web of San Dieguito Lagoon is less complex than that of Tijuana Estuary. Associations among sources and consumers indicated that inputs from intertidal macroalgae, marsh microalgae, and Spartina foliosa provide the organic matter that supports invertebrates, fishes, and the light-footed clapper rail (Rallus longirostris levipes). These three producers occupy tidal channels, low salt marsh, and mid salt marsh habitats. The only consumer sampled that appears dependent upon primary productivity from high salt marsh habitat is the sora (Porzana carolina). Two- and three-source mixing models identified Spartina as the major organic matter source for fishes, and macroalgae for invertebrates and the light-footed clapper rail in Tijuana Estuary. In San Dieguito Lagoon, a system lacking Spartina, inputs of macroalgae and microalgae support fishes. Salicornia virginica, S. subterminalis, Monanthochloe littoralis, sewage- derived organic matter, and suspended particulate organic matter were deductively excluded as dominant, direct influences on the food web. The demonstration of a salt marsh–channel linkage in these systems affirms that these habitats should be managed as a single ecosystem and that the restoration of intertidal marshes for endangered birds and other biota is compatible with enhancement of coastal fish populations; heretofore, these have been considered to be competing objectives. Received: 24 April 1996 / Accepted: 24 October 1996     

Benthic resources are the key to Daphnia middendorffiana survival in a high arctic pond

1. Shallow arctic lakes and ponds have simple and short food webs, but large uncertainties remain about benthic–pelagic links in these systems. We tested whether organic matter of benthic origin supports zooplankton biomass in a pond in NE Greenland, using stable isotope analysis of carbon and nitrogen in the pond itself and in a ¹³C‐enrichment enclosure experiment. In the latter, we manipulated the carbon isotope signature of benthic algae to enhance its isotopic discrimination from other potential food sources for zooplankton. 2. The cladoceran Daphnia middendorffiana responded to the ¹³C‐enrichment of benthic mats with progressively increasing δ¹³C values, suggesting benthic feeding. Stable isotope analysis also pointed towards a negligible contribution of terrestrial carbon to the diet of D. middendorffiana. This agreed with the apparent dominance of autochthonous dissolved organic matter in the pond revealed by analysis of coloured dissolved organic matter. 3. Daily net production by phytoplankton in the pond (18 mg C m^?2 day^?1) could satisfy only up to half of the calculated minimum energy requirements of D. middendorffiana (35 mg C m^?2 day^?1), whereas benthic primary production alone (145 mg C m^?2 day^?1) was more than sufficient. 4. Our findings highlight benthic primary production as a major dietary source for D. middendorffiana in this system and suggest that benthic organic matter may play a key role in sustaining pelagic secondary production in such nutrient‐limited high arctic ponds.     

Food web analysis of an eelgrass (Zostera marina L.) meadow and neighbouring sites in Mitsukuchi Bay (Seto Inland Sea,Japan) using carbon and nitrogen stable isotope ratios

The contribution of benthic microalgal production has been compared both within and outside a coastal eelgrass (Zostera marina L.) meadow. Carbon and nitrogen stable isotope ratios of suspended particulate organic matter (POM), epiphytic and epilithic organic matter (EOM), leaves of Z. marina (inside the meadow only) and two secondary consumer species (small crustaceans and fish) were measured inside and outside a meadow in Mitsukuchi Bay, Northwest Seto Inland Sea, Japan. Inside the meadow, primary producers (epiphyton) and consumers showed higher δ¹³C signatures than outside. Primary and secondary consumers inside the meadow were mainly dependent on epiphyton on the leaves of Z. marina, while consumer species outside the meadow were basically dependent on epilithon.     

