Carbon and nitrogen stable isotope ratios of macroinvertebrates in the littoral zone of Lake Biwa as indicators of anthropogenic activities in the watershed

Carbon and nitrogen stable isotope ratios (δ¹³C and δ¹⁵N) of macroinvertebrates inhabiting littoral zones of lakes can serve as useful indicators of material loading from the watershed. We collected snails (Semisulcospira spp.) and bivalves (Unio douglasiae biwae Kobelt) from 29 littoral sites in Lake Biwa near the mouths of river tributaries with various human population density (HPD) and land-use patterns. The δ¹³C and δ¹⁵N signatures were determined for three potential food sources: particulate organic matter in the pelagic zone (PPOM), riverine particulate organic matter from tributaries (RPOM) and epilithic organic matter in the littoral zone (EOM). The stable isotope mixing model revealed that snails relied mainly on EOM, and bivalves on PPOM and RPOM. Multiple regression analysis showed that intersite variation in δ¹⁵N for snails was best explained by HPD, while variation in δ¹⁵N of EOM and nitrate was explained to a lesser extent by HPD. Comparison with isotope signatures of their food sources and riverine nutrients revealed that snails assimilated anthropogenic nitrogen from wastewater in the watershed. Our results also showed that the δ¹³C value of bivalves was marginally related to the fraction of paddy fields in the watersheds. In conclusion, the isotope signatures of macroinvertebrates inhabiting the littoral zone can be useful indicators of anthropogenic impacts from the watershed.     

Aquatic microfauna alter larval food resources and affect development and biomass of West Nile and Saint Louis encephalitis vector Culex nigripalpus (Diptera: Culicidae)

Ciliate protists and rotifers are ubiquitous in aquatic habitats and can comprise a significant portion of the microbial food resources available to larval mosquitoes, often showing substantial declines in abundance in the presence of mosquito larvae. This top‐down regulation of protists is reported to be strong for mosquitoes inhabiting small aquatic containers such as pitcher plants or tree holes, but the nature of these interactions with larval mosquitoes developing in other aquatic habitats is poorly understood. We examined the effects of these two microbial groups on lower trophic level microbial food resources, such as bacteria, small flagellates, and organic particles, in the water column, and on Culex larval development and adult production. In three independent laboratory experiments using two microeukaryote species (one ciliate protist and one rotifer) acquired from field larval mosquito habitats and cultured in the laboratory, we determined the effects of Culex nigripalpus larval grazing on water column microbial dynamics, while simultaneously monitoring larval growth and development. The results revealed previously unknown interactions that were different from the top‐down regulation of microbial groups by mosquito larvae in other systems. Both ciliates and rotifers, singly or in combination, altered other microbial populations and inhibited mosquito growth. It is likely that these microeukaryotes, instead of serving as food resources, competed with early instar mosquito larvae for microbes such as small flagellates and bacteria in a density‐dependent manner. These findings help our understanding of the basic larval biology of Culex mosquitoes, variation in mosquito production among various larval habitats, and may have implications for existing vector control strategies and for developing novel microbial‐based control methods.     

Feeding ecology of pelagic fish larvae and juveniles in slope waters of the Gulf of Mexico

