How important are terrestrial organic carbon inputs for secondary production in freshwater ecosystems?

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Contribution of Chemoautotrophic Production to Freshwater Macroinvertebrates in a Headwater Stream Using Multiple Stable Isotopes

We estimated the food sources of macroinvertebrates using carbon, nitrogen, and sulfur stable isotopes in a headwater stream. Stream food webs, including macroinvertebrates, rely on production from autochthonous and allochthonous photosynthesis. We found that a freshwater grazer, the snail Semisulcospira libertina, may assimilate different food sources, based on stable carbon and nitrogen isotope evidence from snail muscle and the much lighter sulfur isotope signature than those of other associated macroinvertebrates. Previous studies have shown the importance of methanotrophic and sulfur bacteria in reductive environments (clay and organic‐rich sediments) as food sources for macroinvertebrates. Our results show that the production by chemoautotrophic bacteria contributes to the food sources of a snail in a stream. Thus, the chemoautotrophic bacteria are important in the freshwater food webs, even in mostly aerobic habitats. (© 2006 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Catchment land cover influences macroinvertebrate food‐web structure and energy flow pathways in mountain streams

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Cascading predator control interacts with productivity to determine the trophic level of biomass accumulation in a benthic food web

Large-scale exploitation of higher trophic levels by humans, together with global-scale nutrient enrichment, highlights the need to explore interactions between predator loss and resource availability. The hypothesis of exploitation ecosystems suggests that top–down and bottom–up control alternate between trophic levels, resulting in a positive relationship between primary production and the abundance of every second trophic level. Specifically, in food webs with three effective trophic levels, primary producers and predators should increase with primary production, while in food webs with two trophic levels, only herbivores should increase. We provided short-term experimental support for these model predictions in a natural benthic community with three effective trophic levels, where the number of algal recruits, but not the biomass of gastropod grazers, increased with algal production. In contrast, when the food web was reduced to two trophic levels by removing larger predators, the number of algal recruits was unchanged while gastropod grazer biomass increased with algal production. Predator removal only affected the consumer-controlled early life-stages of algae, indicating that both the number of trophic levels and the life-stage development of the producer trophic level determine the propagation of trophic cascades in benthic systems. Our results support the hypothesis that predators interact with resource availability to determine food-web structure.     

The importance of terrestrial subsidies in stream food webs varies along a stream size gradient

Cross system subsidies of energy and materials can be a substantial fraction of food web fluxes in ecosystems, especially when autochthonous production is strongly limited by light or nutrients. We explored whether assimilation of terrestrial energy varied in specific consumer taxa collected from streams of different sizes and resource availabilities. Since headwater streams are often unproductive, we expected that inputs from surrounding terrestrial systems (i.e. leaf litter, terrestrial invertebrates) would be a more important food source for consumers than in mid‐size rivers that have more open canopies and higher amounts of primary production available for consumers. We collected basal resources, invertebrates, and fish along a gradient in stream size in the Adirondack Mountains (NY, USA) and in Trinidad and Tobago and analyzed all samples for hydrogen isotopes as a means of differentiating biomass derived from allochthonous versus autochthonous sources. We found significant differences in allochthonous energy use within individual consumer taxa, showing that some taxa range from being entirely allochthonous to entirely autochthonous depending on where they were collected on the stream size gradient (grazers and collector–gatherer functional feeding groups), while other taxa are relatively fixed in the source of energy they assimilate (shredder and predator functional feeding groups). Consistent with expectations, allochthonous energy use was positively correlated with canopy cover in both regions for most feeding groups, with individuals from small, shaded streams having a more pronounced allochthonous signal than individuals collected from larger streams with less canopy cover. However, consumers in the shredder/detritivore feeding group did not vary among sites in their allochthonous energy use, and had a mostly allochthonous signal regardless of canopy cover and algal biomass. Our results demonstrate that the importance of energy from terrestrial subsidies can vary markedly but are similar in both temperate and tropical streams, suggesting a widely consistent pattern.     

