Weak Response of Animal Allochthony and Production to Enhanced Supply of Terrestrial Leaf Litter in Nutrient-Rich Lakes

Ecosystems are generally linked via fluxes of nutrients and energy across their boundaries. For example, freshwater ecosystems in temperate regions may receive significant inputs of terrestrially derived carbon via autumnal leaf litter. This terrestrial particulate organic carbon (POC) is hypothesized to subsidize animal production in lakes, but direct evidence is still lacking. We divided two small eutrophic lakes each into two sections and added isotopically distinct maize litter to the treatment sections to simulate increased terrestrial POC inputs via leaf litter in autumn. We quantified the reliance of aquatic consumers on terrestrial resources (allochthony) in the year subsequent to POC additions by applying mixing models of stable isotopes. We also estimated lake-wide carbon (C) balances to calculate the C flow to the production of the major aquatic consumer groups: benthic macroinvertebrates, crustacean zooplankton, and fish. The sum of secondary production of crustaceans and benthic macroinvertebrates supported by terrestrial POC was higher in the treatment sections of both lakes. In contrast, total secondary and tertiary production (supported by both autochthonous and allochthonous C) was higher in the reference than in the treatment sections of both lakes. Average aquatic consumer allochthony per lake section was 27–40%, although terrestrial POC contributed less than about 10% to total organic C supply to the lakes. The production of aquatic consumers incorporated less than 5% of the total organic C supply in both lakes, indicating a low ecological efficiency. We suggest that the consumption of terrestrial POC by aquatic consumers facilitates a strong coupling with the terrestrial environment. However, the high autochthonous production and the large pool of autochthonous detritus in these nutrient-rich lakes make terrestrial POC quantitatively unimportant for the C flows within food webs.     

Zooplankton allochthony is spatially heterogeneous in a boreal lake

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Regional-Level Inputs of Emergent Aquatic Insects from Water to Land

Emergent aquatic insects can provide inputs to terrestrial ecosystems near lentic and lotic waterbodies, producing ecosystem linkages at the aquatic–terrestrial interface. Although aquatic insect emergence has been examined for individual sites, the magnitude and spatial distribution of this phenomenon has not been examined at regional spatial scales. Here, we characterize this cross-habitat linkage for the state of Wisconsin, USA (169,639 km²). We combined GIS hydrological data with empirical data and predictive models of aquatic insect production to estimate annual aquatic emergence for the state of Wisconsin. Total emergence (lentic + lotic) was estimated to be about 6,800 metric tons of C y^?1. Lentic systems comprised 79% of total estimated insect emergence, primarily due to the large amount of lake surface area relative to streams. This is due to both basic ecosystem geometry and the overall abundance of lakes in Wisconsin. Spatial variation was high: insect emergence in southwestern Wisconsin was dominated by streams, whereas for most of the rest of the state insect emergence was dominated by lakes. Lentic inputs to land were highly concentrated (relative to lotic inputs) because lakes have a high ratio of surface area to buffer area. Although less concentrated, the spatial extent of lotic influence was greater: statewide, four times more land area fell within the 100 m buffer zones of streams compared to lakes. Large waterbodies (almost all of which were lakes) were hotspots of insect emergence and input to land. Aquatic insect inputs exceed estimated terrestrial secondary production in 13% of buffer area, and by a factor of 100 or more adjacent to large lakes (>50,000 ha). The model sensitivity analysis showed that the simplifying assumptions and sources of potential error in the input variables had a minor impact on the overall results.     

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.     

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

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Ecosystem Linkages Between Lakes and the Surrounding Terrestrial Landscape in Northeast Iceland

