Climate warming and agricultural stressors interact to determine stream macroinvertebrate community dynamics

Global climate change is likely to modify the ecological consequences of currently acting stressors, but potentially important interactions between climate warming and land‐use related stressors remain largely unknown. Agriculture affects streams and rivers worldwide, including via nutrient enrichment and increased fine sediment input. We manipulated nutrients (simulating agricultural run‐off) and deposited fine sediment (simulating agricultural erosion) (two levels each) and water temperature (eight levels, 0–6°C above ambient) simultaneously in 128 streamside mesocosms to determine the individual and combined effects of the three stressors on macroinvertebrate community dynamics (community composition and body size structure of benthic, drift and insect emergence assemblages). All three stressors had pervasive individual effects, but in combination often produced additive or antagonistic outcomes. Changes in benthic community composition showed a complex interplay among habitat quality (with or without sediment), resource availability (with or without nutrient enrichment) and the behavioural/physiological tendency to drift or emerge as temperature rose. The presence of sediment and raised temperature both resulted in a community of smaller organisms. Deposited fine sediment strongly increased the propensity to drift. Stressor effects were most prominent in the benthic assemblage, frequently reflected by opposite patterns in individuals quitting the benthos (in terms of their propensity to drift or emerge). Of particular importance is that community measures of stream health routinely used around the world (taxon richness, EPT richness and diversity) all showed complex three‐way interactions, with either a consistently stronger temperature response or a reversal of its direction when one or both agricultural stressors were also in operation. The negative effects of added fine sediment, which were often stronger at raised temperatures, suggest that streams already impacted by high sediment loads may be further degraded under a warming climate. However, the degree to which this will occur may also depend on in‐stream nutrient conditions.     

Multiple Stressors in Agricultural Streams: A Mesocosm Study of Interactions among Raised Water Temperature,Sediment Addition and Nutrient Enrichment

Changes to land use affect streams through nutrient enrichment, increased inputs of sediment and, where riparian vegetation has been removed, raised water temperature. We manipulated all three stressors in experimental streamside channels for 30 days and determined the individual and pair-wise combined effects on benthic invertebrate and algal communities and on leaf decay, a measure of ecosystem functioning. We added nutrients (phosphorus+nitrogen; high, intermediate, natural) and/or sediment (grain size 0.2 mm; high, intermediate, natural) to 18 channels supplied with water from a nearby stream. Temperature was increased by 1.4°C in half the channels, simulating the loss of upstream and adjacent riparian shade. Sediment affected 93% of all biological response variables (either as an individual effect or via an interaction with another stressor) generally in a negative manner, while nutrient enrichment affected 59% (mostly positive) and raised temperature 59% (mostly positive). More of the algal components of the community responded to stressors acting individually than did invertebrate components, whereas pair-wise stressor interactions were more common in the invertebrate community. Stressors interacted often and in a complex manner, with interactions between sediment and temperature most common. Thus, the negative impact of high sediment on taxon richness of both algae and invertebrates was stronger at raised temperature, further reducing biodiversity. In addition, the decay rate of leaf material (strength loss) accelerated with nutrient enrichment at ambient but not at raised temperature. A key implication of our findings for resource managers is that the removal of riparian shading from streams already subjected to high sediment inputs, or land-use changes that increase erosion or nutrient runoff in a landscape without riparian buffers, may have unexpected effects on stream health. We highlight the likely importance of intact or restored buffer strips, both in reducing sediment input and in maintaining cooler water temperatures.     

Light availability regulates the response of algae and heterotrophic bacteria to elevated nutrient levels and warming in a northern boreal peatland

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Gastropod grazers and nutrients,but not light,interact in determining periphytic algal diversity

