Relative effects of nutrient enrichment and grazing on epiphyte-macrophyte (Zostera marina L.) dynamics

The independent and interactive effects of nutrient concentration and epiphyte grazers on epiphyte biomass and macrophyte growth and production were examined in Zostera marina L. (eelgrass) microcosms. Experiments were conducted during early summer, late summer, fall, and spring in a greenhouse on the York River estuary of Chesapeake Bay. Nutrient treatments consisted of ambient or enriched (3× ambient) concentrations of inorganic nitrogen (ammonium nitrate) and phosphate. Grazer treatments consisted of the presence or absence of field densities of isopods, amphipods, and gastropods. epiphyte biomass increased with both grazer removal and nutrient enrichment during summer and spring experiments. The effect of grazers was stronger than that of nutrients. There was little epiphyte response to treatment during the fall, a result possibly of high ambient nutrient concentrations and low grazing pressure. Under low grazer densities of early summer, macrophyte production (g m^–2 d^–1) was reduced by grazer removal and nutrient enrichment independently. Under high grazer densities of late summer, macrophyte production was reduced by enrichment only with grazers absent. During spring and fall there were no macrophyte responses to treatment. The relative influence of epiphytes on macrophyte production may have been related to seasonally changing water temperature and macrophyte requirements for light and inorganic carbon.     

Effect of nutrient availability, grazer assemblage and seagrass source population on the interaction between Thalassia testudinum (turtle grass) and its algal epiphytes

Seagrass leaves are often densely covered by epiphytic algae which can suppress seagrass productivity and has been implicated in declines of seagrass meadows worldwide. The net effect of epiphytes on seagrass growth and morphology depends on the independent and interactive effects of a variety of factors, including nutrient availability and the intensity of grazing on epiphytes. Here I report the results of a mesocosm experiment designed to test the effects of nutrient addition and within-functional group variation (grazer species composition and the source population of seagrass) on the strength of the interactions among grazers, epiphytes, and turtle grass (Thalassia testudinum). Turtle grass ramets from two sites in the northern Gulf of Mexico were cleared of epiphytes and transplanted into common-garden mesocosms. Replicate ramets were grown in a split-split plot design with two levels of dissolved nutrients and four different grazer species combinations (Tozeuma carolinense alone, Pagurus maclaughlinae alone, both species together, and no grazers present). As expected, grazers had a significant negative effect on epiphyte biomass/leaf area and a significant positive effect on turtle grass growth in the mesocosms. The two species were more similar in their direct effects on epiphyte biomass than in their indirect effects on turtle grass growth; this may reflect differences in epiphyte community composition under different grazer treatments. The effect of nutrient addition on turtle grass growth depended critically on the intensity of grazing: in the presence of grazers, turtle grass tended to produce a greater biomass of new leaf tissue in the tanks with nutrients added than in the control tanks. However, when grazers were absent, the direction of the effect was reversed, and plants with nutrients added grew less than the control plants. The two source populations of turtle grass differed significantly in epiphyte biomass/leaf area accrued in the mesocosms as well as in the strength of the effect of grazers on turtle grass growth. This suggests that population differentiation in seagrass interactions with epiphytes, as well as spatial and temporal variation in resources and grazer community composition, can greatly effect the role of epiphytes in limiting seagrass productivity.     

The effect of grazing and nutrient supply on periphyton associated bacteria

The effects of nutrient additions and grazing by macro-invertebrates on periphyton-associated algae and bacteria were studied by performing an enclosure experiment on three occasions from early spring to summer at mesotrophic Lake Erken and V?dd?, at the Swedish Baltic coast. There were significant interactions between nutrient additions and grazing on bacterial biomass and specific activity in Lake Erken. Thus, the importance of either bottom-up or top-down effects could not be singled out. Bacterial biomass increased with enrichment only in the absence of grazers. Grazer presence tended to increase bacterial biomass in ambient nutrient conditions, but to decrease bacterial biomass under enrichment. For specific activity the positive response to enrichment was restricted to grazer presence. Hence, grazing by macro-invertebrates may have an indirect positive effect on bacterial activity by enhancing nutrient conditions through their feeding activities and/or fecal pellets production. In addition, we found a significant relationship between bacterial production and chlorophyll a at both sites. This relationship weakened in the presence of macro-invertebrates. Thus, the importance of internal nutrient regeneration by bacteria and algae decreased, possibly due to increased nutrient availability, in the presence of macro-invertebrate grazers.     

