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Increased incorporation of terrestrial organic matter (t-OM) into consumer biomass (allochthony) is believed to reduce growth capacity. In this study, we examined the relationship between crustacean zooplankton allochthony and production in a boreal lake that displays strong seasonal variability in t-OM inputs. Contrary to our hypotheses, we found no effect of allochthony on production at the community and the species levels. The high-frequency seasonal sampling (time-for-space) allowed for estimating the efficiency of zooplankton in converting this external carbon source to growth. From the daily t-OM inputs in the lake (57-3,027 kg C/d), the zooplankton community transferred 0.2% into biomass (0.01-2.36 kg C/d); this level was of the same magnitude as the carbon transfer efficiency for algal-derived carbon (0.4%). In the context of the boundless carbon cycle, which integrates inland waters as a biologically active component of the terrestrial landscape, the use of the time-for-space approach for the quantifying of t-OM trophic transfer efficiency by zooplankton is a critical step toward a better understanding of the effects of increasing external carbon fluxes on pelagic food webs.

Resources and temperature play major roles in determining biological production in lake ecosystems. Lakes have been warming and ‘browning’ over recent decades due to climate change and increased loading of terrestrial organic matter. Conflicting hypotheses and evidence have been presented about whether these changes will increase or decrease fish growth within lakes. Most studies have been conducted in low-elevation lakes where terrestrially derived carbon tends to dominate over carbon produced within lakes. Understanding how fish in high-elevation mountain lakes will respond to warming and browning is particularly needed as warming effects are magnified for mountain lakes and treeline is advancing to higher elevations. We sampled 21 trout populations in the Sierra Nevada Mountains of California to examine how body condition and individual growth rates, measured by otolith analysis, varied across independent elevational gradients in temperature and dissolved organic carbon (DOC). We found that fish grew faster at warmer temperatures and higher nitrogen (TN), but slower in high DOC lakes. Additionally, fish showed better body condition in lakes with higher TN, higher elevation and when they exhibited a more terrestrial δ13C isotopic signature. The future warming and browning of lakes will likely have antagonistic impacts on fish growth, reducing the predicted independent impact of warming and browning alone.

Terrestrial dissolved organic carbon (DOC) provides an external carbon source to lake ecosystems. However, there is ongoing debate about whether external DOC that enters a lake can pass up the food web to support top consumers. We show, from experimental manipulation of a whole lake, that externally loaded DOC can contribute appreciably to fish biomass. Monthly additions of cane sugar with a distinct carbon stable isotope value during 2 years rapidly enriched the 13C content of zooplankton and macroinvertebrates, with a more gradual 13C enrichment of fish. After sugar addition stopped, the 13C content of consumers reverted towards original values. A simple isotope mixing model indicated that by the end of the sugar addition almost 20% of fish carbon in the lake was derived from the added sugar. Our results provide the first direct experimental demonstration at relevant ecological spatial and temporal scales that externally loaded DOC to lakes can indeed transfer to top consumers., final draft, peerReviewed

There is widespread evidence that aquatic consumers use terrestrial resources depending on the features of surrounding catchments., Widespread evidence that organic matter exported from terrestrial into aquatic ecosystems supports recipient food webs remains controversial. A pressing question is not only whether high terrestrial support is possible but also what the general conditions are under which it arises. We assemble the largest data set, to date, of the isotopic composition (δ2H, δ13C, and δ15N) of lake zooplankton and the resources at the base of their associated food webs. In total, our data set spans 559 observations across 147 lakes from the boreal to subtropics. By predicting terrestrial resource support from within-lake and catchment-level characteristics, we found that half of all consumer observations that is, the median were composed of at least 42% terrestrially derived material. In general, terrestrial support of zooplankton was greatest in lakes with large physical and hydrological connections to catchments that were rich in aboveground and belowground organic matter. However, some consumers responded less strongly to terrestrial resources where within-lake production was elevated. Our study shows that multiple mechanisms drive widespread cross-ecosystem support of aquatic consumers across Northern Hemisphere lakes and suggests that changes in terrestrial landscapes will influence ecosystem processes well beyond their boundaries.