The potential interactions of grazing, nutrients and light in influencing autotroph species diversity have not previously been considered. Earlier studies have shown that grazing and nutrients interact in determining autotroph species diversity, since grazing decreases species diversity when nutrients (i.e. N or P) limit autotroph growth, but increases it when nutrients are replete. We hypothesized that increased light intensities would intensify the interactions between grazing and nutrients on algal species diversity, resulting in even stronger reductions in algal species diversity through grazing under nutrient–poor conditions, and to even stronger increases of algal species diversity through grazing under nutrient-rich conditions. We studied the effects of grazing (absent, present), nutrients (ambient, N + P enriched) and light (low light, high light) on benthic algal diversity and periphyton C:nutrient ratios (which can indicate algal nutrient limitation) in a factorial laboratory experiment, using the gastropod grazer Viviparus viviparus. Grazing decreased algal biomass and algal diversity, but increased C:P and N:P ratios of periphyton. Grazing also affected periphyton species composition, by decreasing the proportion of Spirogyra sp. and increasing the proportion of species in the Chaetophorales. Grazing effects on diversity as well as on periphyton N:P ratios were weakened when nutrients were added (interaction between grazing and nutrients). Chlorophyll a (Chl a) per area increased with nutrient addition and decreased with high light intensities. Light did not increase the strength of the interaction between grazing and nutrients on periphytic algal diversity. This study shows that nutrient addition substantially reduced the negative effects of grazing on periphytic algal diversity, whereas light did not interact with grazing or nutrient enrichment in determining periphytic algal diversity.     

Interactive multiple‐stressor effects of the antibiotic monensin,cattle effluent and light on stream periphyton

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Patterns of benthic algae and cyanobacteria along twin‐stressor gradients of nutrients and fine sediment: a stream mesocosm experiment

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MULTIVARIATE ANALYSIS OF PERIPHYTON ASSEMBLAGES IN RELATION TO ENVIRONMENTAL GRADIENTS IN COLORADO ROCKY MOUNTAIN STREAMS 1

We assessed relationships between chemical and physical characteristics and periphyton assemblages in stream reaches of the mineral belt in the Southern Rockies ecoregion of Colorado, United States. Using canonical ordination analyses, we contrasted results wherein assemblage structure was assessed using community metrics or species abundances. Our objectives were to identify community metrics or individual species diagnostic of the primary environmental stressors in these streams, to compare the sensitivity of these two approaches to determining the primary stressors, and to determine how these approaches may be used to differentiate the effects of these environmental stressors from other gradients. For periphyton metrics, the first axis extracted by redundancy analysis correlated with total phosphorus, substrate coarseness and embeddedness, and riparian vegetation condition, whereas the second axis correlated with dissolved metals. For species abundances, the three axes extracted by canonical correspondence analysis (CCA) were correlated with (1) stream size and types of in‐stream habitats; (2) total phosphorus, dissolved ions, and riparian disturbance by agriculture; and (3) sediment coarseness and embeddedness and riparian vegetation condition. Concentrations of dissolved metals were not correlated with the CCA axes. Analyses of species abundances were sensitive to effects associated with nutrients, substrates, and riparian vegetation, whereas analyses of periphyton metrics were sensitive not only to these nutrient and physical habitat effects but also to toxicological effects associated with metals. As a result, the analyses of periphyton metrics may be used to identify which metrics would be useful for a periphyton index of biotic integrity and would also be individually diagnostic of the larger scale stressors in these streams, nutrient, substrate, and riparian vegetation effects of livestock grazing and increased metal concentrations associated with metal mining.     

Disturbance frequency and functional identity mediate ecosystem processes in prairie streams

A major consequence of climate change will be the alteration of precipitation patterns and concomitant changes in the flood frequencies in streams. Species losses or introductions will accompany these changes, which necessitates understanding the interactions between altered disturbance regimes and consumer functional identity to predict dynamics of streams. We used experimental mesocosms and field enclosures to test the interactive effects of flood frequency and two fishes from distinct consumer groups (benthic grazers and water-column minnows) on recovery of stream ecosystem properties (algal form and biomass, invertebrate densities, metabolism and nutrient uptake rates). Our results generally suggest that periphyton communities under nutrient limitation are likely to recover more quickly when grazing and water-column minnows are present and these effects can diminish or reverse with time since the disturbance. We hypothesized that increased periphyton production and biomass was the result of increased nutrient turnover, but decreased light limitation and indirect effects on other trophic levels are alternative explanations. Recovery of stream ecosystem properties after a natural flood differed from mesocosms (e.g. lower algal biomass and no long algal filaments present) and species manipulations did not explain recovery of ecosystem properties; rather, ecosystem processes varied along a downstream gradient of increasing temperature and nutrient concentrations. Different results between field enclosures and experimental mesocosms are attributable to a number of factors including differences in algal and invertebrate communities in the natural stream and relatively short enclosure lengths (mean area=35.8 m²) compared with recirculating water in the experimental mesocosms. These differences may provide insight into conditions necessary to elicit a strong interaction between consumers and ecosystem properties.     

