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

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Low Levels of Allochthony in Consumers Across Three High-Elevation Lake Types

The integration of lakes into watershed-scale energy budgets remains a major goal of aquatic ecosystem ecology. However, this enterprise has focused almost exclusively on temperate and boreal systems and on zooplankton as representatives of system-wide energy flow. We investigated the proportion of consumer biomass derived from terrestrial sources, allochthony, in three classes of high-elevation lakes—alpine, large and small montane—of varying geometry and watershed ecosystem development, and across five taxa, including macrobenthic invertebrates and fish. We used stable isotopes of carbon (¹³C:¹²C), nitrogen (¹⁵N:¹⁴N), and hydrogen (²H:¹H) to fit a modular Bayesian mixing model, which estimated proportional assimilation of phytoplankton, algal periphyton, and terrestrial organic matter by each consumer. The stable isotope analysis was supplemented with a comparison of fatty acid profiles between consumers and producers, and with a Daphnia magna rearing study involving aquatic and terrestrial nutrient sources. The most probable estimate of allochthony across consumer taxa was 41% in small montane lakes (< 0.1 ha) with high terrestrial C loading. For large montane (3–11 ha) and alpine lakes (0.8–3 ha), with substantially less terrestrial influence, allochthony was just 4 and 3%, respectively. Allochthony was also lower on average for benthic grazers than for pelagic consumers. Our results corroborate previous findings that lake size, depth, and light penetration are dominant physical controls on allochthony, but add that it sharply declines at high elevation due to changes in terrestrial primary production near or above tree line.     

Boomerang ecosystem fluxes: organic carbon inputs from land to lakes are returned to terrestrial food webs via aquatic insects

Many ecosystems are linked to their adjacent ecosystems by movements of organisms. For instance, aquatic and terrestrial ecosystems are linked via emerging aquatic insects that serve as prey for terrestrial consumers. However, the role of these organisms in returning recycled carbon to the ecosystem from which it originated is not well known. This is due to the fact that values of carbon isotope signatures from terrestrial leaves and aquatic resources are usually similar and hence results of isotope mixing models need to be considered with caution. We overcame this problem by adding isotopically distinct terrestrial particulate organic carbon (tPOC) as a tracer to the experimental sides of two lakes that were divided in two equal halves with plastic curtains. We focused on aquatic insect larvae (Chironomidae) that fed on maize Zea mays leaves experimentally added to the lakes, and subsequently became prey for terrestrial predators (spiders) after emergence. The carbon isotope values of Chironomidae and spiders were significantly elevated in the lake treatment sides as compared to reference sides, whereas the values of all autochthonous resources were not affected by maize additions. Estimates from stable isotope mixing models indicated a low but demonstrable contribution of maize leaves to the diet of Chironomidae. Overlap between the isotope values of alder leaves, the major natural tPOC source, and autochthonous resources prevented a reliable quantification of allochthony of Chironomidae. However, we qualitatively demonstrated the flow of terrestrial particulate organic carbon to lakes, as leaf fall, and back to terrestrial surroundings via emerging insects. This ‘boomerang’ carbon flux between land and lakes blurs the distinction between autochthonous and allochthonous carbon sources.     

Zooplankton allochthony is spatially heterogeneous in a boreal lake

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Terrestrial subsidies to lake food webs: an experimental approach

Cross-ecosystem movements of material and energy are ubiquitous. Aquatic ecosystems typically receive material that also includes organic matter from the surrounding catchment. Terrestrial-derived (allochthonous) organic matter can enter aquatic ecosystems in dissolved or particulate form. Several studies have highlighted the importance of dissolved organic carbon to aquatic consumers, but less is known about allochthonous particulate organic carbon (POC). Similarly, most studies showing the effects of allochthonous organic carbon (OC) on aquatic consumers have investigated pelagic habitats; the effects of allochthonous OC on benthic communities are less well studied. Allochthonous inputs might further decrease primary production through light reduction, thereby potentially affecting autotrophic resource availability to consumers. Here, an enclosure experiment was carried out to test the importance of POC input and light availability on the resource use in a benthic food web of a clear-water lake. Corn starch (a C(4) plant) was used as a POC source due to its insoluble nature and its distinct carbon stable isotope value (δ(13)C). The starch carbon was closely dispersed over the bottom of the enclosures to study the fate of a POC source exclusively available to sediment biota. The addition of starch carbon resulted in a clear shift in the isotopic signature of surface-dwelling herbivorous and predatory invertebrates. Although the starch carbon was added solely to the sediment surface, the carbon originating from the starch reached zooplankton. We suggest that allochthonous POC can subsidize benthic food webs directly and can be further transferred to pelagic systems, thereby highlighting the importance of benthic pathways for pelagic habitats.     