Stable isotope ratios of carbon (δ¹³C) and nitrogen (δ¹⁵N) were used to investigate feeding patterns of larval and early juvenile pelagic fishes in slope waters of the Gulf of Mexico. Contribution of organic matter supplied to fishes and trophic position within this pelagic food web was estimated in 2007 and 2008 by comparing dietary signatures of the two main producers in this ecosystem: phytoplankton [based on particulate organic matter (POM)] and Sargassum spp. Stable isotope ratios of POM and pelagic Sargassum spp. were significantly different from one another with δ¹³C values of POM depleted by 3–6‰ and δ¹⁵N values enriched by 2 relative to Sargassum spp. Stable isotope ratios were significantly different among the five pelagic fishes examined: blue marlin Makaira nigricans, dolphinfish Coryphaena hippurus, pompano dolphinfish Coryphaena equiselis, sailfish Istiophorus platypterus and swordfish Xiphias gladius. Mean δ¹³C values ranged almost 2 among fishes and were most depleted in I. platypterus. In addition, mean δ¹⁵N values ranged 4–5 with highest mean values found for both C. hippurus and C. equiselis and the lowest mean value for M. nigricans during both years. Increasing δ¹³C or δ¹⁵N with standard length suggested that shifts in trophic position and diet occurred during early life for several species examined. Results of a two‐source mixing model suggest approximately an equal contribution of organic matter by both sources (POM = 55%; pelagic Sargassum spp. = 45%) to the early life stages of pelagic fishes examined. Contribution of organic matter, however, varied among species, and sensitivity analyses indicated that organic source estimates changed from 2 to 13% for a δ¹³C fractionation change of ±0·25‰ or a δ¹⁵N fractionation change of ± 1·0‰ relative to original fractionation values.     

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     

Tracing biogeochemical subsidies from glacier runoff into Alaska's coastal marine food webs

Nearly half of the freshwater discharge into the Gulf of Alaska originates from landscapes draining glacier runoff, but the influence of the influx of riverine organic matter on the trophodynamics of coastal marine food webs is not well understood. We quantified the ecological impact of riverine organic matter subsidies to glacier‐marine habitats by developing a multi‐trophic level Bayesian three‐isotope mixing model. We utilized large gradients in stable (δ¹³C, δ¹⁵N, δ²H) and radiogenic (Δ¹⁴C) isotopes that trace riverine and marine organic matter sources as they are passed from lower to higher trophic levels in glacial‐marine habitats. We also compared isotope ratios between glacial‐marine and more oceanic habitats. Based on isotopic measurements of potential baseline sources, ambient water and tissues of marine consumers, estimates of the riverine organic matter source contribution to upper trophic‐level species including fish and seabirds ranged from 12% to 44%. Variability in resource use among similar taxa corresponded to variation in species distribution and life histories. For example, riverine organic matter assimilation by the glacier‐nesting seabirds Kittlitz's murrelet (Brachyramphus brevirostris) was greater than that of the forest‐nesting marbled murrelet (B. marmoratus). The particulate and dissolved organic carbon in glacial runoff and near surface coastal waters was aged (12100–1500 years BP ¹⁴C‐age) but dissolved inorganic carbon and biota in coastal waters were young (530 years BP ¹⁴C‐age to modern). Thus terrestrial‐derived subsidies in marine food webs were primarily composed of young organic matter sources released from glacier ecosystems and their surrounding watersheds. Stable isotope compositions also revealed a divergence in food web structure between glacial‐marine and oceanic sites. This work demonstrates linkages between terrestrial and marine ecosystems, and facilitates a greater understanding of how climate‐driven changes in freshwater runoff have the potential to alter food web dynamics within coastal marine ecosystems in Alaska.     

Design and management of free water surface constructed wetlands to minimize mosquito production

Constructed wetland technology has broad applications for the treatment of many types of wastewaters and provides an ecological approach to mitigate the release of nutrients and toxic materials into the environment. However, design features, maintenance activities and the characteristics of the wastewater undergoing treatment contribute differentially to potential levels of mosquito production and, consequently, to threats to human and animal health from mosquito-borne pathogens. Of the variables typically considered when designing free-water surface constructed wetlands for the improvement of water quality of municipal and agricultural wastewaters, organic loading (i.e., biochemical oxygen demand, total suspended solids), nutrients (nitrogen and phosphorus), and the configuration and maintenance of emergent vegetation can have strong effects on mosquito production. The production of Culex vectors of encephalitides and filarial worms is directly related to loading rates of organic matter and bottom-up enrichment of larval mosquito resources and their interaction with design features and management practices that reduce the physical and biological factors causing mortality of immature mosquitoes. As loading rates of organic matter and nutrients decline, the diversity of mosquitoes produced by treatment wetlands tends to increase and the relative abundance of Anopheles species, which are capable of vectoring the causative agents of malaria, increases in temperate man-made wetlands. Habitat features and management practices that create intermittently inundated substrate can lead to the production of other mosquitoes (i.e., Aedes, Psorophora) with floodwater life histories. Constructed wetland technology is expected to play an increasing role in water treatment and reclamation in tropical and subtropical countries where virulent mosquito-borne pathogens already cause significant levels of morbidity and mortality.     