Uptake of dissolved organic carbon by biofilms provides insights into the potential impact of loss of large woody debris on the functioning of lowland rivers

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Allochthonous and autochthonous carbon flows in food webs of tropical forest streams

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Ecosystem consequences of diversity depend on food chain length in estuarine vegetation

Biodiversity and food chain length each can strongly influence ecosystem functioning, yet their interactions rarely have been tested. We manipulated grazer diversity in seagrass mesocosms with and without a generalist predator and monitored community development. Changing food chain length altered biodiversity effects: higher grazer diversity enhanced secondary production, epiphyte grazing, and seagrass biomass only with predators present. Conversely, changing diversity altered top‐down control: predator impacts on grazer and seagrass biomass were weaker in mixed‐grazer assemblages. These interactions resulted in part from among‐species trade‐offs between predation resistance and competitive ability. Despite weak impact on grazer abundance at high diversity, predators nevertheless enhanced algal biomass through a behaviourally mediated trophic cascade. Moreover, predators influenced every measured variable except total plant biomass, suggesting that the latter is an insensitive metric of ecosystem functioning. Thus, biodiversity and trophic structure interactively influence ecosystem functioning, and neither factor's impact is predictable in isolation.     

Trophic trickles and cascades in a complex food web: impacts of a keystone predator on stream community structure and ecosystem processes

Chalk streams are among the most species-rich and productive of all temperate ecosystems. Despite this, a few keystone species have the potential to exert disproportionately powerful effects on community structure and ecosystem processes. Two of these are the bullhead Cottus gobio , a small benthic fish that is an extremely abundant, voracious predator, and the freshwater shrimp Gammarus pulex , which dominates the prey assemblage and is the principal detritivore. Field experiments detected a bullhead– Gammarus –detritus trophic cascade, with detrital processing rates slowed dramatically in the presence of the predator. In addition, survey data also revealed strong negative density-dependence between bullhead and brown trout, adding a further link in the cascade. However, although bullhead also depressed the abundance of a dominant grazer, the snail Potamopyrgus antipodarum , there was no cascading effect upon algal production, suggesting that autochthonous inputs were not controlled by top–down effects. This skewed effect of the predator upon autochthonous versus allochthonous basal resources stresses the need to consider both pathways of energy flux into the food web, whereas many previous studies have potentially overemphasized the importance of predator–herbivore–primary producer cascades. The wider community food web contained 142 species and 1383 feeding links. This complex network exhibited "small world" properties, such as high clustering (unlike many other food webs) and shortest path lengths between species were small (in common with many other food webs). In particular, each of the four members of the detrital cascade could be connected to any other species by three links or fewer. Our data revealed that powerful cascading effects can be imbedded within even very complex ecological networks.     

The role of C, N and P in dissolved and particulate organic matter as a nutrient source for phytoplankton growth, including toxic species

Phytoplankton have traditionally been regarded as strictly phototrophic, with a well defined position at the base of pelagic food webs. However, recently we have learned that the nutritional demands of a growing number of phytoplankton species can be met, at least partially, or under specific environmental conditions, through heterotrophy. Mixotrophy is the ability of an organism to be both phototrophic and heterotrophic, in the latter case utilizing either organic particles (phagotrophy) or dissolved organic substances (osmotrophy). This finding has direct implications for our view on algal survival strategies, particularly for harmful species, and energy- and nutrient flow in pelagic food webs. Mixotrophic species may outcompete strict autotrophs, e.g. in waters poor in inorganic nutrients or under low light. In the traditional view of the ‘microbial loop’ DOC is thought to be channeled from algal photosynthesis to bacteria and then up the food chain through heterotrophic flagellates, ciliates and mesozooplankton. Are mixotrophic phytoplankton that feed on bacteria also significantly contributing to this transport of photosynthetic carbon up the food chain? How can we estimate the fluxes of carbon and nutrients between different trophic levels in the plankton food web involving phagotrophic algae? These questions largely remain unanswered. In this review we treat evidence for both osmotrophy and phagotrophy in phytoplankton, especially toxic marine species, and some ecological implications of mixotrophy.     