The effect of wrack composition and diversity on macrofaunal assemblages in intertidal marine sediments

Wrack (dead, washed-up seaweed and seagrass) buried in soft substrata may increase the organic content and alter the physical structure of sediments. These effects may influence the composition and structure of macrofaunal assemblages in the sediment. Such influences can be expected to vary according to the type and amount of wrack as well as the presence of invasive seaweeds in the wrack. In this study, we deliberately buried different amounts of the invasive species Sargassum muticum in isolation or mixed to the native species Ulva sp. and Fucus vesiculosus, in two intertidal sandflats to test some hypotheses about the response of macrofaunal assemblages. We tested whether (1) diversity of detritus (i.e. different mixtures), and (2) the amount of detritus of S. muticum influenced the composition and the relative abundance of macrofaunal assemblages. We also assessed whether the sediment organic carbon and the biomass of benthic microalgae varied depending on the diversity of detritus and the amount of detritus of S. muticum. Finally, we tested if these effects of wrack were consistent across sites. Results indicated that buried wrack affected the composition and structure of macrofaunal assemblages in short-term (i.e. 4 weeks), but there were no differences depending on detritus diversity or the amount of S. muticum. In addition, sediment organic matter and microalgal biomass were not affected by the addition of wrack. They instead varied greatly among small spatial scales (i.e. plots). Wrack composition or abundance of the invasive species S. muticum played thus a small role in shaping the structure of macrofaunal assemblages or the biomass of benthic microalgae in these intertidal sediments, probably because these sediments are frequently affected by various inputs of organic matter and benthic assemblages are already adapted to organically enriched sediments.     

Paleoenvironmental changes across the Cenomanian/Turonian Boundary Event (Oceanic Anoxic Event 2) as indicated by benthic foraminifera from the Demerara Rise (ODP Leg 207)

This study is based on Cenomanian to lower Turonian sediments of Ocean Drilling Program (ODP) Sites 1258, 1259, 1260, and 1261 from the Demerara Rise (Leg 207, western tropical Atlantic, off Suriname) that are oriented along a paleodepth transect. Studied sediments include the Cenomanian/Turonian Boundary Event (CTBE) or Oceanic Anoxic Event 2 (OAE 2) and consist of laminated black shales with TOC values between 5% and 10% below and above OAE 2 and up to 29% within the OAE 2 interval. Benthic foraminiferal assemblages in this eutrophic environment are generally characterized by low diversities and strong fluctuations of abundances, indicating oxygen depletion and high organic matter fluxes. Dominant taxa at all sites are Bolivina anambra, Gavelinella dakotensis, Tappanina sp., Praebulimina prolixa, and Neobulimina albertensis. Based on the positive stable carbon isotope excursion characteristics of OAE 2 we subdivided the studied successions into three intervals: (1) the interval below OAE 2; (2) the carbon isotope excursion reflecting OAE 2; and (3) the interval above OAE 2. In the bathymetrically shallower Sites 1260 and 1261 benthic foraminiferal assemblages indicate anoxic to sometimes slightly dysoxic bottom-water conditions below the OAE 2 interval. The bathymetrically deepest Site 1258, in contrast, reflects more oxygenated bottom waters with an almost continuous occurrence of benthic foraminifera. It is therefore suggested that the shallower sites were located within the oxygen minimum zone (OMZ), whereas Site 1258 was below the OMZ. During OAE 2 anoxic conditions prevailed at the shallower sites. At Site 1258 benthic foraminifera indicate severe dysoxic but not anoxic conditions. This pattern is proposed to reflect a strengthening of the OMZ possibly related to increased primary production during OAE 2. A short-term repopulation event of benthic foraminifera in the lower third of the OAE 2 interval was observed at all sites, reflecting a brief bottom-water oxygenation event. This short-lived event parallels a surface-water cooling and is probably equivalent to the “Plenus Cool Event” in Europe and the “benthonic zone” in the U.S. Western Interior. The benthic foraminifera of a ~0.5 Ma interval following OAE 2 still indicate oxygen depletion of bottom waters. Subsequently, however, a strong increase in benthic foraminiferal abundance and diversity reflects better oxygenation of the bottom-water masses, probably related to a weakening of the OMZ due to decreasing organic matter flux.     

