Recognizing cross‐ecosystem responses to changing temperatures: soil warming impacts pelagic food webs

The energy and materials that move across ecosystem boundaries influence food web structure and key ecosystem functions. Despite the acknowledged importance of such ecological subsidies, surprisingly little information is available regarding the role of environmental temperature in influencing subsidy quality and the response of the recipient ecosystem. We evaluated the impacts of temperature‐mediated changes in leaves from deciduous trees, an important subsidy from terrestrial to freshwater ecosystems, on both the producer‐based and detritivore‐based components of a pelagic pond food web in a field mesocosm experiment. We hypothesized that variation in leaf chemistry driven by increased soil temperature would alter both the quality of leaf subsidies and the pond response. We collected red maple Acer rubrum leaves from heated and ambient temperature plots from the long‐term soil warming experiment at the Harvard Experimental Forest and added them to 167‐l field mesocosms containing established plankton communities, creating ‘no leaf’, ‘ambient leaf’ and ‘heated leaf’ treatments during autumn 2012. We then monitored physical, chemical, and biological responses to treatments until the mesocosms froze six weeks later. Experimental soil warming altered the chemical composition of deciduous leaves, the physical and chemical environment of the aquatic ecosystems to which leaves were added, and the pelagic pond food webs as measured by community composition. Compared to leaves from ambient‐temperature soils, leaves from warmed soils initially resulted in lower water column phosphorus and dissolved organic carbon, reducing bacterial densities. However, the diminished carbon and phosphorus resulting from soil warming also increased light availability that ultimately stimulated cladoceran zooplankton relative to ambient‐temperature leaves. Our results suggest that changes in temperature can alter ecological subsidies in unanticipated ways, and suggest that accurately predicting the potential consequences of climate change will require conducting research across ecosystem boundaries.     

Bottom‐up and top‐down effects of browning and warming on shallow lake food webs

Productivity and trophic structure of aquatic ecosystems result from a complex interplay of bottom‐up and top‐down forces that operate across benthic and pelagic food web compartments. Projected global changes urge the question how this interplay will be affected by browning (increasing input of terrestrial dissolved organic matter), nutrient enrichment and warming. We explored this with a process‐based model of a shallow lake food web consisting of benthic and pelagic components (abiotic resources, primary producers, grazers, carnivores), and compared model expectations with the results of a browning and warming experiment in nutrient‐poor ponds harboring a boreal lake community. Under low nutrient conditions, the model makes three major predictions. (a) Browning reduces light and increases nutrient supply; this decreases benthic and increases pelagic production, gradually shifting productivity from the benthic to the pelagic habitat. (b) Because of active habitat choice, fish exert top‐down control on grazers and benefit primary producers primarily in the more productive of the two habitats. (c) Warming relaxes top‐down control of grazers by fish and decreases primary producer biomass, but effects of warming are generally small compared to effects of browning and nutrient supply. Experimental results were consistent with most model predictions for browning: light penetration, benthic algal production, and zoobenthos biomass decreased, and pelagic nutrients and pelagic algal production increased with browning. Also consistent with expectations, warming had negative effects on benthic and pelagic algal biomass and weak effects on algal production and zoobenthos and zooplankton biomass. Inconsistent with expectations, browning had no effect on zooplankton and warming effects on fish depended on browning. The model is applicable also to nutrient‐rich systems, and we propose that it is a useful tool for the exploration of the consequences of different climate change scenarios for productivity and food web dynamics in shallow lakes, the worldwide most common lake type.     

Nutrients associated with terrestrial dissolved organic matter drive changes in zooplankton:phytoplankton biomass ratios in an alpine lake

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Simulated eutrophication and browning alters zooplankton nutritional quality and determines juvenile fish growth and survival