Seasonal shifts in the importance of bottom–up and top–down factors on stream periphyton community structure

We examined the importance of temporal variability in top–down and bottom–up effects on the accumulation of stream periphyton, which are complex associations of autotrophic and heterotrophic microorganisms. Periphyton contributes to primary production and nutrient cycling and serves as a food resource for herbivores (grazers). Periphyton growth is often limited by the availability of nitrogen and phosphorus, and biomass can be controlled by grazers. In this study we experimentally manipulated nutrients and grazers simultaneously to determine the relative contribution of bottom–up and top–down controls on periphyton over time. We used nutrient diffusing substrates to regulate nutrient concentrations and an underwater electric field to exclude grazing insects in three sequential 16–17 day experiments from August to October in montane Colorado, USA. We measured algal biomass, periphyton organic mass, and algal community composition in each experiment and determined densities of streambed insect species, including grazers. Phosphorus was the primary limiting nutrient for algal biomass, but it did not influence periphyton organic mass across all experiments. Effects of nutrient additions on algal biomass and community composition decreased between August and October. Grazed substrates supported reduced periphyton biomass only in the first experiment, corresponding to high benthic abundances of a dominant mayfly grazer (Rhithrogena spp.). Grazed substrates in the first experiment also showed altered algal community composition with reduced diatom relative abundances, presumably in response to selective grazing. We showed that top–down grazing effects were strongest in late summer when grazers were abundant. The effects of phosphorus additions on algal biomass likely decreased over time because temperature became more limiting to growth than nutrients, and because reduced current velocity decreased nutrient uptake rates. These results suggest that investigators should proceed with caution when extending findings based on short‐term experiments. Furthermore, these results support the need for additional seasonal‐scale field research in stream ecology.     

Functioning of a Shallow-Water Sediment System during Experimental Warming and Nutrient Enrichment

Effects of warming and nutrient enrichment on intact unvegetated shallow-water sediment were investigated for 5 weeks in the autumn under simulated natural field conditions, with a main focus on trophic state and benthic nitrogen cycling. In a flow-through system, sediment was exposed to either seawater at ambient temperature or seawater heated 4°C above ambient, with either natural or nutrient enriched water. Sediment–water fluxes of oxygen and inorganic nutrients, nitrogen mineralization, and denitrification were measured. Warming resulted in an earlier shift to net heterotrophy due to increased community respiration; primary production was not affected by temperature but (slightly) by nutrient enrichment. The heterotrophic state was, however, not further strengthened by warming, but was rather weakened, probably because increased mineralization induced a shortage of labile organic matter. Climate-related warming of seawater during autumn could therefore, in contrast to previous predictions, induce shorter but more intensive heterotrophic periods in shallow-water sediments, followed by longer autotrophic periods. Increased nitrogen mineralization and subsequent effluxes of ammonium during warming suggested a preferential response of organisms driving nitrogen mineralization when compared to sinks of ammonium such as nitrification and algal assimilation. Warming and nutrient enrichment resulted in non-additive effects on nitrogen mineralization and denitrification (synergism), as well as on benthic fluxes of phosphate (antagonism). The mode of interaction appears to be related to the trophic level of the organisms that are the main drivers of the affected processes. Despite the weak response of benthic microalgae to both warming and nutrient enrichment, the assimilation of nitrogen by microalgae was similar in magnitude to rates of nitrogen mineralization. This implies a sustained filter function and retention capacity of nutrients by the sediment.     