Despite a recent emphasis on understanding cross-habitat interactions, few studies have examined the ecological linkages between lakes and surrounding terrestrial habitats. The current paradigm of land–lake interactions is typically unidirectional: the view is that nutrients and matter are transported downslope from the surrounding watershed to their ultimate lacustrine destination. Emergent aquatic insects, which spend their larval stages in lake sediments and emerge as adults to mate over land, can act as vectors of material, energy and nutrients from aquatic to terrestrial habitats. In this study, we document a gradient of midge (Diptera: Chironomidae) infall rates into terrestrial habitats (measured as g dw midges m^?2 d^?1) surrounding eight lakes in Northern Iceland (≈66°N latitude). Lakes ranged from having virtually no midge infall (for example, Helluvaðstjörn, 0.03 g m^?2 d^?1) to extreme levels (for example, Mývatn, 19 g m^?2 d^?1) with abundances of midges decreasing logarithmically with distance from shore. Annual midge input rates are estimated as high as 1200–2500 kg midges ha^?1 y^?1. As midges are approximately 9.2% total N, this can result in a significant fertilization effect of terrestrial habitats with consequences for plant quality and community structure. In addition, we used naturally-occurring δ¹³C and δ¹⁵N isotopes to examine food web structure and diet sources of terrestrial arthropod consumers surrounding lakes with differing amounts of midge input. Terrestrial arthropods showed increased utilization of aquatic-derived (that is, midge) C relative to terrestrial sources as midge infall increased. This pattern was particularly pronounced for predators, such as spiders and opiliones, and some detritivores (Collembola). These findings suggest that, despite being largely ignored, aquatic-to-terrestrial linkages can be large and midges can fuel terrestrial communities by directly serving as resources for predators and decomposers.     

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.     

Terrestrial Contributions to the Aquatic Food Web in the Middle Yangtze River

Understanding the carbon sources supporting aquatic consumers in large rivers is essential for the protection of ecological integrity and for wildlife management. The relative importance of terrestrial and algal carbon to the aquatic food webs is still under intensive debate. The Yangtze River is the largest river in China and the third longest river in the world. The completion of the Three Gorges Dam (TGD) in 2003 has significantly altered the hydrological regime of the middle Yangtze River, but its immediate impact on carbon sources supporting the river food web is unknown. In this study, potential production sources from riparian and the main river channel, and selected aquatic consumers (invertebrates and fish) at an upstream constricted-channel site (Luoqi), a midstream estuarine site (Huanghua) and a near dam limnetic site (Maoping) of the TGD were collected for stable isotope (δ¹³C and δ¹⁵N) and IsoSource analyses. Model estimates indicated that terrestrial plants were the dominant carbon sources supporting the consumer taxa at the three study sites. Algal production appeared to play a supplemental role in supporting consumer production. The contribution from C₄ plants was more important than that of C₃ plants at the upstream site while C₃ plants were the more important carbon source to the consumers at the two impacted sites (Huanghua and Maoping), particularly at the midstream site. There was no trend of increase in the contribution of autochthonous production from the upstream to the downstream sites as the flow rate decreased dramatically along the main river channel due to the construction of TGD. Our findings, along with recent studies in rivers and lakes, are contradictory to studies that demonstrate the importance of algal carbon in the aquatic food web. Differences in system geomorphology, hydrology, habitat heterogeneity, and land use may account for these contradictory findings reported in various studies.     

Combined global change effects on ecosystem processes in nine U.S. topographically complex areas

Concurrent changes in climate, atmospheric nitrogen (N) deposition, and increasing levels of atmospheric carbon dioxide (CO₂) affect ecosystems in complex ways. The DayCent-Chem model was used to investigate the combined effects of these human-caused drivers of change over the period 1980–2075 at seven forested montane and two alpine watersheds in the United States. Net ecosystem production (NEP) increased linearly with increasing N deposition for six out of seven forested watersheds; warming directly increased NEP at only two of these sites. Warming reduced soil organic carbon storage at all sites by increasing heterotrophic respiration. At most sites, warming together with high N deposition increased nitrous oxide (N₂O) emissions enough to negate the greenhouse benefit of soil carbon sequestration alone, though there was a net greenhouse gas sink across nearly all sites mainly due to the effect of CO₂ fertilization and associated sequestration by plants. Over the simulation period, an increase in atmospheric CO₂ from 350 to 600 ppm was the main driver of change in net ecosystem greenhouse gas sequestration at all forested sites and one of two alpine sites, but an additional increase in CO₂ from 600 to 760 ppm produced smaller effects. Warming either increased or decreased net greenhouse gas sequestration, depending on the site. The N contribution to net ecosystem greenhouse gas sequestration averaged across forest sites was only 5–7 % and was negligible for the alpine. Stream nitrate (NO₃ ^?) fluxes increased sharply with N-loading, primarily at three watersheds where initial N deposition values were high relative to terrestrial N uptake capacity. The simulated results displayed fewer synergistic responses to warming, N-loading, and CO₂ fertilization than expected. Overall, simulations with DayCent-Chem suggest individual site characteristics and historical patterns of N deposition are important determinants of forest or alpine ecosystem responses to global change.     

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.     