The first few months of life is the most vulnerable period for fish and their optimal hatching time with zooplankton prey is favored by natural selection. Traditionally, however, prey abundance (i.e., zooplankton density) has been considered important, whereas prey nutritional composition has been largely neglected in natural settings. High‐quality zooplankton, rich in both essential amino acids (EAAs) and fatty acids (FAs), are required as starting prey to initiate development and fast juvenile growth. Prey quality is dependent on environmental conditions, and, for example, eutrophication and browning are two major factors defining primary producer community structures that will directly determine the nutritional quality of the basal food sources (algae, bacteria, terrestrial matter) for zooplankton. We experimentally tested how eutrophication and browning affect the growth and survival of juvenile rainbow trout (Oncorhynchus mykiss) by changing the quality of basal resources. We fed the fish on herbivorous zooplankton (Daphnia) grown with foods of different nutritional quality (algae, bacteria, terrestrial matter), and used GC‐MS, stable isotope labeling as well as bulk and compound‐specific stable isotope analyses for detecting the effects of different diets on the nutritional status of fish. The content of EAAs and omega‐3 (ω‐3) polyunsaturated FAs (PUFAs) in basal foods and zooplankton decreased in both eutrophication and browning treatments. The decrease in ω‐3 PUFA and especially docosahexaenoic acid (DHA) was reflected to fish juveniles, but they were able to compensate for low availability of EAAs in their food. Therefore, the reduced growth and survival of the juvenile fish was linked to the low availability of DHA. Fish showed very low ability to convert alpha‐linolenic acid (ALA) to DHA. We conclude that eutrophication and browning decrease the availability of the originally phytoplankton‐derived DHA for zooplankton and juvenile fish, suggesting bottom‐up regulation of food web quality.     

Browning reduces the availability—but not the transfer—of essential fatty acids in temperate lakes

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Intraspecific variation in a predator drives cascading variation in primary producer community composition

Predation has important cascading impacts on primary producer biomass and community composition in many ecosystems. While most studies have focused on the consequences of interspecific or density differences in predators, it is recognized that phenotypic variation within species can have strong and cascading community and ecosystem consequences at lower trophic levels. In coastal New England lakes, both the presence and life history form of the zooplanktivorous fish alewife, Alosa pseudoharengus, have strong influence on the biomass, size structure and community composition of crustacean zooplankton communities. Here we test the hypothesis that alewife presence and life history will have cascading impacts on phytoplankton biomass and community composition in a mesocosm experiment that previously reported strong biomass and compositional differences of crustacean zooplankton communities among alewife treatments. We show that alewife life history led to small but statistically significant differences in phytoplankton community composition among treatments. This compositional difference was driven primarily by an increase in the density of two edible phytoplankton genera associated with lower zooplankton biomass in the anadromous alewife treatment. Our results show that intraspecific variation in a predator can have cascading effects on primary producer communities. However we did not observe significant differences in total algal biomass.     

Biotic invasions can alter nutritional composition of zooplankton communities

Ecologists and ecosystem managers often base their understanding of trophic dynamics on consumer and resource biomass. However, the factors that alter the relative nutritional value of resources are often poorly understood, despite their potential to decouple trophic interactions. Recent population declines in pelagic fishes of the upper San Francisco Estuary were not accompanied by an equivalent decrease in zooplankton biomass, which are the main resource for the fish and their larvae. It was hypothesized that changes in zooplankton nutritional conditions following the establishment of invasive species caused food‐quality related limitations for these higher‐order consumers. Using stable isotopes, elemental stoichiometry and fatty acid analyses for all dominant invasive and native zooplankton taxa and seston, we characterized the plankton community structure in the estuary and demonstrated taxon‐specific differences in their nutritional value. We then quantified the temporal dynamics in meso‐zooplankton proportions of docosahexaenoic acid (DHA), eicosapentaenoic acid (EPA), polyunsaturated fatty acids (PUFA), and ratio of n3:n6 fatty acids. We found temporal increase in the community‐level DHA, n3 to n6 fatty acid ratio, decrease in the community‐level EPA and PUFA in the brackish water region, but no change in the bulk PUFA proportions in the freshwater region of the estuary. These changes were caused mainly by declines of native cladocerans that are rich in EPA and by an increase in the dominance of invasive taxa with high DHA concentrations, similar to that of native taxa. Although we showed temporal shifts in individual fatty acid classes, the proportion of the essential fatty acids remained relatively high, suggesting that nutritional prey availability for fish remained unchanged with the shift in species composition. We argue that the nutritional content of resource communities should be considered when analyzing the long‐term trophic dynamics and designing effective management and restoration strategies.     