Enhanced production of fatty acids in three strains of microalgae using a combination of nitrogen starvation and chemical inhibitors of carbohydrate synthesis

Microalgae are attracting much attention as superior biodiesel producers. In particular, under stressful conditions, they accumulate organic compounds consisting entirely of carbon and hydrogen. The aim of this work was to increase intracellular fatty acid content in Dunaliella tertiolecta (Chlorophyceae), Nannochloropsis oculata (Eustigmatophyceae), and Porphyridium cruentum (Rhodophyceae) using a combination of nitrogen starvation and chemical inhibitors of carbohydrate biosynthesis. These microalgae were subjected to nitrogen starvation and their physiological changes were then observed over time. In D. tertiolecta, no significant change in total fatty acid content was detected on day 3.5 relative to the initial total fatty acid content (day 0), while total carbohydrate content dramatically increased as the nitrogen starvation period was extended. In N. oculata, total fatty acid content rapidly increased, reaching up to nearly 40% of the DCW at day 3.5. However, total carbohydrate content exhibited a gradual reduction throughout the experiment. In P. cruentum, total carbohydrate content increased up to 43% of DCW on day 3.5 and total fatty acid content increased slightly under nitrogen depletion. These data suggest that different eukaryotic microalgae use different storage products under stressful conditions. Among the three strains, D. tertiolecta showed decreased total carbohydrate content and enhanced total fatty acid content following inhibition of carbohydrate synthesis by dichlorophenyl dimethylurea and cyclohexane diamine tetra acetic acid. The results demonstrate the possibility of furthering our understanding of the fatty acid and carbohydrate biosynthesis metabolic network that responds to environmental changes in microalgae.     

Asymmetric response of sedimentary pool to surface water in organics from a shallow hypereutrophic lake: The role of animal consumption and microbial utilization

Despite of the importance for tracking sources and fates of organics in determining carbon cycling and assessing the overall health of lacustrine ecosystems, little is known of organics behaviors in shallow eutrophic lakes. In this study, for relating the benthic organics to pelagic sources, an ecosystem-level investigation was performed in hypereutrophic Lake Taihu, China, by using source-specific fatty acid biomarkers on the molecular level. Results exhibited the unexpected asymmetric phenomena of organics between surface water and sediments. In the abundance, the dominant organics shifted from cyanobacteria in surface water to terrestrial plants in sediments. In the spatial pattern, as opposed to terrestrial plants, cyanobacteria were found not site-to-site symmetrical, although both of their spatial distributions varied strongly. Essentially, these asymmetric phenomena ascribed to a more considerable loss of cyanobacteria compared to terrestrial plants during sinking, caused by aquatic animal consumption and microbial utilization with different degree. Combined dual stable isotopes (δ¹³C and δ¹⁵N) and fatty acid biomarkers revealed that there were only subtle differences in the diets of benthic and pelagic animals and cyanobacteria were their main food source. Concomitantly, results of δ¹³C of bacteria-specific fatty acids demonstrated that bacteria equally and profoundly affected organics accumulation or preservation in the sediments, because they preferentially utilized labile cyanobacteria as their carbon source instead of terrestrial plants (>95% within these two sources). Consequently, these novel findings clarify that not only in deep lakes, but also shallow eutrophic ones, the extensive losses of autochthonous organic matter can be expected during sedimentation coupling with the dramatical modifications of biogeochemical processes.     

Lipids in Arctic benthos: does the fatty acid and alcohol composition reflect feeding and trophic interactions?