Increased freshwater discharge shifts the trophic balance in the coastal zone of the northern Baltic Sea

Increased precipitation is one projected outcome of climate change that may enhance the discharge of freshwater to the coastal zone. The resulting lower salinity, and associated discharge of both nutrients and dissolved organic carbon, may influence food web functioning. The scope of this study was to determine the net outcome of increased freshwater discharge on the balance between auto‐ and heterotrophic processes in the coastal zone. By using long‐term ecological time series data covering 13 years, we show that increased river discharge suppresses phytoplankton biomass production and shifts the carbon flow towards microbial heterotrophy. A 76% increase in freshwater discharge resulted in a 2.2 times higher ratio of bacterio‐ to phytoplankton production (P_b:P_p). The level of P_b:P_p is a function of riverine total organic carbon supply to the coastal zone. This is mainly due to the negative effect of freshwater and total organic carbon discharge on phytoplankton growth, despite a concomitant increase in discharge of nitrogen and phosphorus. With a time lag of 2 years the bacterial production recovered after an initial decline, further synergistically elevating the microbial heterotrophy. Current climate change projections suggesting increased precipitation may therefore lead to increased microbial heterotrophy, thereby decreasing the transfer efficiency of biomass to higher trophic levels. This prognosis would suggest reduced fish production and lower sedimentation rates of phytoplankton, a factor of detriment to benthic fauna. Our findings show that discharge of freshwater and total organic carbon significantly contributes to the balance of coastal processes at large spatial and temporal scales, and that model's would be greatly augmented by the inclusion of these environmental drivers as regulators of coastal productivity.     

Autochthonous production in shallow littoral zones of five floodplain rivers: effects of flow,turbidity and nutrients

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Stable isotope analysis of production and trophic relationships in a tropical marine hard-bottom community

Seagrasses produce much of the organic carbon in the shallow waters of the Caribbean and it has long been assumed that a substantial portion of this carbon is exported to nearby habitats, contributing substantially to their food webs. In the shallow coastal waters of the Florida Keys (USA), seagrass intersperses with hard-bottom habitat where bushy, red macroalgae are the most prominent primary producers. However, the relative importance of seagrass-derived carbon versus autochthonous algal production or phytoplankton in supporting higher trophic levels within hard-bottom communities has never been investigated. We compared the carbon and nitrogen isotopic values of potential primary producers and representative higher trophic level taxa from hard-bottom sites on the bay-side and ocean-side of the Florida Keys. We also included in our study a set of bay-side sites that experienced significant ecological disturbances over the past decade (e.g., cyanobacteria blooms, seagrass die-off, and sponge die-offs) that may have altered trophic relationships in those regions. We did not detect any differences among regions in the trophic status of hard-bottom taxa that might be associated with ecosystem disturbance. However, our results suggest that autochthonous production of algal detritus is an important source of secondary production in these hard-bottom communities, with seagrass and phytoplankton contributing smaller fractions.     

Tracking the autochthonous carbon transfer in stream biofilm food webs

Food webs in the rhithral zone rely mainly on allochthonous carbon from the riparian vegetation. However, autochthonous carbon might be more important in open canopy streams. In streams, most of the microbial activity occurs in biofilms, associated with the streambed. We followed the autochthonous carbon transfer toward bacteria and grazing protozoa within a stream biofilm food web. Biofilms that developed in a second-order stream (Thuringia, Germany) were incubated in flow channels under climate-controlled conditions. Six-week-old biofilms received either 13C- or 12C-labeled CO?, and uptake into phospholipid fatty acids was followed. The dissolved inorganic carbon of the flow channel water became immediately labeled. In biofilms grown under 8-h light/16-h dark conditions, more than 50% of the labeled carbon was incorporated in biofilm algae, mainly filamentous cyanobacteria, pennate diatoms, and nonfilamentous green algae. A mean of 29% of the labeled carbon reached protozoan grazer. The testate amoeba Pseudodifflugia horrida was highly abundant in biofilms and seemed to be the most important grazer on biofilm bacteria and algae. Hence, stream biofilms dominated by cyanobacteria and algae seem to play an important role in the uptake of CO? and transfer of autochthonous carbon through the microbial food web.     