Differential support of lake food webs by three types of terrestrial organic carbon

Organic carbon inputs from outside of ecosystem boundaries potentially subsidize recipient food webs. Four whole-lake additions of dissolved inorganic ¹³C were made to reveal the pathways of subsidies to lakes from terrestrial dissolved organic carbon (t-DOC), terrestrial particulate organic carbon (t-POC) and terrestrial prey items. Terrestrial DOC, the largest input, was a major subsidy of pelagic bacterial respiration, but little of this bacterial C was passed up the food web. Zooplankton received <2% of their C from the t-DOC to bacteria pathway. Terrestrial POC significantly subsidized the production of both zooplankton and benthic invertebrates, and was passed up the food web to Chaoborus and fishes. This route supplied 33–73% of carbon flow to zooplankton and 20–50% to fishes in non-fertilized lakes. Terrestrial prey, by far the smallest input, provided some fishes with >20% of their carbon. The results show that impacts of cross-ecosystem subsidies depend on characteristics of the imported material, the route of entry into the food web, the types of consumers present, and the productivity of the recipient system.     

Shoreline urbanization interrupts allochthonous subsidies to a benthic consumer over a gradient of lake size

The role of resource subsidies across ecosystem boundaries has emerged as an important concept in contemporary ecology. For lake ecosystems, this has led to interest in quantifying the contribution of terrestrial allochthonous carbon to aquatic secondary production. An inverse relationship between habitat area and the role of allochthonous subsidies has been documented on marine islands and assumed for lakes, yet there have been no tests of this pattern among benthic (lake bottom) consumers. Here, we used carbon stable isotopes to trace terrestrial allochthonous and benthic autochthonous carbon use by the crayfish Pacifastacus leniusculus over a gradient of lake area, productivity and urbanization. Consistent with findings from terrestrial islands, habitat size dictated the importance of allochthonous subsidies, as P. leniusculus transitioned from using predominantly terrestrial carbon in small lakes to an increased reliance on autochthonous production in larger lakes. However, shoreline urbanization interacted with this pattern, particularly for small lakes where greater urbanization resulted in reduced use of allochthonous resources. As such, we provide, to our knowledge, the first confirmation of the predicted relationship between habitat size and importance of allochthonous subsidies to lake benthic consumers, but found that urbanization can interfere with this pattern.     

Carbon sources for lake food webs in the Canadian High Arctic and other regions of Arctic North America

We investigated the role of autochthonous and terrestrial carbon in supporting aquatic food webs in the Canadian High Arctic by determining the diet of the dominant primary consumer, aquatic chironomids. These organisms were studied in fresh waters on 3 islands of the Arctic Archipelago (~74–76°N) including barren polar desert watersheds and a polar oasis with lush meadows. Stomach content analysis of 578 larvae indicated that chironomids primarily ingested diatoms and sediment detritus with little variation among most genera. Carbon and nitrogen stable isotope mixing models applied to 2 lakes indicated that benthic algae contributed 68–95% to chironomid diet at a polar desert site and 70–78% at a polar oasis site. Detritus, originating from either phytoplankton or terrestrial sources, also contributed minor amounts to chironomid diet (0–32%). Radiocarbon measurements for the 2 lakes showed that old terrestrial carbon did not support chironomid production. Carbon stable isotope ratios of chironomids in other High Arctic lakes provided further dietary evidence that was consistent with mixing model results. These findings indicate that, in the Canadian High Arctic, chironomids (and fish that consume them) are supported primarily by benthic algae in both polar desert and oasis lakes. In contrast, our review of carbon flow studies for lakes in other Arctic regions of North America shows that terrestrial carbon and phytoplankton can be important energy sources for consumers. This study provides a baseline to detect future climate-related impacts on carbon pathways in High Arctic lakes.     