How do temperature,dissolved organic matter and nutrients influence the response of Leptodiaptomus ashlandi to UV radiation in a subalpine lake?

1. Ultraviolet radiation (UV) is an important stressor for zooplankton in alpine lake ecosystems. Multiple environmental variables such as dissolved organic matter (DOM), temperature and nutrient availability may alter how UV affects zooplankton. 2. We conducted a week‐long experiment manipulating UV, nutrients and DOM in enclosures suspended at the surface of cold and warm alpine lakes to determine the interactive effects of these variables on ovigerous Leptodiaptomus ashlandi (Marsh, 1893), a calanoid copepod. 3. UV had a negative effect on nauplii and gravid females at the colder temperature and at low, ambient DOM levels, but had no effect at the warmer temperature or when DOM was added. At the warmer temperature, fewer nauplii were produced in the +nutrient compared to ?nutrient treatment. Adult survival was not affected by UV or any other experimental variable. 4. These results demonstrate that the extent of the impact of UV radiation on zooplankton in alpine systems is altered by other environmental variables, and that these effects may not be apparent from experiments that look only at the survival of adult organisms that are better defended against UV.     

Decoupling carbon effects and UV protection from terrestrial subsidies on pond zooplankton

Studies on spatial subsidies have overwhelmingly addressed their trophic nature. However, certain subsidies might affect recipient communities beyond the feeding pathway. Terrestrial colored dissolved organic matter (DOM) is such a subsidy providing a carbon source for bacteria and heterotrophic nanoflagellates which zooplankton can consume (trophic pathway), but also protecting zooplankton against damaging UV radiation through its color (non-trophic pathway). These mechanisms have been quantified separately, but few studies have attempted to decouple them and evaluate their effects in the context of subsidies. In this study, I experimentally isolated the trophic and non-trophic pathways by which DOM (as maple leaf leachate) affects zooplankton, and also addressed how local food resources (i.e., phytoplankton) could mediate the response of zooplankton to DOM subsidies. Terrestrial subsidies (DOM) effectively shielded zooplankton against damaging UV radiation. Local resources (i.e., phytoplankton), however, did not seem to mediate the fitness response of zooplankton to UV radiation. This study also suggested that the carbon in DOM might be slightly detrimental to zooplankton independent of the UV protection effect. High levels of local resources combined with carbon subsidies from DOM did not translate into higher zooplankton survival, presumably because of the detrimental effects of DOM on zooplankton. This study provides further support for the importance of terrestrial subsidies in affecting communities through non-trophic pathways (UV attenuation) and documents that the trophic pathway might be strongly dependent on subsidy quality. Further studies on the role of subsidies that include the less documented non-trophic pathways are needed to improve our mechanistic understanding of how communities and ecosystems respond to spatial subsidies. Handling editor: D. Bade     

Experimental whole‐lake increase of dissolved organic carbon concentration produces unexpected increase in crustacean zooplankton density

The observed pattern of lake browning, or increased terrestrial dissolved organic carbon (DOC) concentration, across the northern hemisphere has amplified the importance of understanding how consumer productivity varies with DOC concentration. Results from comparative studies suggest these increased DOC concentrations may reduce crustacean zooplankton productivity due to reductions in resource quality and volume of suitable habitat. Although these spatial comparisons provide an expectation for the response of zooplankton productivity as DOC concentration increases, we still have an incomplete understanding of how zooplankton respond to temporal increases in DOC concentration within a single system. As such, we used a whole‐lake manipulation, in which DOC concentration was increased from 8 to 11 mg L^?1 in one basin of a manipulated lake, to test the hypothesis that crustacean zooplankton production should subsequently decrease. In contrast to the spatially derived expectation of sharp DOC‐mediated decline, we observed a small increase in zooplankton densities in response to our experimental increase in DOC concentration of the treatment basin. This was due to significant increases in gross primary production and resource quality (lower seston carbon‐to‐phosphorus ratio; C:P). These results demonstrate that temporal changes in lake characteristics due to increased DOC may impact zooplankton in ways that differ from those observed in spatial surveys. We also identified significant interannual variability across our study region, which highlights potential difficulty in detecting temporal responses of organism abundances to gradual environmental change (e.g., browning).     