Biomass Transfer Subsidizes Nitrogen to Offshore Food Webs

We evaluated the potential contribution of allochthonous biomass subsidies to the upper trophic levels of offshore food webs in the northeastern Gulf of Mexico (GOM). We made this evaluation considering nitrogen, an essential and often limiting nutrient in coastal ecosystems, to estimate the potential production of within-ecosystem biomass relative to the known import of biomass from an adjacent seagrass-dominated ecosystem. When adjusted for trophic transfer efficiency, we found the biomass subsidy from a single species (pinfish, Lagodon rhomboides) from nearshore seagrass habitat to the offshore GOM to be greater than the amount of nitrogen exported by two major rivers and local submarine ground water discharge. Our calculations show that seagrass-derived biomass accounts for approximately 25% of the total potential production in the northeastern GOM. This estimate is in agreement with a previous study that found 18.5–25% of the biomass in a predatory reef fish was derived from seagrass biomass inputs. These results indicate that all of the sources we consider account for the majority of the nitrogen available to the food web in the northeastern GOM. Our approach could be adapted to other coupled ecosystems to determine the relative importance of biomass subsidies to coastal ocean food webs.     

Food web pathways for fish communities in small tropical streams

1. It is often assumed that invertebrate consumers in small tropical streams are dependent on allochthonous sources, although recent studies indicate that algae can form the base of food webs in tropical streams. Fish in tropical streams can feed across several trophic levels and the origin and path of energy and nutrient flow is uncertain for many species.
2. We collected fish, insects, periphyton, and leaf litter from 20 streams across four Atlantic Forest catchments. We analysed stomach contents of fish to define trophic guild and fish dietary trophic position. We also analysed stable isotopes of carbon and nitrogen of fish and their resources to identify the main basal resources of the food web and to estimate trophic positions and identify the path of energy flow.
3. We found that autochthonous sources were the primary resource base for fish communities. Trophic positions estimated from diet and isotopes were similar and correlated for insectivore and algivore–insectivore fish, but not for algivore–detritivore or omnivore fish. Using path analysis, fish classified as algivore–detritivores appear to have derived their biomass through a diet of primary consumer insects and periphytic algae and thus, are more likely to play a trophic role as algivore–insectivores in these streams. However, omnivores probably derived much of their biomass from aquatic insects.
4. Our findings support other studies of tropical systems in which the main basal resource is autochthonous, even in small streams. We also show that the assignment to a specific trophic guild for some fish species, based on gut contents, does not reflect what they assimilate into their bodies. In some species, food sources that are uncommon can make a disproportionately important contribution to their biomass.
5. This study affirms the important role of inconspicuous algal resources in aquatic food webs, even in small forested streams, and demonstrates the effectiveness of taking a combined approach of diet analysis, isotopic tracing, and modelling to resolve food web pathways where the level of omnivory is high.

     