Food resources of shredders and other benthic macroinvertebrates in relation to shading conditions in tropical Hong Kong streams

1. Gut content analyses (GCA) of benthic macroinvertebrates, supplemented by carbon and nitrogen stable isotope analyses (SIA), were used to determine the relative contribution of leaf litter and autochthonous food sources to consumer biomass in five shaded and five unshaded streams in tropical Hong Kong. 2. Only four obligate shredders and two facultative shredders were identified out of 58 morphospecies dissected. Non‐shredder taxa consumed little (<23% food eaten) coarse particulate organic matter (CPOM) in spite of its abundance in streams, and GCA revealed that fine particulate organic matter was the major food (25–99%) of most primary consumers. 3. Stable isotope analysis results were in general agreement with the findings of GCA, and confirmed that three of the four obligate shredders had a high dependence (55–78% of assimilated carbon) on CPOM. 4. Autochthonous energy sources were important in all streams: non‐shredding primary consumers examined, which accounted for 72% of total macroinvertebrate abundance in shaded streams, derived (on average) 61% of their biomass from autochthonous foods; the equivalent values for unshaded streams were 72% (abundance) and 71% (biomass).     

Flow intermittency influences the trophic base,but not the overall diversity of alpine stream food webs

Alpine streams can exhibit naturally high levels of flow intermittency. However, how flow intermittency in alpine streams affects ecosystem functions such as food web trophic structure is virtually unknown. Here, we characterized the trophic diversity of aquatic food webs in 28 headwater streams of the Val Roseg, a glacierized alpine catchment. We compared stable isotope (δ¹³C and δ¹⁵N) trophic indices to high temporal resolution data on flow intermittency. Overall trophic diversity, food chain length and diversity of basal resource use did not differ to a large extent across streams. In contrast, gradient and mixing model analysis indicated that primary consumers assimilated proportionally more periphyton and less allochthonous organic matter in more intermittent streams. Higher coarse particulate organic matter (CPOM) C:N ratios were an additional driver of changes in macroinvertebrate diets. These results indicate that the trophic base of stream food webs shifts away from terrestrial organic matter to autochthonous organic matter as flow intermittency increases, most likely due to reduced CPOM conditioning in dry streams. This study highlights the significant, yet gradual shifts in ecosystem function that occur as streamflow becomes more intermittent in alpine streams. As alpine streams become more intermittent, identifying which functional changes occur via gradual as opposed to threshold responses is likely to be vitally important to their management and conservation.     

The functional role of native freshwater mussels in the fluvial benthic environment

1. Freshwater mussels are the dominant consumer biomass in many fluvial systems. As filter feeding grazers, mussels can remove large amounts of particulate matter from the water column and transfer these resources to the substrate as biodeposits (agglutinated mussel faeces and pseudofaeces). Mussel biodeposits are a nutrient rich and easily assimilated food source and therefore may have significant relevance to benthic community structure. This study examines the functional role of Margaritifera falcata in the South Fork Eel River, California. 2. We addressed two main questions: (i) Do mussels increase benthic resources in this system? (ii) If so, does this alter macroinvertebrate community structure? 3. Measurements and enclosure experiments in the South Fork Eel River show that mussels can play a significant role in local food webs by increasing available fine particulate matter (both organic and inorganic) on the substrate. We document increased benthic macroinvertebrate biomass for predators and collectors (Leptophlebidae) in the presence of mussels, but only in late summer.     