Impacts of dreissenid mussel invasions on chlorophyll and total phosphorus in 25 lakes in the USA

1. Invasions of zebra and quagga mussels have had long‐term, large‐scale impacts on lake ecosystems in the USA as characterised by high abundance, broad‐scale spread and effective adaption to new environmental conditions. Due to their high filtering capacity, decreases in chlorophyll a (Chl) and total phosphorus (TP) concentrations have been reported in many affected lakes. 2. In 25 US lakes, we analysed the effects of dreissenid invasions on changes in Chl and TP concentrations, measured as the probability of a concentration decrease in the post‐invasion period and changes in Chl–TP relationships using Bayesian hierarchical regressions. We also examined whether changes in Chl and TP concentrations and in the Chl–TP relationship differed according to lake hydromorphology, such as mean depth or mixing status (mixed versus stratified lakes). 3. Our results showed that dreissenid invasions were often, but not always, associated with subsequent TP and Chl declines. Dreissenid effects on Chl and TP may be influenced by lake thermal structure. Decreases in Chl and TP were consistently found in mixed lakes where benthic–pelagic coupling is tight, while the effects were less predictable in stratified lakes. Within stratified lakes, Chl and TP reductions were more clearly discernible in deeper lakes with long water residence times. 4. Regression results demonstrated that a joint increase in slope and decrease in intercept and a tighter correlation of the Chl–TP relationship were likely to occur in dreissenid‐invaded lakes; this does not support the idea of a shift from bottom‐up to top‐down control of primary production. These results have important implications for management, suggesting that a relaxation of TP standards would be unwarranted. 5. Across lakes, the slope of the Chl–TP relationship for mixed lakes was substantially higher than that for stratified lakes before mussel invasion, indicating an important role of light in limiting primary production. The slope differences between mixed and stratified lakes decreased in the post‐invasion period, possibly because mussel filtration results in a relaxation of light limitation that is more pronounced in deeper, stratified lakes.     

Long‐term nitrogen loading alleviates phosphorus limitation in terrestrial ecosystems

Increased human‐derived nitrogen (N) deposition to terrestrial ecosystems has resulted in widespread phosphorus (P) limitation of net primary productivity. However, it remains unclear if and how N‐induced P limitation varies over time. Soil extracellular phosphatases catalyze the hydrolysis of P from soil organic matter, an important adaptive mechanism for ecosystems to cope with N‐induced P limitation. Here we show, using a meta‐analysis of 140 studies and 668 observations worldwide, that N stimulation of soil phosphatase activity diminishes over time. Whereas short‐term N loading (≤5 years) significantly increased soil phosphatase activity by 28%, long‐term N loading had no significant effect. Nitrogen loading did not affect soil available P and total P content in either short‐ or long‐term studies. Together, these results suggest that N‐induced P limitation in ecosystems is alleviated in the long‐term through the initial stimulation of soil phosphatase activity, thereby securing P supply to support plant growth. Our results suggest that increases in terrestrial carbon uptake due to ongoing anthropogenic N loading may be greater than previously thought.     

Long-term changes in phytoplankton, zooplankton and salmon related to climate

Recently, large‐scale changes in the biogeography of calanoid copepod crustaceans have been detected in the northeastern North Atlantic Ocean and adjacent seas. Strong biogeographical shifts in all copepod assemblages were found with a northward extension of more than ° in latitude of warm‐water species associated with a decrease in the number of colder‐water species. These changes were attributed to regional increase in sea surface temperature. Here, we have extended these studies to examine long‐term changes in phytoplankton, zooplankton and salmon in relation to hydro‐meteorological forcing in the northeast Atlantic Ocean and adjacent seas. We found highly significant relationships between (1) long‐term changes in all three trophic levels, (2) sea surface temperature in the northeastern Atlantic, (3) Northern Hemisphere temperature and (4) the North Atlantic Oscillation. The similarities detected between plankton, salmon, temperature and hydro‐climatic parameters are also seen in their cyclical variability and in a stepwise shift that started after a pronounced increase in Northern Hemisphere Temperature anomalies at the end of the 1970s. All biological variables show a pronounced change which started after circa 1982 for euphausiids (decline), 1984 for the total abundance of small copepods (increase), 1986 for phytoplankton biomass (increase) and Calanus finmarchicus (decrease) and 1988 for salmon (decrease). This cascade of biological events led to an exceptional period, which is identified after 1986 to present and followed another shift in large‐scale hydro‐climatic variables and sea surface temperature. This regional temperature increase therefore appears to be an important parameter that is at present governing the dynamic equilibrium of northeast Atlantic pelagic ecosystems with possible consequences for biogeochemical processes and fisheries.     