From clear lakes to murky waters – tracing the functional response of high‐latitude lake communities to concurrent ‘greening’ and ‘browning’

Climate change and the intensification of land use practices are causing widespread eutrophication of subarctic lakes. The implications of this rapid change for lake ecosystem function remain poorly understood. To assess how freshwater communities respond to such profound changes in their habitat and resource availability, we conducted a space‐for‐time analysis of food‐web structure in 30 lakes situated across a temperature‐productivity gradient equivalent to the predicted future climate of subarctic Europe (temperature +3°C, precipitation +30% and nutrient +45 μg L^?1 total phosphorus). Along this gradient, we observed an increase in the assimilation of pelagic‐derived carbon from 25 to 75% throughout primary, secondary and tertiary consumers. This shift was overwhelmingly driven by the consumption of pelagic detritus by benthic primary consumers and was not accompanied by increased pelagic foraging by higher trophic level consumers. Our data also revealed a convergence of the carbon isotope ratios of pelagic and benthic food web endmembers in the warmest, most productive lakes indicating that the incorporation of terrestrial derived carbon into aquatic food webs increases as land use intensifies. These results, reflecting changes along a gradient characteristic of the predicted future environment throughout the subarctic, indicate that climate and land use driven eutrophication and browning are radically altering the function and fuelling of aquatic food webs in this biome.     

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.     

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

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Terrestrial contributions to Afrotropical aquatic food webs: The Congo River case

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Variation in Detrital Resource Stoichiometry Signals Differential Carbon to Nutrient Limitation for Stream Consumers Across Biomes

Stoichiometric ratios of resources and consumers have been used to predict nutrient limitation across diverse terrestrial and aquatic ecosystems. In forested headwater streams, coarse and fine benthic organic matter (CBOM, FBOM) are primary basal resources for the food web, and the distribution and quality of these organic matter resources may therefore influence patterns of secondary production and nutrient cycling within stream networks or among biomes. We measured carbon (C), nitrogen (N), and phosphorus (P) content of CBOM and FBOM and calculated their stoichiometric ratios (C/N, C/P, N/P) from first- to fourth-order streams from tropical montane, temperate deciduous, and boreal forests, and tallgrass prairie, to compare the magnitude and variability of these resource types among biomes. We then used the ratios to predict nutritional limitations for consumers of each resource type. Across biomes, CBOM had consistently higher %C and %N, and higher and more variable C/N and C/P than FBOM, suggesting that microbial processing results in more tightly constrained elemental composition in FBOM than in CBOM. Biome-specific differences were observed in %P and N/P between the two resource pools; CBOM was lower in %P but higher in N/P than FBOM in the tropical montane and temperate deciduous forest biomes, while CBOM was higher in %P but similar in N/P than FBOM in the grassland and boreal forest biomes. Stable ¹³C isotopes suggest that FBOM likely derives from CBOM in tropical and temperate deciduous forest, but that additional non-detrital components may contribute to FBOM in boreal forests and grasslands. Comparisons of stoichiometric ratios of CBOM and FBOM to estimated needs of aquatic detritivores suggest that shredders feeding on CBOM are more likely to experience nutrient (N and/or P) than C limitation, whereas collector–gatherers consuming FBOM are more likely to experience C than N and/or P limitation. Our results suggest that differences in basal resource elemental content and stoichiometric ratios have the potential to affect consumer production and ecosystem rates of C, N, and P cycling in relatively consistent ways across diverse biomes.     

Use of carbon-13 and carbon-14 natural abundances for stream food web studies

We review the use of stable carbon isotope ratios (δ¹³C) and radiocarbon natural abundances (Δ¹⁴C) for stream food web studies. The δ¹³C value of primary producers (e.g., periphytic algae, hereafter periphyton) in streams is controlled by isotopic fractionation during photosynthesis and variable δ¹³C of dissolved CO₂. When periphyton δ¹³C differs from that of terrestrial primary producers, the relative contribution of autochthony and allochthony to stream food webs can be calculated. Moreover, the variation in periphyton δ¹³C can reveal how much stream consumers rely on local resources because each stream habitat (e.g., riffle vs. pool, open vs. shaded) usually has a distinctive δ¹³C. However, periphyton δ¹³C often overlaps with that of terrestrial organic matter. On the other hand, periphyton Δ¹⁴C is less variable than δ¹³C among habitats, and reflects the Δ¹⁴C of dissolved CO₂, which could be a mixture of “aged” (Δ¹⁴C < 0 ‰) and “modern” (Δ¹⁴C > 0 ‰) carbon. This is because the Δ¹⁴C is corrected by its δ¹³C value for the isotopic fractionation during photosynthesis. Recent studies and our data indicate that many stream food webs are supported by “aged” carbon derived from the watershed via autochthonous production. The combined use of δ¹³C and Δ¹⁴C allows robust estimation of the carbon transfer pathway in a stream food web at multiple spatial scales ranging from the stream habitat level (e.g., riffle and pool) to watershed level (autochthony and allochthony). Furthermore, the Δ¹⁴C of stream food webs will expand our understanding about the time frame of carbon cycles in the watersheds.     