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

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

     

Uncovering trophic positions and food resources of soil animals using bulk natural stable isotope composition

Despite the major importance of soil biota in nutrient and energy fluxes, interactions in soil food webs are poorly understood. Here we provide an overview of recent advances in uncovering the trophic structure of soil food webs using natural variations in stable isotope ratios. We discuss approaches of application, normalization and interpretation of stable isotope ratios along with methodological pitfalls. Analysis of published data from temperate forest ecosystems is used to outline emerging concepts and perspectives in soil food web research. In contrast to aboveground and aquatic food webs, trophic fractionation at the basal level of detrital food webs is large for carbon and small for nitrogen stable isotopes. Virtually all soil animals are enriched in ¹³C as compared to plant litter. This ‘detrital shift’ likely reflects preferential uptake of ¹³C‐enriched microbial biomass and underlines the importance of microorganisms, in contrast to dead plant material, as a major food resource for the soil animal community. Soil organic matter is enriched in ¹⁵N and ¹³C relative to leaf litter. Decomposers inhabiting mineral soil layers therefore might be enriched in ¹⁵N resulting in overlap in isotope ratios between soil‐dwelling detritivores and litter‐dwelling predators. By contrast, ¹³C content varies little between detritivores in upper litter and in mineral soil, suggesting that they rely on similar basal resources, i.e. little decomposed organic matter. Comparing vertical isotope gradients in animals and in basal resources can be a valuable tool to assess trophic interactions and dynamics of organic matter in soil. As indicated by stable isotope composition, direct feeding on living plant material as well as on mycorrhizal fungi is likely rare among soil invertebrates. Plant carbon is taken up predominantly by saprotrophic microorganisms and channelled to higher trophic levels of the soil food web. However, feeding on photoautotrophic microorganisms and non‐vascular plants may play an important role in fuelling soil food webs. The trophic niche of most high‐rank animal taxa spans at least two trophic levels, implying the use of a wide range of resources. Therefore, to identify trophic species and links in food webs, low‐rank taxonomic identification is required. Despite overlap in feeding strategies, stable isotope composition of the high‐rank taxonomic groups reflects differences in trophic level and in the use of basal resources. Different taxonomic groups of predators and decomposers are likely linked to different pools of organic matter in soil, suggesting different functional roles and indicating that trophic niches in soil animal communities are phylogenetically structured. During last two decades studies using stable isotope analysis have elucidated the trophic structure of soil communities, clarified basal food resources of the soil food web and revealed links between above‐ and belowground ecosystem compartments. Extending the use of stable isotope analysis to a wider range of soil‐dwelling organisms, including microfauna, and a larger array of ecosystems provides the perspective of a comprehensive understanding of the structure and functioning of soil food webs.     

Metabolic origin of δ15N values in nitrogenous compounds from Brassica napus L. leaves

Nitrogen isotope composition (δ¹⁵N) in plant organic matter is currently used as a natural tracer of nitrogen acquisition efficiency. However, the δ¹⁵N value of whole leaf material does not properly reflect the way in which N is assimilated because isotope fractionations along metabolic reactions may cause substantial differences among leaf compounds. In other words, any change in metabolic composition or allocation pattern may cause undesirable variability in leaf δ¹⁵N. Here, we investigated the δ¹⁵N in different leaf fractions and individual metabolites from rapeseed (Brassica napus) leaves. We show that there were substantial differences in δ¹⁵N between nitrogenous compounds (up to 30‰) and the content in (¹⁵N enriched) nitrate had a clear influence on leaf δ¹⁵N. Using a simple steady‐state model of day metabolism, we suggest that the δ¹⁵N value in major amino acids was mostly explained by isotope fractionation associated with isotope effects on enzyme‐catalysed reactions in primary nitrogen metabolism. δ¹⁵N values were further influenced by light versus dark conditions and the probable occurrence of alternative biosynthetic pathways. We conclude that both biochemical pathways (that fractionate between isotopes) and nitrogen sources (used for amino acid production) should be considered when interpreting the δ¹⁵N value of leaf nitrogenous compounds.     