EPIPHYTE-GRAZER INTERACTIONS ON ZOSTERA MARINA (ANTHOPHYTA: MONOCOTYLEDONES): EFFECTS OF DENSITY ON COMMUNITY FUNCTION1

Interactions between algal epiphytes and their grazers can have a significant impact on the structure and function of eelgrass (Zostera marina L.) meadows. In Puget Sound, the herbivorous gastropod Lacuna variegata Carpenter and its congeners appear to remove large quantities of the epiphytic community from eelgrass blades. When snails at typical field densities were used in microcosms, Lacuna significantly reduced epiphytic biomass and areal productivity. Biomass-specific productivity of the epiphytic community showed an increasing trend with increasing snail density. Epiphytic productivity increased nonlinearly with increasing epiphytic biomass. The commonly used logistic population growth formula adequately described this relationship. Grazing rate also increased nonlinearly with increasing epiphytic biomass. The Hailing equation adequately described the relationship between grazing rate and epiphytic biomass. The proportion of the epiphytic biomass found on the oldest blade of an eelgrass shoot was related linearly to epiphytic biomass, suggesting that a constant fraction of the epiphytic community is lost regardless of epiphytic density. Lacuna clearly removed large quantities of epiphytic material from eelgrass blades, significantly altering community function. Modified Lotka-Volterra equations, incorporating the logistic growth form and Hailing grazing equation, should prove useful in modeling the epiphyte–grazer interaction.     

Response of seagrass epiphyte loading to field manipulations of fertilization, gastropod grazing and leaf turnover rates

This study evaluates the bottom-up and top-down controls on epiphyte loads under low nutrient additions. Nutrients and gastropod grazers were manipulated in a field experiment conducted within a Thalassia testudinum meadow in Florida Bay, FL, USA. The effect of seagrass leaf turnover rate on epiphyte loading was also evaluated using novel seagrass short-shoot mimics that “grow,” allowing for the manipulation of leaf turnover rates. During the summer growing season and over the course of one seagrass leaf turnover period, low-level water column nutrient additions increased total epiphyte load, epiphyte chlorophyll a, and epiphyte autotrophic index. T. testudinum leaf nutrients (N and P) and leaf productivity also increased. Epiphyte loading and T. testudinum shoot biomass and productivity did not respond to a 60% mean increase in gastropod abundance. Manipulations of seagrass leaf turnover rates at minimum wintertime and maximum summertime rates resulted in a 20% difference in epiphyte loading. Despite elevated grazer abundances and increased leaf turnover rates, epiphyte loads increased with nutrient addition. These results emphasize the sensitivity of T. testudinum and associated epiphytes to low-level nutrient addition in a nutrient-limited environment such as Florida Bay.     

Consumer identity,abundance and nutrient concentration affect epiphyte diversity in an experimental eelgrass system