Temporal shift in contribution of terrestrial organic matter to consumer production in a grassland river

1. We used stable isotopes to study the temporal (early summer versus autumn) pattern of use of terrestrial and aquatic sources of organic carbon by consumers in two bedrock‐confined reaches of a grassland river in New Zealand.
2. The major sources of organic carbon available to primary consumers were expected to be terrestrial leaf‐litter and biofilm from the stream channel. These putative carbon sources showed no significant change in mean δ¹³C between summer and autumn. Leaf litter (mean δ¹³C13C compared to biofilm (mean δ¹³C>?19.92).
3. In contrast to leaf litter and biofilm, the δ¹³C of consumers changed over time, being enriched in ¹³C in the autumn compared with early summer. Both the magnitude (>5‰ in some cases) and rapidity of this shift (< 3 months) was surprising.
4. A two‐source mixing model indicated that, during early summer, terrestrial carbon comprised> 50% of tissue carbon for 15 of the 17 taxa of aquatic consumers analysed. During autumn, terrestrial carbon comprised> 50% of the tissue carbon of only five of 25 taxa. Because the mean δ¹³C of putative food sources was consistent over time, the shift in δ¹³C values for consumers is attributed to a change in relative amounts of terrestrial and aquatic carbon available for consumption.
5. Because seston consists of a mixture of many particles of diverse origin, it may provide an integrated measure of catchment‐wide sources of organic matter entering a stream channel. Like the tissues of most consumers, mean δ¹³C values for seston showed a significant shift toward ¹³C enrichment. This indicated that the relative availability of terrestrial carbon decreased from summer to autumn.
6. The actual quantity of carbon contributed to the stream food‐web by this potential terrestrial–aquatic link is unknown. Although terrestrial carbon may comprise a high proportion of the tissue carbon of consumers prior to summer, the majority of secondary production (and carbon sequestration) probably occurs during early summer as a consequence of rising temperature and high quality food in the form of biofilm.     

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

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Dust mediated transfer of phosphorus to alpine lake ecosystems of the Wind River Range,Wyoming, USA

Alpine lakes receive a large fraction of their nutrients from atmospheric sources and are consequently sensitive to variations in both the amount and chemistry of atmospheric deposition. In this study we explored the spatial changes in lake water chemistry and biology along a gradient of dust deposition in the Wind River Range, Wyoming. Regional differences were explored using the variation in bulk deposition, lake water, sediment, and bedrock geochemistry and catchment characteristics. Dust deposition rates in the Southwestern region averaged 3.34 g m^?2 year^?1, approximately three times higher than deposition rates in the Northwestern region (average 1.06 g m^?2 year^?1). Dust-P deposition rates ranged from 87 µg P m² day^?1 in the Northwestern region to 276 µg P m² day^?1 in the Southwestern region. Subalpine and alpine lakes in the Southwestern region had greater total phosphorus (TP) concentrations (5–13 µg L^?1) and greater sediment phosphorus (SP) concentrations (2–5 mg g^?1) than similar lakes elsewhere in the region (1–8 µg L^?1 TP, 0.5–2 mg g^?1 SP). Lake phosphorus concentrations were related to dissolved organic carbon (DOC) across vegetation gradients, but related to the percent of bare rock, catchment area to lake area, and catchment steepness across dust deposition gradients. Modern phytoplankton and zooplankton biomasses were two orders of magnitude greater in the Southwest than in the Northwest, and alpine lakes in the Southwest had a unique diatom species assemblage with relatively higher concentrations of Asterionella formosa, Pseudostaurosira pseudoconstruens, and Pseudostaurosira brevistriata. These results suggests that catchment controls on P export to lakes (i.e. DOC) are overridden in dominantly bare rock basins where poor soils cannot effectively retain dust deposited P.