Carbon pathways to zooplankton: insights from the combined use of stable isotope and fatty acid biomarkers

1. Numerous studies have quantified the relative contribution of terrestrial‐ and phytoplankton‐derived carbon sources to zooplankton secondary production in lakes. However, few investigated the pathways along which allochthonous and autochthonous carbon (C) was actually conveyed to consumers. 2. We suggest that the combined use of fatty acid and stable isotope biomarkers could solve this issue. We conducted a field study on two oligotrophic lakes, in which primary production increased significantly between 2002 and 2004. We used modelling to estimate the contribution of terrestrial‐ and phytoplankton‐derived C to particulate organic C (POC) and zooplankton production from their δ¹³C values in 2002 and 2004. 3. According to the isotope model, phytoplankton‐derived C accounted for a major part of the POC pool in both lakes and supported more Daphnia sp. production in 2004 than in 2002. Fatty acid data revealed that increased contribution of algal‐C to Daphnia production, although common between both lakes, was achieved through C pathways that were different. In one lake, Daphnia grazed more intensively on phytoplankton, whereas in the other there was greater grazing on bacteria. In the latter case, the increased primary production resulted in greater release of algal‐derived dissolved organic C (DOC), which may have supported extra bacterial and eventually Daphnia, production. 4. This is the first study illustrating that the combination of fatty acid and stable isotope biomarkers could further our understanding of the factors controlling the relative magnitude of food webs pathways conveying organic matter to zooplankton.     

Mass flux calculations show strong allochthonous support of freshwater zooplankton production is unlikely

Many studies have concluded terrestrial carbon inputs contribute 20-70% of the carbon supporting zooplankton and fish production in lakes. Conversely, it is also known that terrestrial carbon inputs are of very low nutritional quality and phytoplankton are strongly preferentially utilized by zooplankton. Because of its low quality, substantial terrestrial support of zooplankton production in lakes is only conceivable when terrigenous organic matter inputs are much larger than algal production. We conducted a quantitative analysis of terrestrial carbon mass influx and algal primary production estimates for oligo/mesotrophic lakes (i.e., TP ≤ 20 μg L(-1)). In keeping with the principle of mass conservation, only the flux of terrestrial carbon retained within lakes can be utilized by zooplankton. Our field data compilation showed the median (inter-quartile range) terrestrial particulate organic carbon (t-POC), available dissolved organic carbon (t-DOC) inputs, and in-lake bacterial and algal production were 11 (8-17), 34 (11-78), 74 (37-165), and 253 (115-546) mg C m(-2) d(-1), respectively. Despite the widespread view that terrestrial inputs dominate the carbon flux of many lakes, our analysis indicates algal production is a factor 4-7 greater than the available flux of allochthonous basal resources in low productivity lakes. Lakes with high loading of t-DOC also have high hydraulic flushing rates. Because t-DOC is processed, i.e., mineralized or lost to the sediments, in lakes at ≈ 0.1% d(-1), in systems with the highest t-DOC inputs (i.e., 1000 mg m(-2) d(-1)) a median of 98% of the t-DOC flux is advected and therefore is not available to support zooplankton production. Further, advection is the primary fate of t-DOC in lakes with hydraulic retention times <3 years. When taking into account the availability and quality of terrestrial and autochthonous fluxes, this analysis indicates ≈ 95-99% of aquatic herbivore production is supported by in-lake primary production.     

Response of plankton to nutrients,planktivory and terrestrial organic matter: a model analysis of whole‐lake experiments

Terrestrial organic matter can be assimilated by aquatic consumers but implications for biomass and production are unresolved. An ecosystem model was fit to estimate effects of phosphorus (P) load, planktivory, and supply rate of terrestrial particulate organic carbon (TPOC) on phytoplankton and zooplankton in five whole‐lake experiments. Phytoplankton biomass increased with P load and planktivory and decreased with TPOC supply rate. Zooplankton biomass increased with P load and responded weakly to planktivory and TPOC supply rate. Zooplankton allochthony (proportion of carbon from terrestrial sources) decreased with P load and planktivory and increased with TPOC supply rate. Lakes with low allochthony (< 0.3) had wide ranges of phytoplankton and zooplankton biomass and production, depending on P load and planktivory. Lakes with high allochthony (> 0.3) had low biomass and production of both phytoplankton and zooplankton. In summary, terrestrial OC inhibits primary production and is a relatively low‐quality food source for zooplankton.     