Conceptual models predict a unimodal effect of consumer abundance on prey diversity with the highest diversity at intermediate consumer abundance (intermediate disturbance hypothesis). Consumer selectivity and prey productivity are assumed to be further important determinants. Preferential grazing on dominant prey species favoured by high nutrient supply is supposed to increase prey diversity, whereas the effect of consumers on prey diversity may be negative under low nutrient conditions (grazer reversal hypothesis). We tested the effect of four common consumers the isopod Idotea baltica, the amphipod Gammarus oceanicus, and the gastropods Littorina littorea and Rissoa membranacea on diversity and composition of epiphytes growing on eelgrass Zostera marina. Consumer density was manipulated (four levels: grazer free control, low, medium, high) based on abundances observed in eelgrass systems. Additionally, we manipulated nutrient supply (three levels) and the presence of Idotea in a factorial experiment. The impact of consumer abundance on epiphyte diversity varied depending on consumer identity and epiphyte evenness was affected rather than species number in this short‐term experiment. Idotea reduced epiphyte diversity (Shannon‐Wiener index H') and Gammarus increased epiphyte diversity. Littorina had no effect at low and medium abundance, but a negative effect in the high density treatment. Only Rissoa supported the conceptual models as it caused the proposed unimodal pattern in epiphyte diversity. The varying species‐specific selectivity of the studied consumers is likely to explain their diverse impact on epiphyte diversity. Nutrients enhanced epiphyte diversity at medium enrichment, whereas higher nutrient supply reduced epiphyte diversity. The effect of Idotea changed from negative at low nutrient concentration to positive at higher nutrient supply, supporting the grazer reversal hypothesis. This study implies that consumer species identity and nutrient concentrations are important in controlling prey diversity and composition. Different consumer selectivity and changes in selectivity with growing consumer abundance and nutrient concentration are the causal factors for this effect.     

Effects of eutrophication on the interaction between algae and grazers in an Andean stream

Nutrient excess is a common disturbance that affects biological interactions in river ecosystems. The response of nutrient supply on primary producers and Tricorythodes sp., a common mayfly grazer, was determined in experimental chambers set in a tropical, high Andean stream. Chambers in an experimentally fertilized reach developed higher amount of both benthic and detached chlorophyll than chambers in an upstream control reach. Fertilization produced a slight increase in grazer biomass, and reduced algal biomass compared to grazer-free chambers. These results show that nutrient excess spread bottom-up effects through the food web, and that relevant top-down effects could also be detected. Eutrophication may produce relevant changes in the food web of tropical high-mountain streams.     

Epiphyte grazing enhances productivity of remnant seagrass patches

Anthropogenic nutrient enrichment is increasingly modifying community structure and ecosystem functioning in terrestrial and aquatic ecosystems. In marine ecosystems, the paradigm is that nutrient enrichment leads to a decline of seagrasses by stimulating epiphytic algal growth, which shades and overgrows seagrasses. This ignores the potential for herbivores, which graze upon epiphytic algae, to partially or wholly counter such nutrient effects. We conducted a field experiment to assess the role that the trochid gastropod Calthalotia fragum plays in reducing nutrient impacts on the seagrass, Posidonia australis, in an urbanized Australian estuary, Botany Bay, Sydney. In a field experiment, where nutrient loading and grazer density were orthogonally manipulated, nutrient enrichment failed to promote epiphyte biomass or diminish growth and primary productivity of P. australis. To the contrary, nutrient enrichment enhanced photosynthesis of the seagrass in plots where the grazer was present at higher density. Epiphytic growth was negatively affected by increased C. fragum density, while P. australis shoot growth was positively influenced. Thus, in this study system, grazing appears to play a much greater role in determining seagrass primary productivity and above‐ground growth than moderate nutrient loading, suggesting that the interaction between grazers and nutrients depends on the relative levels of each. Our study contributes to a growing body of literature suggesting that effects of nutrient loading on benthic assemblages are not universally negative, but are dependent on the biotic and abiotic setting.     

Macrophyte-epiphyte biomass and productivity in an eelgrass (Zostera marina L.) community

The biomass, productivity (¹⁴C), and photosynthetic response to light and temperature of eelgrass, Zostera marina L. and its epiphytes was measured in a shallow estuarine system near Beaufort, North Carolina, during 1974. The maximum of the biomass (above-ground) was measured in March; this was followed by a general decline throughout the rest of the year. The average biomass was 105.0 g dry wt m^?2; 80.3 g dry wt m^?2 was eelgrass and 24.7 g dry wt m^?2 was epiphytes. The productivity of eelgrass averaged 0.88 mg C g^?1 h^?1 which was similar to that of the epiphytes, 0.65 mg C g^?1 h^?1. Eelgrass and epiphyte productivity was low during the spring and early summer, gave a maximum during late summer and fall, and declined during the winter; this progression was probably due to environmental factors associated with tidal heights. On an areal basis, the average annual productivity was 0.9 g C m^?2 day^?1 for eelgrass and 0.2 g C m^?2 day^?1 for the epiphytes. Rates of photosynthesis of both eelgrass and epiphytes increased with increasing temperature to an asymptotic value at which the system was light saturated. Both eelgrass and epiphytes had a temperature optimum of < 29 °C. A negative response to higher temperatures was also reflected in biomass measurements which showed the destruction of eelgrass with increasing summer temperatures. The data suggest that the primary productivity cycles of macrophytes and epiphytes are closely interrelated.     