Effects of nutrient loading,temperature regime and grazing pressure on nutrient limitation of periphyton in experimental ponds

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Nutrient loading and grazing by the minnow Phoxinus erythrogaster shift periphyton abundance and stoichiometry in mesocosms

1. Anthropogenic activities in prairie streams are increasing nutrient inputs and altering stream communities. Understanding the role of large consumers such as fish in regulating periphyton structure and nutritional content is necessary to predict how changing diversity will interact with nutrient enrichment to regulate stream nutrient processing and retention. 2. We characterised the importance of grazing fish on stream nutrient storage and cycling following a simulated flood under different nutrient regimes by crossing six nutrient concentrations with six densities of a grazing minnow (southern redbelly dace, Phoxinus erythrogaster) in large outdoor mesocosms. We measured the biomass and stoichiometry of overstory and understory periphyton layers, the stoichiometry of fish tissue and excretion, and compared fish diet composition with available algal assemblages in pools and riffles to evaluate whether fish were selectively foraging within or among habitats. 3. Model selection indicated nutrient loading and fish density were important to algal composition and periphyton carbon (C): nitrogen (N). Nutrient loading increased algal biomass, favoured diatom growth over green algae and decreased periphyton C : N. Increasing grazer density did not affect biomass and reduced the C : N of overstory, but not understory periphyton. Algal composition of dace diet was correlated with available algae, but there were proportionately more diatoms present in dace guts. We found no correlation between fish egestion/excretion nutrient ratios and nutrient loading or fish density despite varying N content of periphyton. 4. Large grazers and nutrient availability can have a spatially distinct influence at a microhabitat scale on the nutrient status of primary producers in streams.     

Multiple anthropogenic stressors exert complex,interactive effects on a coral reef community

Multiple natural and anthropogenic stressors impact coral reefs across the globe leading to declines of coral populations, but the relative importance of different stressors and the ways they interact remain poorly understood. Because coral reefs exist in environments commonly impacted by multiple stressors simultaneously, understanding their interactions is of particular importance. To evaluate the role of multiple stressors we experimentally manipulated three stressors (herbivore abundance, nutrient supply, and sediment loading) in plots on a natural reef in the Gulf of Panamá in the Eastern Tropical Pacific. Monitoring of the benthic community (coral, macroalgae, algal turf, and crustose coralline algae) showed complex responses with all three stressors impacting the community, but at different times, in different combinations, and with varying effects on different community members. Reduction of top–down control in combination with sediment addition had the strongest effect on the community, and led to approximately three times greater algal biomass. Coral cover was reduced in all experimental units with a negative effect of nutrients over time and a synergistic interaction between herbivore exclosures and sediment addition. In contrast, nutrient and sediment additions interacted antagonistically in their impacts on crustose coralline algae and turf algae so that in combination the treatments limited each other’s effects. Interactions between stressors and temporal variability indicated that, while each stressor had the potential to impact community structure, their combinations and the broader environmental conditions under which they acted strongly influenced their specific effects. Thus, it is critical to evaluate the effects of stressors on community dynamics not only independently but also under different combinations or environmental conditions to understand how those effects will be played out in more realistic scenarios.     

Interactions among irradiance,nutrients, and herbivores constrain a stream algal community

Using stream-side, flow-through channels, I tested for the effects of nutrients (NU) (nitrogen plus phosphorus), irradiance (L), and snail grazing (G) on a benthic algal community in a small, forested stream. Grazed communities were-dominated by a chlorophyte (basal cells ofStigeoclonium) and a cyanophyte (Chamaesiphon investiens), whereas ungrazed communities were comprised almost entirely of diatoms, regardless of nutrient and light levels. Snails maintained low algal biomass in all grazed treatments, presumably by consuming increased algal production in treatments to which L and NU were increased. When nutrients were increased, cellular nutrient content increased under ambient conditions (shaded, grazed) and biomass and productivity increased when snails were removed and light was increased. Together, nutrients and light had positive effects and grazing had negative effects on biomass (chlorophylla, AFDM, algal biovolume) and chlorophyll-and areal-specific productivity in ANOVAs. However, in most cases, only means from treatments in which all three factors were manipulated (ungrazed, +NU&L treatments) were significantly different from controls; effects of single factors were generally undetectable. These results indicate that all three factors simultaneously limited algal biomass and productivity in this stream during the summer months. Additionally, the effects of these factors in combination were in some cases different from the effects of single factors. For example, light had slight negative effects on some biomass parameters when added at ambient snail densities and nutrient concentrations, but had strong positive effects in conjunction with nutrient addition and snail removal. This study demonstrates that algal biomass and productivity can be under multiple constraints by irradiance, nutrients, and herbivores and indicates the need to employ multifactor experiments to test for such interactive effects.     