Biogenic methane in freshwater food webs

1. It has long been known that substantial amounts of methane are produced in anoxic lake sediments, and the components of the methane cycle in lakes have been well described. At oxic–anoxic interfaces, methane‐oxidising bacteria (MOB) convert methane to microbial biomass and can be highly productive. However, only recently has methane been recognised as a potentially important carbon and energy source for lake food webs, and some instances have also been reported of methane contribution to river food webs. Stable isotope analysis (SIA) has provided compelling evidence in this respect and has been supplemented by other lines of evidence. 2. In the benthic food webs of lakes, profundal chironomid larvae appear to be the main conduits for trophic transfer of biogenic methane via grazing on MOB. The mode of feeding of these larvae and the microhabitats they generate both promote larval ability to exploit MOB production. Support to chironomid larvae from methane is rather widespread, but its degree is highly variable; estimates suggest that in some lakes methane‐carbon might contribute more than 60% of chironomid carbon biomass. 3. Evidence of crustacean zooplankton in lakes deriving part of their carbon from methane is currently more limited. Reports from some lakes have indicated Daphnia with a substantial (>50%) contribution of methane‐carbon in their biomass. However, for this to happen, an oxic–anoxic interface where sufficient MOB production can occur needs to be within the range of vertical migrations by zooplankton, which may only rarely be the case. Hence, a significant methane subsidy of pelagic food webs in lakes is probably much less widespread than for benthic food webs. 4. There is also recent and currently very limited evidence that some stream benthos derives biomass carbon (reported values up to 30%) from methane. This can occur in stagnant backwater pools where conditions can be analogous to those in lake sediments. However, groundwater aquifers can also supply water supersaturated with methane to some rivers, providing a basis for a microbially‐mediated transfer of methane‐carbon to river benthos. 5. Evidence for significant transfer of methane‐derived carbon to higher trophic levels is still very limited. Within some lakes, those fish species that feed extensively on chironomid larvae can derive a substantial part (perhaps up to 20%) of their carbon biomass from methane. It is also likely that methane‐carbon produced in lakes or rivers is exported to riparian ecosystems when emerging chironomids or other insects are eaten by invertebrate or avian predators. 6. We argue that conceptual models of freshwater food webs, and especially those for lakes, need to be modified to enable incorporation of biogenic methane as a carbon and energy source. For some types of lakes, carbon and energy budgets certainly need to take account of the production and utilisation of biogenic methane, and the accumulating evidence indicates that this is a more widespread phenomenon that has generally been acknowledged hitherto.     

Net heterotrophy in Faroe Islands clear-water lakes: causes and consequences for bacterioplankton and phytoplankton

1. Five oligotrophic clear‐water lakes on the Faroe Islands were studied during August 2000. Algal and bacterial production rates, community respiration, and CO₂ saturation were determined. In addition, we examined the plankton community composition (phytoplankton and heterotrophic nanoflagellates) and measured the grazing pressure exerted by common mixotrophic species on bacteria. 2. High respiration to primary production (6.6–33.2) and supersaturation of CO₂ (830–2140 μatm) implied that the lakes were net heterotrophic and that the pelagic heterotrophic plankton were subsidised by allochthonous organic carbon. However, in spite of the apparent high level of net heterotrophy, primary production exceeded bacterial production and the food base for higher trophic levels appeared to be mainly autotrophic. 3. We suggest that the observed net heterotrophy in these lakes was a result of the oligotrophic conditions and hence low primary production in combination with an input of allochthonous C with a relatively high availability. 4. Mixotrophic phytoplankton (Cryptomonas spp., Dinobryon spp. and flagellates cf. Ochromonas spp.) constituted a large percentage of the plankton community (17–83%), possibly as a result of their capacity to exploit bacteria as a means of acquiring nutrients in these nutrient poor systems.     

Resolving the predator first paradox: Arthropod predator food webs in pioneer sites of glacier forelands

Primary succession on bare ground surrounded by intact ecosystems is, during its first stages, characterized by predator‐dominated arthropod communities. However, little is known on what prey sustains these predators at the start of succession and which factors drive the structure of these food webs. As prey availability can be extremely patchy and episodic in pioneer stages, trophic networks might be highly variable. Moreover, the importance of allochthonous versus autochthonous food sources for these pioneer predators is mostly unknown. To answer these questions, the gut content of 1,832 arthropod predators, including four species of carabid beetles, two lycosid and several linyphiid spider species caught in early and late pioneer stages of three glacier forelands, was screened molecularly to track intraguild and extraguild trophic interactions among all major prey groups occurring in these systems. Two‐thirds of the 2,310 identified food detections were collembolans and intraguild prey, while one‐third were allochthonous flying insects. Predator identity and not successional stage or valley had by far the strongest impact on the trophic interaction patterns. Still, the variability of prey spectra increased significantly from early to late pioneer stage, as did the niche width of the predators. As such the structure of pioneer arthropod food webs in recently deglaciated Alpine habitats seems to be driven foremost by predator identity while site and early successional effects contribute to a lesser extent to food web variability. Our findings also suggest that in these pioneer sites, predatory arthropods depend less on allochthonous aeolian prey but are mainly sustained by prey of local production.