Reducing dissolved phosphorus loading to the Salton Sea with aluminum sulfate

Terrestrial support of aquatic food webs is becoming well established in the science of ecology. However, while terrestrial subsidies of energy have been shown to exert strong effects on aquatic food webs, it is less clear how variations in these subsidies, via natural or anthropogenic factors, will affect recipient ecosystems. To assess the influence of variations in terrestrial subsidies on an aquatic food web, we manipulated leaf-litter inputs in artificial ponds. Decreasing litter inputs did not affect any of the response variables in artificial ponds. This may be because the minimal amount of terrestrial carbon present combined with autochthonous production was enough to sustain the food web and/or the food web was altered in ways not detected by the experimental design. However, increasing leaf-litter inputs increased the percent survival and developmental rate of larval wood frogs (Rana sylvatica). Conversely, increasing litter input appeared to have no influence on zooplankton or salamander larvae. Increasing litter inputs also increased the dissolved organic carbon content and decreased the percent saturation of dissolved oxygen in artificial ponds. As system respiration in aquatic systems is frequently dominated by microbial respiration, we hypothesize that the effects of increasing litter input on wood frogs were the result of an increase food resources (i.e., microbes) for tadpoles. The lack of a response by salamander larvae and zooplankton may be due to the densities of zooplankton in tanks providing enough food for salamanders in all treatments, variation among specific zooplankton species in their ability to exploit these resources and transfer energy to salamanders, or omnivory among zooplankton offsetting the affects of leaf-litter inputs. Additional work is needed to determine the influence of litter inputs on zooplankton and salamanders in this community. These data demonstrate that variations in leaf-litter inputs can influence food web structure; however, the importance of these variations will likely be dependent upon the trophic position of various consumers. Handling editor: J. Cole     

Experimental dietary manipulations for determining the relative importance of allochthonous and autochthonous food resources in tropical streams

1. Autochthonous sources of organic matter appear to make a minor contribution to food webs in temperate forest streams, but their roles in supporting consumer biomass in tropical lotic environments have received little attention. We investigated the importance of autochthonous and allochthonous food sources to Brotia hainanensis (Pachychilidae), a detritivorous and algivorous snail common in Hong Kong hillstreams, using experimental dietary manipulations and assimilation-based analyses, including stoichiometry, carbon (C) and nitrogen (N) stable isotopes and fatty acid (FA) profiles.
2. Juvenile B. hainanensis collected in Pak Ngau Shek Stream were cultured under controlled laboratory conditions and fed for 2 months with either conditioned Liquidambar formosana (Hamamelidaceae) leaf litter or periphyton. Samples of B. hainanensis were also collected from the stream at the end of the experiment for comparison with snails reared in the laboratory.
3. Periphyton and leaf litter exhibited marked differences in C/N ratios, δ ¹³C and δ ¹⁵N values and FA profiles. Stable isotope analysis and FA profiling of laboratory-reared and field-collected B. hainanensis both confirmed that snails relied primarily on autochthonous foods, especially periphytic diatoms and cyanobacteria. Stoichiometry results indicated that periphyton was a more nutritious food (with lower C/N ratio) than leaf litter.
4. This is the first study demonstrating that the combined use of stable isotopes and FA profiles is an effective diagnostic tool to trace the basal food sources of consumers in natural stream habitats. Our findings further support the hypothesis that primary production in tropical streams is generally more important to aquatic consumers than inputs of terrestrial detritus.     

Impact of eutrophication on the carbon stable‐isotopic baseline of benthic invertebrates in two deep soft‐water lakes

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Sources of organic carbon supporting the food web of an arid zone floodplain river