Long‐term nutrient addition alters arthropod community composition but does not increase total biomass or abundance

A simple bottom–up hypothesis predicts that plant responses to nutrient addition should determine the response of consumers: more productive and less diverse plant communities, the usual result of long‐term nutrient addition, should support greater consumer abundances and biomass and less consumer diversity. We tested this hypothesis for the response of an aboveground arthropod community to an uncommonly long‐term (24‐year) nutrient addition experiment in moist acidic tundra in arctic Alaska. This experiment altered plant community composition, decreased plant diversity and increased plant production and biomass as a deciduous shrub, Betula nana, became dominant. Consistent with strong effects on the plant community, nutrient addition altered arthropod community composition, primarily through changes to herbivore taxa in the canopy‐dwelling arthropod assemblage and detritivore taxa in the ground assemblage. Surprisingly, however, the loss of more than half of plant species was accompanied by negligible changes to diversity (rarefied richness) of arthropod taxa (which were primarily identified to family). Similarly, although long‐term nutrient addition in this system roughly doubles plant production and biomass, arthropod abundance was either unchanged or decreased by nutrient addition, and total arthropod biomass was unaffected. Our findings differ markedly from the handful of terrestrial studies that have found bottom‐up diversity cascades and productivity responses by consumers to nutrient addition. This is probably because unlike grasslands and salt marshes (where such studies have historically been conducted), this arctic tundra community becomes less palatable, rather than more so, after many years of nutrient addition due to increased dominance of B. nana. Additionally, by displacing insulating mosses and increasing the cover of shrubs that cool and shade the canopy microenvironment, fertilization may displace arthropods keenly attuned to microclimate. These results indicate that terrestrial arthropod assemblages may be more constrained by producer traits (i.e. palatability, structure) than they are by total primary production or producer diversity.     

Changes in behavior are unable to disrupt a trophic cascade involving a specialist herbivore and its food plant

Changes in ecological conditions can induce changes in behavior and demography of wild organisms, which in turn may influence population dynamics. Black brant (Branta bernicla nigricans) nesting in colonies on the Yukon–Kuskokwim Delta (YKD) in western Alaska have declined substantially (~50%) since the turn of the century. Black brant are herbivores that rely heavily on Carex subspathacea (Hoppner's sedge) during growth and development. The availability of C. subspathacea affects gosling growth rates, which subsequently affect pre‐ and postfledging survival, as well as size and breeding probability as an adult. We predicted that long‐term declines in C. subspathacea have affected gosling growth rates, despite the potential of behavior to buffer changes in food availability during brood rearing. We used Bayesian hierarchical mixed‐effects models to examine long‐term (1987–2015) shifts in brant behavior during brood rearing, forage availability, and gosling growth rates at the Tutakoke River colony. We showed that locomotion behaviors have increased (β = 0.05, 95% CRI: 0.032–0.068) while resting behaviors have decreased (β = ?0.024, 95% CRI: ?0.041 to ?0.007), potentially in response to long‐term shifts in forage availability and brood density. Concurrently, gosling growth rates have decreased substantially (β = ?0.100, 95% CRI: ?0.191 to ?0.016) despite shifts in behavior, mirroring long‐term declines in the abundance of C. subspathacea (β = ?0.191, 95% CRI: ?0.355 to ?0.032). These results have important implications for individual fitness and population viability, where shifts in gosling behavior putatively fail to mitigate long‐term declines in forage availability.     