The shredding activity of gammarids facilitates the processing of organic matter by the subterranean amphipod Niphargus rhenorhodanensis

1. The functional feeding group approach has been widely used to describe the community structure of benthic invertebrates in relation to organic matter resources. Based on this functional framework, positive interactions between feeding groups (especially shredders and collector‐gatherers) were postulated in the River Continuum Concept. However, relationships with organic matter have been poorly documented for invertebrates living in the hyporheic zone. 2. We hypothesised that the common subterranean amphipod Niphargus rhenorhodanensis would feed on fine particulate organic matter (FPOM), which is more abundant than coarse particulate organic matter (CPOM) in hyporheic habitats, and should be favoured by the occurrence of shredders that produce FPOM from CPOM. 3. We used laboratory experiments to quantify leaf litter processing by N. rhenorhodanensis and a common shredder, the surface amphipod Gammarus roeselii. We estimated rates of feeding and assimilation (using nitrogen stable isotopes) of the two species separately and together to reveal any potential shredder–collector facilitation between them. 4. Measured leaf litter mass loss showed that N. rhenorhodanensis did not act as a shredder, unlike G. roeselii. Organic matter dynamics and ¹⁵N/¹⁴N ratios in tissues of niphargids indicated that N. rhenorhodanensis was a collector‐gatherer feeding preferentially on FPOM. We also found a positive influence of the gammarid shredders on the assimilation rate of N. rhenorhodanensis, which fed on FPOM produced by the shredders, supporting the hypothesis of a positive interaction between surface shredders and hyporheic collector‐gatherers.     

Carbon sources and trophic structure in a macrophyte‐dominated polyculture pond assessed by stable‐isotope analysis

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Habitat and resource use by larval and juvenile fishes in an arid-land river (Rio Grande, New Mexico)

1. Studies of mesic temperate and tropical rivers suggest an important role for floodplain habitats as nursery areas for larval and juvenile fishes. In arid‐land rivers the extent and duration of flooding is diminished and habitats and resources used by larval fishes are poorly known. Our study documented habitat and resource use of larval fishes in the Rio Grande, New Mexico, an arid‐land river. 2. Spatial and temporal distribution of larval and juvenile fishes and their inferred microhabitat preferences were studied during spring, summer and autumn, 2003. Stable carbon (¹³C : ¹²C) and nitrogen (¹⁵N : ¹⁴N) isotope ratios were measured to identify nutrient sources and characterise trophic positions of young‐of‐year fishes in this system. 3. Some fishes recruited during high flows (in spring), whereas others recruited during low‐flow periods in late summer. Regardless of the timing of reproduction, microhabitats with lower current velocity and higher temperature appeared to serve as vital nursery grounds for Rio Grande fishes. Ephemeral backwaters and disconnected side channels held the highest abundance and diversity of larvae and juveniles. 4. Stable isotope analyses revealed that fish larvae obtained carbon predominately from algal production in early summer, but used organic carbon derived from emergent macrophytes as river discharge decreased in mid‐summer. This shift may have been facilitated by microinvertebrate prey that grazed down edible algae and then switched to macrophytes in mid‐summer. Nitrogen isotope ratios did not differ among species or early life stages, suggesting that larval and juvenile fishes use similar food resources, especially when restricted to isolated pools in summer.     

Small reductions in forest cover weaken terrestrial-aquatic linkages in headwater streams