The snail <Emphasis Type="Italic">Potamopyrgus antipodarum</Emphasis> grows faster and is more active in the shade,independent of food quality

Ecological stoichiometry has advanced food web ecology by emphasising the importance of food quality over food quantity for herbivores. Here, we focus on the effects of abiotic factors such as nutrients and light (known to influence food quality) on grazer growth rates. As model organism we used the mudsnail Potamopyrgus antipodarum that is native to New Zealand but invasive elsewhere. In a stream channel experiment in New Zealand, we manipulated light (two levels) and nutrients (four levels) to achieve a range of primary producer carbon: nutrient ratios and added mudsnails (3 densities + ungrazed control) to 128 periphyton covered stream channels in a 2 × 4 × 4 full factorial design. We measured snail growth rate and activity, food quality and nutritional imbalance, to test the predictions that (1) less light and more nutrients increase periphyton food quality and thus snail growth rates, and (2) less crowding leads to higher food availability and thus higher snail growth rates. We found that snail growth rates were higher under low light than under high light intensities and this difference increased with increasing nutrient addition. These changes in growth rate were not mediated by food quality in terms of periphyton nutrient ratios. Furthermore, experimental treatments strongly affected snail behaviour. Snails grazed more actively in the low light treatments, and thus it is more likely that snail growth rates were directly affected by light levels, maybe as a result of innate predator avoidance behaviour or as a reaction to high UV intensities. We conclude that in our stream channels snail growth rate was limited by factors other than food quality and quantity such as UV exposure, algal defences or the relatively low ambient water temperature.     

Effects of light, dissolved organic carbon, and inorganic nutrients [2pt] on the relationship between algae and heterotrophic bacteria in stream periphyton

Depending on the chemical and physical environment, algae and heterotrophic bacteria in stream periphyton communities likely engage in both positive and negative interactions. We tested the hypothesis that bacteria are more closely associated with algae when allochthonous sources of labile DOC are low and algae are not light limited. Secondly, we tested the hypothesis that, under extremely oligotrophic conditions, bacteria will out-compete algae for inorganic nutrients if their carbon requirements are met by allochthonous sources. Experiments were carried out using in situ light manipulations and nutrient diffusing substrates (releasing inorganic nutrients and /or glucose) in Harts Run, an oligotrophic stream located in north central Kentucky. Although we found that both algal and bacterial biomass were higher under ambient light, bacteria did not respond to glucose in the dark. This may indicate that bacteria were associated with algae not as a carbon source, but as a substrate for colonization. In the nutrient × glucose manipulation, we found that bacteria were co-limited by inorganic nutrients. There was no evidence of algae being negatively affected by competition with bacteria for nitrogen and phosphorus. Although low temperatures might have played a role in preventing inorganic nutrient competition between these two groups of organisms, the results of both experiments may indicate that the quantitative link between periphytic bacteria and algae is stronger under oligotrophic conditions.     

Stressor intensity determines antagonistic interactions between species invasion and multiple stressor effects on ecosystem functioning

Biological invasions, nutrient enrichment and ocean warming are known to threaten biodiversity and ecosystem functioning. The independent effects of these ecological stressors are well studied, however, we lack understanding of their cumulative effects, which may be additive, antagonistic or synergistic. For example, the impacts of biological invasions are often determined by environmental context, which suggests that the effects of invasive species may vary with other stressors such as pollution or climate change. This study examined the effects of an invasive seaweed (Sargassum muticum) on the structure and functioning of a synthetic macroalgal assemblage and tested explicitly whether these effects varied with nutrient enrichment and ocean warming. Overall, the presence of S. muticum increased assemblage productivity rates and warming altered algal assemblage structure, which was characterised by a decrease in kelp and an increase in ephemeral green algae. The effects of S. muticum on total algal biomass accumulation, however, varied with nutrient enrichment and warming, producing antagonistic cumulative effects on total algal biomass accumulation. These findings show that the nature of stressor interactions may vary with stressor intensity and among response variables, which leads to less predictable consequences for the structure and functioning of communities.     