Arctic benthic organisms of various taxa (Anthozoa, Polychaeta, Pantopoda, Crustacea, Echinodermata) were collected on the shelves off northeast Greenland, Spitsbergen and the western Barents Sea. Their fatty acid compositions were generally characterised by the predominance of the polyunsaturated fatty acids 20:5(n-3) and 22:6(n-3) together with the saturated fatty acid 16:0, which reflect the dominance of phospholipids. The fatty acid compositions of most benthic specimens were influenced by fatty acids of dietary origin. High amounts of the fatty acid 16:1(n-7), typical of diatoms, were found in different taxa from the northeast Greenland shelf. The 18:4(n-3) fatty acid, often typical of non-diatom input, was only dominant in Ophiopholis aculeata from the Spitsbergen shelf. In some taxa small amounts of wax esters were detected with alcohol moieties similar to those of the dominant Arctic copepods. The occurrence of intact wax esters, as well as the wax ester typical fatty acids 20:1(n-9) and 22:1(n-11), also suggested ingestion of large herbivorous copepods. An unusual fatty acid composition was found for most brittle stars, due to a ratio of the 18:1(n-9) and (n-7) fatty acid isomers below 1 with lowest ratios of 0.1. A similar low ratio was also detected in the polychaete Onuphis conchylega. The extremely low portions of the 18:1(n-9) fatty acid are striking, since carnivores are generally characterised by high levels of this fatty acid. A clear gradient from low 18:1(n-9) to (n-7) ratios in suspension feeders, via predatory decapods, to higher ratios in the scavenging amphipods was a major characteristic of the benthic species. Our investigations showed that lipid analyses can give important hints on trophic relationships of benthic species and may serve as means to establish the intensity of pelagic-benthic coupling.     

Stable carbon isotope ratios in Fundulus heteroclitus (L.) muscle tissue and gut contents from a North Carolina spartina marsh

Recent food habit and productivity research on the killifish, Fundulus heteroclitus (L.), suggests that this species is important in the movement of organic material within and out of the salt marsh ecosystem. The stable carbon isotope ratios of Fundulus muscle tissue and gut contents were measured seasonally to determine the carbon source of the killifish. For muscle δ¹³C values ranged from -13.9% to -15.8% and -15.8% to -18.4% for gut contents, suggesting differential assimilation of ingested material. Muscle tissue indicate that assimilated carbon probably originated from a mixture of benthic algae and Spartina ingested by major prey species: the filddle: crab, Uca pugnax (Smith): the polychaete, Nereis (Neanthes) succinea Frey and Leuckart; the tanaid, Leptochelia rapax Harger; and other small crustaceans. Small fish (age0–1 yr) showed slightly lower δ¹³C values than larger fish (age 1–3 yr), consistent with differences in their feeding habits. The only seasonal pattern was an enrichment in tissue ¹³C that occurred for both small and large fish in the spring, corresponding to the peak spawning period. The δ¹³C determination is a useful tool in identifying and tracing carbon sources through the salt marsh food web, detecting differences in feeding habits between age classes of organisms, and assessing the relative nutritive value of ingested dietary items if differential assimilation is suspected.     

Isotopic analysis of three food web theories in constricted and floodplain regions of a large river

Analyses of stable isotope (δ¹³C and δ¹⁵N) and C:N ratios of food webs within a floodplain and a constricted-channel region of the Ohio River during October 1993 and July 1994 indicate that the increasingly influential flood pulse concept (FPC) does not, for either location, adequately address food web structure for this very large river. Furthermore, results of this study suggest that the riverine productivity model (RPM) is more appropriate than the widely known river continuum concept (RCC) for the constricted region of this river. These␣conclusions are based on stable isotope analyses of potential sources of organic matter (riparian C₃ trees, riparian C₄ grasses and agricultural crops, submerged macrophytes, benthic filamentous algae, benthic particulate organic matter, and transported organic matter containing detritus and phytoplankton) and various functional feeding groups of invertebrate and fish consumers. The FPC, which stresses the key contribution of organic matter, particularly terrestrial organic matter, originating from the floodplain to riverine food webs, was judged inappropriate for the floodplain region of the Ohio River for hydrodynamic and biotic reasons. The rising limb and peak period of discharge typically occur in November through March when temperatures are low (generally much less than 10°C) and greater than bank-full conditions are relatively unpredictable and short-lived. The major food potentially available to riverine organisms migrating into the floodplain would be decaying vegetation because autotrophic production is temperature and light limited and terrestrial insect production is minimal at that time. It is clear from our data that terrestrial C₄ plants contribute little, if anything, to the consumer food web (based on δ¹³C values), and δ¹⁵N values for C₃ plants, coarse benthic organic matter, and fine benthic organic matter were too depleted (∼7–12‰ lower than most invertebrate consumer values) for this organic matter to be supporting the food web. The RPM, which emphasizes the primary role of autotrophic production in large rivers, is the most viable of the remaining two ecosystem models for the constricted-channel region of the Ohio based on stable isotope linkage between sources and consumers of organic matter in the food web. The most important form of food web organic matter is apparently transported (suspended) fine (FTOM) and ultra-fine particulate organic matter. We propose that phytoplankton and detritus of an autochthonous origin in the seston would represent a more usable energy source for benthic (bivalve molluscs, hydropsychid caddisflies) and planktonic (microcrustaceans) suspension feeders than the more refractory allochthonous materials derived from upstream processing of terrestrial organic matter. Benthic grazers depend heavily on nonfilamentous benthic algae (based on gut analysis from a separate study), but filamentous benthic algae have no apparent connection to invertebrate consumers (based on δ¹³C values). Amphipod and crayfish show a strong relationship to aquatic macrophytes (possibly through detrital organic matter rather than living plant tissue). These observations contrast with the prediction of the RCC that food webs in large rivers are based principally on refractory FTOM and dissolved organic matter from upstream inefficiencies in organic-matter processing and the bacteria growing upon these suspended or dissolved detrital compounds. The conclusions drawn here for the Ohio River cannot yet be extended to other floodplain and constricted-channel rivers in temperate and tropical latitudes until more comparable data are available on relatively pristine and moderately regulated rivers. Received: 3 January 1997 / Accepted: 28 August 1998     