How humans alter dissolved organic matter composition in freshwater: relevance for the Earth’s biogeochemistry

Dissolved organic matter (DOM) is recognized for its importance in freshwater ecosystems, but historical reliance on DOM quantity rather than indicators of DOM composition has led to an incomplete understanding of DOM and an underestimation of its role and importance in biogeochemical processes. A single sample of DOM can be composed of tens of thousands of distinct molecules. Each of these unique DOM molecules has their own chemical properties and reactivity or role in the environment. Human activities can modify DOM composition and recent research has uncovered distinct DOM pools laced with human markers and footprints. Here we review how land use change, climate change, nutrient pollution, browning, wildfires, and dams can change DOM composition which in turn will affect internal processing of freshwater DOM. We then describe how human-modified DOM can affect biogeochemical processes. Drought, wildfires, cultivated land use, eutrophication, climate change driven permafrost thaw, and other human stressors can shift the composition of DOM in freshwater ecosystems increasing the relative contribution of microbial-like and aliphatic components. In contrast, increases in precipitation may shift DOM towards more relatively humic-rich, allochthonous forms of DOM. These shifts in DOM pools will likely have highly contrasting effects on carbon outgassing and burial, nutrient cycles, ecosystem metabolism, metal toxicity, and the treatments needed to produce clean drinking water. A deeper understanding of the links between the chemical properties of DOM and biogeochemical dynamics can help to address important future environmental issues, such as the transfer of organic contaminants through food webs, alterations to nitrogen cycling, impacts on drinking water quality, and biogeochemical effects of global climate change.

     

A matter of timing: heat wave impact on crustacean zooplankton

1. Climate change has affected zooplankton phenology and abundance in many freshwater ecosystems. The strong temperature anomalies that characterise summer heat waves make these events particularly suitable to study the effects of different seasonal warming patterns on zooplankton. Since heat waves are expected to occur more frequently under continuing climate change, they may also allow us to investigate how freshwater systems will be affected in the future. 2. Using a long‐term data set (1991–2007) from a shallow, eutrophic lake in Germany, we identify time periods in spring and summer during which cyclopoid copepods and bosminids are particularly sensitive to changes in water temperature. Based on this knowledge, we consider why summer populations responded differently to recent heat wave events that occurred at different times in the season. 3. Linear regressions of moving averages suggested that water temperatures shortly before and shortly after the clear‐water phase (CWP) were crucial for summer development of bosminids and cyclopoid copepods, respectively. Algal food availability (diatoms and cryptophytes) in the first weeks after the CWP also strongly influenced the summer populations of the two zooplankton groups. 4. Inter‐annual differences in water temperature during the critical time periods at least partly explained the contrasting responses of cyclopoid copepods and bosminids to heat wave events. 5. Our findings indicate that the zooplankton response to climate warming, particularly to heat wave events, is critically dependent on the temporal pattern of elevated water temperatures. Beyond that, we show that the summer zooplankton populations react to periods of warming in relation to events in the plankton annual cycle (such as the CWP in eutrophic lakes) rather than to warming at a fixed time in the season.     

Roles of parasitic fungi in aquatic food webs: a theoretical approach

1. Parasites are ubiquitous in ecosystems, but their roles in material transfer are poorly understood. Fungal parasites in freshwater ecosystems are of major importance to small heterotrophic eukaryotes and consume large phytoplankton that are resistant to zooplankton grazing. 2. To evaluate their ecosystem‐level effects, we developed a simple food web model that incorporates competition between small and large phytoplankton for nutrients, zooplankton grazing on small phytoplankton, fungal parasitism on large phytoplankton and includes a newly discovered trophic link from fungal zoospores to zooplankton (F‐Z link). 3. Our model demonstrates the likely occurrence of an indirect mutualism between fungi and zooplankton, in which fungal parasitism increases zooplankton production by reducing the biomass of inedible large phytoplankton. Contradicting the expectation from a previous short‐term experiment that the F‐Z link may benefit zooplankton, the presence of the F‐Z link can reduce material transfer from phytoplankton to zooplankton because of the negative effect of the indirect mutualism. The model indicates that high growth efficiency of fungi on host tissue and their high nutrient status for zooplankton are crucial for the F‐Z link to increase zooplankton production. 4. The model also indicates that the contribution of material transfer via F‐Z link to zooplankton increases with nutrient availability. Our results suggest that parasitic fungi may be a key player in material transfer, especially in eutrophic ecosystems.     

Decadal‐scale regional changes in Canadian freshwater zooplankton: the likely consequence of complex interactions among multiple anthropogenic stressors

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