1. We assessed the impacts of deforestation on the energy base of headwater food webs in seven headwater streams in the Upper Chattahoochee basin, GA, U.S.A where percentage forest in catchments ranged from 82 to 96%. We measured terrestrial organic matter standing crop and determined consumer (crayfish and insectivorous fish) dependence on terrestrial versus aquatic energy sources via gut content and stable isotope analyses. 2. Standing crop of coarse particulate organic matter (CPOM) declined with deforestation at large scales (i.e. catchment deforestation and riparian deforestation at the entire stream network scale). Terrestrial plant matter, the dominant component of crayfish guts, declined in crayfish guts with reductions in CPOM standing crop and with deforestation. 3. Crayfish and insectivorous fish δ¹³C showed enrichment trends with deforestation, indicating isotopic divergence from CPOM, the most ¹³C‐depleted basal resource, with reductions in catchment and riparian forest cover. Crayfish δ¹³C also exhibited enrichment with decreased instream CPOM standing crop. 4. A concentration‐dependent mixing model was used to calculate the relative dependence of crayfish and fish on terrestrial versus aquatic basal resources. Results suggested that both allochthonous CPOM and autochthonous production were important basal resources. Consumer dependence on CPOM decreased with reductions in canopy cover. 5. Our data suggest the importance of forest cover to headwater food webs at multiple scales, and that relatively low levels of riparian deforestation along headwater streams can lead to reductions in stream food web dependence on terrestrial subsidies.     

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

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

     

Significance of instream autotrophs in trophic dynamics of the Upper Mississippi River

Trophic dynamics of large river–floodplain ecosystems are still not well understood despite development of several conceptual models over the last 25 years. To help resolve questions about the relative contribution of algal and detrital organic matter to food webs in the Upper Mississippi River, we (1) separated living and detrital components of ultrafine and fine transported organic matter (UTOM and FTOM, respectively) by colloidal silica centrifugation; (2) identified stable isotope signatures (δ¹³C and δ¹⁵N) for these two portions of transported organic matter and other potential organic matter sources; and (3) employed a multiple source, dual-isotope mixing model to determine the relative contribution of major energy sources to primary consumers and the potential contribution of basal sources to the biomass of secondary consumers. The δ¹³C and δ¹⁵N of living and detrital fractions of UTOM and FTOM were distinct, indicating clear differences in isotopic composition of the algal and detrital fractions of transported organic matter. Living and detrital transported organic matter also differed from other potential organic matter sources by either δ¹³C or δ¹⁵N. A six-source mixing model using both δ¹³C and δ¹⁵N indicated that algal transported organic matter was the major resource assimilated by primary consumers. The contribution of detrital transported organic matter was small in most cases, but there were a small number of taxa for which it could potentially contribute to more than half the assimilated diet. Colloidal dissolved organic matter, which includes heterotrophic bacteria, accounted for only a small fraction of the organic matter assimilated by most primary consumers, indicating that coupling between microbial processes and metazoan production is minimal. Terrestrial C₃ litter from the floodplain forest floor and aquatic macrophytes were also relatively unimportant to the assimilated diet of primary consumers. Application of the mixing model to compare basal source isotopic ratios to secondary consumers revealed that most organic matter moving from primary to secondary consumers originated from algal TOM. Our findings indicate that autochthonous organic matter is the major energy source supporting metazoan production in the main channel of this large river, at least during the summer. This study joins a number of other investigations performed globally that indicate organic matter originating from instream production of sestonic and benthic microalgae is a major driver in the trophic dynamics of large river ecosystems.     

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     

Wetland isolation facilitates larval mosquito density through the reduction of predators

• 1 Wetlands harbour high biodiversity and offer important ecosystem services, but they are also a habitat for mosquito larvae (Diptera: Culicidae), which are important disease vectors.
• 2 Isolation among remnant, or newly created wetlands and ponds, and their consequent density in the landscape, is a key factor that can influence a variety of food web processes, including effects on mosquitoes which are important prey to many predators.
• 3 We assess the impact of habitat isolation on the density of pond‐breeding mosquitoes (several Anopheles and Culex species) both directly and indirectly through the food web.
• 4 Results from structural equation modelling of survey data shows that larval mosquitoes are denser in ponds that are more isolated from one another, and that this result was primarily driven indirectly by a reduction of larval mosquito predators (e.g. predaceous insects and amphibians). Furthermore, results from a long‐term mesocosm experiment factorially manipulating isolation and predator reduction show that the effect of isolation on mosquito density was eliminated when predators were experimentally reduced.
• 5 It is concluded that metacommunity processes, both directly and indirectly mediated through predators, can play an important role in the local abundance of wetland breeding mosquitoes and possibly the diseases they spread.