Nutrient impacts on epifaunal density and species composition in a subtropical seagrass bed

The capacity of epifauna to control algal proliferation following nutrient input depends on responses of both grazers and upper trophic level consumers to enrichment. We examined the responses of Thalassia testudinum (turtle grass) epifaunal assemblages to nutrient enrichment at two sites in Florida Bay with varying levels of phosphorus limitation. We compared epifaunal density, biomass, and species diversity in 2 m² plots that had either ambient nutrient concentrations or had been enriched with nitrogen and phosphorus for 6 months. At the severely P-limited site, total epifaunal density and biomass were two times higher in enriched than in unenriched plots. Caridean shrimp, grazing isopods, and gammarid amphipods accounted for much of the increase in density; brachyuran crabs, primary predatory fish, and detritivorous sea cucumbers accounted for most of the increase in biomass. At the less P-limited site, total epifaunal density and biomass were not affected by nutrient addition, although there were more caridean shrimp and higher brachyuran crab and pink shrimp biomass in enriched plots. At both sites, some variation in epifaunal density and biomass was explained by features of the macrophyte canopy, such as T. testudinum and Halodule wrightii percent cover, suggesting that enrichment may change the refuge value of the macrophyte canopy for epifauna. Additional variation in epifaunal density and biomass was explained by epiphyte pigment concentrations, suggesting that enrichment may change the microalgal food resources that support grazing epifauna. Increased epifaunal density in enriched plots suggests that grazers may be able to control epiphytic algal proliferation following moderate nutrient input to Florida Bay. Electronic supplementary material Electronic supplementary material is available for this article at and accessible for authorised users.     

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.     

Non‐consumptive effects of avian predators on fish behavior and cascading indirect interactions in seagrasses

Top–down impacts of avian predators are often overlooked in marine environments despite evidence from other systems that birds significantly impact animal distribution and behavior; instead, birds are typically recognized for the impacts of their nutrient rich guano. This is especially true in shallow seagrass meadows where restoration methods utilize bird perches or stakes to attract birds as a passive fertilizer delivery system that promotes the regrowth of damaged seagrasses. However, this method also increases the local density of avian piscivores that may have multiple unexplored non‐consumptive effects on fish behavior and indirect impacts to seagrass communities. We utilized laboratory and field experiments to investigate whether visual cues of avian predators impacted the behavior of the dominant demersal fish in seagrass habitats, the pinfish Lagodon rhomboides, and promoted cascading interactions on seagrass‐associated fauna and epiphytes. In laboratory mesocosms, pinfish displayed species specific responses to models of avian predators, with herons inducing the greatest avoidance behaviors. Avoidance patterns were confirmed in field seagrass meadows where heron models significantly reduced the number of fish caught in traps. In a long term field experiment, we investigated whether avian predators caused indirect non‐consumptive effects on seagrass communities by monitoring fish abundances, invertebrate epiphyte grazers, and the seagrass epiphytes in response to heron models, bird exclusions, and bird stakes. On average, more fish were recovered under bird exclusions and fewer fish under heron models. However, we found no evidence of cascading effects on invertebrate grazers or epiphytes. Bird stake treatments only displayed a simple nutrient effect where higher bird abundances resulted in higher epiphyte biomass. Our results indicate that although birds and their visual cues can affect fish and epiphyte abundance through non‐consumptive effects and nutrient enrichment, these impacts do not propagate beyond one trophic level, most likely because of dampening by omnivory and larger scale processes.     