SUMMARY 1. Many Australian inland rivers are characterised by vast floodplains with a network of anastomosing channels that interconnect only during unpredictable flooding. For much of the time, however, rivers are reduced to a string of disconnected and highly turbid waterholes. Given these features, we predicted that aquatic primary production would be light-limited and the riverine food web would be dependent on terrestrial carbon from floodplain exchanges and direct riparian inputs.
2. To test these predictions, we measured rates of benthic primary production and respiration and sampled primary sources of organic carbon and consumers for stable isotope analysis in several river waterholes at four locations in the Cooper Creek system in central Australia.
3. A conspicuous band of filamentous algae was observed along the shallow littoral zone of the larger waterholes. Despite the high turbidity, benthic gross primary production in this narrow zone was very high (1.7–3.6 g C m⁻² day⁻¹); about two orders of magnitude greater than that measured in the main channel.
4. Stable carbon isotope analysis confirmed that the band of algae was the major source of energy for aquatic consumers, ultimately supporting large populations of crustaceans and fish. Variation in the stable carbon and nitrogen isotope signatures of consumers suggested that zooplankton was the other likely major source.
5. Existing ecosystem models of large rivers often emphasise the importance of longitudinal or lateral inputs of terrestrial organic matter as a source of organic carbon for aquatic consumers. Our data suggest that, despite the presence of large amounts of terrestrial carbon, there was no evidence of it being a significant contributor to the aquatic food web in this floodplain river system.     

Terrestrial support of pelagic consumers: patterns and variability revealed by a multilake study

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Habitat structure determines resource use by zooplankton in temperate lakes

While the importance of terrestrial linkages to aquatic ecosystems is well appreciated, the degree of terrestrial support of aquatic consumers remains debated. Estimates of terrestrial contributions to lake zooplankton have omitted a key food source, phytoplankton produced below the mixed layer. We used carbon and nitrogen stable isotope data from 25 Pacific Northwest lakes to assess the relative importance of particulate organic matter (POM) from the mixed layer, below the mixed layer and terrestrial detritus to zooplankton. Zooplankton and deep POM were depleted in 13C relative to mixed layer POM in lakes that can support deep primary production. A Bayesian stable isotope mixing model estimated that terrestrial detritus contributed <5% to zooplankton production, and confirms the role of lake optical and thermal properties; deep POM accounted for up to 80% of zooplankton production in the clearest lakes. These results suggest terrestrial support of lake zooplankton production is trivial.     

Intraspecific Autochthonous and Allochthonous Resource Use by Zooplankton in a Humic Lake during the Transitions between Winter,Summer and Fall

Seasonal patterns in assimilation of externally produced, allochthonous, organic matter into aquatic food webs are poorly understood, especially in brown-water lakes. We studied the allochthony (share biomass of terrestrial origin) in cladoceran, calanoid and cyclopoid micro-crustacean zooplankton from late winter to fall during two years in a small humic lake (Sweden). The use of allochthonous resources was important for sustaining a small population of calanoids in the water column during late winter. However, in summer the calanoids shifted to 100% herbivory, increasing their biomass several-fold by making efficient use of the pelagic primary production. In contrast, the cyclopoids and cladocerans remained at high levels of allochthony throughout the seasons, both groups showing the mean allochthony of 0.56 (range in mean 0.17-0.79 and 0.34-0.75, for the respective group, depending on model parameters). Our study shows that terrestrial organic matter can be an important resource for cyclopoids and cladocerans on an annual basis, forming a significant link between terrestrial organic matter and the higher trophic levels of the food web, but it can also be important for sustaining otherwise herbivorous calanoids during periods of low primary production in late winter.     

Allochthonous Organic Matter Supports Benthic but Not Pelagic Food Webs in Shallow Coastal Ecosystems

Rivers transport large amounts of allochthonous organic matter (OM) to the ocean every year, but there are still fundamental gaps in how allochthonous OM is processed in the marine environment. Here, we estimated the relative contribution of allochthonous OM (allochthony) to the biomass of benthic and pelagic consumers in a shallow coastal ecosystem in the northern Baltic Sea. We used deuterium as a tracer of allochthony and assessed both temporal variation (monthly from May to August) and spatial variation (within and outside river plume). We found variability in allochthony in space and time and across species, with overall higher values for zoobenthos (26.2 ± 20.9%) than for zooplankton (0.8 ± 0.3%). Zooplankton allochthony was highest in May and very low during the other months, likely as a result of high inputs of allochthonous OM during the spring flood that fueled the pelagic food chain for a short period. In contrast, zoobenthos allochthony was only lower in June and remained high during the other months. Allochthony of zoobenthos was generally higher close to the river mouth than outside of the river plume, whereas it did not vary spatially for zooplankton. Last, zoobenthos allochthony was higher in deeper than in shallower areas, indicating that allochthonous OM might be more important when autochthonous resources are limited. Our results suggest that climate change predictions of increasing inputs of allochthonous OM to coastal ecosystems may affect basal energy sources supporting coastal food webs.