Multiple stressor effects on coral reef ecosystems

Global climate change has profound implications on species distributions and ecosystem functioning. In the coastal zone, ecological responses may be driven by various biogeochemical and physical environmental factors. Synergistic interactions can occur when the combined effects of stressors exceed their individual effects. The Red Sea, characterized by strong gradients in temperature, salinity, and nutrients along the latitudinal axis provides a unique opportunity to study ecological responses over a range of these environmental variables. Using multiple linear regression models integrating in situ, satellite and oceanographic data, we investigated the response of coral reef taxa to local stressors and recent climate variability. Taxa and functional groups responded to a combination of climate (temperature, salinity, air‐sea heat fluxes, irradiance, wind speed), fishing pressure and biogeochemical (chlorophyll a and nutrients ‐ phosphate, nitrate, nitrite) factors. The regression model for each species showed interactive effects of climate, fishing pressure and nutrient variables. The nature of the effects (antagonistic or synergistic) was dependent on the species and stressor pair. Variables consistently associated with the highest number of synergistic interactions included heat flux terms, temperature, and wind speed followed by fishing pressure. Hard corals and coralline algae abundance were sensitive to changing environmental conditions where synergistic interactions decreased their percentage cover. These synergistic interactions suggest that the negative effects of fishing pressure and eutrophication may exacerbate the impact of climate change on corals. A high number of interactions were also recorded for algae, however for this group, synergistic interactions increased algal abundance. This study is unique in applying regression analysis to multiple environmental variables simultaneously to understand stressor interactions in the field. The observed responses have important implications for understanding climate change impacts on marine ecosystems and whether managing local stressors, such as nutrient enrichment and fishing activities, may help mitigate global drivers of change.     

Evaluating the multiple mechanisms underlying herbivore-algal interactions in streams

Stream algal responses to herbivory were investigated under different environmental conditions. Snail densities and nutrient concentrations were manipulated in experimental enclosures to document the influence of nutrient availability on the magnitude of algal responses to herbivory. Periphyton mats in other enclosures were subjected to physical disruption by artificial means to evaluate the influence of disturbance on algal abundance. The impact of herbivory on algal abundance decreased substantially with increase in water column nutrient concentrations. This result was explained by findings that: (1) algal accumulation was constrained by nutrient availability under ambient water quality conditions. (2) accumulation of most algal populations was stimulated by nutrient enrichment only under grazed conditions. Thus, snail grazing simultaneously exerted a negative impact on algal abundance, by removal and consumption of a portion of the periphyton mat, and a stimulative effect, by increasing the availability of nutrients to remaining cells. Algal responses to artificial disturbance indicated that stimulative effects of herbivory were caused by the physical disruption of thr assemblage rather than by other processes (e. g., nutrient regeneration). However, consumptive losses far outweighed stimulative effects on algal abundance under ambient nutrient conditions. The magnitude of these two antagonistic effects was comparable under enriched conditions because grazing had only slight effects on algal abundance. Thus, the importance of different mechanisms of algal-herbivore interactions is strongly influenced by ambient environmental conditions, a finding that has important implications for predicting the outcome of herbivore-algal interactions in ecosystems with pronounced temporal and spatial variation in biotic and abiotic conditions.     

Macroinvertebrate colonization dynamics on artificial substrates along an algal resource gradient

Riparian canopy removal and land use may introduce multiple stressors that can alter food and habitat for stream organisms, but the influence of these alterations on macroinvertebrate colonization dynamics is less well known. A field study involving the simultaneous placement and removal of artificial substrates was performed to examine how macroinvertebrate colonization rates might vary with algal accumulation within a perennial stream segment in eastern Ohio, USA. The study was conducted over a 60-day summer colonization period in three reaches that were selected to represent an algal resource gradient according to canopy cover and agricultural nutrient sources in the riparian corridor. Total nitrogen, water temperatures, and mean algal biomass from substrates increased along the resource gradient represented by the study sites. Total macroinvertebrate biomass and the abundance and biomass of scrapers also increased according to the gradient. Correlation results indicated that chlorophyll a biomass, rather than time or temperature, was better related to the abundance and biomass of most primary consumers on substrates. These results suggest that the combined effects of elevated temperatures and nutrients can result in relatively rapid algal accrual that may alter the colonization and establishment of macroinvertebrate communities in streams subjected to gradients of riparian disturbances.