Nutrient control of phytoplankton production in Lake Naivasha,Kenya

Lake Naivasha, a shallow tropical lake in Kenya's Rift Valley, has an unstable water column and is moderately eutrophic. Nutrient (bottom-up) control of primary production is more important than grazing (top-down) control. Experimental nutrient enrichment was used to investigate bottom-up control in more detail. Minor nutrients were not found to be limiting, whilst nitrogen was more limiting than phosphorus with an algal preference for ammonium over nitrate. Sediments form a phosphorus sink but there is hypolimnetic release from the one area showing regular temporary stratification. This indicates that the rate of primary production in the water column could double if conditions change to allow lake-wide nutrient release from sediments. Both external and recycled nutrient regeneration are important.     

Experimental evidence for enhanced top-down control of freshwater macrophytes with nutrient enrichment

The abundance of primary producers is controlled by bottom-up and top-down forces. Despite the fact that there is consensus that the abundance of freshwater macrophytes is strongly influenced by the availability of resources for plant growth, the importance of top-down control by vertebrate consumers is debated, because field studies yield contrasting results. We hypothesized that these bottom-up and top-down forces may interact, and that consumer impact on macrophyte abundance depends on the nutrient status of the water body. To test this hypothesis, experimental ponds with submerged vegetation containing a mixture of species were subjected to a fertilization treatment and we introduced consumers (mallard ducks, for 8 days) on half of the ponds in a full factorial design. Over the whole 66-day experiment fertilized ponds became dominated by Elodea nuttallii and ponds without extra nutrients by Chara globularis. Nutrient addition significantly increased plant N and P concentrations. There was a strong interactive effect of duck presence and pond nutrient status: macrophyte biomass was reduced (by 50 %) after the presence of the ducks on fertilized ponds, but not in the unfertilized ponds. We conclude that nutrient availability interacts with top-down control of submerged vegetation. This may be explained by higher plant palatability at higher nutrient levels, either by a higher plant nutrient concentration or by a shift towards dominance of more palatable plant species, resulting in higher consumer pressure. Including nutrient availability may offer a framework to explain part of the contrasting field observations of consumer control of macrophyte abundance.     

Effects of macrograzers and light on periphyton stoichiometry

Ecological stoichiometry describes the biochemical constraints of trophic interactions emerging from the different nutrient content and nutrient demand of producers and consumers, respectively. Most research on this topic originates from well-mixed pelagic food webs, whereas the idea has received far less attention in spatially structured habitats. Here, we test how light as well as grazing and nutrient regeneration by consumers affects growth and biomass of benthic primary producers. In the first laboratory experiment, we manipulated grazer presence (two different snail species plus ungrazed control), in the second experiment we factorially combined manipulation of grazer presence and light intensity. We monitored snail and periphyton biomass as well as dissolved and particulate nutrients (nitrogen and phosphorus) over time. Grazers significantly reduced algal biomass in both experiments. Grazers affected periphyton nutrient content depending on the prevailing nutrient limitation and their own body stoichiometry. In the nitrogen (N-) limited first experiment, grazers increased N both in the periphyton and in the water column. The effect was stronger for grazers with lower N-content. In the phosphorus (P-) limited second experiment, grazers increased the P-content of the periphyton, but the grazer with lower N-content had additionally positive effects on algal N. Light reduction did not affect periphyton biomass, but increased chlorophyll-, N- and P-content of the periphyton. These experiments revealed that the indirect effects of grazers on periphyton were bound by stoichiometric constraints of nutrient incorporation and excretion.     