Sources of organic matter in Ria Formosa revealed by stable isotope analysis

The aim of this study is to assess the major sources of organic matter for macroconsumers in the Ria Formosa tidal lagoon. The C, S and N isotopic natural abundances of abundant primary producers of particulate organic matter (POM) and Mytilus galloprovincialis muscle and digestive gland were analysed. The chlorophyll a (Chl a), the suspended particulate matter (SPM) and the POM were measured along the Faro-Olhão channel. The range of variation of stable isotope values among primary producers in Ria Formosa was low suggesting difficulties in the assessment of their relative contribution to higher levels of the food web. Chl a values decreased from outer station to inner station, while SPM and POM values increased. The multiple isotope approach illustrates that POM values along the Faro-Olhão channel, may result from a mixture of upland plants, benthic plants and phytoplankton. Mussel values indicate a selective diet of benthic macrophytes and phytoplankton, with the relative proportions of each determined by the location in the channel. During winters, the upland plants may be an important source of organic matter in the inner lagoon while phytoplankton was an important source of organic matter in the outer lagoon.     

Isotope ratios of cellulose from plants having different photosynthetic pathways

Hydrogen and carbon isotope ratios of cellulose nitrate and oxygen isotope ratios of cellulose from C₃, C₄, and Crassulacean acid metabolism (CAM) plants were determined for plants growing within a small area in Val Verde County, Texas. Plants having CAM had distinctly higher deuterium/hydrogen (D/H) ratios than plants having C₃ and C₄ metabolism. When hydrogen isotope ratios are plotted against carbon isotope ratios, each photosynthetic mode separates into a distinct cluster of points. C₄ plants had many D/H ratios similar to those of C₃ plants, so that hydrogen isotope ratios cannot be used to distinguish between these two photosynthetic modes. Portulaca mundula, which may have a modified photosynthetic mode between C₄ and CAM, had a hydrogen isotope ratio between those of the C₄ and CAM plants. When oxygen isotope ratios are plotted against carbon isotope ratios, no distinct clustering of the C₄ and CAM plants occurs. Thus, oxygen isotope ratios are not useful in distinguishing between these metabolic modes. A plot of hydrogen isotope ratios versus oxygen isotope ratios for this sample set shows considerable overlap between oxygen isotope ratios of the different photosynthetic modes without a concomitant overlap in the hydrogen isotope ratios of CAM and the other two photosynthetic modes. This observation is consistent with the hypothesis that higher D/H ratios in CAM plants relative to C₃ and C₄ plants are due to isotopic fractionations occurring during biochemical reactions.