Aquatic ecosystems are hotspots of decomposition and a globally significant source of carbon dioxide to the atmosphere. Carbon fixed in the terrestrial biosphere (allochthonous) supplements the carbon fixed in aquatic ecosystems (autochthonous), forming the organic matter (OM) available to aquatic consumers. Recent studies have re-emphasized the potential importance of allochthonous inputs to lake food webs and their dynamics. Specifically, the variation among lake ecosystems in the amount of allochthonous inputs and importance of those resources to aquatic consumers is not well understood. The aim of this dissertation is to quantify the patterns of allochthonous resource availability and use among and within lakes. Using hydrogen and carbon stable isotopes, I found that the organic matter in 39 north temperate lakes is dominated by allochthonous material. Additionally, within a lake, the composition of organic matter does not vary vertically in the water column. I demonstrated using oxygen and argon saturations that metalimnetic oxygen maxima are not solely the result of biological production or indicative of increased autochthonous resource availability for consumers compared to surface waters. This finding was confirmed with a whole-lake metalimnetic 13C addition in Peter Lake which was also used to track metalimnetic resource use by zooplankton. As hypothesized, metalimnetic resource use was low for Peter Lake consumers with allochthonous resources being the most important. This is consistent with the high but variable allochthonous resource use by zooplankton quantified using hydrogen stable isotopes in the same lakes that allochthonous resource availability was measured. Allochthonous resource use increases with allochthonous resource availability in these lakes and is highest in small lakes. Finally, I evaluate three key issues regarding the use of hydrogen stable isotopes in mixing models; 1) correction for environmental water in consumer tissues, 2) consideration of differential fractionation among biochemical constituents, and 3) consideration of differential H-exchange among samples and standards. Despite these complicating factors, analysis of consumer resource use is possible using whole organisms’ δ2H. My dissertation quantifies the variability of allochthonous resource availability and use by zooplankton in lakes and related it to ecosystem characteristics. Based on my findings and given the numerical dominance of small lakes globally, allochthonous resources are likely important for consumers in most lakes. Allochthonous resource use in aquatic food webs indicates interdependence and connectivity between ecosystems. Therefore, being able to predict the allochthonous resource availability and use in a food web is necessary for ecosystem management, restoration, and identifying lakes that could be susceptible to watershed, regional, or global change.

We investigated the terrestrial influence on two chained deep ultraoligotrophic lakes of North Patagonia (Argentina) through the seasonal analysis of two pelagic allochthonous indicators: i) water color, as a proxy of allochthonous dissolved organic matter in lakes; and ii) the color to chlorophyll a ratio (Color:Chla), as an indicator of the relationship between allochthonous and autochthonous carbon pools. We also evaluated the potential transfer pathways of the allochthonous dissolved organic matter into the pelagic food webs of these deep lakes, including the natural zooplankton δ 13C in the analysis. The dynamics of the allochthonous indicators were related to the precipitation regime, water level fluctuations, and hydrogeomorphic and catchment features of lakes Moreno East and Moreno West. The water color (absorbance at 440nm) was extremely low (

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 clearability 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 (? 13C). 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. � 2011 Springer-Verlag.

Lake ecosystems are strongly linked to their terrestrial surroundings by material and energy fluxes across ecosystem boundaries. However, the contribution of terrestrial particulate organic carbon (tPOC) from annual leaf fall to lake food webs has not yet been adequately traced and quantified. In this study, we conducted whole-lake experiments to trace artificially added tPOC through the food webs of two shallow lakes of similar eutrophic status, but featuring alternative stable regimes (macrophyte rich vs. phytoplankton dominated). Lakes were divided with a curtain, and maize (Zea mays) leaves were added, as an isotopically distinct tPOC source, into one half of each lake. To estimate the balance between autochthonous carbon fixation and allochthonous carbon input, primary production and tPOC and tDOC (terrestrial dissolved organic carbon) influx were calculated for the treatment sides. We measured the stable isotope ratios of carbon (d13C) of about 800 samples from all trophic consumer levels and compared them between lake sides, lakes, and three seasons. Leaf litter bag experiments showed that added maize leaves were processed at rates similar to those observed for leaves from shoreline plants, supporting the suitability of maize leaves as a tracer. The lake-wide carbon influx estimates confirmed that autochthonous carbon fixation by primary production was the dominant carbon source for consumers in the lakes. Nevertheless, carbon isotope values of benthic macroinvertebrates were significantly higher with maize additions compared to the reference side of each lake. Carbon isotope values of omnivorous and piscivorous fish were significantly affected by maize additions only in the macrophytedominated lake and d13C of zooplankton and planktivorous fish remained unaffected in both lakes. In summary, our results experimentally demonstrate that tPOC in form of autumnal litterfall is rapidly processed during the subsequent months in the food web of shallow lakes and is channeled to secondary and tertiary consumers predominantly via the benthic pathways. A more intense processing of tPOC seems to be connected to a higher structural complexity in littoral zones, and hence may differ between shallow lakes of alternative stable states. peerReviewed