Effect of grazing by isopods and amphipods on growth of Ulva spp. (Chlorophyta)

Eutrophication of shallow coastal waters often leads to blooms of macroalgae. Grazing by crustaceans, such as amphipods and isopods, can reduce macroalgal biomass accumulation. At the same time, growth of the macroalgae can be stimulated by epiphyte removal. The role of grazing by isopods and amphipods on Ulva spp. biomass development was investigated in the Veerse Meer, a brackish lagoon situated in the southwest Netherlands. Exclusion of grazing in the field did not stimulate Ulva spp. growth. In fact, growth rates were higher in exclosures that allowed grazers to enter. Edibility tests identified the amphipod Gammarus locusta, and the isopods Idotea chelipes and Sphaeroma hookeri as potential grazers on Ulva spp. However, when epiphytic diatoms were present on the Ulva spp. thalli, Gammarus and Sphaeroma grazed on ephiphytes and not on Ulva tissue. Only Idotea continued to graze on Ulva spp. A laboratory growth experiment revealed a positive effect of Gammarus presence on Ulva spp. growth, probably caused by preferential removal of epiphytic diatoms from the Ulva spp. thalli. The growth stimulation by epiphyte removing grazers such as Gammarus may explain the higher growth rates in the presence of grazers observed in the field. When determining the potential role of invertebrate grazers in controlling macroalgal biomass accumulation, it is important to include an assessment of the epiphyte abundance on the macroalgae, as preferential removal of epiphytes may stimulate growth and thus have the opposite effect.     

Effects of increased temperature and nutrient enrichment on the stoichiometry of primary producers and consumers in temperate shallow lakes

1. We studied the effects of increased water temperatures (0–4.5 °C) and nutrient enrichment on the stoichiometric composition of different primary producers (macrophytes, epiphytes, seston and sediment biofilm) and invertebrate consumers in 24 mesocosm ecosystems created to mimic shallow pond environments. The nutrient ratios of primary producers were used as indicative of relative nitrogen (N) or phosphorus (P) limitation. We further used carbon stable isotopic composition (δ¹³C) of the different primary producers to elucidate differences in the degree of CO₂ limitation. 2. Epiphytes were the only primary producer with significantly higher δ¹³C in the enriched mesocosms. No temperature effects were observed in δ¹³C composition of any primary producer. Independently of the treatment effects, the four primary producers had different δ¹³C signatures indicative of differences in CO₂ limitation. Seston had signatures indicating negligible or low CO₂ limitation, followed by epiphytes and sediment biofilm, with moderate CO₂ limitation, while macrophytes showed the strongest CO₂ limitation. CO₂ together with biomass of epiphytes were the key variables explaining between 50 and 70% of the variability in δ¹³C of the different primary producers, suggesting that epiphytes play an important role in carbon flow of temperate shallow lakes. 3. The ratio of carbon to chlorophyll a decreased with increasing temperature and enrichment in both epiphytes and seston. The effects of temperature were mainly attributed to changes in algal Chl a content, while the decrease with enrichment was probably a result of a higher proportion of algae in the seston and epiphytes. 4. Macrophytes, epiphytes and seston decreased their C : N with enrichment, probably as an adaptation to the different N availability levels. The C : N of epiphytes and Elodea canadensis decreased with increasing temperature in the control mesocosms. Sediment biofilm was the only primary producer with lower C : P and N : P with enrichment, probably as a result of higher P accumulation in the sediment. 5. Independently of nutrient level and increased temperature effects the four primary producers had significantly different stoichiometric compositions. Macrophytes had higher C : N and C : P and, together with epiphytes, also the highest N : P. Seston had no N or P limitation, while macrophytes and epiphytes may have been P limited in a few mesocosms. Sediment biofilm indicated strong N deficiency. 6. Consumers had strongly homeostatic stoichiometric compositions in comparison to primary producers, with weak or no significant treatment effects in any of the groups (insects, leeches, molluscs and crustaceans). Among consumers, predators had significantly higher N content and lower C : N than grazers.     

Warming shifts top-down and bottom-up control of pond food web structure and function

The effects of global and local environmental changes are transmitted through networks of interacting organisms to shape the structure of communities and the dynamics of ecosystems. We tested the impact of elevated temperature on the top-down and bottom-up forces structuring experimental freshwater pond food webs in western Canada over 16 months. Experimental warming was crossed with treatments manipulating the presence of planktivorous fish and eutrophication through enhanced nutrient supply. We found that higher temperatures produced top-heavy food webs with lower biomass of benthic and pelagic producers, equivalent biomass of zooplankton, zoobenthos and pelagic bacteria, and more pelagic viruses. Eutrophication increased the biomass of all organisms studied, while fish had cascading positive effects on periphyton, phytoplankton and bacteria, and reduced biomass of invertebrates. Surprisingly, virus biomass was reduced in the presence of fish, suggesting the possibility for complex mechanisms of top-down control of the lytic cycle. Warming reduced the effects of eutrophication on periphyton, and magnified the already strong effects of fish on phytoplankton and bacteria. Warming, fish and nutrients all increased whole-system rates of net production despite their distinct impacts on the distribution of biomass between producers and consumers, plankton and benthos, and microbes and macrobes. Our results indicate that warming exerts a host of indirect effects on aquatic food webs mediated through shifts in the magnitudes of top-down and bottom-up forcing.     

Nutrient Enrichment and Food Web Composition Affect Ecosystem Metabolism in an Experimental Seagrass Habitat

Background

Food web composition and resource levels can influence ecosystem properties such as productivity and elemental cycles. In particular, herbivores occupy a central place in food webs as the species richness and composition of this trophic level may simultaneously influence the transmission of resource and predator effects to higher and lower trophic levels, respectively. Yet, these interactions are poorly understood.

Methodology/Principal Findings

Using an experimental seagrass mesocosm system, we factorially manipulated water column nutrient concentrations, food chain length, and diversity of crustacean grazers to address two questions: (1) Does food web composition modulate the effects of nutrient enrichment on plant and grazer biomasses and stoichiometry? (2) Do ecosystem fluxes of dissolved oxygen and nutrients more closely reflect above-ground biomass and community structure or sediment processes? Nutrient enrichment and grazer presence generally had strong effects on biomass accumulation, stoichiometry, and ecosystem fluxes, whereas predator effects were weaker or absent. Nutrient enrichment had little effect on producer biomass or net ecosystem production but strongly increased seagrass nutrient content, ecosystem flux rates, and grazer secondary production, suggesting that enhanced production was efficiently transferred from producers to herbivores. Gross ecosystem production (oxygen evolution) correlated positively with above-ground plant biomass, whereas inorganic nutrient fluxes were unrelated to plant or grazer biomasses, suggesting dominance by sediment microbial processes. Finally, grazer richness significantly stabilized ecosystem processes, as predators decreased ecosystem production and respiration only in the zero- and one- species grazer treatments.

Conclusions/Significance

Overall, our results indicate that consumer presence and species composition strongly influence ecosystem responses to nutrient enrichment, and that increasing herbivore diversity can stabilize ecosystem flux rates in the face of perturbations.     

The role of light availability and herbivory on algal responses to nutrient enrichment in a riparian wetland,Alaska

We investigated how the relative availability of solar radiation in the presence or absence of grazing alters the ability of benthic algae to respond to nutrient enrichment in an Alaskan marsh. We used a factorial mesocosm experiment that included nutrient enrichment (enriched or control), grazing (grazed or ungrazed), and light (unshaded or shaded) to simulate shading by macrophytes early and late in the growing season, respectively. We found stronger effects of grazers and nutrients compared to light on benthic algal biomass and taxonomic composition. Algal biomass increased in nutrient‐enriched treatments and was reduced by grazing. Shading did not have an effect on algal biomass or taxonomic composition, but the concentration of chl a per algal biovolume increased with shading, demonstrating the ability of algae to compensate for changes in light availability. Algal taxonomic composition was more affected by grazer presence than nutrients or light. Grazer‐resistant taxa (basal filaments of Stigeoclonium) were replaced by diatoms (Nitzschia) and filamentous green algae (Ulothrix) when herbivores were removed. The interacting and opposing influences of nutrients and grazing indicate that the algal community is under dual control from the bottom‐up (nutrient limitation) and from the top‐down (consumption by herbivores), although grazers had a stronger influence on algal biomass and taxonomic composition than nutrient enrichment. Our results suggest that low light availability will not inhibit the algal response to elevated nutrient concentrations expected with ongoing climate change, but grazers rapidly consume algae following enrichment, masking the effects of elevated nutrients on algal production.